european journal of taxonomy 527: 1–2 issn 2118-9773 https://doi.org/10.5852/ejt.2019.527 www.europeanjournaloftaxonomy.eu 2019 · trietsch c. et al. this work is licensed under a creative commons attribution license (cc by 4.0). c o r r i g e n d u m 1 the following corrections have been made to paper no. 502 (https://doi.org/10.5852/ejt.2019.502) a photographic catalog of ceraphronoidea types at the muséum national d’histoire naturelle, paris (mnhn), with comments on unpublished notes from paul dessart – corrigendum carolyn trietsch 1,*, istván mikó 2 & andrew r. deans 3 1,3 frost entomological museum, the pennsylvania state university, university park, pa, 16802 usa. 2 university of new hampshire collection of insects and other arthropods, department of biological sciences, university of new hampshire, spaulding hall, durham, nh, 03824 usa. * corresponding author: carolyntrietsch@gmail.com 2 email: istvan.miko@gmail.com 3 email: adeans@psu.edu trietsch c., mikó i. & deans a.r. 2019. a photographic catalog of ceraphronoidea types at the muséum national d’histoire naturelle, paris (mnhn), with comments on unpublished notes from paul dessart – corrigendum . european journal of taxonomy 527: 1–2. https://doi.org/10.5852/ejt.2019.527 at the beginning of the chapter ‘acknowledgments’ should be added the following paragraph that was accidentally forgotten. the correct ‘acknowledgments’ are given here. acknowledgments the authors would like to thank agnièle touret-alby for all of her help in contributing historical documents and allowing ct to visit the mnhn and access the ceraphronoidea collections, and for going above and beyond to track down missing specimens so that they could be included in this work. the authors would also like to thank all of the other curators and staff present at the mnhn collections; without access to these collections, this work would not exist. the authors would furthermore like to thank david g. notton for his insight and expertise on the history of j.j. kieffer. special thanks to norman johnson and luciana musetti for their immensely helpful catalogue of ceraphronoidea, which has no doubt advanced research efforts on the superfamily. this material is based upon work supported by the u.s. national science foundation, under grant number deb-1353252. any opinions, findings, and conclusions or recommendations expressed in this material are those of the authors and do not necessarily reflect the views of the national science foundation. published on: 23 may 2019 https://doi.org/10.5852/ejt.2019.527 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ https://doi.org/10.5852/ejt.2019.502 mailto:carolyntrietsch@gmail.com mailto:istvan.miko@gmail.com mailto:adeans@psu.edu https://doi.org/10.5852/ejt.2019.527 european journal of taxonomy 527: 1–2 (2019) 2 printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. 1 european journal of taxonomy 717: 1–2 issn 2118-9773 https://doi.org/10.5852/ejt.2020.717.1093 www.europeanjournaloftaxonomy.eu 2020 · petzold a. et al. this work is licensed under a creative commons attribution license (cc by 4.0). c o r r i g e n d u m urn:lsid:zoobank.org:pub:129c82f8-325f-40ea-9f29-06940057c24a first insights into past biodiversity of giraffes based on mitochondrial sequences from museum specimens – corrigendum alice petzold 1, anne-sophie magnant 2, david edderai 3, bertrand chardonnet 4, jacques rigoulet 5, michel saint-jalme 6 & alexandre hassanin 7,* 1,2,7 institut de systématique, évolution, biodiversité (isyeb), sorbonne université, mnhn, cnrs, ephe, ua, muséum national d’histoire naturelle, 55 rue buffon cp 51 75005 paris, france. 3 5 chemin du bas d’anville 17750 etaules, france. 4 92210 saint cloud, france. 5 deceased [18 may 2020]. former address: direction générale déléguée aux musées, jardins botaniques et zoologiques, muséum national d’histoire naturelle, 57 rue cuvier 75005 paris, france. 6 centre d’ecologie et des sciences de la conservation, umr 7204 mnhn cnrs-upmc, muséum national d’histoire naturelle 75005 paris, france. * corresponding author: alexandre.hassanin@mnhn.fr 1 email: alice.petzold1@mnhn.fr 2 email: annesophie.magnant@gmail.com 3 email: davidedderai@hotmail.fr 4 email: bertrand.chardonnet@gmail.com 6 email: michel.saint-jalme@mnhn.fr 1 urn:lsid:zoobank.org:author:e7fdae2b-14a3-4b47-bd66-4e607ac5cefb 2 urn:lsid:zoobank.org:author:c202cfc1-8e73-460d-b0b8-6b00964cf40b 3 urn:lsid:zoobank.org:author:5edaf19c-7e3e-44de-b5de-510b60fd81c7 4 urn:lsid:zoobank.org:author:3e4dc1db-5af4-4574-a14c-5a19bb78e231 5 urn:lsid:zoobank.org:author:ec4bdeb1-b643-4d46-a960-d0bc44d0d9b3 6 urn:lsid:zoobank.org:author:7754b40e-deb5-4133-b91e-0c0341d2211d 7 urn:lsid:zoobank.org:author:0dcc3e08-b2ba-4a2c-ada5-1a256f24daa1 petzold a., magnant a.-s., edderai d., chardonnet b., rigoulet j., saint-jalme m. & hassanin a. 2020. first insights into past biodiversity of giraffes based on mitochondrial sequences from museum specimens – corrigendum. european journal of taxonomy 717: 1–2. https://doi.org/10.5852/ejt.2020.717.1093 the present corrigendum corrects errors that occurred in petzold et al. (2020). page 2, table 2: the voucher name of the tenth specimen in table 2 should be changed from “mhnt-1996.121.2” to “mhnt.ost.1996.121.2”. https://doi.org/10.5852/ejt.2020.717.1093 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:129c82f8-325f-40ea-9f29-06940057c24a mailto:alexandre.hassanin@mnhn.fr mailto:alice.petzold1@mnhn.fr mailto:annesophie.magnant@gmail.com mailto:davidedderai@hotmail.fr mailto:bertrand.chardonnet@gmail.com mailto:michel.saint-jalme@mnhn.fr http://zoobank.org/urn:lsid:zoobank.org:author:e7fdae2b-14a3-4b47-bd66-4e607ac5cefb http://zoobank.org/urn:lsid:zoobank.org:author:c202cfc1-8e73-460d-b0b8-6b00964cf40b http://zoobank.org/urn:lsid:zoobank.org:author:5edaf19c-7e3e-44de-b5de-510b60fd81c7 http://zoobank.org/urn:lsid:zoobank.org:author:3e4dc1db-5af4-4574-a14c-5a19bb78e231 http://zoobank.org/urn:lsid:zoobank.org:author:ec4bdeb1-b643-4d46-a960-d0bc44d0d9b3 http://zoobank.org/urn:lsid:zoobank.org:author:7754b40e-deb5-4133-b91e-0c0341d2211d http://zoobank.org/urn:lsid:zoobank.org:author:0dcc3e08-b2ba-4a2c-ada5-1a256f24daa1 https://doi.org/10.5852/ejt.2020.717.1093 european journal of taxonomy 717: 1–2 (2020) 2 page 10, fig. 3: the voucher name of the specimen from abyssinia (ethiopia) clustering in the haplogroup ʻnubiaʼ should be changed from “mhnt-1996.121.2” to “mhnt.ost.1996.121.2”. page 11, line 9: the voucher name of the specimen from abyssinia (ethiopia) should be changed from “mhnt1996.121.2” to “mhnt.ost.1996.121.2”. page 12, fig. 4: the voucher name of the specimen from abyssinia (ethiopia) clustering in the haplogroup ʻnubiaʼ should be changed from “mhnt-1996.121.2” to “mhnt.ost.1996.121.2”. page 17, the two last lines: the voucher name of the specimen from abyssinia (ethiopia) should be changed from “mhnt1996.121.2” to “mhnt.ost.1996.121.2”. page 21, in ʻtype material examinedʼ: the ʻtype material examinedʼ section for giraffa camelopardalis should be amended as follows: type material examined neotype (here designated) ethiopia • 1 specimen (skin and complete skeleton); abyssinia; mhnt.ost.1996.121.2 (skull), mhnt.ost.1996.121.1 (postcranial skeleton), mhnt.art.4 (skin). reference petzold a., magnant a.-s., edderai d., chardonnet b., rigoulet j., saint-jalme m. & hassanin a. 2020. first insights into past biodiversity of giraffes based on mitochondrial sequences from museum specimens. european journal of taxonomy 703: 1–33. https://doi.org/10.5852/ejt.2020.703 published on: 17 september 2020 printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. https://doi.org/10.5852/ejt.2020.703 1 european journal of taxonomy 607: 1–2 issn 2118-9773 https://doi.org/10.5852/ejt.2020.607 www.europeanjournaloftaxonomy.eu 2020 · stöhr s. et al. this work is licensed under a creative commons attribution license (cc by 4.0). c o r r i g e n d u m urn:lsid:zoobank.org:pub:2b06a2fb-f503-4c36-a11b-f16ad1295544 resolving the ophioderma longicauda (echinodermata: ophiuroidea) cryptic species complex: five sisters, three of them new – corrigendum sabine stöhr 1,*, alexandra anh-thu weber 2, emilie boissin 3 & anne chenuil 4 1 swedish museum of natural history, department of zoology, frescativägen 40, s-10405 stockholm, sweden. 2 marine invertebrates, museum victoria, gpo box 666, melbourne vic 3001, australia. 2 ifremer, centre de bretagne, rem / eep, laboratoire environnement profond, 29280 plouzané, france. 3 psl research university: ephe-upvd-cnrs, usr3278 criobe, université de perpignan, 52 avenue paul alduy, 66860 perpignan cedex, france. 4 aix-marseille université, institut méditerranéen de biodiversité et d’ecologie marine et continentale (imbe) cnrs ird uapv, station marine d’endoume, chemin de la batterie des lions, f-13007 marseille, france. * corresponding author: sabine.stohr@nrm.se 2 email: aweber@museum.vic.gov.au 3 email: emilie.boissin@univ-perp.fr 4 email: anne.chenuil@imbe.fr 1 urn:lsid:zoobank.org:author:412800eb-aace-4313-9810-61f89b740405 2 urn:lsid:zoobank.org:author:0c152e34-0cd1-4f4f-81ae-818ac5018f1c 3 urn:lsid:zoobank.org:author:b35ff544-ae44-43b2-b1ca-3769a281f386 4 urn:lsid:zoobank.org:author:8b07d296-065b-44d4-b4b1-d744b9d9546a stöhr s., weber a.a.-t., boissin e. & chenuil a. 2020. resolving the ophioderma longicauda (echinodermata: ophiuroidea) cryptic species complex: five sisters, three of them new – corrigendum. european journal of taxonomy 607: 1–2. https://doi.org/10.5852/ejt.2020.607 the last paragraph of the introduction (pp 4–5, stöhr et al. 2020) should be amended as follows: “....the nomenclatural problem was resolved by decision of the international commission of zoological nomenclature (melville 1980), which also erroneously decided that ophioderma is of feminine gender. this was corrected a year later (international commission of zoological nomenclature 1981) and ophioderma is of neuter gender. unfortunately, this correction has gone largely unnoticed until now, leading to confusion, with some authors using the feminine form and others the neuter form of the species epithets in this genus. from now on, the neuter form alone should be used. to stabilize a species concept, .....” all occurrences of ophioderma longicauda should be changed to ophioderma longicaudum. all occurrences of ophioderma africana sp. nov. should be changed to ophioderma africanum sp. nov., and all occurrences of ophioderma hybrida should be changed to ophioderma hybridum sp. nov. https://doi.org/10.5852/ejt.2020.607 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:2b06a2fb-f503-4c36-a11b-f16ad1295544 mailto:sabine.stohr%40nrm.se?subject= mailto:aweber%40museum.vic.gov.au?subject= mailto:emilie.boissin%40univ-perp.fr?subject= mailto:anne.chenuil%40imbe.fr?subject= http://zoobank.org/urn:lsid:zoobank.org:author:412800eb-aace-4313-9810-61f89b740405 http://zoobank.org/urn:lsid:zoobank.org:author:0c152e34-0cd1-4f4f-81ae-818ac5018f1c http://zoobank.org/urn:lsid:zoobank.org:author:b35ff544-ae44-43b2-b1ca-3769a281f386 http://zoobank.org/urn:lsid:zoobank.org:author:8b07d296-065b-44d4-b4b1-d744b9d9546a https://doi.org/10.5852/ejt.2020.607 european journal of taxonomy 607: 1–2 (2020) 2 the title should be changed to “resolving the ophioderma longicaudum (echinodermata: ophiuroidea) cryptic species complex: five sisters, three of them new”. the following reference should be added to the list of references: international commission of zoological nomenclature. 1981. corrigenda. bulletin of zoological nomenclature 38 (4): 317. reference stöhr s., weber a.a.-t., boissin e. & chenuil a. 2020. resolving the ophioderma longicauda (echinodermata: ophiuroidea) cryptic species complex: five sisters, three of them new. european journal of taxonomy 600: 1–37. https://doi.org/10.5852/ejt.2020.600 published on: 21 february 2020 printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. https://doi.org/10.5852/ejt.2020.600 review of the swedish species of ophion (hymenoptera: ichneumonidae: ophioninae), with the description of 18 new species and an illustrated key to swedish species – corrigendum niklas johansson 1,* & björn cederberg 2 1 fredriksberg/baskarp 566 92 habo, sweden. 2 uppsala-näs, asplunda 51, 775 91 uppsala, sweden. * corresponding author: chrysis32@yahoo.se 2 email: bjornceder@gmail.com 1 urn:lsid:zoobank.org:author:b0031f47-99f7-4fee-b545-b6f1e79e1a4e 2 urn:lsid:zoobank.org:author:011c3e1f-eb9e-4bda-b368-1cc50bb525d7 johansson n. & cederberg b. 2019. review of the swedish species of ophion (hymenoptera: ichneumonidae: ophioninae), with the description of 18 new species and an illustrated key to swedish species – corrigendum. european journal of taxonomy 560: 1–3. https://doi.org/10.5852/ejt.2019.560 the present corrigendum corrects errors that occurred in the ‘abstract’, ‘table of contents’ and on several other pages in johansson & cederberg (2019). the species name on page 106 also needs to be emended. page 1, in ‘abstract’, lines 13–14: “ophion slaviceki kriechbaumer, 1892 is excluded from synonymy with ophion luteus linnaeus, 1758 stat. rev.” should read: “ophion slaviceki kriechbaumer, 1892 stat. rev. is excluded from synonymy with ophion luteus linnaeus, 1758.” page 1, in ‘abstract’, line 18: “ophion albistylus szépligeti, 1905 (syn. nov.) is synonymized with ophion pteridis kriechbaumer, 1879 and ophion frontalis strobl, 1904 (syn. nov.) is synonymized with ophion areolaris brauns, 1889 syn. nov.” should read: “ophion albistylus szépligeti, 1905 syn. nov. is synonymized with ophion pteridis kriechbaumer, 1879 and ophion frontalis strobl, 1904 syn. nov. is synonymized with ophion areolaris brauns, 1889.” page 2, in ‘table of contents’: “ophion luteus (linnaeus, 1758) stat. rev.” should read: “ophion luteus (linnaeus, 1758)” european journal of taxonomy 560: 1–3 issn 2118-9773 https://doi.org/10.5852/ejt.2019.560 www.europeanjournaloftaxonomy.eu 2019 · johansson n. & cederberg b. this work is licensed under a creative commons attribution license (cc by 4.0). c o r r i g e n d u m urn:lsid:zoobank.org:pub:e8c6917e-ad9c-4e9a-a5fb-5cd21b7138e9 1 mailto:chrysis32%40yahoo.se?subject= mailto:bjornceder%40gmail.com?subject= http://zoobank.org/urn:lsid:zoobank.org:author:b0031f47-99f7-4fee-b545-b6f1e79e1a4e http://zoobank.org/urn:lsid:zoobank.org:author:011c3e1f-eb9e-4bda-b368-1cc50bb525d7 https://doi.org/10.5852/ejt.2019.560 https://doi.org/10.5852/ejt.2019.560 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:e8c6917e-ad9c-4e9a-a5fb-5cd21b7138e9 page 43, ‘remarks’ for ophion areolaris, lines 5–6: “ophion frontalis strobl, 1904 is therefore to be regarded as a junior synonym of ophion areolaris brauns, 1889 syn. nov.” should read: “ophion frontalis strobl, 1904 syn. nov. is therefore to be regarded as a junior synonym of ophion areolaris brauns, 1889.” page 76, heading: “ophion luteus (linnaeus, 1758) stat. rev.” should read: “ophion luteus (linnaeus, 1758)” page 94, remarks for ophion pteridis, lines 6–8: “the holotype of ophion albistylus in nhmh was studied and this species is synonymous with ophion pteridis syn. nov., a fact already noted by scaramozzino by the labels attached to the specimen.” should read: “the holotype of ophion albistylus in nhmh was studied and this species is found to be synonymous with ophion pteridis, a fact already noted by scaramozzino by the labels attached to the specimen.” page 104, paratypes for ophion tenuicornis, line 13: “22‒24 2017; r. isaksson leg.” should read: “22‒24 sep. 2017; r. isaksson leg.” page 104, paratypes for ophion tenuicornis, line 20: “1 aug.‒28 jul. 2016; b. andersson leg.” should read: “28 jul.‒1 aug. 2016; b. andersson leg.” page 106, species name: “ophion vardali” should read in the text and figure captions: ophion vardalae johansson, 2019 (emendation of name): this species was originally described as ophion vardali johansson, 2019 (in johansson & cederberg 2019). the ending in ‘i’, however, is incorrect and therefore the specific name is here corrected as ophion vardalae johansson, 2019 (in johansson & cederberg 2019), in compliance with article 31.1.2 of the iczn (the ending in ‘ae’ is for a woman, ‘i’ for a man). page 121, in ‘the ophion obscuratus aggregate’, second paragraph, last sentence: “based on this, o. polyguttator is excluded from synonymy with o. obscuratus stat. rev. and regarded as a nomen inquirendum.” should read: “based on this, o. polyguttator stat. rev. is excluded from synonymy with o. obscuratus and regarded as a nomen inquirendum.” page 128, in ‘the ophion pteridis aggregate’, second paragraph, lines 8–9: “studies of the type material show that ophion pteridis (fig. 52a‒f) is the correct name for one of the two infuscate species and that ophion albistylus is a junior synonym of that species, syn. nov.” should read: “studies of the type material show that ophion pteridis (fig. 52a‒f) is the correct name for one of the two infuscate species and that ophion albistylus syn. nov. is a junior synonym of that species.” european journal of taxonomy 560: 1–3 (2019) 2 reference johansson n. & cederberg b. 2019. review of the swedish species of ophion (hymenoptera: ichneumonidae: ophioninae), with the description of 18 new species and an illustrated key to swedish species. european journal of taxonomy 550: 1–136. https://doi.org/10.5852/ejt.2019.550 published on: 3 october 2019 printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. johansson n. & cederberg b., swedish species of ophion – corrigendum 3 https://doi.org/10.5852/ejt.2019.550 european journal of taxonomy 165: 1–3 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2015.165 www.europeanjournaloftaxonomy.eu 2015 · segers et al. this work is licensed under a creative commons attribution 3.0 license. n o t i c e 1 period of public commentary begins on the revised proposal of species-group level names, and on the proposal of genus-group level names of the candidate part of list of available names (lan) in the phylum rotifera hendrik segers 1,*, willem h. de smet 2, diego fontaneto 3, claus hinz 4, charles hussey 5, evangelia michaloudi 6, robert l. wallace 7 & christian d. jersabek 8 1 royal belgian institute of natural sciences, vautierstraat 29, b 1000 brussels, belgium. 2 department of biology, ecobe, university of antwerp, campus drie eiken, universiteitsplein 1, b-2610 wilrijk, belgium. 3 national research council, institute of ecosystem study, largo tonolli 50, 28922 verbania pallanza, italy. 4 systematics and evolutionary biology, institute of biology and environmental sciences, university of oldenburg, carl von ossietzky-str. 9-11, 26129 oldenburg, germany. 5 natural history museum, cromwell road,london sw7 5bd, united kingdom. 6 department of zoology, school of biology, aristotle university, gr-54124 thessaloniki, greece. 7 department of biology, ripon college, ripon, wi, usa. 8 department of organismal biology, university of salzburg, a–5020 salzburg, austria, and academy of natural sciences of drexel university, center for systematic biology & evolution, 19103 philadelphia,pa, usa. * corresponding author: hendrik.segers@naturalsciences.be segers h., de smet w.h., fontaneto d., hinz c., hussey c., michaloudi e., wallace r.l. & jersabek c.d. 2015. period of public commentary begins on the revised proposal of species-group level names level, and on the proposal of genus-group level names of the candidate part of list of available names (lan) in the phylum rotifera. european journal of taxonomy 165: 1–3. http://dx.doi.org/10.5852/ejt.2015.165 following iczn (1999) article 79 chapter 17 (http://www.nhm.ac.uk/hosted-sites/iczn/code/), we, with the full support of the international community of rotifer researchers as expressed during subsequent international rotifera symposia, developed a candidate part of the list of available names for species and genera of rotifera from the start of zoological nomenclature to the year 2000. regarding species-group level names, details of how the candidate part was assembled are described in segers et al. (2012). following article 79, a notice was published in bzn 69: 2 march 2012 and the candidate part was made available from 31 march 2012 to 30 march 2013, to receive public comment. at the close of this period the proposal was referred to the international commission on zoological nomenclature’s ad hoc committee for it to receive comments (as per article 79.2.2.3 of the code). in april 2014 the ad hoc committee informed us that they have recommended considering a revised proposal. the conditions of article 79.2.2.4 have been met. in conformity with article 79.2.2.5, http://dx.doi.org/10.5852/ejt.2015.165 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/3.0/ mailto:hendrik.segers%40naturalsciences.be?subject= http://dx.doi.org/10.5852/ejt.2015.165 http://www.nhm.ac.uk/hosted-sites/iczn/code/ european journal of taxonomy 165: 1–3 (2015) 2 a request has been made by the ad hoc committee that a section about how we have responded to reviewers’ comments made on the original candidate part were included on a website. the revised proposal, taking into account the comments received, is now being made available at http://rotifera. hausdernatur.at/home/rotiferlan and http://iczn.org/content/lan-committees, and comments may be sent to the rotifer committee (rotiferalan@gmail.com), or to the commission secretariat (iczn@nhm. ac.uk). this second year of comment will begin on 20 december 2015 and will end on 19 december 2016. the candidate part is divided into two sections. section a includes names that are proposed to be included in the list of available names; section b includes names of rotifers that are proposed not to be included in the lan for reasons described in segers et al. (2012). comments are invited on both sections. regarding genus-group level names, the candidate part contains the following information: • correct spelling of the name; • its bibliographic reference, authorship, date of publication (availability), and, if appropriate, its nomenclatural status; • the name, status, and way of fixation of its name-bearing type; and • if the name has been subject of a commission ruling, the relevant opinion(s). for information and ease of retrieval, the taxonomic status and gender of each genus name, and, if appropriate, additional information on the name is added between square brackets. the latter elements do not form part of the list of available names proper and would not be covered under article 79.4 of the code. the candidate part contains a single section and includes all names established, or in use for taxa in rotifera. an appendix listing names of genera that have been established for use in rotifera but that are based on type species that do not pertain to the phylum is added. this includes the name eretmia gosse 1886, suppressed for the purposes of the principle of priority but not for those of the principle of homonymy and attributed, with reservation, to rotifera (sub rotatoria) in opinion 816 bzn 24: 149150 (iczn, 1967). as far as can be judged from the drawing of the type species (eretmia pentathrix gosse 1886 gosse in hudson & gosse 1886), the organism is likely to be a filose amoeba (?euglypha or ?scutiglypha; rhizaria), and not a rotifer. following article 79, the candidate part will now become available for a first period of one year to receive public comment. it can be downloaded here: http://rotifera.hausdernatur.at/home/rotiferlan http://iczn.org/content/lan-committees# and comments may be sent to rotiferalan@gmail.com, or by direct correspondence with the iczn secretariat (iczn@nhm.ac.uk). the year of comment will begin on 20 december 2015 and will end on 19 december 2016. references hudson c.t., gosse p.h. 1886. the rotifera; or wheel-animalcules, both british and foreign, vol. ii. longmans, green & co., london. international commission on zoological nomenclature 1967. opinion 816. eretmia gosse, 1886 (?rotatoria): suppressed under the plenary powers. bulletin of zoological nomenclature 24: 149–150. international commission on zoological nomenclature 1999. international code of zoological nomenclature. fourth edition. international trust for zoological nomenclature, london. segers h., de smet w.h., fischer c., fontaneto c., michaloudi e., wallace r.l. & jersabek c.d. 2012. towards a list of available names in zoology, partim phylum rotifera. zootaxa 3179: 61–68. http://rotifera.hausdernatur.at/home/rotiferlan http://rotifera.hausdernatur.at/home/rotiferlan http://iczn.org/content/lan-committees mailto:rotiferalan%40gmail.com?subject= mailto:iczn%40nhm.ac.uk?subject= mailto:iczn%40nhm.ac.uk?subject= http://rotifera.hausdernatur.at/home/rotiferlan http://iczn.org/content/lan-committees# mailto:rotiferalan%40gmail.com?subject= mailto:iczn%40nhm.ac.uk?subject= segers h. et al., period of public commentary in the phylum rotifera 3 manuscript received: 10 november 2015 manuscript accepted: 2 december 2015 published on: 17 december 2015 topic editor: koen martens desk editor: kristiaan hoedemakers printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark. published on 17 december 2015. european journal of taxonomy www.europeanjournaloftaxonomy.eu http://dx.doi.org/10.5852/ejt.2011.1 2011 · the edit / ejt task force this work is licensed under a creative commons attribution 3.0 license. editorial 1 launch of the european journal of taxonomy (ejt) "the publication of results and the accessibility of publications are a precondition for the efficiency of the research process." (sietmann 2008). the edit / ejt task force laurence bénichou1, koen martens2, graham higley3, isabelle gérard4, steven dessein5 & daphné duin6 1muséum national d’histoire naturelle, paris, france / managing editor ejt 2royal belgian institute of natural sciences, brussels, belgium / editor in chief ejt 3natural history museum, london, uk 4royal museum for central africa, tervuren, belgium 5national botanic garden of belgium, meise, belgium 6edit office, paris, france we are very pleased and proud to announce the launch of the european journal of taxonomy. the ejt is an international, online, fast-track, peer-reviewed, open access journal in descriptive taxonomy, covering subjects in zoology, entomology, botany, and palaeontology, owned and run by a consortium of european natural history institutes. ejt is a collaborative project outcome of the edit network. the foundation of this new journal will inevitably raise some questions, not the least of which is why does a scientific field such as taxonomy need a new journal? surely there are enough taxonomic journals already, some of them well-known and many well-established! a core mission of natural history institutions (nhis) is to contribute to the understanding of the natural world and our place in it, and to disseminate this knowledge. correspondingly, many nhis have been publishers of journals and monographic series since their foundation, for some over 200 years ago. a key aspect of nhi-publications is the description of species (and higher taxa) and the unravelling of their relationships to other species, i.e. taxonomy and classification. biodiversity description and the formal naming of species is scattered across hundreds of journals, many of which are inaccessible and with small print runs. today, several of these publications are at risk of extinction. yet, dissemination of original research on biodiversity is crucial to the mission of nhis. in today’s digital era, the field of scholarly publishing is rapidly changing and the journals of nhis are faced with complex strategic and technical issues concerning visibility, access, format, and financial structure of their titles. the real challenge for institutional journals is to keep up with the rapid www.europeanjournaloftaxonomy.eu www.europeanjournaloftaxonomy.eu http://dx.doi.org/10.5852/ejt.2011.1 http://creativecommons.org/licenses/by/2.0/ european journal of taxonomy 1: 1-3 (2011) 2 developments in online services and standards. those journals that miss the digital boat may become obscure, user unfriendly and may disappear in the near future. in parallel, ever more institutional journals have been outsourced to private publishers and, with the need to be ever mindful of the 'isi impact factor', have shifted their editorial focus from descriptive taxonomy to phylogenetic and molecular analytical research. the result is that today there are fewer nhi-driven communication channels for alpha (descriptive) taxonomy, and those that still exist usually do not have a wide circulation. to reverse this downward spiral, a consortium of nhis has decided to launch a new, co-published journal of taxonomy that will play an important role in moving the field to open-access publishing and digital archiving, while enabling the institutions to take a greater control of the dissemination of scientific information. nhis are both producers and consumers of taxonomic research and it is in their interest to continue production and to provide access to high quality work in this domain. a jointly published journal of taxonomy, funded by several institutions, sends a strong political message about the importance of descriptive taxonomic research within nhis in europe and elsewhere. descriptive taxonomy is needed by a wide user base. traditionally, public funds are used to subsidise either reader-based subscriptions (libraries) or authorpaid fees for open-access. as scientific publications are the fruit of public-funded research, we strongly believe that scholarly publications should be made accessible to all, at no charge. this is why the european journal of taxonomy (ejt) is applying an alternative public open-access business model where neither readers, nor authors have to pay fees for subscriptions or publication. ejt endorses the view, that ability to use and reuse data as freely as possible is the key to innovation and further advancement of science (i2010 digital libraries initiative). therefore, the most suitable economic model for ejt is open access without a charge-processing fee. this economic model will take away potential barriers for researchers to publish their work. our objective is to enhance coordination between nhis aimed at setting up a cross-institutional strategy at an international level in order to promote the adoption of common standards and ways of working, wherever possible. an extensive, international team of scientific editors will guard the scientific quality of papers published in ejt, so that high level taxonomic papers of interest to a wide and international scientific audience will be published (see www.europeanjournaloftaxonomy.eu). ejt will accept major taxonomic contributions, taxonomic revisions, checklists of taxa on at least (sub) continental scale and opinion papers related to progress made in descriptive taxonomy. all submissions will be handled fully electronically through the open journal system, which we will use in line with our philosophy of open access publishing. peer reviewing will be rigorous, and both scientific and technical standards of the journal will be high. skilled publishing staff in the institutions constitute the production team of ejt. they will build a strong cross-institutional team, able to implement ongoing technical advances in the field of scientific publishing. coordinating institutional resources into a single journal contributes to excellence, prevents repetition, and increases efficiency in the dissemination of taxonomic data, while providing a secure, long-term publishing and archiving platform at minimal cost. although the title of the journal might hint of geographic restriction, the scope of ejt is global: authorship and region of study are not intended to be exclusively european. authors are, however, encouraged to involve european natural history collections by consulting extant material, or by depositing (type-) material related to the published papers in the collection of a european nhi. www.europeanjournaloftaxonomy.eu the edit / ejt task force, editorial: launch of ejt 3 ejt aims to offer all the modern interactive web-based facilities of high-level, high impact journals. it will provide links to all leading biodiversity-related databases in which new names of species and genera will automatically be included. ejt sets a high standard in taxonomic publishing. we hope that the members of the scientific community will find this new journal valuable and useful and will welcome it, both as authors and as readers. founding members of the ejt consortium are: • muséum national d’histoire naturelle (paris, france) • natural history museum (london, uk) • royal belgian institute of natural sciences (brussels, belgium) • royal museum for central africa (tervuren, belgium) • national botanic garden of belgium (meise, belgium). other potential members are presently being considered. we plan to enlarge the consortium to at least 10-15 european members, more if possible. this will enable us to continue enlarging the publication platform for taxonomy. because of its focus, ejt belongs to taxonomists, and we encourage taxonomists to engage with this initiative. we thank all members of european distributed institute of taxonomy (edit) who have shown an interest in the creation of this journal, and in particular simon tillier, the edit coordinator, for endowing this initiative both in principle and in practice. our gratitude goes also to patrick grootaert, from rbins (brussels, belgium) and michèle ballinger from cnrs (paris, france) who have both actively supported the project from the start. we are also grateful for the enthusiasm and professionalism of the team of journal managers endlessly working to improve ejt: natacha beau, kristiaan hoedemakers and charlotte thionois. pkp (public knowledge project) (http://pkp.sfu.ca/) kindly assisted us with the adaptation and implementation of their ojs (open journal system) system. references i20 10 digital libraries initiative, december 2009. final report: digital libraries: recommendations and challenges for the future. high level expert group on digital libraries: 17 p. sietmann r. 2008. introduction: quo vadis, knowledge society? in: openaccess– opportunities and challenges – a handbook by the european commission and the german commission for unesco. published on: 9 september 2011 http://pkp.sfu.ca/ http://ec.europa.eu/information_society/activities/digital_libraries/doc/hleg/reports/hlg_final_report09.pdf http://ec.europa.eu/research/science-society/document_library/pdf_06/open-access-handbook_en.pdf european journal of taxonomy 10: 1-3 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2012.10 www.europeanjournaloftaxonomy.eu 2012 · yves samyn & olivier de cock this work is licensed under a creative commons attribution 3.0 license. o p i n i o n p a p e r 1 no name, no game samyn y. & de clerck o. 2012. no name, no game. european journal of taxonomy 10: 1-3. http://dx.doi. org/10.5852/ejt.2012.10 yves samyn1 & olivier de clerck2 1 belgian national focal point to the global taxonomy initiative; royal belgian institute of natural sciences, vautierstraat 29, 1000 brussels, belgium. email: yves.samyn@naturalsciences.be (corresponding author). 2 phycology research group, ghent university, krijgslaan 281, building s8 9000 gent, belgium. “the most efficient solution to speeding taxonomic production, ultimately, is to train and hire more taxonomists.” (deans et al. 2012: 81) in an interesting contribution joppa et al. (2011) revisit some aspects of the taxonomic impediment (evenhuis 2007; http://www.cbd.int/gti/) and come to the conclusion that, contrary to the generally accepted idea, both the rates of species description and the number of taxonomists have increased exponentially since the 1950’s. joppa et al. (2011) also note a marked decline in the number of species described per taxonomist which they attribute to the difficulty of finding new species in an ever declining ‘missing species pool’. therefore, their results might be interpreted that today’s taxonomic workforce is sufficient to describe the remaining (shallow) ‘pool of missing species’. in this contribution, we question if this is indeed the case and propose a solution for speeding up taxonomic descriptions. we feel that joppa et al. (2011) are overly enthusiastic, probably because their test cases represent a selection of the better-studied taxa (flowering plants, conus snails, spiders, amphibians, birds and mammals). contradicting the findings of joppa et al. (2011) and using a much broader taxon sampling, which includes many poorly studied groups, costello et al. (2011) observe that species description has roughly remained constant since the second world war, at least for terrestrial species. these results are congruent with other counts of species descriptions (bacher 2012; chapman 2009; may 2011; zhang 2010; http://species.asu.edu/sos). so it seems that the increasing rate of species description as reported by joppa et al. (2011) is taxon specific rather than general. in stark contrast to the constant rate of species description is the rate with which species are discovered by using dna sequence data. exhaustive bio-inventory initiatives to map diversity of poorly explored areas around the world (e.g. http://laplaneterevisitee.org/; http://mooreabiocode.org) in combination with large-scale barcoding efforts (hajibabei et al. 2007; valentini et al., 2009) result in an explosion of species/sequences in repositories such as genbank and bold systems that are not linked to known species. this growing number of what have been called ‘dark taxa’ by page (http://iphylo.blogspot.com/2011/04/ dark-taxa-genbank-in-post-taxonomic.html), makes one question the use of proper taxonomic names at all. indeed, it can be argued that a lot of biology does not per se require formally described taxon names. however, the value of scientific names exceeds the field of biology sensu stricto and matters http://dx.doi.org/10.5852/ejt.2012.10 http://www.europeanjournaloftaxonomy.eu/index.php/ejt http://creativecommons.org/licenses/by/3.0/ http://dx.doi.org/10.5852/ejt.2012.10 http://dx.doi.org/10.5852/ejt.2012.10 mailto:yves.samyn@naturalsciences.be http://www.cbd.int/gti/ http://species.asu.edu/sos http://laplaneterevisitee.org/ http://mooreabiocode.org http://iphylo.blogspot.com/2011/04/dark-taxa-genbank-in-post-taxonomic.html http://iphylo.blogspot.com/2011/04/dark-taxa-genbank-in-post-taxonomic.html european journal of taxonomy 10: 1-3 (2012) 2 for society as a whole. for instance, the international legislation that controls the trade of endangered species (cites) uses species names as currency, just as the convention on migratory species and the convention on biological diversity and its associated access and benefit sharing protocol. the widening gap between discovered and described biodiversity and the difficulties of linking this new diversity to existing names could very well represent the true achilles’ heel of the taxonomic system. descriptive taxonomy is slow, despite cybertaxonomy and biodiversity informatics. 250 years of modern taxonomy makes that we now recognize nearly 2 million species (chapman 2009), but in many groups a multitude of synonyms exists for every accepted name. godfray (2002) clearly exaggerated when he compared the past to a dead weight choking every serious attempt of a modern revision. but, it is certainly true that linking clades, discovered through modern day molecular phylogenies, to often long forgotten names and their type material, consumes a too disproportionate amount of time for taxonomists. such is unacceptable and therefore curators of natural history collections have started to firmly embrace and invest in the possibilities of the internet to make their collections available and searchable digitally. taxonomists have also started to explore new ways in making available their research findings faster and more transparently for their peers. examples of good practice include the eu-funded edit platform for cybertaxonomy (http://wp5.e-taxonomy.eu/) and scratchpads (http://scratchpads.eu/) projects and the us nsf funded morphbank (http://www.morphbank.net/) and morphobank (http://www.morphobank. org/) projects. some (deans et al. 2012) dwell on roughly the same ideas and propose to go even further by establishing a database (‘phenobank’) that stores standardized, machine-readable semantic descriptions of specimens, not taxa. it remains to be tested if such approach would outcompete classical descriptions of taxa, not specimens, especially if these are framed into an integrative taxonomic approach (padial et al. 2010). another underestimated part of the problem is that most taxonomists sensu joppa et al., that is ‘those individuals who describe new species’ can only devote a fraction of their time on the description of new species. apart from the multitude of tasks that refrains many from inventorying and describing biodiversity, taxonomists also face a hypothesis driven scientific system that is ruled by impact factors or other metrics. this reality seduces many taxonomists to publish an ecological, evolutionary or biogeographic story rather than descriptive taxonomic papers. so given that species names matter, and that we need to close the gap between discovery and description of diversity, what to do? the answer, just as ‘anecdotally’ hinted by joppa et al. (2011), might be to increasingly involve those developing countries that have pro-actively chosen to document their biodiversity. this could be done by installing operational hubs in these countries, where local taxonomists as well as specialized technical staff are trained. development cooperation agencies from all over the world might be interested in providing financial aid to such initiatives, given of course, that they can be convinced that ’no name means no game’. the belgian development cooperation has, through its taxonomic capacity building program (http://www.taxonomy.be), realized this already several years ago. the european journal of taxonomy (ejt) which publishes descriptive taxonomy, (including redescriptions of taxa) regional or global checklists as well as taxonomic revisions and monographs, might serve as the ideal forum to release the obtained results, as long as the quality of the descriptions is rigorously assured through thorough peer review and firm editorial policy. if ejt accepts this challenge, it will become instrumental in speeding up the delivery of taxonomic names. by doing so, ejt will drive the implementation of legally binding international treaties in the field of biodiversity conservation and sustainable management. references bacher s. 2012. still not enough taxonomists: reply to joppa et al. trends in ecology and evolution 27: 65-66. http://dx.doi.org/10.1016/j.tree.2011.11.003 chapman a.d. 2009. numbers of living species in australia and the world. 2nd edition. australian biodiversity information services: toowoomba, australia. http://wp5.e-taxonomy.eu/ http://scratchpads.eu/ http://www.morphbank.net/ http://www.morphobank.org/ http://www.morphobank.org/ http://www.taxonomy.be/ http://dx.doi.org/10.1016/j.tree.2011.11.003 samyn y. & de clerck o, no name, no game 3 costello m.j., wilson s. & houlding. b. 2011. predicting total global species richness using rates of species description and estimates of taxonomic effort. systematic biology, first published online 18 aug. 2011 http://dx.doi.org/10.1093/sysbio/syr080 deans a.r.,yoder m.j. & balhoff j.p. 2012. time to change how we describe biodiversity. trends in ecology and evolution 27: 78-84. http://dx.doi.org/10.1016/j.tree.2011.11.007 evenhuis n.l. 2007. helping solve the “other” taxonomic impediment: completing the eight steps to total enlightenment and taxonomic nirvana. zootaxa 1407: 3-12. godfray h.ch. 2002. challenges for taxonomy. nature 417: 17-19. http://dx.doi.org/10.1038/417017a hajibabaei m., singer g.a.c., hebert p.d.n. & hickey d.a. 2007. dna barcoding: how it complements taxonomy, molecular phylogenetics and population genetics. trends in genetics 23: 167-172. http:// dx.doi.org/10.1016/j.tig.2007.02.001 joppa l.n., roberts d.l. & pimm s.l. 2011. the population ecology and social behavior of taxonomists. trends in ecology and evolution 26: 551-553. http://dx.doi.org/10.1016/j.tree.2011.07.010 may r.m. 2011. why worry about how many species and their loss? plos biology 9: e1001130. http://dx.doi.org/10.1371/journal.pbio.1001130 padial j.m., miralles a., de la riva i. & vences m. 2010. the integrative future of taxonomy. frontiers in zoology 7:16. http://dx.doi.org/10.1186/1742-9994-7-16 valentini a., pompanon f. & taberlet p. 2009. dna barcoding for ecologists. trends in ecology and evolution 24: 110-117. http://dx.doi.org/10.1016/j.tree.2008.09.011 zhang z-q. 2010. describing unexplored biodiversity: zootaxa in the international year of biodiversity. zootaxa 2768: 1-4. manuscript received: 5 march 2012 manuscript accepted: 23 march 2012 published on: 27 march 2012 topic editor: koen martens in compliance with article 8.6 of the iczn, printed versions of all papers are deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; national botanic garden of belgium, meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark. http://dx.doi.org/10.1093/sysbio/syr080 http://dx.doi.org/10.1016/j.tree.2011.11.007 http://dx.doi.org/10.1038/417017a http://dx.doi.org/10.1016/j.tig.2007.02.001 http://dx.doi.org/10.1016/j.tig.2007.02.001 http://dx.doi.org/10.1016/j.tree.2011.07.010 http://dx.doi.org/10.1371/journal.pbio.1001130 http://dx.doi.org/10.1186/1742-9994-7-16 http://dx.doi.org/10.1016/j.tree.2008.09.011 european journal of taxonomy 172: 1–6 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2016.172 www.europeanjournaloftaxonomy.eu 2016 · ustjuzhanin p. et al. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:3c881225-2a0b-42c1-8db0-57803281ac87 1 new species of “giant” plume moths of the genus platyptilia (lepidoptera, pterophoridae) from uganda peter ustjuzhanin 1, vasiliy kovtunovich 2 & anna ustjuzhanina 3,* 1 altai state university, lenina 61, barnaul 656049, russia. 2 moscow society of nature explorers. home address: malaya filevskaya str. 24/1, app. 20, moscow 121433, russia. 3 national research tomsk polytechnic university, lenina 30, tomsk 634050, russia. 1 e-mail: petrust@mail.ru 2 e-mail: vasko-69@mail.ru * corresponding author: uak@tpu.ru 1 urn:lsid:zoobank.org:author:60f67ccf-f6c9-4cd4-a2df-f3216dc46958 2 urn:lsid:zoobank.org:author:1959492d-b1a5-45f7-9c11-f079d399b42b 3 urn:lsid:zoobank.org:author:4a20cb2f-2fa9-4fb6-9983-82e203ffb0e1 abstract. this paper describes two new species of plume moths from the group of the so-called “giant” platyptilia hubner, 1825: platyptilia fletcheri ustjuzhanin & kovtunovich sp. nov. and p. stanleyi ustjuzhanin & kovtunovich sp. nov. both species were collected in the rwenzori mountains in uganda and rwanda, respectively. platyptilia stanleyi ustjuzhanin & kovtunovich sp. nov. exceeds all the known african species of pterophoridae in its wingspan of 49 mm. key words. pterophoridae, platyptilia, uganda, rwenzori mountains, new species. ustjuzhanin p., kovtunovich v. & ustjuzhanina a. 2016. new species of “giant” plume moths of the genus platyptilia (lepidoptera, pterophoridae) from uganda. european journal of taxonomy 172: 1–6. http://dx.doi. org/10.5852/ejt.2016.172 introduction the genus platyptilia hübner, 1825 is widespread all over the world. it is represented by 80 species, more than 40 of them known from africa (gielis 2003, 2011). among them are very large species, having about five centimeters in wingspan. they have been described by meyrick (1924, 1932, 1938), gielis (2008, 2011) and kovtunovich & ustjuzhanin (2014) and include: platyptilia aarviki gielis, 2008, p. daemonica meyrick, 1932, p. melitroctis meyrick, 1924, p. postbarbata meyrick, 1938, p. rhyncholoba meyrick, 1924, p. nyungwea gielis, 2011 and p. mugesse kovtunovich & ustjuzhanin, 2014. when examining material at the natural history museum, london, we discovered two more new species, belonging to the group of “giant platyptilia”: platyptilia fletcheri ustjuzhanin & kovtunovich sp. nov. and p. stanleyi ustjuzhanin & kovtunovich sp. nov. both species were collected during the famous british expeditions to the rwenzori mountains in 1934–1935 and 1952, and three specimens were found in rwanda (t.a. barns) in 1921. http://dx.doi.org/10.5852/ejt.2016.172 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:3c881225-2a0b-42c1-8db0-57803281ac87 mailto:petrust%40mail.ru?subject= mailto:vasko-69%40mail.ru?subject= mailto:uak%40tpu.ru?subject= http://zoobank.org/urn:lsid:zoobank.org:author:60f67ccf-f6c9-4cd4-a2df-f3216dc46958 http://zoobank.org/urn:lsid:zoobank.org:author:1959492d-b1a5-45f7-9c11-f079d399b42b http://zoobank.org/urn:lsid:zoobank.org:author:4a20cb2f-2fa9-4fb6-9983-82e203ffb0e1 http://dx.doi.org/10.5852/ejt.2016.172 http://dx.doi.org/10.5852/ejt.2016.172 european journal of taxonomy 172: 1–6 (2016) 2 material and methods this paper is based on material from the natural history museum, london, uk (bmnh). holotypes and paratypes of the new species are stored in bmnh, and two paratypes of platyptilia fletcheri ustjuzhanin & kovtunovich sp. nov. are stored in the collections of p. ustjuzhanin and v. kovtunovich (cuk, novosibirsk and moscow, russia). the preparation of genitalia is necessary for the identification of pterophoridae. the dissection was performed with standard methods. results class hexapoda blainville, 1816 order lepidoptera linnaeus, 1758 superfamily pterophoroidea kuznetsov & stekolnikov, 1979 family pterophoridae zeller, 1841 subfamily platyptilinae tutt, 1906 tribe platyptilini bigot, gibeaux, nel & picard, 1998 genus platyptilia hübner, 1825 platyptilia fletcheri ustjuzhanin & kovtunovich sp. nov. urn:lsid:zoobank.org:act:785f8238-9629-4217-8417-13973f95cc30 figs 1–4 diagnosis platyptilia fletcheri ustjuzhanin & kovtunovich sp. nov. can be distinguished by the large wings and variegated coloring: from dark-brown without any pattern to brown-grey with the pattern of pale areas, spots and dots. in the male genitalia the new species resembles platyptilia aarviki gielis, 2008, but clearly differs from it by the shape of the sacculus, the long anellus arms and also the wedge-shaped cut figs 1-4. platyptilia fletcheri ustjuzhanin & kovtunovich sp. nov. 1. adult, holotype, ♂, habitus (bmnh 21796). 2. adult, paratype, ♂, melanistic color (bmnh 21801). 3. male genitalia, holotype. 4. female genitalia, paratype (bmnh 21802). http://zoobank.org/urn:lsid:zoobank.org:act:785f8238-9629-4217-8417-13973f95cc30 ustjuzhanin p. et al., “giant” plume moths from uganda 3 of the saccus. in the female genitalia the new species resembles platyptilia sabia (felder & rogenhofer, 1875) by the shape of the antrum, but differs from it by the more narrow ostium and shorter ductus. etymology the species is named after the famous british entomologist david stephen fletcher, member of the 1952 expedition to the rwenzori mountains. material examined holotype uganda: ♂, ruwenzori range, nyamaleju, 10,530 ft, 14–19 jul. 1952, d.s. fletcher, ruwenzori exped. b.m. 1952-566 (bmnh 21796). paratypes uganda: 2 ♂♂, same data as holotype (bmnh 21800, cuk 244); 1 ♂, ruwenzori range, lake mahoma, 9,600 ft, 12 jul. 1952, d.s. fletcher (bmnh 21797); 1 ♀ kigezi dist., mt mgahinga, 8,000 ft, 22–27 nov. 1934, f.w. edwards (bmnh 21802); 1 ♂, 2 ♀♀, ruwenzori range, namwamba valley, 6,500 ft, dec. 1934–jan. 1935, f.w. edwards, b.m.e. afr. exp. b.m. 1935-203 (bmnh 21801, bmnh 21799, cuk 245). rwanda: 1 ♂, kisiba, bugoie forest, w kivu, 8,500 ft, nov. 1921, t.a. barns (bmnh 21798); 2 ♀♀, virunga mts, lake kivu, 10,000 ft, oct. 1921, t.a. barns (bmnh 22764). description external characters. wingspan 38–40 mm (holotype: 40 mm). head, thorax and tegulae brown-grey. labial palpi straight, brown, three times as long as than eye diameter; third segment narrowed at tip and bent down. antennae thin, brown. fore wings brown-grey. costal margin significantly darker than medial area. two brown spots near wing base. two fuzzy brown spots not reaching cleft. first lobe with yellowish longitudinal stripes, apical part tapered. second lobe pale-grey with dusting of brown scales in form of shapeless brushstrokes. fringe pale grey inside cleft, brown-grey at outer wing margin, darkbrown in basal part. hind wings unicolorous, pale grey. third lobe fringe with well defined black hairs at outer margin, most intensively expressed in basal part. abdomen grey, rather long. hind legs long, pale-brown, noticeably darkened at base of spurs. male genitalia. valves symmetrical. аpex isolated and drawn downwards. sacculus wide in basal part, then narrowing and slightly widened in distal part. tegumen bilobed. uncus narrow, slightly curved, tapered at apex. juxta narrow, arched. anellus arms long, wide in basal part, strongly narrowed and slightly bent inward in distal part. saccus with proximate wedge-like outer edge and with triangular process from above, at its inner side. phallus curved in centre, bluntly rounded, basal process straight, equal to length of coecum. vesica with thin spinous cornutus in distal part of phallus. female genitalia. papillae anales oval, narrow. apophyses posteriores thin, long, three times longer than papillae anales. apophyses anteriores also long, slightly shorter than posteriores ones, with small, spinous process in distal part. ostium flat. antrum wide, its length twice the length of papillae anales. ductus bursae narrow, 1.5 times longer than antrum, with thin sclerite in middle part. ductus seminalis near junction with bursa copulatrix. bursa copulatrix large, oval, with two short, straight signa tapered to apices. remarks the new species is characterized by the bright variation in the color of the wings. half of the 11 type specimens have melanistic color. the species inhabits the mountainous areas of 1980–3210 m above sea level. european journal of taxonomy 172: 1–6 (2016) 4 distribution uganda, rwanda. flight period july, october–january. platyptilia stanleyi ustjuzhanin & kovtunovich sp. nov. urn:lsid:zoobank.org:act:5429545b-1fa2-469c-bf69-a81605adaea2 figs 5–6 diagnosis platyptilia stanleyi ustjuzhanin & kovtunovich sp. nov. can be distinguished by its giant wingspan, which is unique among all the species of the genus platyptilia as well as among all the african pterophoridae. in the male genitalia the new species resembles platyptilia fletcheri ustjuzhanin & kovtunovich sp.nov. in the shape of the phallus and uncus, but clearly differs from it by the short and wide anellus arms and the fold on the inner side of the sacculus. etymology the species is named after the famous british traveler, journalist and explorer of africa, henry morton stanley, the first european who visited the rwenzori mountains in 1876. material examined holotype uganda: ♂, ruwenzori range, nyamaleju, 10,530 ft, 14–19 jul. 1952, ruwenzori exped. b.m. 1952-566, d.s. fletcher (bmnh 21803). description male external characters. wingspan 49 mm. thorax and tegulae brown-grey, head noticeably paler. labial palpi straight, twice as long as eye diameter; third segment narrowed at top and bent down. antennae thin, brown. fore wings brown. medial area slightly clarified. dark brown spot in middle part of first quarter of wing. two other spots (poorly expressed and slightly fuzzy) near cleft. first lobe dark brown in apical part. second lobe significantly darker than basic wing color. fringe inside cleft brown-grey; grey at outer margin of wing, with bunches of dark brown cilia. hind wings unicolorous, pale grey. third lobe fringe with well expressed black cilia at outer margin, from middle of wing to base of lobe. abdomen brown. hind legs long, brown, with portions of dark scales at base of spurs and in distal part. figs 5-6. platyptilia stanleyi ustjuzhanin & kovtunovich sp. nov., holotype, ♂ (bmnh 21803). 5. adult, habitus. 6. male genitalia. http://zoobank.org/urn:lsid:zoobank.org:act:5429545b-1fa2-469c-bf69-a81605adaea2 ustjuzhanin p. et al., “giant” plume moths from uganda 5 male genitalia. valves symmetrical. аpex isolated and slightly tapered. sacculus wide in basal and middle part; distal part significantly narrowed. fold on inner side of sacculus. tegumen bilobed. uncus narrow, rather long, slightly widened in middle part and tapered at apex. anellus arms short, wide, tapered only at apex. saccus with proximate wedge-like outer edge and with triangular process from above, on its inner side. phallus short, curved in center and rounded at base; basal process long, equal to length of coecum. vesica with thin, needle-shaped cornutus in distal part of phallus. female unknown. distribution uganda. flight period july. acknowledgments the authors are grateful to kevin tuck, the former curator of the microlepidoptera collection at the natural history museum (london, uk) for access to the pterophoridae collections and granting a loan of the museum material for examination. references gielis c. 2003. pterophoroidea & alucitoidea (lepidoptera). world catalogue of insects 4: 1–198. http://dx.doi.org/10.1002/mmnd.20030500211 gielis c. 2008. ten new species of afrotropical pterophoridae. zoologische mededelingen 82: 43–57. gielis c. 2011. notes on some african pterophoridae, with description of new species (lepidoptera). boletin de la sociedad entomologica aragonesa (s.e.a.) 49: 33–63. kovtunovich v.n., ustjuzhanin p.ya. & murphy r. 2014. plume moths of malawi (lepidoptera: pterophoridae). zootaxa 3847: 451–494. http://dx.doi.org/10.11646/zootaxa.3847.4.1 meyrick e. 1924. zoological results of the swedish expedition to central africa, 1921. insecta 4: microlepidoptera. arkiv för zoologi 16 (14): 1–2. meyrick e. 1932. entomological expedition to abyssinia 1926–27. microlepidoptera. transactions of the entomological society of london 80: 107–109. meyrick e. 1938. exploration du parc national albert. mission g.f. witte (1933–1935). fasc. 14. pterophoridae, tortricina and tineina: 3–28. institut des parcs nationaux du congo belge, brussels. manuscript received: 8 october 2015 manuscript accepted: 10 november 2015 published on: 18 january 2016 topic editor: koen martens desk editor: kristiaan hoedemakers http://dx.doi.org/10.1002/mmnd.20030500211 http://dx.doi.org/10.11646/zootaxa.3847.4.1 european journal of taxonomy 172: 1–6 (2016) 6 printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark. european journal of taxonomy 31: 1-8 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2012.31 www.europeanjournaloftaxonomy.eu 2012 · parmentier e. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:7acaf744-b665-4008-82bb-339e41808dd4 1 echiodon prionodon, a new species of carapidae (pisces, ophidiiformes) from new zealand eric parmentier laboratoire de morphologie fonctionnelle et evolutive, institut de chimie, bât. b6c, université de liège, b-4000 liège, belgium. email: e.parmentier@ulg.ac.be urn:lsid:zoobank.org:author:18447bb6-9cd7-49d3-8269-553783c3eb2c abstract. a new species of pearlfish, echiodon prionodon, is described from three specimens. this species is diagnosed by having a serrated margin on the posterior edge of the fangs, expanded thoracic plates on some abdominal vertebrae and ventral swimbladder tunic ridges. this species was only found in coastal waters around the north island of new zealand. the diagnosis of eurypleuron is revised. key words. echiodon, eurypleuron, pearlfish, new zealand. parmentier e. 2012. echiodon prionodon, a new species of carapidae (pisces, ophidiiformes) from new zealand. european journal of taxonomy 31: 1-8. http://dx.doi.org/10.5852/ejt.2012.31 introduction the carapidae, which include pyramodontinae and carapinae, are eel-like fishes. they range from shallow water to moderately deep waters of the continental slope (nielsen et al. 1999). several species belonging to the genera onuxodon, carapus and encheliophis are well known for their unusual behavior of entering and living inside invertebrate hosts such as sea cucumbers, sea stars, or bivalves (trott 1981). some echiodon species could be commensals with sponges or polychaete worm tubes (nielsen et al. 1999), but it is not proven. the echiodontini tribe (carapinae, carapidae) comprises three genera (markle & olney 1990). 1) onuxodon smith, 1955, with three species, diagnosed by the presence of a rocker bone in front of the swimbladder and a commensal relationships with species of mollusca (tyler 1970; parmentier et al. 2000; amaoka & yoseda 2005). 2) echiodon thompson, 1837, comprising a group of 11 valid species (and 1 unnamed larva) found over a depth range of 18–2000 m (williams 1984a, b; markle & olney 1990; williams & machida 1992; anderson 2005). this taxon is supported by one synapomorphy: a ventral patch of tunic ridges on the posterior swimbladder (markle & olney 1990). 3) eurypleuron markle & olney, 1990, characterized by having thoracic plates formed by expanded parapophyses (transverse processes) on the fifth through 18-20th vertebrae (males only) and an exterilium gut supported by elongate, cartilaginous, ventral processes of the coracoid in vexillifer larvae. markle & olney (1990) considered the genus monospecific, eurypleuron owasianum (matsubara, 1953), with two major disjunct populations. however, these two populations have differences in d30 counts, precaudal http://dx.doi.org/10.5852/ejt.2012.31 http://www.europeanjournaloftaxonomy.eu/index.php/ http://creativecommons.org/licenses/by/3.0/ http://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:7acaf744-b665-4008-82bb-339e41808dd4 mailto:e.parmentier%40ulg.ac.be?subject=e.parmentier%40ulg.ac.be http://zoobank.org/urn:lsid:zoobank.org:author:18447bb6-9cd7-49d3-8269-553783c3eb2c http://dx.doi.org/10.5852/ejt.2012.31 european journal of taxonomy 31: 1-8 (2012) 2 vertebral numbers and stomach color. on the basis of these characters and the antitropical distributions (the northern population is found above 20° north and southern population occurs below 20° south), williams & machida (1992) proposed that these populations correspond to two different species: eurypleuron owasianum in the north and eurypleuron cinereum (smith, 1955) in the south. according to the authors, additional specimens are required to better resolve this issue. examination of the collection of carapids held at the museum of new zealand te papa tongarewa (te papa) revealed three specimens with characters shared by echiodon and eurypleuron. the three specimens are here described as a new species and comments on the diagnosis of these genera are given. material and methods measurements were made with digital calipers to the nearest 0.1 mm. information of the axial skeleton was obtained from x-ray digital photographs. the following abbreviations for meristic characters are from markle & olney (1990): a30 = anal-fin rays anterior to vertical through articulation between vertebrae 30 and 31 ardo = anal-fin rays anterior to dorsal-fin origin d30 = dorsal-fin rays anterior to vertical through vertebrae 30 and 31 p1 = pectoral-fin rays pcv = precaudal vertebrae vao = vertebrae to anal-fin origin vdo = vertebrae to dorsal-fin origin vpb = vertebra number under predorsal bone other abbreviations are: hl = head length tl = total length faku = faculty of agriculture, kyoto university nmnz = museum of new zealand te papa tongarewa rusi = rhodes university, j.l.b. smith institute of ichthyology, grahamstown zmuc = zoological museum of the university of copenhagen the present specimens were compared with echiodon cryomargarites markle, williams & olney, 1983 (nmnz p016530, nmnz p016537), echiodon pegasus markle & olney, 1990 (nmnz p016529, nmnz p020999), echiodon rendahli (whitley, 1941) (nmnz p014855) and with radiographs of echiodon neotes markle & olney, 1990 (zmuc p77815), eurypleuron cinereum (rusi 309) and eurypleuron owasianum (faku 34517, 34518, 34519). parmentier e., echiodon prionodon sp. nov. from new zealand 3 results class actinopterygii klein, 1885 order ophidiiformes berg, 1937 family carapidae jordan & fowler, 1902 genus echiodon thompson, 1837 echiodon prionodon sp. nov. urn:lsid:zoobank.org:act:44a01315-f569-4553-b342-69c5d1c51acf figs 1-4 diagnosis a species of echiodon, with a serrated posterior margin on the fangs, expanded thoracic plates on some abdominal vertebrae, pcv 33–35, d30 42–45, a30 40–41. etymology from the greek priôn meaning saw, and odous (odon) meaning tooth, in reference to the unique morphology of the fang at the tip of the jaw. type material holotype ♂, 165 mm tl, in nmnz (p.041833), rv tangaroa, sta. tan 0413/119, 13 nov. 2004. paratypes nmnz p.003281 (97 mm tl), off kapiti island, 40°51.0’s – 174°52.0’e, fred abernethy, 37 m, apr. 1958, badly cleared and stained specimen; nmnz p.052493 (57+ mm, anterior portion only), outer north taranaki bight, rv tangaroa, sta. tan 1105/137, 38°26.70’s – 173°18.97’e, 170-240 m epibenthic sled, 5 apr. 2011. type locality off white island, outer bay of plenty, new zealand, 37° 33.57’s – 176° 59.1-58.8’e, 313 m bottom trawl. fig 1. echiodon prionodon sp. nov., left lateral view (holotype nmnz p.041833, 165 mm tl). http://zoobank.org/urn:lsid:zoobank.org:act:44a01315-f569-4553-b342-69c5d1c51acf european journal of taxonomy 31: 1-8 (2012) 4 fig. 2. echiodon prionodon sp. nov., left lateral view of the left premaxillary fang showing the posterior serrated margin. nmnz p.041833 holotype nmnz p.003281 paratype nmnz p.052493 paratype total length (mm) 165 97 head length 17.8 12.8 12.5 preanus length 21.1 (118) 14.9 (116) 15.4 (123) predorsal length 26.3 (147) 15.8 (123) 17.8 (142) snout length 2.5 (14) 2.0 (15) 2.2 (17) upper jaw length 7.8 (44) 5.9 (46) 6.0 (48) lower jaw length 9.3 (52) 6.4 (49) 6.8 (54) horizontal eye diameter 4.1 (23) 3.3 (25) 2.9 (23) head depth 6.9 (39) 5.3 (41) depth at anus 8.4 (47) 5.8 (45) 6.3 (50) interorbital length 2.2 (12) 1.4 (11) 1.5 (12) pectoral length 8.7 (49) 8.2 (65) meristic pcv 33 33 35 d30 45 42 a30 40 41 vdo 8 8 8 vao 7 6 7 vpb 7 7 p1 15 14 13 table 1. morphometric (mm) and meristic data for echiodon prionodon sp. nov. numbers in brackets refer to measurements in % head length. parmentier e., echiodon prionodon sp. nov. from new zealand 5 description selected counts and measurements are given in table 1 and the holotype is shown in figure 1. body slender, much higher than wide, tapering into a pointed tail; greatest body depth (at anus) approximately 5% of total length; no caudal fin; dorsal fin origin posterior to anal fin origin, anus ends in a tube at half length of the pectoral fin. dorsal profile of head slightly convex. snout rounded in lateral view, slightly projecting beyond upper jaw. olfactory lobe approximately 70% of snout length, closer to the eye than to the snout tip; anterior nostril developed in a small tube; posterior nostril directly in front of anterior margin of eye, elliptical, being higher than long. eye elongate, longer than high. mouth oblique, upper jaw extending beyond posterior margin of eye; posterior portion of maxilla unsheathed. anterior tip of lower jaw behind tip of snout, lower jaw occlusion with palatine bones. short opercular spine exposed through an elliptical slit in skin. gill opening extends from upper end of pectoral fin base to below rear end of maxillary. seven branchiostegal rays. ceratobranchials 1 with three slender gill-rakers, with tooth pads on upper limb; other gill-rakers tubercular. two enlarged caniniform teeth near symphysis in premaxilla and in dentary (one tooth missing on left lower jaw in holotype). each of these caniniform teeth has a serrated margin posteriorly (fig. 2). upper and lower jaw fangs are separated from the posterior teeth rows by a pronounced diastema. eight to nine rows of small, minute, straight, conical teeth on dentary, 3-4 outer rows with smaller teeth. four to five rows of small conical teeth on the upper jaw, teeth on inner rows somewhat longer and curved inwardly, teeth of outer rows similar to lower jaw teeth. palatine with 4-5 rows of villiform teeth. vomer a small oblong bump with irregular disposition of small conical teeth, posterior one being somewhat bigger. in the holotype (fig. 3), vertebral centra 6 to 24 with parapophyses expanded to form lateral plates (vertebral centra 6 to 14 in nmnz p.052493, 6 to 17 in nmnz p.003281). holotype swimbladder extends to 30th vertebral centra (57.9 mm from the snout tip), slight central constriction separating anterior part, brown with dark spots, and shorter posterior part, unpigmented and with ventral tunic ridges. color pattern after eight years in alcohol, the holotype has cream-colored body and head. melanophores highly concentrated at the level of the geniohyoideus (throat), but not on the lower jaws, at the oro-branchial cavity on the tongue, the palate, branchial arches, on the inner face of the opercula, on the parietals and on the posterior parts of the frontal. stomach and peritoneal cavity black, anus unpigmented. melanophores concentrated at the base of dorsal and anal fins and forming lines on the lateral lines and on different myosepta. pterygiophores of anal and dorsal fins black from the tip to approximately 1/5 of the body length. differential diagnosis echiodon prionodon sp. nov. is unique in having serrated posterior margin on the fangs. moreover, it differs from all others echiodon species by the thoracic plates on some abdominal vertebrae and it differs from eurypleuron species by having tunic ridges on the swimbladder. fig. 3. echiodon prionodon sp. nov., left lateral x-ray (holotype nmnz p.041833, 165 mm tl). european journal of taxonomy 31: 1-8 (2012) 6 distribution endemic to coastal waters around the north island of new zealand (fig. 4). the species seems to be benthic from 30 to 315 m depth. the holotype was caught along with other fish species which usually are associated with shelly-gravel to sandy bottoms with patch reefs as gnathophis umbrellabius (whitley, 1948) (congridae), hoplostethus mediterraneus cuvier, 1829 (trachichthyidae) and paraulopus nigripinnis (günther, 1878) (paraulopidae). discussion echiodon and eurypleuron are sister genera with similar morphologies (markle & olney 1990). both have eel-like bodies, one to several large symphyseal fangs on dentary and premaxilla. they lack cardiform teeth, pelvic bones and rocker bone (nielsen et al. 1999). in the carapid cladogram, markle & olney (1999: fig. 45) noted two non-homoplastic characters in support of eurypleuron: greatly expanded plate-like parapophyses on centre 5 to 18-22 (males only) and an exterilium gut in larvae. the genus is currently monotypic but williams & machida (1992) recommended more research. they argued, that the genus consists of two species: a northern pacific population with eurypleuron owasianum (matsubara, 1953) and a southern pacific population with eu. cinereum (smith, 1955). in eurypleuron fig. 4. distribution of echiodon prionodon. the red circle depicts the locality where the holotype was caught. parmentier e., echiodon prionodon sp. nov. from new zealand 7 from both hemispheres, the only autapomorphic character (ventral tunic ridges on the swimbladder) that supports echiodon was not found. echiodon prionodon sp. nov. shows characters of both genera: ventral tunic ridges on the swimbladder and expanded parapophyses. the number of plate-like parapophyses is, however, more variable than in eurypleuron owasianum (from 8 to 18), which seems related to the size of the fish. i was unable to confirm the sex of the three specimens and therefore cannot confirm if it is a sex-related character. in addition, radiographs do not show the parapophyses expanded in the same way as in eurypleuron and i conclude that this character results from either convergence or is a homoplastic character. many ophidiiformes use their swimbladder and associated bones to produce sounds (courtenay & mckittrick 1970; parmentier et al. 2002; fine et al. 2007; parmentier et al. 2010). previous anatomical studies showed all the carapids have sound-producing muscles and therefore are able to produce sounds (parmentier et al. 2002). it is noteworthy that all the characters separating adult echiodon from eurypleuron are related to the sound-producing mechanism (parmentier & diogo 2006). this system could play an important role in the speciation process of the group. because swimbladder ventral tunic ridges are common to all the echiodon species, it is more parsimonious to place ec. prionodon sp. nov. in this genus; otherwise, the genus would be deprived of autapomorphic characters. moreover, ec. prionodon sp. nov. shares additional characters with other echiodon species. it has in common with ec. drummondi thompson, 1837, ec. dentatus (cuvier, 1829) and ec. rendahli a central constriction of the swimbladder, with ec. rendahli a high number of pcv, the same kind of dentition (minute teeth in several rows on both lower and upper jaws) and the same geographic range (williams 1984a; markle & olney 1990). more research is required to define the diagnosis of eurypleuron. the former diagnosis was: thoracic plates formed by expanded parapophyses (transverse processes) on the fifth through 18-20th vertebrae (males only); an exterilium gut supported by elongate cartilaginous ventral processes of the coracoid in vexillifer larvae; lack of ventral tunic ridges on the posterior portion of the swim bladder; dorsal-fin origin over anal-fin origin resulting in equivalent (or almost so) a30 and d30 values (markle & olney 1990). two characters are now invalid: expanded parapophysis (this study) and the equal d30 and a30 counts, recently also found in the newly described species echiodon atopus anderson, 2005 from the south atlantic (anderson 2005). acknowledgements i am grateful to the fishes team of the museum of new zealand te papa tongerewa for providing facilities and access to the museum collection. carl struthers photographed the holotype and jeremy barker made x-rays . clive roberts and andrew stewart commented on the early draft of this paper. references amaoka k. & yoseda y. 2005. two species of pearl fishes, onuxodon parvibrachium and o. fowleri (ophidiiformes, carapidae) from the honeycomb oyster (hyotissa hyotis), from ishigaki island, okinawa prefecture. bulletin of the biogeographical society of japan 60: 5-12. anderson m.e. 2005. description of a new species of echiodon (teleostei: carapidae) from the south atlantic ocean. zootaxa 809: 1-5. courtenay w.r. & mckittrick f.a. 1970. sound-producing mechanisms in carapid fishes, with notes on phylogenetic implications. marine biology 7: 131-137. european journal of taxonomy 31: 1-8 (2012) 8 fine m.l., lin h., nguyen b.b., rountree r.a., cameron t.m. & parmentier e. 2007. functional morphology of the sonic apparatus in the fawn cusk-eel lepophidium profundorum (gill, 1863). journal of morphology 268: 953-966. http://dx.doi.org/10.1002/jmor.10551 markle d.f. & olney j.e. 1990. systematics of the pearlfish (pisces: carapidae). bulletin of marine science 47: 269-410. nielsen j.g., cohen d.m., markle d.f. & robins c.r. 1999. fao species catalogue. volume 18. ophidiiform fishes of the world (order ophidiiformes). an annotated and illustrated catalogue of pearlfishes, cusk-eels, brotulas and other ophidiiform fishes known to date. fao fisheries synopsis, 125. fao, rome. parmentier e., castro-aguirre j.l. & vandewalle p. 2000. morphological comparison of the buccal apparatus in two bivalve commensal teleostei: encheliophis dubius and onuxodon fowleri (carapidae, ophidiiformes). zoomorphology 120: 29-37. http://dx.doi.org/10.1007/s004359900020 parmentier e., chardon m. & vandewalle p. 2002. preliminary study on the ecomorphological signification of the sound-producing complex in carapidae. in: aerts p., d’août k., herrel a. & van damme r. (eds) topics in functional and ecological vertebrate morphology: 139-151. shaker publishing, maastricht. parmentier e. & diogo r. 2006. evolutionary trends of swimbladder sound mechanisms in some teleost fishes. in: ladich f., collin s.p., moller p. & kapoor b.g. (eds) communication in fishes, vol. i: 43-68. science publishers, enfield. parmentier e., bouillac g., dragičević b., dulčić j. & fine m.l. 2010. call properties and morphology of the sound-producing organ in ophidion rochei (ophidiidae). journal of experimental biology 213: 3230-3236. http://dx.doi.org/10.1242/jeb.044701 trott l.b. 1981. a general review of the pearlfishes (pisces, carapidae). bulletin of marine science. 31: 623-629. tyler j.c. 1970. a redescription of the inquiline carapid fish onuxodon parvibrachium, with a discussion of the skull structure and the host. bulletin of marine science 20: 148-164. williams j.t. 1984a. studies on echiodon (pisces: carapidae), with description of two new indo-pacific species. copeia 1984 (2): 410-422. williams j.t. 1984b. synopsis and phylogenetic analysis of the pearlfish subfamily carapinae (pisces: carapidae). bulletin of marine science 34: 386-397. williams j.t. & machida y. 1992. echiodon anchipterus: a valid western pacific species of the pearlfish family carapidae with comments on eurypleuron. japanese journal of ichthyology 38: 367-373. manuscript received: 29 august 2012 manuscript accepted: 21 november 2012 published on: 7 december 2012 topic editor: rudy jocqué printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; national botanic garden of belgium, meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark http://dx.doi.org/10.1002/jmor.10551 http://dx.doi.org/10.1007/s004359900020 http://dx.doi.org/10.1242/jeb.044701 european journal of taxonomy 175: 1–9 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2016.175 www.europeanjournaloftaxonomy.eu 2016 · grichanov i.ya. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:cadca890-3440-49f6-afc8-ffbe9fb22d2d 1 two new species of dactylonotus parent, 1934 (diptera: dolichopodidae) from south africa and a key to afrotropical species igor ya. grichanov all-russian institute of plant protection, podbelskogo 3, st.petersburg, pushkin, 196608, russia. e-mail: grichanov@mail.ru urn:lsid:zoobank.org:author:5320ad3a-92d8-4820-8091-24802f8c8c06 abstract. two new species from south africa, dactylonotus nigricorpus sp. nov. and dactylonotus tsitsikamma sp. nov., are described and illustrated. d. nigricorpus sp. nov. differs from all other species of the genus in the black body, the smaller size and the shorter antenna. d. tsitsikamma sp. nov. is peculiar in the genus in bearing a flag of long setae on the fifth segment of the fore tarsus. an identification key to 6 afrotropical species of the genus is provided. key words. diaphorinae, new species, identification key, afrotropical region. grichanov i.ya. 2016. two new species of dactylonotus parent, 1934 (diptera: dolichopodidae) from south africa and a key to afrotropical species. european journal of taxonomy 175: 1–9. http://dx.doi.org/10.5852/ ejt.2016.175 introduction based on recent results of a cladistics analysis, capellari (2013) demonstrated the monophyly of the generic group including anepsiomyia bezzi, 1902, argyra macquart, 1834, dactylonotus parent, 1934, somillus brèthes, 1924 and symbolia becker, 1921 (composing the tribe argyrini negrobov, 1986, placed currently within the subfamily diaphorinae, but deserving subfamily rank). the genus dactylonotus is peculiar in its disjunctive distribution, with six species known from southern africa and one species, d. formosus (parent, 1933), inhabiting new zealand (grichanov 1998, 2000, this paper; bickel 1999). the recent references dealing with this genus in the afrotropical region include grichanov (2011) and grichanov et al. (2011a, 2011b). in this paper two new species from south africa are described, and an identification key to afrotropical species of dactylonotus is presented. material and methods the holotypes of the new species are housed at the national museum, bloemfontein, south africa (bmsa). they were studied with a zeiss discovery v12 stereo microscope and photographed with an axiocam mrc5 camera. morphological terminology and abbreviations follow cumming & wood (2009). body length is measured from the base of the antenna to the tip of genital capsule. wing length is measured from the base to the wing apex. male genitalia were macerated in 10% koh. the figures showing the hypopygium in lateral view (figs 4, 10–12) are oriented as it appears on the intact specimen, http://dx.doi.org/10.5852/ejt.2016.175 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:cadca890-3440-49f6-afc8-ffbe9fb22d2d mailto:grichanov%40mail.ru?subject= http://zoobank.org/urn:lsid:zoobank.org:author:5320ad3a-92d8-4820-8091-24802f8c8c06 http://dx.doi.org/10.5852/ejt.2016.175 http://dx.doi.org/10.5852/ejt.2016.175 european journal of taxonomy 175: 1–9 (2016) 2 with the morphologically ventral surface of the genitalia facing up, dorsal surface down, anterior end facing right and posterior end facing left. results class hexapoda blainville, 1816 order diptera linnaeus, 1758 suborder brachycera schiner, 1862 superfamily empidoidea latreille, 1804 epifamily dolichopodoidae latreille, 1809 family dolichopodidae latreille, 1809 subfamily diaphorinae schiner, 1864 tribe argyrini negrobov, 1986 genus dactylonotus parent, 1934 dactylonotus parent, 1934: 136. type species dactylonotus grandicornis parent, 1934 (by monotypy). diagnosis the genus can be recognised by the finger-like projection or conus of the antennal pedicel, which overlaps the postpedicel dorsally in both sexes (figs 2, 7); the postpedicel with distinct apex, with relatively short dorsal arista-like stylus, either median or subapical in position; the occiput convex or flat; male frons and face broad; legs with an anterior preapical seta on the mid and hind femora; the wing costa extending beyond tip of r4+5, ending at apex of vein m; vein m unbroken (figs 3, 8); and male sternite 8 with strong projecting setae (figs 4, 10). remarks two species of the genus were formerly associated with the genera syntormon loew, 1857 (sympycninae), neurigona rondani, 1856 (neurigoninae) and tenuopus curran, 1924 (neurigoninae or genus incertae sedis), showing the uncertain position of dactylonotus within the subfamily diaphorinae. such characters previously unknown for the genus as the black body (d. nigricorpus sp. nov.) and ornamented fore tarsus (d. tsitsikamma sp. nov.) confirm the close relation of dactylonotus with argyra and somillus and their combination in an independent taxon of the subfamily rank. dactylonotus nigricorpus sp. nov. urn:lsid:zoobank.org:act:6a263872-6460-45cf-8cdf-387c0d610111 figs 1–4, 11 diagnosis the new species is close to d. rudebecki vanschuytbroeck, 1960 as described by grichanov (1998) under the name d. meuffelsi, differing from the latter and all other species of the genus in the black body, smaller size, shorter antenna, the hypopygium morphology, the arista-like stylus apical, the mid femur without long setae, the simple tarsi with short claws and the small pulvilli. etymology the species name is composed from latin “niger” and “corpus” (black body). . http://zoobank.org/urn:lsid:zoobank.org:act:6a263872-6460-45cf-8cdf-387c0d610111 grichanov i.ya., dactylonotus species from south africa 3 type material holotype republic of south africa: ♂, western cape, table mountain n.p., clovelly, sweeping grasses & mature fynbos, 34°07.465 s, 18°26.094 e, 24 oct. 2012, 64 m, a.h. kirk-spriggs (bmsa). description male (figs 1–4, 11) measurements. body length without antennae 3.8 mm, antenna length 1.4 mm, wing length 3.2 mm, wing width 1.2 mm. head. frons small, black, grey pollinose, with medial trapezoid depression, slightly prominent around base of antennae; face densely whitish pollinose, weakly narrowed in upper part, with parallel sides in lower part, 2 × as high as wide at clypeus; occiput flat, black, grey pollinose; one pair of long ocellar and short postvertical setae; no vertical setae; postocular setae black in upper part and white in middle and lower parts of head; eyes with short white hairs; antennae (fig. 2) inserted at upper fifth of head, black, 2 × longer than height of head; scape long, microscopically haired; pedicel with long, slightly widened at apex, dorso-lateral (inner view) process, covered by dorsal, lateral and ventral setulae; postpedicel figs 1–4. dactylonotus nigricorpus sp. nov., ♂. 1. habitus. 2. antenna, inner lateral view. 3. wing. 4. hypopygium after maceration, left lateral view. european journal of taxonomy 175: 1–9 (2016) 4 very long, widest at apex of pedicel, with acute apex, 3 × as long as high in middle, entirely covered by microscopic hairs; arista apical, with microscopic hairs. length ratio of scape to pedicel to postpedicel (dorsal to ventral sides) to arista-like stylus (1st and 2nd segments) in mm, 0.27/0.40/0.49/0.74/0.05/0.32. palpus and proboscis short, black, with black hairs. thorax. mostly black, with black bristles; mesonotum weakly pollinose; pleura densely grey pollinose; metanotum brownish below; propleuron with 2 strong black setae in lower part; 6 pairs of dorsocentral bristles decreasing in size anteriorly, with several scattered hairs on anterior slope; acrostichals biseriate, extending to 5th pair of dorsocentrals; scutellum with two long strong bristles and two short fine lateral setae, 1/4 as long as medians, dorsally bare. legs. including coxae yellow, with black bristles, tarsi brown-black from tip of basitarsus; fore coxa anteriorly with short black hairs and five or six black lateral and apical setae of various length in one row; mid coxa anteriorly with rather long black hairs and setae; mid and hind coxae with black bristle at base. femora without long setae; mid and hind femora with one strong subapical anterior seta; fore tibia with 3 strong anterodorsal, 2 short posterodorsal, 3 short ventral setae; mid tibia with 3 strong anterodorsal, 1 small dorsal, 1 strong and 2 small posterodorsal, 1 anteroventral, 1–2 small posteroventral setae; hind tibia with 3 anterodorsal, 3 posterodorsal, 4–5 short ventral setae; all tibiae with strong apical setae; all tarsi simple, with short claws and small pulvilli. femur, tibia and tarsomere (from first to fifth) length ratio: fore leg: 65/75/38/22/16/11/10, mid leg: 88/102/51/27/21/12/12, hind leg: 95/120/39/39/28/17/12. wing (fig. 3). ovate, fumous; veins brown; r4+5 and m1+2 slightly curved posteriad at apex; m1+2 nearly straight; crossvein dm-cu straight; ratio of dm-cu to apical part of cua1 (in mm), 0.28/0.66; anal vein foldlike; anal angle obtuse; lower calypter yellow, with black cilia; halter yellow. abdomen. mostly black, with black hairs and marginal setae; 6th segment reduced; 7th segment small, bare; 8th segment with short hairs and row of strong setae; hypopygium (figs 4, 11) black; epandrium subtriangular, strongly projected ventrally; hypandrium simple, broad, curved, with subapical spinelike process; phallus expanded at apex and biapicate; two epandrial setae positioned on distal side of epandrium, one of them (dorsal) pedunculate; epandrial lobe long, fingerlike, with rounded apex and short apical seta; surstylus yellow-brown, bilobate, with almost straight lobes, with several setae as figured; ventral lobe thinner than dorsal lobe, clavate; postgonite small, slightly curved ventrally; cercus short, yellow, with black setae. female unknown. distribution south africa (western cape). dactylonotus tsitsikamma sp. nov. urn:lsid:zoobank.org:act:9d497d9a-dfc2-4856-857a-86c75c373bb6 figs 5–10, 12 diagnosis the new species is close to d. univittatus (loew, 1858) as described by grichanov (2000), differing in the presence of the posterior brush of long setae on the fifth segment of the fore tarsus, the brownish yellow hypopygium and the morphology of the hypopygial appendages. the arista-like stylus is middorsal; the fore coxa are with black hairs; the mid femur is without long hairs and setae. http://zoobank.org/urn:lsid:zoobank.org:act:9d497d9a-dfc2-4856-857a-86c75c373bb6 grichanov i.ya., dactylonotus species from south africa 5 etymology the species is named after the tsitsikamma national park in south africa, where the type material was collected. type material holotype republic of south africa: ♂, [eastern cape,] tsitsikamma n.p., below sleepkloof, malaise trap, indigenous forest, 33°56.974 s, 23°54.926 e, 20–22 jan. 2009, 64 m, a.h. kirk-spriggs & s. otto (bmsa). description male (figs 5–10) measurements. body length without antennae 5.5 mm, antenna length 1.6 mm, wing length 5.0 mm, wing width 1.7 mm. head (somewhat shrunken). frons as wide as high, black, grey pollinose, with medial rhomboid depression; face densely whitish pollinose, narrow; occiput flat, black, grey pollinose; one pair of long figs 5–10. dactylonotus tsitsikamma sp. nov., ♂. 5. habitus. 6. antenna, outer lateral view. 7. antenna, inner lateral view. 8. wing. 9. apical segments 4–5 of fore tarsus. 10. hypopygium after maceration, left lateral view. european journal of taxonomy 175: 1–9 (2016) 6 ocellar and short postvertical setae; no vertical setae; postocular setae black in upper part and white in middle and lower parts of head; eyes with short white hairs; antennae (figs 6–7) inserted at upper fourth of head, dirty-yellow, brownish dorsally and apically, 1.5 × longer than height of head; scape long, bare; pedicel long, slightly widened at apex, dorso-lateral (inner view) process, covered by dorsal, lateral and ventral setulae; postpedicel very long, widest at apex of pedicel, with narrow rounded apex, 2.8 × as long as high in middle, entirely covered by microscopic hairs; arista-like stylus middorsal, with short hairs. length ratio of scape to pedicel to postpedicel (dorsal to ventral sides) to arista-like stylus (1st and 2nd segments) in mm, 0.33/0.38/0.40/0.73/0.43/0.60. palpus and proboscis short, yellow, with black hairs. thorax. mostly brownish yellow, with black bristles; mesonotum with broad median metallic blue violet stripe embracing area between 2nd–6th pairs of dorsocentrals; scutellum dorsally blue-violet with yellow margin; pleura with a black spot below calypter; thoracic pollination weak; proepisternum with 2 long brownish ventral setae and 4 yellow hairs above; 6 pairs of dorsocentral bristles decreasing in size anteriorly, with several scattered hairs on anterior slope; acrostichals biseriate, extending to 5th pair of dorsocentrals; scutellum with two long strong bristles and two short fine lateral setae, 1/4 as long as medians, dorsally bare. legs. including coxae yellow, with black bristles, apical segments of all tarsi brownish; fore coxa anteriorly with short black hairs and row of six black lateral and apical setae of various length; mid coxa anteriorly with rather long black hairs and setae; mid and hind coxae with black bristle at base; hind coxa with additional small seta below middle; femora without long setae; mid and hind femora each with one strong preapical anterior bristle; fore tibia with 3 strong anterodorsal, 2 short posterodorsal setae; mid tibia with 3 strong anterodorsal, 1 small dorsal, 3 strong posterodorsal setae; hind tibia with 3 anterodorsal, 5 posterodorsal, 4–5 short ventral setae; all tibiae with strong apical setae; 4th and 5th segments of fore tarsus slightly swollen; 5th segment (fig. 9) of same tarsus with posterior brush of flattened setae, as long as 5th segment, with slightly enlarged claws and pulvilli; mid and hind tarsi simple, with short claws and small pulvilli. femur, tibia and tarsomere (from first to fifth) length ratio (in mm): fore leg: 1.65/1.63/1.02/0.41/0.28/0.2/0.22, mid leg: 1.82/2.05/1.14/0.55/0.43/0.28/0.19, hind leg: 2.27/2.5/0.7/0.71/0.42/0.27/0.19. wing (fig. 8). ovate, almost hyaline; veins brown; r2+3 and r4+5 slightly curved posteriad at apex; m1+2 nearly straight; crossvein dm-cu straight; ratio of dm-cu to apical part of cua1 (in mm) 0.36/0.92; anal vein foldlike; anal angle obtuse; lower calypter yellow, with black cilia; halter yellow. figs 11–12. hypopygium, left lateral view. 11. dactylonotus nigricorpus sp. nov. 12. dactylonotus tsitsikamma sp. nov. abbreviations: cer = cercus; ep = epandrium; epl = epandrial lobe; hyp = hypandrium; ph = phallus; dsur, vsur = dorsal and ventral lobes of surstylus. grichanov i.ya., dactylonotus species from south africa 7 abdomen. mostly yellow, with black hairs and marginal setae; 1st segment brownish anteriorly; 2nd segment with brown t-shaped spot dorsally; 3rd–5th segments each with triangular dark-brown spot dorsally; 6th segment reduced, brown; 7th segment small, bare; 8th segment and epandrium mostly dirty yellow, partly brown, 8th segment with short hairs and row of strong setae; hypopygium (figs 10, 12) yellow-brown; epandrium ovate; hypandrium broad, with 2 large ventral projections; phallus pointed at apex; two epandrial setae positioned on distal side of epandrium, one of them (dorsal) pedunculate; epandrial lobe long, fingerlike, with rounded apex and 2 short apical setae; surstylus yellow-brown, bilobate, with almost straight lobes, with several apical setae as figured; ventral lobe thinner than dorsal lobe, clavate; postgonite thick, slightly curved ventrally; cercus short, yellow, with black setae. female unknown. distribution south africa (eastern cape). key to the afrotropical species of dactylonotus (males) 1 thorax and abdomen almost entirely black (fig. 1); postpedicel 3 times longer than high; aristalike stylus apical (fig. 2); body 3.8 mm ……………………………………d. nigricorpus sp. nov. – thorax and abdomen mostly yellow; postpedicel various in length; arista-like stylus subapical or middorsal ……………………………………………………………………………………………2 2 arista-like stylus middorsal, located near apex of pedicel process ………………………………3 – arista-like stylus subapical …………………………………………………………………………5 3 fore coxa with yellow-brown bristles; mid femur with two ventral rows of black setae in basal half, half as long as femur diameter; body 5 mm ……………………d. grandicornis parent, 1934 – fore coxa with black bristles; mid femur without long setae ……………………………………4 4 fifth segment of fore tarsus without posterior brush of long setae; hypopygium brown-black; body 4.6–5.3 mm ……………………………………………………………d. univittatus (loew, 1858) – fifth segment of fore tarsus with posterior brush of long setae (fig. 9); hypopygium (fig. 10) brownish yellow; body 5.5 mm. ……………………………………………d. tsitsikamma sp. nov. 5 fore tarsus with enlarged pulvilli; postpedicel 3 times longer than high; arista-like stylus as long as postpedicel; body 4 mm …………………………………………………d. frater parent, 1939 – tarsal pulvilli normal; postpedicel 4.5 times longer than high; arista-like stylus half as long as postpedicel; body 4.7 mm ………………………………………d. rudebecki vanschuytbroeck, 1960 discussion unfortunately, for each of the newly described species only one male was found despite intensive collecting with malaise trap and sweeping. because the secondary sexual characters of these species (including the structure of their genitalia) are unique, their identification will not be problematic in the future. dactylonotus nigricorpus sp. nov. is the only species with a black body, whereas the other species have mainly yellow bodies. d. tsitsikamma sp. nov. is remarkable in the genus in bearing a flag of long setae on the fifth segment of the fore tarsus. i think their descriptions are important due to our incomplete knowledge of the southern african dolichopodids and our poor understanding of the diaphorine genera. the two new species and d. frater seem to be extremely rare in nature, all known by a single male only. summarising my work with the genus over the last 20 years, i could find 319 specimens of d. grandicornis, 37 specimens of d. univittatus and 9 specimens of d. rudebecki in the collections of european journal of taxonomy 175: 1–9 (2016) 8 dozens european and african museums. thus, classical patterns of relative species abundance (e.g., mcgill et al. 2007) are here confirmed. expanded knowledge of the genus will probably be essential to improve our understanding of the phylogenetic relations among diaphorine flies of the world. acknowledgements the author is sincerely grateful to dr. ashley h. kirk-spriggs (bloemfontein, south africa) for his kindness in providing specimens for study. the editorial team of the ejt and anonymous reviewers kindly commented on earlier drafts of the manuscript. the work was partly supported by the russian foundation for basic research grant n 14-04-00264-a. references bickel d. 1999. australian sympycninae ii: syntormon loew and nothorhaphium, gen. nov., with a treatment of the western pacific fauna, and notes on the subfamily rhaphiinae and dactylonotus parent (diptera: dolichopodidae). invertebrate taxonomy 13: 179–206. capellari r.s. 2013. phylogeny of the diaphorinae (diptera: dolichopodidae). in: klass k.-d. (main organizer) 6th dresden meeting on insect phylogeny, september 27-29, 2013. abstracts – poster presentations: 56. cumming j.m. & wood d.m. 2009. chapter 2. adult morphology and terminology. in: brown b.v., borkent a., cumming j.m., wood d.m., woodley n.e. & zumbado m.a. (eds) manual of central american diptera, volume 1: 9–50. nrc research press, ottawa. grichanov i.ya. 1998. a new species of dactylonotus parent (diptera: dolichopodidae) from south africa. international journal of dipterological research 9 (1): 27–29. grichanov i.ya. 2000. afrotropical neurigoninae and notes on the diaphorine genus dactylonotus parent (diptera: dolichopodidae). belgian journal of entomology 2: 257–271. grichanov i.ya 2011. an illustrated synopsis and keys to afrotropical genera of the epifamily dolichopodoidae (diptera: empidoidea). priamus serial publication of the centre for entomological studies ankara supplement 24: 1–99. grichanov i.ya., mostovski m.b. & muller b. 2011a. new records of afrotropical dolichopodidae (diptera) from the collection of natal museum (1). international journal of dipterological research 22 (1): 3–9. grichanov i.ya., mostovski m.b. & muller b. 2011b. new records of afrotropical dolichopodidae (diptera) from the collection of natal museum (2). international journal of dipterological research 22 (2): 81–98. mcgill b.j., etienne r.s., gray j.s., alonso d., anderson m.j., benecha h.k., dornelas m., enquist b.j., green j.l., he f., hurlbert a.h., magurran a.e., marquet p.a., maurer b.a., ostling a., soykan c.u., ugland k.i. & white e.p. 2007. species abundance distributions: moving beyond single prediction theories to integration within an ecological framework. ecology letters 10 (10): 995–1015. http://dx.doi. org/10.1111/j.1461-0248.2007.01094.x parent o. 1934. additions à la faune éthiopienne (diptères: dolichopodidés). bulletin de la société entomologique d’égypte 18: 112–138. manuscript received: 9 october 2015 manuscript accepted: 12 january 2016 http://dx.doi.org/10.1111/j.1461-0248.2007.01094.x http://dx.doi.org/10.1111/j.1461-0248.2007.01094.x grichanov i.ya., dactylonotus species from south africa 9 published on: 16 february 2016 topic editor: koen martens desk editor: kristiaan hoedemakers printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark. european journal of taxonomy 27: 1-9 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2012.27 www.europeanjournaloftaxonomy.eu 2012 · teresa darbyshire this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:09359e10-277b-4018-bc69-7944d8de6fd4 1 re-description of dysponetus joeli olivier et al., 2012 (polychaeta, chrysopetalidae), with a new key to species of the genus teresa darbyshire department of biodiversity & systematic biology, amgueddfa cymru – national museum wales, cathays park, cardiff cf10 3np, wales, u.k. email: teresa.darbyshire@museumwales.ac.uk urn:lsid:zoobank.org:author:b7103b9c-2a1a-4adc-bc28-55ea9b8850c9 abstract. dysponetus is a genus of the family chrysopetalidae with twelve currently described species. specimens are fragile and easily damaged or broken during sampling making identification difficult. the most recently described species, dysponetus joeli olivier et al., 2012, from the bay of biscay and the english channel, was described from a few small, damaged and poorly preserved specimens. new specimens from the isles of scilly, in much better condition, resembled d. joeli except for the absence of ventral cirri on segment 3. examination of the type material of d. joeli showed it to be identical to these new specimens and highlighted errors in the original description of the species. the present paper corrects the errors and a revised key to species is produced. the differences between d. joeli and the two most similar species, d. bipapillatus dahlgren, 1996 and d. macroculatus dahlgren, 1996 are also detailed. keywords. dysponetus, tentacular cirri, ventral cirri, cirrophores. darbyshire t. 2012. re-description of dysponetus joeli olivier et al., 2012 (polychaeta, chrysopetalidae), with a new key to species of the genus. european journal of taxonomy 27: 1-9. http://dx.doi.org/10.5852/ejt.2012.27 introduction there are currently twelve described species of the genus dysponetus levinsen, 1879 from around the world. most occur in shallow water and, with a few exceptions, are distributed in the atlantic region. all are small and fragile and rarely recorded from surveys, most likely because they are easily overlooked or found in fragments not identifiable to species level. several species are known only from their original description and, even where multiple records do exist, these number in single figures. five species have been described for the northeast atlantic region: d. caecus (langerhans, 1880), d. gracilis hartman, 1965, d. paleophorus hartmann-schröder, 1974, d. pygmaeus levinsen, 1879 and d. joeli olivier et al., 2012. all of these have potential to be found in uk waters although only two currently are. dysponetus gracilis is, as yet, only recorded from deep water (>400 m) samples outside of the uk (hartman 1965; hartman & fauchald 1971; aguirrezabalaga et al. 1999), d. paleophorus has not been recorded since its first description off norway and records for d. pygmaeus are restricted to the arctic and japanese pacific (levinsen 1879; annenkova 1935; imajima & hartman 1964). dysponetus http://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:09359e10-277b-4018-bc69-7944d8de6fd4 mailto:teresa.darbyshire%40museumwales.ac.uk?subject= http://zoobank.org/urn:lsid:zoobank.org:author:b7103b9c-2a1a-4adc-bc28-55ea9b8850c9 http://dx.doi.org/10.5852/ejt.2012.27 european journal of taxonomy 27: 1-9 (2012) 2 caecus, the most recorded dysponetus species, has been identified from shallow littoral samples from the uk (scotland: dahlgren & pleijel 1995) as well as from other european locations (laubier 1964; o’ connor 1981; kirkegaard 1992; parapar & san martín 1993; dahlgren & pleijel 1995; brito et al. 1996). an anomalous addition to these records would appear to be the records of böggemann (2009) from the abyssal southeast atlantic. however, böggemann addresses this anomaly in his paper by suggesting that the lack of previous records from this region and depth is related to its small size and fragility (probably the scarcity of previous samples too). most recently, d. joeli was described from four shallow sites in the uk and france. in 2009, a national museum wales subtidal survey of the isles of scilly, an island archipelago off the southwest tip of the uk, collected five specimens of dysponetus in coarse sandy sediments. initial examination of the specimens showed them to possess four eyes, a character possessed by half of the dysponetus species, none of which, at that time, were described for the uk. other characters, including the absence of ventral cirri on segment 3, were found that differentiated it from all but two other dysponetus species, d. bipapillatus dahlgren, 1996 from the mediterranean and d. macroculatus dahlgren, 1996 from papua new guinea. the description of d. joeli was published shortly before the description was finished and the two species showed such great similarity that the type specimens of the latter were obtained and examined in order to determine whether the differences were indeed real or might be due to the damaged nature of the specimens used. examination of the holotype of d. joeli revealed that the ventral cirri of segment 3 were in fact entirely absent rather than merely detached and lost, and thus the specimens were indeed the same species. consequently, this required distinguishing characters between d. joeli, d. bipapillatus and d. macroculatus to also be detailed. a full re-description is therefore made to clarify the species characters and differentiate it from the two most similar species. materials and methods specimens of dysponetus joeli were collected during a subtidal survey of the isles of scilly in june 2009. samples were collected using a 0.1m2 van veen grab from the r.v. sepia of the marine biological association uk. sediment was elutriated gently with seawater and the resulting sample sorted live the same day. animals were relaxed in a 7% magnesium chloride solution and fixed in 4% formaldehyde. final preservation was in 80% alcohol with 2% propylene glycol added. the type specimens of d. joeli were collected from shallow maerl beds by day, van veen or smithmcintyre grab and sieved through a 0.5 or 1.0 mm mesh (olivier et al. 2012). all drawings and measurements were made using a camera lucida attachment on a nikon labophot-2 compound microscope or a nikon eclipse e400 binocular microscope. microscope photographs were taken using automontage™ software and sem images were obtained using a neoscope sem. type specimens of d. joeli are held in the collections of the muséum national d’histoire naturelle in paris (mnhn) and the new specimens from the isles of scilly are in the national museum wales collections (nmw.z). specimens of d. bipapillatus and d. macroculatus were borrowed from the swedish museum of natural history (smnh) to enable a full comparison of characters. results phylum annelida lamarck, 1809 class polychaeta grube, 1850 family chrysopetalidae ehlers, 1864 genus dysponetus levinsen, 1879 darbyshire t., redescription of dysponetus joeli olivier et al., 2012 3 dysponetus joeli olivier et al., 2012 figs 1a–d, 2a–d dysponetus joeli olivier et al., 2012: 989–996, figs 1–3. material examined chausey, france, sta. ssmm01 (48° 55.570’ n, 001° 48.270’ w), maerl, 10.0 m, holotype (mnhn poly type 1533), paratype (mnhn poly type 1534), 18 apr. 2006; off st mary’s, isles of scilly, sta. 3b (49° 54.20’ n, 006° 18.94’ w), coarse sand/shell/gravel with some silt/clay, 25 m, 2 specimens (nmw.z.2009.027.0001–0002), 25 jun. 2009; west of st martin’s, isles of scilly, sta. 14b (49° 57.86’ n, 006° 15.21’ w), shell gravel in muddy sand, 35.1 m, 2 specimens (nmw.z.2009.027.0003–0004), 27 jun. 2009; west of isles of scilly, sta. 24b (49° 55.22’ n, 006° 23.91’ w), silty coarse sand/shell gravel, 47.1 m, 1 specimen used for sem (nmw.z.2009.027.0005), 29 jun. 2009. fig. 1. dysponetus joeli olivier et al., 2012. a. nmw.z.2009.027.0003, anterior end, dorsal view. b. nmw.z.2009.027.0004, anterior end, ventral view. c. nmw.z.2009.027.0003, posterior end, ventral view. d. holotype (mnhn poly type 1533), anterior end, ventral view. only a few chaetae drawn in each case for clarity. – ac = accessory chaeta, d = dorsal (cirrus/cirrophore), la = lateral antenna, ma = median antenna, m = mouth appendage, p = palp, ps = palp scar, v = ventral (cirrus/cirrophore); numbers indicate the segment. european journal of taxonomy 27: 1-9 (2012) 4 description holotype in 4 pieces, posteriorly incomplete, 3.1 mm long for 19 chaetigers. paratype in 3 pieces, anterior dissected fragment of 5 chaetigers with 2 further detached chaetigers. 3 non-type specimens up to 6 mm long, 0.35 mm wide (between segments, not including parapodia or chaetae) with 25–33 chaetigers. two other incomplete non-type specimens examined. the following description is based on entire specimens from the isles of scilly, except where specified as pertaining to the type material. body shape cylindrical, ventrally flattened, tapered slightly at posterior. body pale cream in colour (fixed, unstained), eyes dark reddish brown (fig. 2a) but pigment may degrade in alcohol. eyes of type specimens already barely discernable. prostomium oblong, wider anteriorly (fig. 1a). four large, rounded eyes visible in fixed specimens, anterior pair larger and more widely separated than posterior. median antenna small, bottle-shaped, arising anterodorsally from prostomium. lateral antennae bottle-shaped, slightly smaller than median, arising immediately dorsal to palps. antennae without distinct ceratophores. palps directed posteriorly, stout, oval, more than twice as long as wide. no antennae or palps remaining on type specimens examined. nuchal organs not observed. distinct single mouth appendage on lower lip, anteriorly directed, digitiform with blunt tip (fig. 1b, 2b). single pair of stylet-shaped jaws, visible through body wall with methyl green staining. proboscis not observed. first two segments slightly elevated dorsally with four pairs tentacular cirri, longer than but with same shape as dorsal cirri of third and following segments, anteriorly directed. first segment achaetous, second segment with notochaetae only, situated slightly anterior to dorsal tentacular cirrus. third segment biramous; dorsal cirri present, ventral cirri absent (figs 1a, b, d, 2b). holotype lacking all cirri on anterior three segments, but presence indicated by cirrophores (fig. 1d). following segments all biramous with both dorsal and ventral cirri. single notoand neuroacicula present in each parapodium. notopodial lobes reduced. dorsal cirri long, slender, longer than chaetae (210–430 µm, longest on median chaetigers), cirrophores present. styles slightly proximally swollen, distally tapering, tips blunt. notoacicula difficult to detect. notochaetae inserted dorsal to cirrus, densely packed, directed posteriorly leaving middle part of dorsum exposed. chaetae d-shaped in cross-section (fig. 2c) with denticles sharply pointed, in two parallel rows, 15–20 on each side. notochaetal count, mid-body segments, up to 26. neuropodia well-developed, conical mounds. compound neurochaetae, with heterogomph shafts and fine bidentate falcigerous blades (fig. 2d). neurochaetal count, mid-body segments, at least 20–26 (chaetae densely packed and difficult to accurately count). up to two accessory simple chaetae, similar to but smaller than notochaetae, inserted distally and anteriorly on neuropodial lobe (fig. 1c). ventral cirri fusiform, shorter than dorsal cirri (length 110–270 µm, longest on median chaetigers), arising posteroventrally on neuropodial lobe (fig. 1c). final segment lacking notoand neurochaetae, cirrophores of dorsal and ventral cirri observed although only a single, rounded dorsal cirrus observed on 1 specimen (nmw.z.2009.027.0003). pygidium conical with single projection (length 40 µm), cylindrical, slightly distally tapering, inserted posteroventrally (fig. 1c), anus terminal. eggs visible within one specimen (nmw.z.2009.027.0001), flattened oval in shape, maximum width approx. 50 µm, possibly immature. darbyshire t., redescription of dysponetus joeli olivier et al., 2012 5 habitat the species was originally described from maerl beds in the bay of biscay and the english channel. the new specimens are all from coarse sand and shell sediments collected around the isles of scilly. remarks the original description of d. joeli was based on small, damaged specimens and this may explain why the absence of the ventral cirri on chaetiger 3 was attributed to loss of the cirri rather than actual absence. although cirri are easily lost from specimens, the cirrophores can still be seen under light microscopy at x400 magnification or greater. application of methyl green staining can help distinguish these features. under sem conditions the absence of cirrophores on chaetiger 3 is obvious (fig. 2b). an additional complication in determining the described characters of d. joeli arises from the character matrix scores in the original paper (olivier et al. 2012). in their species description, the authors stated that there were 2 pairs of tentacular cirri on the first 2 segments but made no mention whether segment 3 possessed or lacked ventral cirri. they then later scored ventral cirri as absent for the first 2 segments (no separate score for tentacular cirri) and present for the third in the character matrix however, in the fig. 2. dysponetus joeli olivier et al., 2012. a. nmw.z.2009.027.0003, whole specimen, dorsal view (scale bar 1 mm). b. nmw.z.2009.027.0005, anterior end, ventrolateral view (scale bar 50 µm). c. notochaetae (scale bar 10 µm). d. neurochaetae (scale bar 20 µm). – d = dorsal (cirrophore), la = lateral antenna, m = mouth appendage, p = palp, ps = palp scar, v = ventral (cirrophore); numbers indicate the segment. european journal of taxonomy 27: 1-9 (2012) 6 species key itself, d. joeli is keyed out with ventral cirri being present on the 3rd segment (i.e. chaetiger 2). it is apparent from the character states scored for the other species in the matrix that the authors have used the term ‘ventral cirrus’ interchangeably with ‘tentacular cirrus’. earlier authorities have also been variable in their use of the terms ‘tentacular cirrus’ versus ‘cirrus’ with regards to the appendages of the first 3 segments in species of dysponetus. in this paper, only the cirri of the first 2 segments, which lack parapodia and have cirrophores directly attached to the body wall, are regarded as being tentacular. a later paper will discuss the different characters of dysponetus species in more detail. the lack of ventral cirri on segment 3 distinguishes d. joeli from most other dysponetus species. the only other species with this characteristic are d. bidentatus day, 1954, d. bipapillatus and d. macroculatus, although d. bidentatus lacks ventral cirri on segment 2 also (day 1954). the presence or absence of ventral cirri on segment 3 is unknown for d. hebes (webster & benedict, 1887); however, this species differs considerably by having a double mouth appendage as opposed to single, and sphaerical not elongated palps. with the revised character for segment 3, the species key in olivier et al. would place d. joeli with both d. bipapillatus and d. macroculatus. the latter two species are distinguished in the key according to the relative size of the eyes; however, this is not a good character as the eye pigments degrade in alcohol and disappear over time – as is already evident in the type specimens. a new revised key to the 12 species of the genus is provided below. dysponetus joeli is a much larger species than both d. bipapillatus and d. macroculatus. even the type specimens, described as ‘small’ (ranging from 3.5–4 mm in length, incomplete) and smaller than those from the isles of scilly, were 2–3 times the size of each of the latter species, respectively. dysponetus joeli is most similar to d. bipapillatus from the mediterranean, sharing all of the obvious characteristics. the paired papillae on segment 8 described for d. bipapillatus were not observed on any of the d. joeli specimens; however, these appendages were only seen on a few specimens of d. bipapillatus and are thus not a good character for comparison. in his paper, dahlgren (1996) stated that the specimens, though small, were considered mature adults due to the presence of the paired papillae, interpreted as genital organs, and the development of the anterior segments. a comparison of the individual characters shows d. joeli to possess twice as many neurochaetae as d. bipapillatus across all segments (20–26 compared to 9–13), more numerous denticles on the notochaetae (15–20 as opposed to 10–15) and a shorter pygidial projection (40 µm versus 50 µm), the last character particularly noticeable in relation to the larger body size of d. joeli. similarly, the number of neurochaetae in d. joeli is greater than the 19–22 found on d. macroculatus, and, in general, the former species is a much larger animal than the latter, being 2–3 times as long at maturity (as evidenced by the presence of eggs in specimens of both). key to dysponetus species 1 ventral tentacular cirri absent on segment 1 ………………………………………………………2 – ventral tentacular cirri present on segment 1 ………………………………………………………6 2 elongate palps; 4 eyes ……………………………………d. bulbosus* hartmann-schröder, 1982 – sphaerical palps; 2 eyes or eyes absent ……………………………………………………………3 3 anterior median antenna; double mouth appendage on lower lip; 2 eyes ………………………… ………………………………………………………………d. hebes (webster & benedict, 1887) – dorsal median antenna; eyes absent ………………………………………………………………4 4 ventral tentacular cirri present on segment 2 …………………………d. gracilis hartman, 1965 – ventral tentacular cirri absent on segment 2 ………………………………………………………5 darbyshire t., redescription of dysponetus joeli olivier et al., 2012 7 5 mouth appendage absent; paleae present among notochaetae …………………………………… …………………………………………………………d. paleophorus hartmann-schröder, 1974 – double mouth appendage on lower lip; paleae absent ……………d. pygmaeus levinsen, 1879 6 mouth appendage absent …………………………………………………………………………7 – single mouth appendage on lower lip ……………………………………………………………8 7 elongate palps, ventral tentacular cirri present on segment 2 ……d. hesionides böggemann, 2009 – sphaerical palps, ventral tentacular cirri absent on segment 2 ……d. profundus böggemann, 2009 8 eyes absent, ventral cirri present on segment 3 ……………………d. caecus (langerhans, 1880) – eyes present, ventral cirri absent on segment 3 …………………………………………………9 9 16 or less notochaetae in mid-body segments …………d. bulbosus* hartmann-schröder, 1982 – 20 or more notochaetae in mid-body segments …………………………………………………10 10 ventral tentacular cirri absent on segment 2 ……………………………d. bidentatus day, 1954 – ventral tentacular cirri present on segment 2 ……………………………………………………11 11 few neurochaetae, only 9–13 in mid-body segments ……………d. bipapillatus dahlgren, 1996 – numerous neurochaetae in mid-body segments, 19 or more …………………………………12 12 small body size, 2–3 mm; 19–22 neurochaetae …………………d. macroculatus dahlgren, 1996 – large body size, >3 mm; 20–26 neurochaetae ……………………d. joeli olivier et al., 2012 * descriptions of d. bulbosus (hartmann-schröder 1982, 1986) contradict each other over the presence or absence of ventral cirri on each of the first few segments. for this reason, d. bulbosus is included twice in the key to account for the discrepancy in descriptions. dysponetus bulbosus will be reviewed in a forthcoming paper. acknowledgements the work in the isles of scilly was part of the assessment of marine biodiversity linked to ecosystems project funded by the department for environment, food & rural affairs (defra). thanks must go to dr richard warwick and dr paul somerfield who jointly led the project, the captain and crew of the marine biological association’s research vessel sepia and everyone else involved with the sampling. elin sigvaldodottír and tarik meziane are also thanked for the loan of specimens from the swedish museum of natural history, stockholm and muséum national d’histoire naturelle in paris that enabled this paper to be written. dr andrew mackie is thanked for his comments and improvements on the paper. references aguirrezabalaga f., san martín g., petersen m.e. & ceberio a. 1999. presencia de dysponetus gracilis hartman, 1965 (polychaeta, chrysopetalidae) en las costas europeas, golfo de vizcaya. boletín de la real sociedad española de historia natural (sección biológica) 95: 23–27. annenkova n.p. 1935. über dysponetus pygmaeus levinsen und euzonus arcticus grube. comptes rendus (doklady) de l’académie des sciences de l’ussr 1935 3(5): 233–236. böggemann m. 2009. polychaetes (annelida) of the abyssal se atlantic. organisms diversity & evolution 9: 252–428. http://www.sciencedirect.com/science/journal/14396092 brito m.c., núñez j., bacallado j.j. & ocaña, o. 1996. anélidos poliquetos de canarias: orden phyllodocida (chrysopetalidae, pisionidae, glyceridae, sphaerodoridae, hesionidae y pilargidae). in: http://www.sciencedirect.com/science/journal/14396092 european journal of taxonomy 27: 1-9 (2012) 8 llinas o., gonzález j.a. & rueda m.j. 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(polychaeta: chrysopetalidae) from the north-east atlantic, with a cladistic analysis of the genus and a key to species. journal of the marine biological association of the uk 92: 989–996. http://dx.doi.org/10.1017/ s0025315412000562 http://www.biodiversitylibrary.org/item/107744#page/599/mode/1up http://www.biodiversitylibrary.org/item/107744#page/599/mode/1up http://dx.doi.org/10.5962/bhl.title.2081 http://dx.doi.org/10.5962/bhl.title.2081 http://www.biodiversitylibrary.org/item/110689#page/1/mode/1up http://www.biodiversitylibrary.org/item/110689#page/1/mode/1up http://dx.doi.org/10.1017/s0025315412000562 http://dx.doi.org/10.1017/s0025315412000562 darbyshire t., redescription of dysponetus joeli olivier et al., 2012 9 parapar j. & san martín g. 1993. anélidos poliquetos mesopsámmicos sublitorales de la costa de ceuta (españa). cahiers de biologie marine 34: 363–381. webster h.e. & benedict j.e. 1887. the annelida chaetopoda, from eastport, maine. annual report of the united states commission of fish and fisheries, washington, vol. for 1885: 707–758, pls 1–8. http://docs.lib.noaa.gov/rescue/cof/cof_1885.pdf manuscript received: 17 august 2012 manuscript accepted: 19 october 2012 published on: 8 november 2012 topic editor: rudy jocqué in compliance with the iczn, printed versions of all papers are deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; national botanic garden of belgium, meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark. http://docs.lib.noaa.gov/rescue/cof/cof_1885.pdf european journal of taxonomy 693: 1–12 issn 2118-9773 https://doi.org/10.5852/ejt.2020.693 www.europeanjournaloftaxonomy.eu 2020 · wang y.-r. et al. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:0acf3bb2-8df5-43ae-adf9-75345d94d5b8 1 bathya brevicarpus gen. et sp. nov. (amphipoda: senticaudata: calliopiidae), from hydrothermal vents, okinawa trough, north-west pacific yan-rong wang 1, chao-dong zhu 2, zhong-li sha 3,* & xian-qiu ren 4 1,2 key laboratory of zoological systematics and evolution, institute of zoology, chinese academy of sciences, beijing 100101, china. 1,3,4 institute of oceanology, chinese academy of sciences, qingdao 266071, china. 3 laboratory for marine biology and biotechnology, qingdao national laboratory for marine science and technology, qingdao, china. 3 center for ocean mega-science, chinese academy of sciences, qingdao 266071, china. 1,2,3 college of biological sciences, university of chinese academy of sciences, beijing 100049, china. * corresponding author: shazl@qdio.ac.cn 1 email: wangyr@qdio.ac.cn 2 email: zhucd@ioz.ac.cn 4 email: 1278593370@qq.com 1 urn:lsid:zoobank.org:author:30ee7948-0a00-431b-8a67-b52f531fec66 2 urn:lsid:zoobank.org:author:0ddf9891-c900-47a5-9e44-978388c41181 3 urn:lsid:zoobank.org:author:ecb423ca-544c-4765-9f40-484db8ba1001 4 urn:lsid:zoobank.org:author:6aa3884a-5be7-4c2d-a597-9c270bc37a7d abstract. one individual referable to calliopiidae g.o. sars, 1893 was collected from a chemically reduced habitat, the hydrothermal vent systems in okinawa trough, and was identified as a new genus and species belonging to this family after a morphological examination. a formal description of this new species and a discussion of the relationship of the new genus within calliopiidae are presented. keywords. taxonomy, morphology, calliopiidae, new genus, deep sea. wang y.-r., zhu c.-d., sha, z.-l. & ren x.-q. 2020. bathya brevicarpus gen. et sp. nov. (amphipoda: senticaudata: calliopiidae), from hydrothermal vents, okinawa trough, north-west pacific. european journal of taxonomy 693: 1–12. https://doi.org/10.5852/ejt.2020.693 introduction the calliopiids have been placed in the eusiroidea stebbing, 1888 (e.g., bousfield & hendrycks 1995, 1997; de broyer et al. 2007). a phylogenetic study based on morphology by lowry & myers (2013) showed that the calliopiidae g.o. sars, 1893 sit in the hadziidira s. karaman, 1943 and have a close relationship with pontogeneiidae stebbing, 1906. this family currently contains 27 genera and more https://doi.org/10.5852/ejt.2020.693 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:0acf3bb2-8df5-43ae-adf9-75345d94d5b8 mailto:shazl@qdio.ac.cn mailto:wangyr@qdio.ac.cn mailto:zhucd@ioz.ac.cn mailto:1278593370@qq.com http://zoobank.org/urn:lsid:zoobank.org:author:30ee7948-0a00-431b-8a67-b52f531fec66 http://zoobank.org/urn:lsid:zoobank.org:author:0ddf9891-c900-47a5-9e44-978388c41181 http://zoobank.org/urn:lsid:zoobank.org:author:ecb423ca-544c-4765-9f40-484db8ba1001 http://zoobank.org/urn:lsid:zoobank.org:author:6aa3884a-5be7-4c2d-a597-9c270bc37a7d https://doi.org/10.5852/ejt.2020.693 european journal of taxonomy 693: 1–12 (2020) 2 than 100 species, most of which are described from arctic and subarctic regions (e.g., barnard & karaman 1991; ren & huang 1991; weisshappel 2001; d’udekem d’acoz 2007, 2010, 2012). the calliopiids reported from vent fields are represented by two species: bouvierella curtirama bellansantini & thurston, 1996 collected at a depth of 1636 m from the mid-atlantic ridge and leptamphopus fragilis larsen & krapp-schickel, 2007 collected at a depth of 2656 m near wuzza bare mount, northeast pacific (bellan-santini & thurston 1996; larsen & krapp-schickel 2007). the chinese research vessel “kexue”, using the remote operated vehicle (rov) “faxian” to survey the biodiversity of hydrothermal vents in okinawa trough, collected one unusual specimen referable to calliopiidae. a morphological examination shows that this individual is a new calliopiid. however, the new species does not belong to any of the described genera within this family based on the combination of the following morphological characters: a small rostrum; the absence of the antennae calceoli; the article 3 of antenna 1 not produced apicoventrally; the inner plate of maxilla 1 bearing only one apical seta; the gnathopod 2 not linear or greatly elongate and the carpus of the gnathopods 1–2 much shorter than the propodus; pereopod 5 slightly longer than pereopod 7; epimeron 3 smooth; the outer ramus of uropod 3 half-length of the inner ramus; the entire telson, longer than broad. hence, a new genus, bathya gen. nov., is erected to accomodate this new species herein. a key to all genera of calliopiidae is also presented. material and methods the present material was collected by rov “faxian”, during expeditions to the okinawa trough hydrothermal vents by the institute of oceanology, chinese academy of sciences (iocas) in april 2014. the type specimen examined is deposited in the marine biological museum, chinese academy of sciences (mbmcas), qingdao, china, and preserved in 75% ethanol. it was examined and dissected under a dissecting microscope (zeiss discovery v20). line drawings were made with a tablet (wacom intuos pro pth-851) and adobe photoshop (ver. cs6). the length measurements are made along the outline of the animals, beginning from the anterior margin of head to the end of the telson. abbreviations (used in figs 1–2) a = antenna c = coxa e = epimeron g = gnathopod h = head l = left md = mandible mx1 = maxilla 1 mx2 = maxilla 2 mxp = maxilliped p = pereopod r = right t = telson u = uropod wang y.-r. et al., a new genus and species of calliopiidae (amphipoda) from okinawa trough 3 results order amphipoda latreille, 1816 suborder senticaudata lowry & myers, 2013 superfamily calliopioidea g.o. sars, 1893 family calliopiidae g.o. sars, 1893 genus bathya gen. nov. urn:lsid:zoobank.org:act:aa9c504c-ce7e-426a-a988-27098bd64d25 type species bathya brevicarpus gen. et sp. nov. diagnosis antennae calceoli absent. antenna 1 longer than antenna 2; accessory flagellum absent. mandible molar triturative, columnar; palp 3-articulate, extremely elongated, article 3 longer than article 2. maxilla 1 inner plate bearing one apical seta; palp article 1 shorter than article 2. maxilla 2 inner plate without oblique setal row. maxilliped outer plate not reaching end margin of palp article 2. gnathopod 1 similar in shape to gnathopod 2, slightly smaller than gnathopod 2; subchelate; carpus much shorter than propodus; with posterior lobe. coxae 5–6 bilobate, posterior lobe larger than anterior lobe; coxa 7 rounded. pereopod 5 slightly longer than pereopod 7; anterior margin of propodus with bifid robust setae; dactylus simple, with small nail. epimerons 2–3 posterior margin smooth, posteroventral corner of epimeron 2 subacute. uropods 1–2 inner ramus longer than outer ramus, both inner and outer ramus with marginal robust setae. uropod 3 inner ramus about twice longer than outer ramus, both inner and outer ramus with marginal robust setae, without simple or plumose setae. telson entire, linguiform, longer than broad. etymology the generic name bathya refers to the type species designated herein that was collected from bathyal waters. distribution north-west pacific, okinawa trough, the hydrothermal vents at a depth of 996.9 m. remarks actually, it is a little questionable that bathya gen. nov. is placed under the calliopiidae g.o. sars, 1893 for having the pereopod 5 longer than pereopod 7. the phylogenetic study by lowry & myers (2013) based on morphology analyzed showed that the calliopiidae and pontogeneiidae stebbing, 1906 cluster together and form a clade with hornelliidae d’udekem d’acoz, 2010 and cheirocratidae d’udekem d’acoz, 2010. the new genus is not listed under cheirocratidae or hornelliidae, which are also not having antennae calceoli, for having the following characteristics: the inner plate of maxilla 1 only bearing apical seta, maxilla 2 without oblique setal row, the rami of uropod 3 unequal in length, and the entire telson. the new genus is not suited to be placed in pontogeneiidae for the propodus of gnathopod 2 having robust setae along the palmar margin. moreover, sanchoidae lowry, 2006, which are known as endemic to australia and associated with sponges (lowry & barnard 2001), have been listed under the calliopiidae. however, the present genus cannot be incorporated in this family for the lack of a nonrecessed head and a dorsoventrally flattened urosome (lowry 2006). hence, it is more reasonable that this new genus is to be placed under calliopiidae with an emendation of the diagnosis of this family. http://zoobank.org/urn:lsid:zoobank.org:act:aa9c504c-ce7e-426a-a988-27098bd64d25 european journal of taxonomy 693: 1–12 (2020) 4 besides having the pereopod 5 longer than pereopod 7, bathya gen. nov. can be distinguished its congeners of the calliopiidae by having the combination of the following characters: eyes present but not well pigmented; antenna 1 longer than antenna 2; the absence of an accessory flagellum; the inner plate of maxilla 1 only bearing one apical seta; maxilla 2 without oblique setal row; gnathopod 1 similar in shape and size to gnathopod 2; pleonites 1–3 without dorsal carinae; the rami of uropod 3 unequal in length; the entire telson. bathya brevicarpus gen. et sp. nov. urn:lsid:zoobank.org:act:2a8520b1-c0c6-4bab-9c8d-a7dfb57b111d figs 1–2 diagnosis as for the genus. etymology the new species name contains the latin ‘brev’ (= short), referring to the carpus of the gnathopods 1 and 2 being shorter than the propodus. material examined holotype north-west pacific • ♂, 5.6 mm, dissected; okinawa trough; depth 996.9 m; 17 apr. 2014; ry0108, rov-4; mbm 286556. description body. dorsally smooth. head. nearly subequal in length to pereonites 1 and 2 combined, rostrum small, anterior lobe rounded, lower margin with large and acute projecting tooth anterodistally; eyes present, but not pigmented, hardly visible in ethanol material. antenna. antenna 1 longer than antenna 2; peduncular article 1 wider and longer than article 2; article 2 less than twice longer than article 3; article 3 not produced apicoventrally; primary flagellum 23-articulate, few very thin and short setae scattered along flagellum; accessory flagellum absent. antenna 2 slender than antenna 1, peduncular article 5 distinctly longer than article 4; flagellum longer than peduncle, 28-articulate. mouth parts. upper lip and lower lip broken. mandible symmetrical, with incisor dentate, bearing 7 teeth; lacinia mobilis dentate, with 8 teeth; with 12 accessory spines; molar well developed, columnar, triturative; palp elongate, slender, 3-articulate, article 3 longer than article 2, ventral face bearing short stout setae. maxilla 1 with inner plate bearing one apical stout seta; palp 2-articulate, article 2 longer than article 1, bearing 6 small robust and three long simple setae. maxilla 2 inner plate slightly narrower than outer, only bearing sparse slender subapical marginal setae, without oblique row of slender setae. gnathopods. coxae 1–4 longer than broad, coxa 4 broader than coxae 1–3. gnathopod 1 subchelate, slightly smaller than gnathopod 2; coxa weakly produced anteroventrally; merus subrectangular, bearing row of slender setae along distal margin; carpus shorter than propodus, cup-shaped, with dense long slender setae on posterior margin; propodus suboval, palm acute, posterior margin with robust and short simple setae; dactylus evenly tapering, with acute tip. gnathopod 2 similar in shape to ganthopod 1; coxa subrectangular, subequal in length to coxa 1, anterior margin slightly convex; carpus distinctly http://zoobank.org/urn:lsid:zoobank.org:act:2a8520b1-c0c6-4bab-9c8d-a7dfb57b111d wang y.-r. et al., a new genus and species of calliopiidae (amphipoda) from okinawa trough 5 fig. 1. bathya brevicarpus gen. et sp. nov., holotype, ♂ (mbm 286556). european journal of taxonomy 693: 1–12 (2020) 6 fig. 2. bathya brevicarpus gen. et sp. nov., holotype, ♂ (mbm 286556). wang y.-r. et al., a new genus and species of calliopiidae (amphipoda) from okinawa trough 7 shorter than propodus, weakly lobed; propodus with tooth-like protruded sub-distally, bearing short simple and robust setae along posterior margin; dactylus with acute tip, evenly tapering. pereopods. pereopod 3 (distal three articles broken in holotype) slender, merus bearing large produced lobe anterodistally. pereopod 4 similar to pereopod 3, but coxa much broader. pereopod 5 slightly longer than pereopod 7, coxa bilobate, anterior lobe smaller than posterior lobe; merus broader than distal three articles, with posterodistal strongly produced as lobe, anterior margin with 4+ groups of 2–3 robust setae; propodus 1.4 times longer than carpus, anterior margin bearing 8 groups of 2–3 bifid robust setae; dactylus slender, with acute tip, evenly tapering, 0.7 length of propodus, with small nail. pereopod 6 (distal three articles broken in holotype) gill smaller than that of pereopod 5; coxa bilobate, anterior lobe much smaller than posterior lobe; basis slight longer than that of pereopod 5. pereopod 7 coxa unilobate, rounded; basis broader than that of pereopods 5 and 6, anterior margin bearing robust setae; propodus subequal in length to carpus; dactylus half-length of propodus, with small acute nail. gills. present on coxae 2–6, small, not pleated. epimeral plates 1 and 3. without tooth on posterior margin, postero-corner of epimeral 2 being subacute. uropods. uropod 1 overreaching distal end of uropod 2, without interramal spur; peduncle shorter than rami, with 19 marginal and one large distal robust setae; outer ramus shorter than inner ramus, but subequal in broad to inner, with 9 marginal and three distal robust setae; inner ramus with 14 marginal and 3 distal robust setae. uropod 2 peduncle distinctly shorter than rami, with 5 marginal and two distal robust setae; outer ramus 0.6 times shorter than inner ramus, with 10 marginal and three distal robust setae; inner ramus with 15 marginal and three distal robust setae. uropod 3 shorter than uropod 2, peduncle shorter than rami, with one robust seta distally; outer ramus 0.55 times shorter than inner ramus, with 7 marginal robust setae; inner ramus with 15 marginal robust setae. telson. entire, longer than broad (length 1.7 times as long as width at base), distal margin convex. discussion the calliopiids are cosmopolitan (barnard 1964; bousfield & hendrycks 1997; lowry & myers 2012), and have been reported from various environments including fresh water, bipolar, shallow and deep waters and hydrothermal vents (e.g., barnard & karaman 1991; bellan-santini & thurston 1996; krappschickel & sorbe 2006; d’udekem d’acoz 2007; lowry & myers 2012; ringvold & tandberg 2014). for example, species of the most species-rich genus of calliopiidae, apherusa walker, 1891, are mostly recorded from the neritic environment (barnard & karaman 1991; krapp-schickel & sorbe 2006) and 13 of 17 known species of oradarea walker, 1903 are found in the antarctic and sub-antarctic islands (alonso 2012). two calliopiids, bouvierella curtirama bellan-santini & thurston, 1996 and leptamphopus fragilis larsen & krapp-schickel, 2007, have been reported from vent fields. however, the present new species belongs to neither bouvierella chevreux, 1900 nor leptamphopus g.o. sars, 1893 (see table 1). this is the third calliopiid from a hydrothermal vent habitat. as the new species cannot be assigned to any genus of calliopiidae, a new genus is erected herein. additionally, a key to all genera of the calliopiidae is presented below. moreover, the morphological differences between several genera of calliopiidae, such as halirages boeck, 1871, haliragoides g.o. sars, 1893 and apherusa, are blurred (ringvold & tandberg 2014). the phylogenetic analyses based on a comparison of 18s and 28s rdna sequences by verheye et al. (2016) shows that the calliopiidae are not monophyletic. hence, the entire calliopiid family is in need of revision. european journal of taxonomy 693: 1–12 (2020) 8 identification key to genera of the calliopiidae key based on original or amended descriptions of genera and adapted from previous keys given by barnard (1964), barnard & karaman (1991) and bousfield & hendrycks (1997). all genera included in this key are according to lowry & myers (2013) and worms (2020). 1. mandibular molar not triturative, usually conical ............................................................................ 2 – mandibular molar triturative, columnar, rather conical .................................................................... 3 2. carpus of gnathopods 1–2 shorter than propodus, weakly lobed; antenna 2 elongate ....................... ...............................................................................................................harpinioides stebbing, 1888 – carpus of gnathopods 1–2 as long as propodus, unlobed; antenna 1 elongate ................................... .............................................................................................................. calliopiurus bushueva, 1986 3. inner plate of maxilla 2 much broader than outer plate .................... pontogeneoides nicholls, 1938 – inner plate of maxilla 2 not much broader than outer plate .............................................................. 4 4. coxae very short and progressively longer towards coxa 7 ............................................................. 5 – coxae not as greatly shortened and not progressively lengthened towards coxa 7 .......................... 6 5. carpus and propodus of pereopods 6–7 extremely elongate (planktonic); body carinate; coxa 1 not or scarcely produced anteriorly ........................... stenopleuroides birstein & m. vinogradov, 1964 – carpus and propodus of pereopods 6–7 not extremely elongate; body smooth; coxa 1 produced anteriorly ................................................................................................. stenopleura stebbing, 1888 6. palp of maxilla 1 reduced, not exceeding apex of outer plate, article 1 longer than article 2 ............. ......................................................................................................................... laothoes boeck, 1871 – palp of maxilla 1 ordinary, article 1 shorter than article 2 ................................................................ 7 7. rami of uropods 1–2 without marginal robust setae, only with distal robust setae ........................... ...........................................................................................................calliopiella schellenberg, 1925 – rami of uropods 1–2 with both marginal and distal robust setae ..................................................... 8 8. gnathopods 2 very slender, linear, carpus very slender and elongate, unlobed, propodus generally elongate and linear (except in amphithopsis) ................................................................................... 9 – gnathopod 2 not very slender nor linear nor greatly elongate (propodus not especially elongate) .... ......................................................................................................................................................... 18 9. gnathopod 1 ordinary, neither linear nor elongate ......................................................................... 10 – gnathopod 1 linear, elongate .......................................................................................................... 12 table 1. synoptic table comparing bathya brevicarpus gen. et sp. nov. and congeneric calliopiids reported from vent fields on the basis of literature data. species accessory flagellum eyes gnathopod 2 telson bathya brevicarpus absent present not linear entire bouvierella curtirama present absent linear truncate leptamphopus fragilis present absent linear cleft wang y.-r. et al., a new genus and species of calliopiidae (amphipoda) from okinawa trough 9 10. propodus of gnathopod 2 not linear; dactylus of pereopods 3–7 serrated .......................................... ..................................................................................................................amphithopsis boeck, 1861 – propodus of gnathopod 2 linear; dactylus of pereopod 3–7 not serrated ........................................11 11. accessory flagellum absent; lower lip with inner lobe absent ..............bouvierella chevreux, 1900 – accessory flagellum present; lower lip with inner lobe present ................... oradarea walker, 1903 12. dactylus of pereopods 3–7 with one or more superior robust setae ............. cleippides boeck, 1871 – dactylus of pereopods 3–7 without superior robust setae .............................................................. 13 13. carpus of gnathopods 1–2 much longer than propodus ................................................................. 16 – carpus of gnathopods 1–2 scarcely longer and usually shorter than propodus .............................. 14 14. gnathopod 2 much longer than gnathopod 1, carpus and propodus extremely slender like that of pereopod 3–7 ................................................................................... leptamphopus g.o. sars, 1893 – gnathopod 2 as long as or slightly longer than ganthopod 1, carpus and propodus normal .......... 15 15. telson entire, linguiform ....................................................membrilopus barnard & karaman, 1987 – telson with shallow apical notch on distal margin ....................frigora ren in ren & huang, 1991 16. antennae calceolate; antenna 1, peduncular article 3 with posterodistal process; uropod3, rami margin only with simple setae ....................................................................... halirages boeck, 1871 – antennae usually lacking calceoli; antenna 1, peduncular article 3 unmodified; uropod 3, margins with robust setae and inner marginal setae only ............................................................................. 17 17. propodus of gnathopods expanded; pereopods 5–7 elongated ........... haliragoides g.o. sars, 1893 – propodus of gnathopods not expanded; pereopods 5–7 normal, not extremely elongated ................ ....................................................................................................................... apherusa walker, 1891 18. peduncular article 3 of antenna 1 produced apicoventrally ............................................................ 19 – peduncular article 3 of antenna 1 not or weakly produced apicoventrally ..................................... 21 19. carpus on either of gnathopods 1–2 much shorter than propodus ................................................. 20 – carpus on either of gnathopods 1–2 scarcely shorter than or longer than propodus .......................... .....................................................................................................................lopyastis thurston, 1974 20. carpus of gnathopods not lobate ................................................................tylosapis thurston, 1974 – carpus of gnathopods strongly lobate .....................................................calliopius lilljeborg, 1865 21. epimeron 3 serrate .......................................................................................................................... 22 – epimeron 3 smooth ......................................................................................................................... 23 22. accessory flagellum well developed, 3+ articulate ...............................weygrechita stuxberg, 1880 – accessory flagellum scale-like ............................................................... oligochinus barnard, 1969 23. rostrum small ................................................................................................................................. 24 – rostrum large .................................................................................................................................. 27 24. accessory flagellum absent ............................................................................................................ 25 – accessory flagellum 1-articulate, scale-like ........ paracalliopiella tzvetkova & kudrjaschov, 1975 25. gnathopod 1 larger in size than gnathopod 2 .................... whangarusa barnard & karaman, 1987 – gnathopod 1 similar in size to gnathopod 2 ................................................................................... 26 european journal of taxonomy 693: 1–12 (2020) 10 26. carpus of gnathopods 1–2 much shorter than propodus; outer ramus of uropod 3 ½ length of inner ramus; telson longer than broad, entire .................................................................... bathya gen. nov. – carpus of gnathopods 1–2 subequal to propodus; rami of uropod 3 subequal in length; telson broader than long, emarginated ...............................................lutriwita lowry & myers, 2012 (fresh water) 27. dactylus of pereopods 3–7 bifid ......................................manerogeneia barnard & karaman, 1987 – dactylus of pereopods 3–7 pectinate .........................................metaleptamphopus chevreux, 1911 acknowledgments many thanks to ed a. hendrycks from the department of research and collections, canadian museum of nature for offering some papers, which were crucial during the identification process of the new species. this work was funded by the national key r&d program of china (2018yfc0310702; 2018yfc0310802), the key research program of frontier sciences, chinese academy of sciences (qyzdb-sswdqc036), the strategic priority research program of the chinese academy of sciences (xda22050302) and the national natural science foundation of china (31625024). references alonso g.m. 2012. amphipod crustaceans (corophiidea and gammaridea) associated with holdfasts of macrocystis pyrifera from the beagle channel (argentina) and additional records from the southwestern atlantic. journal of natural history 46 (29–30): 1799–1894. https://doi.org/10.1080/00222933.2012.692825 barnard j.l. 1964. revision of some families, genera and species of gammaridean amphipoda. crustaceana 7 (1): 49–74. https://doi.org/10.1163/156854064x00263 barnard j.l. & karaman g.s. 1991. the families and genera of marine gammaridean amphipoda (except marine gammaroids). records of the australian museum supplement 13 (1 & 2): 1–866. part 1: https://doi.org/10.3853/j.0812-7387.13.1991.91 part 2: https://doi.org/10.3853/j.0812-7387.13.1991.367 bellan-santini d. & thurston m.h. 1996. amphipoda of the hydrothermal vents along the mid-atlantic ridge. journal of natural history 30: 685–702. https://doi.org/10.1080/00222939600770381 bousfield e.l. & hendrycks e.a. 1995. the amphipod superfamily eusiroidea in the north american pacific region. i. family eusiridae: systematics and distributional ecology. amphipacifica 1: 3–59. bousfield e.l. & hendrycks e.a. 1997. the amphipod superfamily eusiroidea in the north american pacific region. ii. family calliopiidae. systematics and distributional ecology. amphipacifica 2 (3): 3–66. de broyer c., lowry j.k., jażdżewski k. & robert h. 2007. synopsis of the amphipoda of the southern ocean. part 1. catalogue of the gammaridean and corophiidean amphipoda (crustacea) of the southern ocean with distribution and ecological data. bulletin de l’institut royal des sciences naturelles de belgique 77 (suppl. 1): 1–325. d’udekem d’acoz c. 2007. the genera haliragoides and neohela in the north atlantic, with the description of two new deepwater species from norway and svalbard (crustacea: amphipoda). cahiers de biologie marine 48 (1): 17–35. d’udekem d’acoz c. 2010. contribution to the knowledge of european liljeborgiidae (crustacea, amphipoda), with considerations on the family and its affinities. bulletin de l’institut royal des sciences naturelles de belgique, biologie 80: 127–259. https://doi.org/10.1080/00222933.2012.692825 https://doi.org/10.1163/156854064x00263 https://doi.org/10.3853/j.0812-7387.13.1991.91 https://doi.org/10.3853/j.0812-7387.13.1991.367 https://doi.org/10.1080/00222939600770381 wang y.-r. et al., a new genus and species of calliopiidae (amphipoda) from okinawa trough 11 d’udekem d’acoz c. 2012. on the genus halirages (crustacea, amphipoda), with the description of two new species from scandinavia and arctic europe. european journal of taxonomy 7: 1–32. https://doi.org/10.5852/ejt.2012.7 krapp-schickel t. & sorbe j.c. 2006. apherusa delicata n. sp., a new suprabenthic amphipod (crustacea, eusiroidea, calliopiidae) from the northern bay of biscay, with a discussion of the genus. organisms diversity & evolution 6: 57–65. https://doi.org/10.1016/j.ode.2005.05.002 larsen k. & krapp-schickel t. 2007. amphipoda (crustacea: peracarida) from chemically reduced habitats; 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[in chinese.] ringvold h. & tandberg a.h.s. 2014. a new deepwater species of calliopiidae, halirages helgae (crustacea, amphipoda), with a synoptic table to halirages species from the northeast atlantic. european journal of taxonomy 98: 1–13. https://doi.org/10.5852/ejt.2014.98 verheye m.l., martin p., backeljau t. & d’udekem d’acoz c. 2016. dna analyses reveal abundant homoplasy in taxonomically important morphological characters of eusiroidea (crustacea, amphipoda). zoologica scripta 45: 300–321. https://doi.org/10.1111/zsc.12153 weisshappel j.b. 2001. distribution and diversity of the hyperbenthic amphipod family calliopiidae in the different seas around the greenland-iceland-faeroe-ridge. sarsia 86: 143–151. https://doi.org/10.1080/00364827.2001.10420469 worms editorial board 2020. world register of marine species. available from http://www.marinespecies.org [accessed 6 jun. 2020]. https://doi.org/10.14284/170 manuscript received: 5 september 2019 manuscript accepted: 18 may 2020 published on: 29 july 2020 topic editor: rudy jocqué desk editor: pepe fernández printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, https://doi.org/10.5852/ejt.2012.7 https://doi.org/10.1016/j.ode.2005.05.002 https://doi.org/10.1017/s002531540705672x https://doi.org/10.1651/0278-0372(2001)021[0231:roteag]2.0.co;2 https://doi.org/10.11646/zootaxa.3499.1.2 http://dx.doi.org/10.11646/zootaxa.3610.1.1 https://doi.org/10.5852/ejt.2014.98 https://doi.org/10.1111/zsc.12153 https://doi.org/10.1080/00364827.2001.10420469 http://www.marinespecies.org https://doi.org/10.14284/170 european journal of taxonomy 693: 1–12 (2020) 12 brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. ole_link5 ole_link6 ole_link7 _goback european journal of taxonomy 201: 1–12 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2016.201 www.europeanjournaloftaxonomy.eu 2016 · kochaiphat p. et al. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e 1 two new species of tetrastigma (miq.) planch. (vitaceae) from thailand phongsakorn kochaiphat1, anna trias-blasi2 & pimwadee pornpongrungrueng3,* 1,3 applied taxonomic research center, department of biology, faculty of science, khon kaen university, khon kaen 40002, thailand. 2 royal botanic gardens kew, richmond, surrey tw9 3ae, england, u.k. * corresponding author: ppimwa@kku.ac.th 1 email: phongsakorn.s@gmail.com 2 email: a.triasblasi@kew.org abstract. two new species of tetrastigma from thailand, t. calcicola kochaiph. & trias-blasi sp. nov. and t. jaichagunii c.l.li ex kochaiph. & trias-blasi sp. nov. are described and illustrated. tetrastigma calcicola sp. nov. is a slender climber restricted to the open areas on limestone mountains at high elevation in the northern part of thailand. the other species, t. jaichagunii sp. nov., is similar to t. harmandii planch., but differs from it by having more densely verrucose young branches, broader leaflets, 4-lobed thick discs, bigger globose berries and oblongoid seeds. this species occurs along streams or in forest margins in evergreen forest and it is widely distributed in several parts of thailand. keywords. tetrastigma, taxonomy, thailand, revision, vitaceae. kochaiphat p., trias-blasi a. & pornpongrungrueng p. 2016. two new species of tetrastigma (miq.) planch. (vitaceae) from thailand. european journal of taxonomy 201: 1–12. http://dx.doi.org/10.5852/ejt.2016.201 introduction tetrastigma planch. (vitaceae) is a genus comprising approximately 95 species, distributed in tropical and subtropical asia with a few of them reaching australia (planchon 1887; latiff 1983; chen et al. 2011; trias-blasi et al. 2012; wen 2007). it can be easily distinguished from other genera of vitaceae by its polygamo-dioecy, 4-merous flowers, 4-lobed or 4-parted stigma, and absent or very short styles. twenty-six species have been recorded in thailand (kochaiphat et al. 2014), of which five are endemic. during preparation of the revision of tetrastigma for the flora of thailand account, the first author came across several specimens collected from doi tung and tham luang khun nam nang non forest park (chiang rai province) with a unique combination of characters that did not match any previously described species. in addition, a number of specimens representing a distinct taxon were found. some were annotated as tetrastigma jaichagunii by the late prof. c.l. li, however, no formal description had been published http://dx.doi.org/10.5852/ejt.2016.201 https://creativecommons.org/licenses/by/3.0/ http://dx.doi.org/10.5852/ejt.2016.201 european journal of taxonomy 201: 1–12 (2016) 2 for this name. therefore, this species is described here as new species and the name t. jaichagunii is applied. material and methods this study is based on the investigation of the herbarium specimens from relevant major herbaria as well as field collections and observations in thailand. herbarium specimens from the following herbaria were examined: aau, abd, bcu, bk, bkf, bm, c, cmu, cmub, e, k, kku, l, p, psu, qbg and tcd. the herbarium abbreviations follow thiers (continuously updated). voucher specimens collected from field surveys were made following the method of bridson & forman (1989) and were deposited at kku, bkf and qbg. the vegetative parts were measured in a dry state. reproductive parts were rehydrated by boiling in water. measurements were made under light microscope. the morphological terminology generally follows beentje (2010), wen (2007) and jackes (1989); and for berry and seed terminology follows latiff (1983) and chen & manchester (2011). taxonomy class equisitopsida c.agardh (agardh et al. 1825) subclass magnoliidae novák ex takht. (takhtajan 1967) superorder rosanae takht. (takhtajan 1967) order vitales juss. ex bercht. & j.presl (berchtold & presl 1820) family vitaceae juss. (jussieu 1789) nom. cons. genus tetrastigma (miq.) planch. (planchon 1887) two tetrastigma from thailand, t. calcicola sp. nov. and t. jaichagunii sp. nov., are described herein as new species. accordingly, the key to tetrastigma in thailand (kochaiphat et al. 2014) has been revised from couplet 16 as follows. key to the species of tetrastigma in thailand 1. tendrils 5–7 palmately branched; leaves palmately 3-foliolate ........ t. triphyllum (gagnep.) w.t.wang – tendrils simple or bifurcate; leaves simple, palmately 3‒5(‒7)-foliolate or pedately 5‒7(‒9)-foliolate ........................................................................................................................................................... 2 2. inflorescences on old stems (cauliflorous plants), more than 5 inflorescences per nodes ..................... ............................................................................................................................. t. cauliflorum merr. – inflorescences on young branches, 1‒3 inflorescences per nodes ...................................................... 3 3. young branches with 4‒5 sharp ridges ........................................ t. quadrangulum gagnep. & craib – young branches round or nearly round .............................................................................................. 4 4. leaves palmately 3‒5(‒7)-foliolate or mixed with simple leaves or all leaves simple ..................... 5 – leaves pedately 5‒7(‒9)-foliolate rarely mixed with 3-foliolate ..................................................... 16 5. stems verrucose or with corky protuberances ................................................................................... 6 – stems smooth, pubescent or with flaky cork ...................................................................................... 8 6. stems with corky protuberances; leaves ovate to elliptic; stigma distinctly 4-lobed ......................... 7 – stems verrucose; leaves lanceolate; stigma peltate ............................................. t. harmandii planch. 7. leaves fleshly; stigma cruciform; berries ellipsoid when dry ............... t. cruciatum craib & gagnep. – leaves coriaceous or papyraceous; stigma pointed lobes; berries pyriform when dry ......................... ......................................................................... t. assimile (kurz) c.l.li ex kochaiph. & trias-blasi kochaiphat p. et al., new species of tetrastigma from thailand 3 8. shrubs, creeping, erect or decumbent ................................................................................................ 9 – lianas .............................................................................................................................................. 10 9. leaves simple or mixed with palmately 3(‒5)-foliolate; pedicel 0.4‒0.6 cm long ............................... ....................................................................................................................... t. bambusetorum craib – leaves palmately 3-foliolate; pedicel 0.8–1.2 cm long .................................... t. apiculatum gagnep. 10. leaflets glabrous .............................................................................................................................. 11 – leaflets pubescent or hirsute at least along the midrib on lower leaf surface ................................... 15 11. female flower disc conspicuous, thick and adnate to lower part of ovary ........................................ 12 – female flower disc inconspicuous ................................................................................................... 14 12. terminal leaflets broadly elliptic to rhombic; berries pyriform when dry ............................................. .................................................................................. t. pedunculare (wall. ex m.a.lawson) planch. – terminal leaflets elliptic, ovate or lanceolate; berries ellipsoid when dry ........................................ 13 13. leaves coriaceous or subcoriaceous .................................................................. t. erubescens planch. – leaves papyraceous ........................................................................ t. dubium (m.a.lawson) planch. 14. leaflets broadly ovate or broadly elliptic, apex caudate ................. t. campylocarpum (kurz) planch. – leaflets lanceolate, apex attenuate ..................................................... t. planicaule (hook.f.) gagnep. 15. ovary brownish hirsute; leaves palmately 5-foliolate ................ t. obovatum (m.a.lawson) gagnep. – ovary glabrous; leaves palmately 3-foliolate ................................. t. dubium (m.a.lawson) planch. 16. leaves 5-foliolate, rarely mixed with 7-foliolate ............................................................................. 17 – leaves 7(‒9)-foliolate, rarely mixed with 5-foliolate ...................................................................... 29 17. branches glabrous; leaflets glabrous on both surfaces ..................................................................... 18 – branches more or less pubescent; leaflets pubescent at least along the midvein on lower leaf surface ......................................................................................................................................................... 26 18. secondary branches of peduncle 2 ....................................................................... t. baenzigeri c.l.li – secondary branches of peduncle more than 2 .................................................................................. 19 19. berries pyriform or triangular when dry .......................................................................................... 20 – berries globose, ovoid or ellipsoid when dry ................................................................................... 21 20. terminal leaflets ovate–lanceolate, 3–4 × 6–9 cm; berries pyriform when dry ... t. pyriforme gagnep. – terminal leaflets obovate or elliptic, 4‒5.5 × 9‒12 cm; berries triangular when dry ............................ .......................................................................................... t. rumicispermum (m.a.lawson) planch. 21. inflorescences compound umbel on pseudo-terminal branch or axile, umbelliform, loose; tendrils bifurcate ............................................................................................... t. serrulatum (roxb.) planch. – inflorescences compound umbel on axile, globose, compact; tendrils unbranched ......................... 22 22. female flower disc inconspicuous .................................................... t. pachyphyllum (hemsl.) chun – female flower disc conspicuous, thick and adnate to lower part of ovary ........................................ 23 23. stigma peltate, rounded or slightly 4-lobed ..................................................................................... 24 – stigma cruciform, 4-lobed, obtuse, acute or pointed apex ............................................................... 25 european journal of taxonomy 201: 1–12 (2016) 4 24. berries 1.8–3.0 × 1.5–2.5 cm; seed oblongoid; female flower disc 4-angled ........................................ .................................................................. t. jaichagunii c.l.li ex kochaiph. & trias-blasi sp. nov. – berries 0.8–1.2 × 0.8–1.2 cm; seed ellipsoid; female flower disc rounded ......... t. harmandii planch. 25. stigmas 4 acute or pointed lobes, style tubular 0.5–0.8 mm long ... t. dubium (m.a.lawson) planch. – stigmas 4 obtuse lobes, style absent ............................. t. calcicola kochaiph. & trias-blasi sp. nov. 26. leaflets broadly elliptic or ovate ..................................................................................................... 27 – leaflets narrowly elliptic or lanceolate ........................................... t. dubium (m.a.lawson) planch. 27. leaflets pubescent along nerve or at least along the midrib on the lower surface ............................ 28 – leaflets pubescent on both surfaces ........................................ t. macrocorymbum gagnep. ex j.wen 28. berries up to 1.5 cm in diameter .............................................................................. t. pilosum c.l.li – berries more than 2 cm in diameter ....................................................... t. siamense gagnep. & craib 29. berries 2‒2.5 cm in diameter ................................................................................ t. teaniatum c.l.li – berries 0.8‒1.5 cm in diameter ........................................................................................................ 30 30. outer petals densely papillose ..................................... t. leucostaphylum (dennst.) alston ex mabb. – outer petals glabrous ....................................................................................................................... 31 31. secondary vein almost 90° with midrib; old branches verrucose, light brown; stigma 4 pointed lobes .............................................................................................................................. t. delavayi gagnep. – secondary vein angle nearly 45° with midrib; old branches flaky, grey; stigma 4 rounded lobes or nearly globose, not lobed ............................................................................. t. godefroyanum planch. tetrastigma calcicola kochaiph. & trias-blasi sp. nov. table 1, figs 1–2 urn:lsid:ipni.org:names:77155114-1 diagnosis tetrastigma calcicola sp. nov. is most closely aligned to t. pachyphyllum (hemsl.) chun, but differs in having a 4-lobed disc adnate to the lower part of the ovary, and small berries with a smooth surface (table 1). etymology the specific epithet refers to the habitat of this species, which is only found on limestone mountains. type materials holotype thailand: chiang rai, tham luang khun nam nang non forest park, 21 mar. 2011, m. norsaengsri & n. tathana 7810, ♀ fl. (holo-: qbg!). paratype thailand: chiang rai, tham luang khun, nam nang non forest park, 10 jul. 2012, m. norsaengsri & n. tathana 9736, ♀ fr. (qbg). http://www.ipni.org/urn:lsid:ipni.org:names:77155114-1 kochaiphat p. et al., new species of tetrastigma from thailand 5 description slender climber. stems terete, young branches terete, verrucose; mature stems corky; tendrils simple; stipules 2, deltate, c. 0.2 × 0.2 cm long, scale-like with age. leaves compound, 3-foliolate or pedately 5-foliolate; petiole 0.8–5.0 cm long, glabrous, base slightly pulvinate; leaflets coriaceous; terminal leaflet petiolule 0.2–2.0 cm long, glabrous, terminal leaflet blade elliptic, obovate to oblanceolate, 2.0–8.0 × 1.0–3.5 cm, base cuneate, margins serrate with minute protruding vein tip, apex acute to acuminate, petiolules of lateral leaflet complex 0.1–1.0 cm long, glabrous, lateral leaflet petiolules, sessile to 0.6 cm long, glabrous, lateral leaflet blade elliptic, obovate to oblanceolate, 1.0–5.5 × 0.5–2.8 cm, base cuneate or asymmetrical, margins and apex as for terminal leaflet; veins conspicuous on lower sides, midrib protruding on both surfaces, glabrous, 1 main basal vein, 4–7 pairs of lateral veins. inflorescences axillary on young stem, 1–2 per node, compound umbel, male plant not seen; female plant 1.0–2.5 cm in diameter, compact, globose; peduncles 0.3–1.0 cm, rarely puberulous. female flowers’ bud ovoid, 1.0–1.5 mm long; pedicels 0.5–1.0 mm long, papillose; calyx disciform, margins entire, papillose; petals ovate, 1.5–1.8 × 0.8 mm, apex slightly corniculate, outer surface densely papillose, margins entire; disc 4-lobed adnate to lower part of ovary; staminode clavate, c. 1.2 mm long; ovary conical, c. 0.8 × 0.8 mm; style sessile; stigma cruciform, 4-lobed, lobes obtuse, ciliate. berries globose to ellipsoid, 0.8–1.0 × 0.7–0.8 cm, surface smooth, yellow or white when ripe, 1–3-seeded. seeds ellipsoid, 0.7–0.8 × 0.4–0.5 cm, testa transversely rugulose on both sides, adaxial surface with a y shaped furrow, abaxial surface with an oblong chalaza, endosperm ‘ ’ shaped in cross-section. distribution known only from thailand (fig. 2). ecology and phenology in open area on limestone mountain, dry evergreen forest, altitude 1,300–1,450 m; flowering: february– march; fruiting: july–october. characters t. calcicola kochaiph. & trias-blasi sp. nov. t. pachyphyllum (hemsl.) chun female flowers pedicels length 0.5–1.0 mm 2.0–3.0 mm petals size 1.5–1.8 × 0.8 mm 2.5–3.0 × 1.2–1.7 mm disc 4-lobed adnate to lower part of ovary inconspicuous berries shape globose to ellipsoid globose size 0.8–1.0 × 0.7–0.8 cm 1.4–1.7 × 1.4–1.8 cm surface smooth coarse seeds size 0.7–0.8 × 0.4–0.5 cm 0.9 × 0.5–0.7 cm endosperm shape in cross-section table 1. main morphological differences between t. calcicola kochaiph. & trias-blasi sp. nov. and t. pachyphyllum (hemsl.) chun. european journal of taxonomy 201: 1–12 (2016) 6 fig. 1. tetrastigma calcicola kochaiph. & trias-blasi sp. nov. a. branch and female inflorescence. b. infructescence. c. ovary. d. corolla, dorsal view. e. corolla, lateral view. f. corolla, ventral view. g. seed, dorsal view. h. seed, ventral view. i. seed, transversal section. drawn by woranart thammarong. a, c–f from m. norsaengsri & n. tathana 7810; b from r. pooma 1186; g–i from m. norsaengsri & n. tathana 9736. kochaiphat p. et al., new species of tetrastigma from thailand 7 fig. 2. distribution of tetrastigma calcicola kochaiph. & trias-blasi sp. nov. (■) and t. jaichagunii c.l.li ex kochaiph. & trias-blasi sp. nov. (●). european journal of taxonomy 201: 1–12 (2016) 8 additional specimens examined thailand. chiang rai:doi tung, 22 oct. 1995, r. pooma 1186, ♀ fr. (bkf, cmub); ibid., 18 feb. 2005, j.f. maxwell 05-147, ♀ fl. (cmub). conservation status we suggest to treat this species as data deficient (dd) according to iucn (2012), as this species is only known from a few herbarium specimens and only two localities have been recorded: doi tung and tham luang khun nam nang non forest park in chiang rai province. although the species seems to occur only in limestone mountains at high elevation, the number of populations and their sizes are not known so far. tetrastigma jaichagunii c.l.li ex kochaiph. & trias-blasi sp. nov. table 2, figs 2–3 urn:lsid:ipni.org:names:77155116-1 diagnosis this species is similar to t. harmandii planch., but differs from it by having more densely verrucose young branches, broader leaflets, 4-lobed thick discs, bigger globose berries and oblongoid seeds (table 2). some specimens were misidentified as t. hookeri planch., an indian species, but t. hookeri has narrow leaves and an inconspicuous disc in female flowers. etymology we think that prof. c.l. li selected the specific epithet ‘jaichagunii’ in honour of mr manit jaichagun, from the cites scientific authority in thailand. type materials holotype thailand. trang, khao chong, 12 mar. 1974, k. larsen & s.s. larsen 33238, ♀ fl. (holo-: aau!, iso-: bkf!, l!, p!). paratypes thailand. phetchaburi, kaeng krachan national park, 3 aug. 1995, k. larsen, s.s. larsen, c. tange, r. moran & p. puudjaa 45370, ♀ fr. (aau); satun, ta le bun, 26 nov. 1985, j.f. maxwell 851060, ♀ fr. (l). description large climber. stems flattened with age, young branches terete, verrucose; mature stems corky; tendrils simple; stipules 2, deltate, c. 0.4 × 0.3 cm, reduced to scale-like with age. leaves compound, 3-foliolate or pedately 5(–7)-foliolate; petioles 4.0–19.0 cm long, verrucose, base pulvinate; leaflets papyraceous to subcoriaceous; terminal leaflet petiolule, 1–5.0 cm long, glabrous, terminal leaflet blade lanceolate, elliptic, to oblanceolate, 11.5–33.0 × 3.5–12.0 cm, base obtuse to cuneate, margins coarsely serrate with c. 0.5 protruding vein tip, apex acuminate to caudate, petiolules of lateral leaflet complex 1.0–3.5 cm long, glabrous, lateral leaflet petiolules, sessile–2.5 cm long, glabrous, lateral leaflet blade lanceolate, elliptic to broadly elliptic, oblong, 8.5–30 × 2.5–11.5 cm, base obtuse to cuneate or asymmetrical, margins and apex as terminal leaflet; veins conspicuous on lower sides, midrib protruding on both surfaces, glabrous, 1 main basal vein, 6–13 pairs of lateral veins. inflorescences axillary on young stem, single, compound umbel; 2–3 cm in diameter, compact, umbelliform; peduncles 0.5–1.0 cm, puberulent, male plant not seen. male flowers not seen. female flowers’ bud ovoid, 2.0–3.0 mm long; pedicels 2.0–3.0 mm long, papillose; calyx disciform to cupuliform, margins undulate, ciliate; petals ovate to oblong, 2.2–3.0 × http://www.ipni.org/urn:lsid:ipni.org:names:77155116-1 kochaiphat p. et al., new species of tetrastigma from thailand 9 1.5–1.8 mm, apex hooded, outer surface densely papillose, margins entire; disc thick, 4-angled, adnate to ovary; staminode clavate–trullate, 0.5–1.0 mm long; ovary conical, 1.0–1.2 × 1.0–1.5 mm; style cylindrical, c. 0.5 mm long; stigma peltate, round or slightly 4-lobed, ciliate. berries globose, 1.8–3.0 × 1.5–2.5 cm, surface smooth, green when young, yellow when ripe, 1–4-seeded. seeds oblongoid, 1.2–1.4 × 1.0–1.2 cm, testa transversely rugose on both sides, adaxial surface with an oblong furrow, abaxial surface with a linear chalaza, apex bilobed, apical notch 1.0–3.0 mm, endosperm ‘ ’ shaped in cross-section. distribution known only from thailand (fig. 2). ecology and phenology along streams or in forest margins in evergreen forest; altitude 0–1,500 m; flowering: january–april; fruiting: june–january. additional specimens examined thailand. northern: chiang mai: chiang mai, along the road, 40 km to pai, 17 sep. 1995, k. larsen, s.s. larsen, c. tange & d. sookchaloem 46583, ♀ fl. (aau); chiang dao, 26 dec. 1940, h.b.g. garrett 1214, ♀ fr. (tcd, l, p); doi chiang dao, 13 jul. 1950, h.b.g. garrett 1336 (p, l, k); ibid., 18 dec. 1951, h.b.g. garrett 1374 (p, l, k); ibid., 22 jan. 1989, j.f. maxwell 89-0081, ♀ fr. (l); hang dong, mae khanin, 7 mar. 1999, s. watthana & w. pongamornkul wat. 240, ♀ fl. (qbg); mae tang, 19 nov. 1990, j.f. maxwell 90-1259, ♀ fr. (cmu); mae tang, pa pae, 17 sep. 1995, bgo. staff. 4507 (qbg); south-western: kanchanaburi: between kriti and mueang cha, 9 jul. 1973, r. geesink & c. phengklai 6220, ♀ fr. (aau, bkf, e, l, p); thung yai naresuan, 11 aug. 1992, j.f. maxwell 93-869, ♀ fr. (cmub); thong pha phum, thung yai naresuan, 10 jun. 2002, m. van de bult 565, ♀ fr. (bkf, cmub); phetchaburi: kaeng krachan national park, 26 aug. 1995, j.a.n. parnell, j. pendry & t. boonthavikoon 95-450, ♀ fr. (k); ibid., 25 jun. 2000, m.f. newman, t. boonthavikoon, c. hemrat & d.j. middleton 1040, ♀ fr. (aau, bkf, l, p); ibid., 9 aug. 2002, d.j. middleton, s. suddee, s.j. davies & c. hemrat 888, ♀ fr. (bkf, cmub); prachuap khiri khan: kaeng krachan national park, pa la-u waterfalls, 14 aug. 2002, d.j. middleton, s. suddee, s.j. davies & c. hemrat 1060, ♀ characters t. jaichagunii c.l.li ex kochaiph. & trias-blasi sp. nov. t. harmandii planch. female flowers petals shape ovate to oblong ovate disc thick, 4-angled thick, rounded berries size 1.8–3.0 × 1.5–2.5 cm 0.8–1.2 × 0.8–1.2 cm seeds shape oblongoid ellipsoid size 1.2–1.4 × 1.0–1.2 cm 0.7–0.8 × 0.6–0.7 cm endosperm shape in cross-section table 2. main morphological differences between t. jaichagunii c.l.li ex kochaiph. & trias-blasi sp. nov. and t. harmandii planch. european journal of taxonomy 201: 1–12 (2016) 10 fr. (bkf, cmub); peninsular: ranong: klong naka national park, 24 nov. 1974, r. geesink, p. hiepko & c. charoenpol 7519, ♀ fr. (bkf, k); ibid., 7 jan. 1990, s. hoover 5048, ♀ fr. (e); ibid., 8 jan. 1990, s. hoover 5481, ♀ fr. (e); ibid., 13 jan. 1990, s. hoover 6329, ♀ fr. (e); khao po ta luang kaeo, 31 jan. 1929, a.f.g. kerr 16912, ♀ fl. (l, p); no locality, 22 jan. 1929, a.f.g. kerr 16805a (bk, k); kapoe, 15 jul. 1979, c. niyomdham et al. 320, ♀ fr. (aau, bkf, k, l, p); khao po ta luang kaeo, 11 dec. 1979, t. shimizu, h. toyokuni, h. koyama, t. yahara & c. niyomdham t-26922 (aau, bkf, l); surat thani: klong phanom national park, 21 mar. 2005, s. gardner, p. sidisunthorn & p. tippayasri st1875, ♂ fl. (qbg); phangnga: khao po ta luang kaeo, 2 feb. 1929, a.f.g. kerr fig. 3. tetrastigma jaichagunii c.l.li ex kochaiph. & trias-blasi sp. nov. a. branch and female inflorescence. b. infructescence. c. ovary. d. corolla, ventral view. e. corolla, dorsal view. f. corolla, lateral view. g. seed, ventral view. h. seed, dorsal view. i. seed, transversal section. drawn by woranart thammarong. a, c–f from k. larsen & s.s. larsen 33238; b from k. larsen et al. 4537; g–i from j.f. maxwell 85-1060. kochaiphat p. et al., new species of tetrastigma from thailand 11 16990, ♀ fl. (bk, bm, k, l); phuket: kathu, 12 mar. 1929, a.f.g. kerr 17463 (bk, k); nakhon si thammarat: ka rom waterfalls, 14 apr. 1985, j.f. maxwell 85-0396, ♀ fl. (bkf, l); ibid., 30 oct. 1993, k. larsen, s.s. larsen, c.t. norgaard, k. pharsen, p. puudjaa & w. ueachirakan 44163, ♀ fr. (aau); tha sala, 2 mar. 1986, j.f. maxwell 86-0111, ♀ fl. (bkf, l); thung song, 19 jul. 1929, rabil 93, ♀ fr. (bk, l); phatthalung: khao pu khao ya national park, 16 nov. 1990, k. larsen, s.s. larsen, w. nanakorn, w. ueachirakan & p. sirirugsa 41534, ♀ fl. (aau, k); trang: khao chong, 13 jun. 1974, r. geesink, t. hattink & c. charoenpol 7181, ♀ fr. (bkf, k); ibid., 10 jul. 2000, d.j. middleton, t. boonthavikoon, s.j. davies, c. hemrat & m.f. newman 373, ♀ fr. (aau, bkf); pa liang, 24 apr. 1931, m.c. laksanakarn 791, ♀ fl. (l, k); sai rung waterfalls, 15 mar. 2006, s. gardner & v. chamchaumroon st2480, ♀ fl. (qbg); satun: ko ta ru tao, 19 jan. 1918, a.f.g. kerr 14188, ♀ fl. (bm, k); klong ton, 11 mar. 1928, a.f.g. kerr 14469, ♀ fl. (l, k); yala: bang lang national park, 12 feb. 2004, d.j. middleton, m. phuphat, r. pooma & k. williams 3011, ♂ fl. (bkf, e); no locality, 25 mar. 1998, c. niyomdham 5331, ♀ fl. (bkf, k). conservation status this species has rather large populations and is widely distributed in several parts of the country. therefore, it is considered least concern (lc). discussion tetrastigma calcicola sp. nov. and t. jaichagunii sp. nov. are distinct from the other previously described species. the major characters for recognising the species are female flowers and fruits. tetrastigma calcicola sp. nov. differs from its closest relative, t. pachyphyllum, by its 4-lobed disc that is adnate to the lower part of the ovary, and small berries with a smooth surface. tetrastigma jaichagunii sp. nov. can be separated from t. harmandii by its more densely verrucose young branches, broader leaflets, 4-lobed thick discs, bigger globose berries and oblongoid seeds. currently, these two new species are known only from thailand. tetrastigma calcicola sp. nov. is restricted to limestone mountains at high elevation in the northern part of thailand, while t. jaichagunii sp. nov. has a wider range of distribution. the latter species has only been recorded from evergreen forest in several parts of thailand in large populations. however, it seems possible that it might be found in neighbouring countries such as myanmar or laos if a more extensive field survey is carried out in the future. acknowledgements this work was supported by the graduate school, khon kaen university and science achievement scholarship of thailand (sast). the first author would like to thank prof. dr. john a.n. parnell and other staff at tdc for their kind help and also grateful to the staff of aau, abd, bcu, bk, bkf, bm, c, cmu, cmub, e, k, kku, l, p, psu, qbg and tcd for their help and for the loan of/or access to specimens. thanks to mr. woranart thammarong for the illustrations. references agardh c.a., holmberg l.p. & lundstrom p.m. 1825. classes plantarum. literis berlingianis, lundae [lund]. http://dx.doi.org/10.5962/bhl.title.7657 beentje h. 2010. the kew plant glossary. kew publishing, kew, uk. berchtold b.v. von & presl j.s. 1820. o přirozenosti rostlin, aneb rostlinár, obsahugjcj: gedanj on žiwobytj rostlinném pro sebe a z ohledu giných žiwoků, podlé stawu nyněgssjbo znánj; k rozssjřenj http://dx.doi.org/10.5962/bhl.title.7657 european journal of taxonomy 201: 1–12 (2016) 12 přirodnictwj; w potaženj na užitećnost w rolnictwj, hospodářstwj, řemestech, uměnj i obchodu a w wztahowánj obzwlásstnjm na lekařstwj. enders, prague. bridson d. & forman l. 1989. the herbarium handbook. royal botanic gardens, kew, london. chen p., chen l. & wen j. 2011. the first phylogenetic analysis of tetrastigma (miq.) planch, the host of rafflesiaceae. taxon 60 (2): 499–512. chen i. & manchester s.r. 2011. seed morphology of vitaceae. international journal of plant science 172 (1): 1–35. http://dx.doi.org/10.1086/657283 iucn 2012. iucn red list categories and criteria: version 3.1. second edition. iucn, gland, switzerland and cambridge, uk. jackes b.r. 1989. revision of the australian vitaceae, 5. tetrastigma (miq.) planchon. austrobaileya 3 (1): 149–158. jussieu a.l. de 1789. genera plantarum, secundum ordines naturales disposita, juxta methodum in horto regio parisiensi exaratam. herissant & barrois, paris. http://dx.doi.org/10.5962/bhl.title.7762 kochaipat k., trias-blasi a. & pornpongrungrueng p. 2014. a new combination and new records of tetrastigma (vitaceae) from thailand. phytotaxa 183 (4): 272–278. http://dx.doi.org/10.11646/ phytotaxa.183.4.6 latiff a. 1983. studies in malesian vitaceae vii. the genus tetrastigma in the malay peninsula. gardens’ bulletin singapore 36 (2): 213–228. planchon j.e. 1887. monographie des ampélidées vrais. in: de candolle a.f.p.p. & de candolle c. 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(ed.) the families and genera of vascular plants. vol. 9: 467– 479. springer-verlag, berlin. manuscript received: 14 october 2015 manuscript accepted: 8 january 2016 published on: 30 may 2016 topic editor: koen martens desk editor: natacha beau printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark. http://dx.doi.org/10.1086/657283 http://dx.doi.org/10.5962/bhl.title.7762 http://dx.doi.org/10.11646/phytotaxa.183.4.6 http://dx.doi.org/10.11646/phytotaxa.183.4.6 http://sweetgum.nybg.org/ih/ http://sweetgum.nybg.org/ih/ http://dx.doi.org/10.1600/036364412x656437 european journal of taxonomy 570: 1–14 issn 2118-9773 https://doi.org/10.5852/ejt.2019.570 www.europeanjournaloftaxonomy.eu 2019 · shipunov a. et al. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e 1 mysterious chokeberries: new data on the diversity and phylogeny of aronia medik. (rosaceae) alexey shipunov 1,*, sofia gladkova 2, polina timoshina 3, hye ji lee 4, jinhee choi 5, sarah despiegelaere 5 & bryan connolly 5 1,4,5,6 minot state university, biology, 500 university ave, minot, nd, usa. 2,3 department of biology, moscow state university, russia. 7 framingham state university, biology, 100 state st, framingham, ma, usa. * corresponding author: dactylorhiza@gmail.com 2 email: mgu@univer.msu.ru 3 email: mgu@univer.msu.ru 4 email: msu@minotstateu.edu 5 email: msu@minotstateu.edu 6 email: msu@minotstateu.edu 7 email: bconnolly@framingham.edu abstract. aronia medik. (chokeberry, rosaceae) is a genus of woody shrubs with two or three north american species. species boundaries and relationships between species of aronia are frequently under question. the only european species in the genus, a. mitschurinii a.k.skvortsov & maitul., is suggested to be an inter-generic hybrid. in order to clarify the relationships between species of aronia, we performed several morphometric and molecular analyses and found that the molecular and morphological diversity within data on american aronia is low, and species boundaries are mostly not clearly expressed. whereas morphology is able to separate american species from a. mitschurinii, there is no support for such discrimination from the molecular data; our analyses did not reveal evidence of a. mitschurinii hybrid origin. we believe that higher-resolution markers are needed to resolve species boundaries and putative hybridization events. keywords. aronia, rosaceae, dna, morphology, hybridity. shipunov a., gladkova s., timoshina p., lee h.j., choi j., despiegelaere s. & connolly b. 2019. mysterious chokeberries: new data on the diversity and phylogeny of aronia medik. (rosaceae). european journal of taxonomy 570: 1–14. https://doi.org/10.5852/ejt.2019.570 introduction chokeberry, aronia medik. (medikus 1789), is a genus of woody shrubs in the family rosaceae. the boundaries of species of aronia are unclear and typically three (hardin 1973; persson hovmalm et al. 2004) or two (robertson et al. 1991) species are recognized in north america. the following north american species are recognized:1) aronia melanocarpa (michx.) elliott or black chokeberry (with https://doi.org/105852/ejt.2019.570 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ mailto:dactylorhiza%40gmail.com?subject= mailto:mgu%40univer.msu.ru?subject= mailto:mgu%40univer.msu.ru?subject= mailto:msu%40minotstateu.edu?subject= mailto:msu%40minotstateu.edu?subject= mailto:msu%40minotstateu.edu?subject= mailto:bconnolly%40framingham.edu?subject= https://doi.org/10.5852/ejt.2019.570 european journal of taxonomy 570: 1–14 (2019) 2 diploid and tetraploid forms; connolly 2014), 2) aronia arbutifolia (l.) pers. or red chokeberry, 3) aronia × floribunda (lindl.) spach (synonyms: aronia × prunifolia (marshall) rehder, aronia atropurpurea britton) or purple chokeberry (likely of hybrid origin with possible subsequent introgression; connolly 2014). the only european species in the genus, a. mitschurinii (skvortsov & majtulina 1982; skvortsov et al. 1983) or mitschurin’s chokeberry is suggested to be an inter-generic hybrid × sorbaronia mitschurinii (skvortsov & maitul.) sennikov, but its origin is not yet clarified (leonard 2011; smolik et al. 2011; leonard et al. 2013; sennikov & phipps 2013; vinogradova & kuklina 2014). chokeberries are agriculturally promising for fruit and nutraceuticals, and also have applications as ornamental landscape plants (brand 2010; taheri et al. 2013; connolly 2014). aronia mitschurinii has bigger and tastier fruits then other aronias and was adopted as valuable agricultural fruit plant across north america (kask 1987). we believe that instability in aronia taxonomy is largely the result of insufficient sampling; thus, our goal was to increase sampling, use expert knowledge to identify or re-identify our samples, and then to employ dna sequence analyses and advanced morphometry methods in order to perform a comprehensive analysis of aronia diversity which will clarify species relationships. material and methods dna sequencing and molecular analysis we sequenced the nuclear rdna internal transcribed spacer 2 (its2), the chloroplast encoded rbcl gene and trnl-f spacer (kuzmina & ivanova 2011) from several representatives of each of five aronia species (34, 38 and 15, respectively). these markers are commonly used for studying the phylogeny and origins of angiosperms species including rosaceae (campbell et al. 2007; li et al. 2012; lo & donoghue 2012; dluzewska et al. 2013; zarrei et al. 2014). in addition, we amplified and sequenced the second intron of the nuclear low-copy leafy gene which has been considered a useful marker in phylogenetic analysis of rosaceae (oh & potter 2003; lo et al. 2007; li et al. 2017). we obtained 42 tissue samples from herbarium collections in the usa (cas, f, huh, jeps, mo, ny, uc, us) and russia (mha). we also used 31 freshly collected leaf tissue samples taken in the field and from the rosaceae living collection of the university of connecticut (usa). field collected material (mostly of a. mitschurinii) was from the central russian tverj and moscow regions, where tissue samples were obtained from the living plants growing in the cultivation and the living plants that have escaped from the cultivation. the list of samples, all sequences, datasets and scripts are available on zenodo (https://doi.org/10.5281/zenodo.3276157). all sequences were also submitted to genbank. in addition to aronia, we used samples of sorbus aucuparia l. (linnaeus 1753) and of the intergeneric hybrid, × sorbaronia fallax (c.k.schneid.) c.k.schneid. (sorbus aucuparia × aronia melanocarpa: schneider 1906; connolly 2009). genbank sequences of s. aucuparia were also used. the first sample is phylogenetically distant from both aronia and sorbus s.str. (sun et al. 2018) and therefore was used as an outgroup in most phylogenetic analyses. considering problems with aronia species identification (typical issue is that proper determination requires checking the autumn coloration of fruits: hardin 1973), genbank sequences of this genus were not thought to be reliably identified. dna was extracted using either a mo bio powerplant dna isolation kit (mo bio laboratories, carlsbad, california, usa), or nucleospin plant ii kit (macherey-nagel gmbh & co. kg, düren, germany). dry plant leaf material (typically, 0.05–0.09 g) was powdered using a sterile mortar and pestle and then processed in accordance with the supplied protocol. we increased the lysis time to 30 minutes and used thermomixer on the slow rotation speed (350 rpm) instead of water bath. https://doi.org/10.5281/zenodo.3276157 shipunov a. et al., new data on the phylogeny of aronia 3 nanodrop 1000 spectrophotometer (thermo scientific, wilmington, de, usa) was used to assess the concentration and purity (the 260/280 nm ratio of absorbance) of dna samples. we sequenced the markers mentioned above using primers and protocols in accordance with recommendations of the barcoding of life consortium (kuzmina & ivanova 2011) and the lfy1/lfy2 primers (oh & potter 2003). pcr was carried out as follows: the reaction mixture in a total volume of 20 μl contained 5.2 μl of pcr master mix (components from qiagen corporation, germantown, maryland), 1 μl of 10 μm forward and reverse primers, 1 μl of dna solution from the extraction above and 11.8 μl purified water. samples were incubated in a thermal cycler: 94° for 5 min, then 35 cycles of 94° for 1 min; 51° for 1 min, 72° for 2 min, and finally 72° for 10 min. regardless to the marker and species, we always received single band pcr products. they were sent for purification and sequencing to functional biosciences, inc. (madison, wyoming) and sequenced there in accordance with standard protocol. sequences were obtained, assembled and edited using sequenchertm 4.5 (genes codes corporation, ann arbor, michigan, usa) and then aligned with aliview / muscle (larsson aronia mitschurinii a−20−1−3 aronia mitschurinii a−201 aronia mitschurinii a−202 aronia melanocarpa4 a−203 aronia melanocarpa2 a−204 aronia floribunda a−205 aronia melanocarpa4 a−206 aronia melanocarpa2 a−208 aronia arbutifolia a−211 aronia mitschurinii a−212 aronia melanocarpa4 a−213 sorbaronia fallax a−215 aronia mitschurinii a−23−2−1 aronia arbutifolia a−402 aronia arbutifolia a−602 aronia arbutifolia a−604 aronia melanocarpa4 a−609 aronia floribunda a−207 aronia melanocarpa2 a−214 aronia melanocarpa4 a−408 aronia melanocarpa2 a−502 aronia floribunda a−503 aronia arbutifolia a−603 aronia floribunda a−606 aronia arbutifolia a−608 aronia melanocarpa4 a−610 aronia floribunda a−703 aronia floribunda a−707 aronia arbutifolia a−801 aronia arbutifolia a−802 aronia melanocarpa a−804 aronia mitschurinii s11 aronia mitschurinii s9 sorbus aucuparia s6 kf718372 100 56 91 82 0.005 fig. 1. ml tree resulted from the analysis of concatenated its2 and rbcl sequences. european journal of taxonomy 570: 1–14 (2019) 4 2014) and clustalx (thompson et al. 1997) using gap opening cost 9, gap extension cost 0.05 and iub weight matrix. in total, we produced 309 dna sequences, and after the preliminary analysis, selected the 113 longest and minimally noisy sequences for further research. those selected sequences belonged to 42 individual plants from all species of aronia (12–24 sequences per species). in terms of the markers, most successful were the amplification and subsequent sequencing of rbcl (35 sequences) and the least successful the amplification of the first variant of leafy second intron (6 sequences). phylogenetic analyses were implemented with the help of the r ape and phangorn packages (paradis et al. 2004; schliep 2011). we used the kimura distance, neighbor-joining and maximum likelihood (under a gtr model with gamma distribution selected using the akaike information criterion implemented in r) trees for the most of our calculations. counts i0 i1 i12 i2 i23 r0 r1 r2 t0 t1 2 4 6 8 a. arbutifolia a. floribunda 2 4 6 8 a. melanocarpa 2n i0 i1 i12 i2 i23 r0 r1 r2 t0 t1 a. melanocarpa 4n 2 4 6 8 a. mitschurinii fig. 2. abundance of haplotypes per species. each rectangle is a species. the first letter of haplotype name represent the marker: i for its2, r for rbcl and t for trnl-f. dots correspond with haplotype counts. shipunov a. et al., new data on the phylogeny of aronia 5 morphometric analyses we gathered morphometric data from 436 herbarium samples and living plants of aronia (including 187 samples of a. mitschurinii). we applied several methods of multivariate analyses using both ‘classical’ and geometric morphometrics approaches. as the dried herbarium samples might suffer from shrinking (volkova et al. 2011), we compared the fresh and the dried herbarium samples of a. mitschurinii to calculate the shrinking coefficient. this coefficient was less than 4%, therefore shrinking was not taken into account in our geometric morphometrics analyzes. several leaf-related morphological characters were measured, including ‘classic’ petiole and leaf length, leaf maximal width and position of maximal width. in geometric morphometrics analyses, we used the standard thin plate spline approach (zelditch et al. 2012) with 10 landmarks located on the tip and base of the leaf plus on the points of maximal curvature on the leaf contour (see fig. 7 for the examples). coordinates of the landmarks were written to the data file with tpsdig (rohlf 2010). 8 6 4 2 0 dissimilarity a. mitschurinii a. mitschurinii a. melanocarpa a. arbutifolia a. floribunda a. arbutifolia a. floribunda a. floribunda a. melanocarpa2 a. mitschurinii a. melanocarpa2 a. mitschurinii a. floribunda a. floribunda a. melanocarpa4 a. melanocarpa2 a. mitschurinii a. mitschurinii a. mitschurinii a. mitschurinii a. floribunda a. arbutifolia a. arbutifolia a. arbutifolia a. melanocarpa4 a. arbutifolia a. arbutifolia a. arbutifolia a. arbutifolia a. arbutifolia a. melanocarpa a. floribunda a. melanocarpa2 a. melanocarpa4 a. melanocarpa4 a. melanocarpa4 a. melanocarpa4 fig. 3. cluster analysis (binary distance, ward clustering method) of the occurrence of eight basic rbcl and its2 haplotypes. european journal of taxonomy 570: 1–14 (2019) 6 consensus configuration, values of principal relative and partial warps (which characterize the degree of differences between the specimen and consensus configuration), were calculated with the r geomorph package (adams & otarola-castillo 2013) which implements tps analysis in a way similar to principal component analysis (pca). principal component analysis of the relative warps matrix was employed for the classification of leaf shapes, similarly to the way described elsewhere (shipunov & bateman 2005). many statistical methods have the ability to combine different types of data, and we used this to study the joint matrix that includes morphological characters and haplotype data of exactly the same samples (similar to shipunov et al. 2004). we employed model-based clustering (scrucca et al. 2016) and t-sne algorithm (t-sne is ‘t-distributed stochastic neighbor embedding’) (van der maaten & hinton 2008). the latter is a nonlinear dimensionality reduction algorithm, popular for representing high-dimensional data. more simple clustering of haplotype occurrence data used binary distances and ward linkage. all statistical analyses were performed in r (r core team 2018). results dna sequencing and molecular analysis most of the analyses were based on its2 and rbcl alignments. analysis of the trnl-f and leafy data was more problematic, since they did not amplify well from many of the studied herbarium samples; sorbaronia fallax a−215 leafy2 sorbus aucuparia kt834297 leafy2 aronia mitschurinii s9 leafy2 aronia arbutifolia a−403 leafy2 aronia floribunda a−606 leafy2 aronia melanocarpa4 a−609 leafy2 aronia floribunda a−210 leafy2 aronia floribunda a−707 leafy2 aronia melanocarpa4 a−213 a−610 leafy2 aronia arbutifolia a−603 a−801 leafy2 aronia floribunda a−205 leafy2 aronia melanocarpa4 a−203 a−206 leafy2 aronia melanocarpa2 a−204 leafy2 aronia melanocarpa4 a−408 leafy2 aronia melanocarpa a−804 leafy2 0.02 fig. 4. ml tree from the leafy2 sequences. shipunov a. et al., new data on the phylogeny of aronia 7 therefore, these markers were analyzed separately. the resulting trees always had extremely short branches (fig. 1). bootstrap was high enough (> 97%) to support only the whole aronia group. within aronia, there were no clades receiving high statistical support. within each locus, most of our sequences differ only by several nucleotides. across all sequences analyzed, we found ten haplotypes. we counted only three haplotypes from a. mitschurinii samples whereas the more diverse sets were found in a. melanocarpa and a. arbutifolia (fig. 2). as our phylogeny trees are not highly informative, we employed the cluster analysis of the haplotype occurrence (fig. 3) which revealed the tendency of a. arbutifolia samples to group in one cluster (7 out of 10 are in one group). aronia mitschurinii samples did not form a cluster, and grouped together with the bulk of other aronia. data from our trnl-f sequences were generally in agreement with rbcl data. only two haplotypes were found, one the most frequent and the other specific to a. arbutifolia. we were able to sequence both variants (potential paralogs) of leafy second intron (burgess et al. 2015), and therefore constructed two separate datasets, one for each variant. from × sorbaronia fallax, we amplified the second copy, which was identical to the same copy of sorbus aucuparia (genbank id kt834297). each leafy dataset contained at least one a. mitschurinii sample. in the analyses of both −60 −40 −20 0 20 40 − 3 0 − 2 0 − 1 0 0 1 0 2 0 pc1 p c 2 a. arbutifolia a. floribunda a. melanocarpa a. mitschurinii fig. 5. ordination on the plane of first two principal components from the analysis of morphological data. european journal of taxonomy 570: 1–14 (2019) 8 datasets, a. mitschurinii samples always robustly grouped with a. melanocarpa and did not group with sorbus. this pattern is illustrated on fig. 4 which reflects relationships within the second, most samplerich leafy dataset. morphometric and combined analyses the principal component analysis (pca) of morphometric data (fig. 5) was able to clearly separate the a. mitschurinii samples. samples of other species did not form clusters. in the case of joint matrix (morphological characters + haplotype data), the scarcity of a. mitschurinii data did not allow for robust conclusions, but it is remarkable that these two a. mitschurinii samples occupy a separate branch on the minimum spanning tree and therefore distinctive (fig. 6) from all other aronia. geometric morphometrics was first used to produce the consensus shapes for each of our species. these shapes demonstrated the bending required to make imaginary thin plate splines to converge. figure 7 shows that a. mitschurinii samples require the smaller deformations, whereas a. arbutifolia −60 −40 −20 0 20 40 − 4 0 − 2 0 0 2 0 4 0 6 0 a. arbutifolia a. floribunda a. melanocarpa a. mitschurinii fig. 6. ordination from multidimensional scaling of the gower distance matrix from the combined dataset of haplotypes occurrence and morphology. shipunov a. et al., new data on the phylogeny of aronia 9 and a. floribunda require more modifications of their average shapes. pca ordination of relative warps provided another view on the diversity of our samples. it was similar to the morphology ordination, and separated a. mitschurinii whereas other species were intermixed (fig. 8). we also tried two combined approaches which included geometric morphometrics data together with (1) morphology and (2) morphology + haplotype data from the same samples. pca was not really helpful for the analysis of geometric morphometrics + morphology data, but t-sne (van der maaten & hinton 2008) allowed to arrange the samples in more understandable way (fig. 9); here most of a. mitschurinii samples were clearly separated from the rest of aronia. on the other hand, the model-based clustering (scrucca et al. 2016) returned the clustering model with only one component (i.e., no internal clusters). discussion our phylogenetic analyses show that the overall genetic and morphological diversity within aronia is low, and no clear species boundaries could be detected based on dna data. in our morphological ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● a. arbutifolia ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● a. melanocarpa ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● a. mitschurinii ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● a. floribunda fig. 7. transformation grids of leaf shape from four aronia species, with deformations required to reach the overall average shape. european journal of taxonomy 570: 1–14 (2019) 10 and combined analyses, only a. mitschurinii could be separated with confidence. these results are in contrast to connolly (2014) where four aronia species could be morphologically distinguished. this discrepancy between morphological and molecular data could mean that aronia still awaits more advanced methods of data collection and data analysis. however, considering the diversity of our approaches which include phylogeny trees, haplotype analysis, classic and geometric morphometry and combined integrative analysis, the possible conclusion is that at least some borders within aronia are not expressed as it is typical in neighboring genera (guo et al. 2012, li et al. 2012) but may delimit only subspecific or lower level variation; it is even possible that most of the aronia samples represent one polymorphic species. no clear similarities, morphological or genetic, between any aronia and sorbus aucuparia were discovered. the × sorbaronia fallax sample has the its2 and second copy of leafy second intron identical to sorbus aucuparia, indicative of a hybridization event between sorbus aucuparia and aronia melanocarpa. however, this pattern was not present in our samples of a. mitschurinii. therefore, we believe that the hybrid origin of a. mitschurinii is under question. this hypothesis has an independent support from observations on a. mitschurinii propagation which almost always results in the uniform progeny (vinogradova & kuklina 2014). in addition, whereas morphology alone allows for the separation −0.1 0.0 0.1 0.2 − 0 .1 0 − 0 .0 5 0 .0 0 0 .0 5 0 .1 0 pc1 p c 2 a. arbutifolia a. floribunda a. melanocarpa a. mitschurinii fig. 8. ordination from the principal component analysis of relative warps. shipunov a. et al., new data on the phylogeny of aronia 11 of a. mitschurinii from other species, there is no support of such dissimilarity from molecular data. dna-wise, a. mitschurinii does not differ from the other aronia species. the apparent conflict between aflp (leonard et al. 2013) and our sequencing analysis could be due to over-sensitivity of the first analysis (pelser et al. 2003). it is important to note that leonard et al. (2013) did not show the affinity of a. mitschurinii to the sorbus, they have had only found similarities with × sorbaronia hybrids. it is also possible that a. mitschurinii is a backcross to aronia, and the sorbus genome is largely lost. therefore, we cannot fully exclude the possibility that thorough cloning of multiple markers or highthroughput sequencing, might reveal the traces of others genomes in a. mitschurinii. thus, the next step in aronia phylogeny research should involve more high resolution molecular markers, especially multilocus nuclear data. acknowledgements we are grateful to the curators of cas, huh, jeps, mha, mo, ny, uc and us herbaria for the kind permission to analyze herbarium samples. aronia samples from russia were collected mostly around −4 −2 0 2 4 − 1 0 − 5 0 5 a. arbutifolia a. floribunda a. melanocarpa a. mitschurinii fig. 9. t-sne ordination of the combined morphology + geometric morphometrics data. european journal of taxonomy 570: 1–14 (2019) 12 ‘lake 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[in russian.] smolik m., ochmian i. & smolik b. 2011. rapd and issr methods used for fingerprinting selected, closely related cultivars of aronia melanocarpa. notulae botanicae horti agrobotanici cluj-napoca 39: 276–284. https://doi.org/10.15835/nbha3926268 sun j., shi s., li j., yu j., wang l., yang x., guo l. & zhou s. 2018. phylogeny of maleae (rosaceae) based on multiple chloroplast regions: implications to genera circumscription. biomed research international 2018: 7627191. https://doi.org/10.1155/2018/7627191 taheri r., connolly b.a., brand m.h. & bolling b.w. 2013. underutilized chokeberry (aronia melanocarpa, aronia arbutifolia, aronia prunifolia) accessions are rich sources of anthocyanins, flavonoids, hydroxycinnamic acids, and proanthocyanidins. journal of agricultural and food chemistry 61: 8581–8588. https://doi.org/10.1021/jf402449q thompson j.d., gibson t.j., plewniak f., jeanmougin f. & higgins d.g. 1997. the clustal_x windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. nucleic acids research 25 (24): 4876–4882. https://doi.org/10.1093/nar/25.24.4876 vinogradova y.k. & kuklina a.g. 2014. aronia mitschurinii: from origination to naturalization. geos, moscow. [in russian.] volkova p., kasatskaya s., boiko a. & shipunov a. 2011. stability of leaf form and size during specimen preparation of herbarium specimens. feddes repertorium 121: 219–225. https://doi.org/10.1002/fedr.201000021 zarrei m., stefanovic s. & dickinson t.a. 2014. reticulate evolution in north american black-fruited hawthorns (crataegus section douglasia; rosaceae): evidence from nuclear its2 and plastid sequences. annals of botany 114: 253–269. https://doi.org/10.1093/aob/mcu116 zelditch m.l., swiderski d.l. & sheets h.d. 2012. geometric morphometrics for biologists: a primer. academic press. manuscript received: 26 august 2018 manuscript accepted: 22 august 2019 published on: 23 october 2019 topic editor: frederik leliaert desk editor: connie baak printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. https://doi.org/10.15835/nbha3926268 https://doi.org/10.1155/2018/7627191 https://doi.org/10.1021/jf402449q https://doi.org/10.1093/nar/25.24.4876 https://doi.org/10.1002/fedr.201000021 https://doi.org/10.1093/aob/mcu116 136 european journal of taxonomy 778: 136–137 issn 2118-9773 https://doi.org/10.5852/ejt.2021.778.1573 www.europeanjournaloftaxonomy.eu 2021 · machado m. et al. this work is licensed under a creative commons attribution license (cc by 4.0). c o r r i g e n d u m urn:lsid:zoobank.org:pub:59bfbdda-bd99-42a9-8d4f-a81cb2df8011 kryptochroma: a new genus of bark-dwelling crab spiders (araneae, thomisidae) – corrigendum miguel machado 1,*, rafaela viecelli 2, catherine guzati 3, cristian j. grismado 4 & renato a. teixeira 5 1,2,3,5 laboratório de aracnologia, escola de ciências, pontifícia universidade católica do rio grande do sul (pucrs), porto alegre, rio grande do sul, brazil. 4 división aracnología, museo argentino de ciencias naturales “bernardino rivadavia”, buenos aires, argentina. *corresponding author: machadom.arachno@gmail.com 2 email: rafaviecelli1807@gmail.com 3 email: c.guzati@edu.pucrs.br 4 email: grismado@macn.gov.ar 5 email: renato.teixeira@pucrs.br 1 urn:lsid:zoobank.org:author:9c99df66-7481-4de2-bfc2-fe73d29f8a6e 2 urn:lsid:zoobank.org:author:78ca106b-2cfc-4b8b-8e05-168350da95f8 3 urn:lsid:zoobank.org:author:9a03ff9f-07d8-4f15-9735-fac96fe23015 4 urn:lsid:zoobank.org:author:a1d00976-57ef-417e-ae28-df3bf2bab6b1 5 urn:lsid:zoobank.org:author:ba923a63-8230-40c0-819a-588ee3815ca9 machado m., viecelli r., guzati c., grismado c.j. & teixeira r.a. 2021. kryptochroma: a new genus of barkdwelling crab spiders (araneae, thomisidae) – corrigendum. european journal of taxonomy 778: 136–137. https://doi.org/10.5852/ejt.2021.778.1573 the present corrigendum corrects errors that occured in machado et al. (2021). page 48, in ʻmaterial examinedʼ: the voucher name of the holotype of kryptochroma quadrata machado & viecelli, 2021 should be changed from “ufmg 22673” to “mpeg 22673” page 65, in ʻnoteʼ: the species stephanopis furcillata keyserling, 1880 is not only a senior synonym of the species sidymella multispinulosa (mello-leitão, 1944) but also should be combined to the genus sidymella. therefore, this taxonomic act results in the proposition of sidymella fucillata (keyserling, 1880) comb. nov. https://doi.org/10.5852/ejt.2021.778.1573 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:59bfbdda-bd99-42a9-8d4f-a81cb2df8011 https://orcid.org/0000-0003-3193-9830 https://orcid.org/0000-0002-1260-7964 http://orcid.org/0000-0002-1756-9821 mailto:machadom.arachno@gmail.com mailto:rafaviecelli1807@gmail.com mailto:c.guzati@edu.pucrs.br mailto:grismado@macn.gov.ar mailto:renato.teixeira@pucrs.br http://zoobank.org/urn:lsid:zoobank.org:author:9c99df66-7481-4de2-bfc2-fe73d29f8a6e http://zoobank.org/urn:lsid:zoobank.org:author:78ca106b-2cfc-4b8b-8e05-168350da95f8 http://zoobank.org/urn:lsid:zoobank.org:author:9a03ff9f-07d8-4f15-9735-fac96fe23015 http://zoobank.org/urn:lsid:zoobank.org:author:a1d00976-57ef-417e-ae28-df3bf2bab6b1 http://zoobank.org/urn:lsid:zoobank.org:author:ba923a63-8230-40c0-819a-588ee3815ca9 https://doi.org/10.5852/ejt.2021.778.1573 machado m. et al., kryptochroma: a new genus of thomisidae – corrigendum 137 references machado m., viecelli r., guzati c., grismado c.j. & teixeira r.a. 2021. kryptochroma: a new genus of barkdwelling crab spiders (araneae, thomisidae). european journal of taxonomy 778: 26–70. https://doi.org/10.5852/ejt.2021.778.1565 published on: 18 november 2021 printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. https://doi.org/10.5852/ejt.2021.778.1565 186 european journal of taxonomy 746: 186–187 issn 2118-9773 https://doi.org/10.5852/ejt.2021.746.1329 www.europeanjournaloftaxonomy.eu 2021 · wang w.y. et al. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:66f27603-7c26-4492-aad6-64a41e249d7e revision of the elusive ant genus rhopalomastix (hymenoptera, formicidae, myrmicinae) in thailand based on morphology and dna barcodes, with descriptions of three new species – corrigendum wendy y. wang 1,*, gordon w.j. yong 2 & weeyawat jaitrong 3 1 lee kong chian natural history museum, national university of singapore, 2 conservatory drive, singapore 117377. 2 department of biological sciences, faculty of science, national university of singapore, 16 science drive 4, singapore 117558. 3 natural history museum, national science museum, technopolis, khlong 5, khlong luang, pathum thani, 12120 thailand. * corresponding author: nhmwyw@nus.edu.sg – wywang24@gmail.com 2 email: gordonyongwj@gmail.com 3 email: polyrhachis@yahoo.com 1 urn:lsid:zoobank.org:author:41730b76-d515-42d0-a3be-2788adfdaa8a 2 urn:lsid:zoobank.org:author:3d3752df-3522-4dee-b4a3-7e85ca774263 3 urn:lsid:zoobank.org:author:e456f06f-486f-4bd5-aee6-6dca639c3e27 1 https://orcid.org/0000-0003-0745-4702 3 https://orcid.org/0000-0003-1362-0754 wang w.y., yong g.w.j. & jaitrong w. 2021. revision of the elusive ant genus rhopalomastix (hymenoptera, formicidae, myrmicinae) in thailand based on morphology and dna barcodes, with descriptions of three new species – corrigendum. european journal of taxonomy 746: 186–187. https://doi.org/10.5852/ejt.2021.746.1329 the present corrigendum corrects errors that occurred in wang et al. (2021). page 117, citation: the year of publication should be 2021 instead of 2012: wang w.y., yong g.w.j. & jaitrong w. 2012. revision of the elusive ant genus rhopalomastix (hymenoptera, formicidae, myrmicinae) in thailand based on morphology and dna barcodes, with descriptions of three new species. european journal of taxonomy 739: 117–157. should read: wang w.y., yong g.w.j. & jaitrong w. 2021. revision of the elusive ant genus rhopalomastix (hymenoptera, formicidae, myrmicinae) in thailand based on morphology and dna barcodes, with descriptions of three new species. european journal of taxonomy 739: 117–157. http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:66f27603-7c26-4492-aad6-64a41e249d7e mailto:nhmwyw@nus.edu.sg mailto:wywang24@gmail.com mailto:gordonyongwj@gmail.com mailto:polyrhachis@yahoo.com http://zoobank.org/urn:lsid:zoobank.org:author:41730b76-d515-42d0-a3be-2788adfdaa8a http://zoobank.org/urn:lsid:zoobank.org:author:3d3752df-3522-4dee-b4a3-7e85ca774263 http://zoobank.org/urn:lsid:zoobank.org:author:e456f06f-486f-4bd5-aee6-6dca639c3e27 https://orcid.org/0000-0003-0745-4702 https://orcid.org/0000-0003-1362-0754 https://doi.org/10.5852/ejt.2021.746.1329 wang w.y. et al., rhopalomastix in thailand 187 page 128, distribution: thailand (central, nakhon nayok province) (fig. 16a). should read: thailand (northeast, sakhon nakhon province) (fig. 16a). reference wang w.y., yong g.w.j. & jaitrong w. 2021. revision of the elusive ant genus rhopalomastix (hymenoptera, formicidae, myrmicinae) in thailand based on morphology and dna barcodes, with descriptions of three new species. european journal of taxonomy 739: 117–157. https://doi.org/10.5852/ejt.2021.739.1271 published on: 27 april 2021 printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. https://doi.org/10.5852/ejt.2021.739.1271 1 european journal of taxonomy 712: 1–15 issn 2118-9773 https://doi.org/10.5852/ejt.2020.712 www.europeanjournaloftaxonomy.eu 2020 · insisiengmay o. et al. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e two new species of kaempferia l. (zingiberaceae) from cambodia and lao pdr oudomphone insisiengmay 1,*, mark fleming newman 2 & thomas haevermans 3 1,3 institut de systématique évolution biodiversité (isyeb), muséum national d’histoire naturelle, centre national de la recherche scientifique, école pratique des hautes études, université des antilles, sorbonne université, 57 rue cuvier, cp 39, 75005 paris, france. 1 cabinet of the lao academy of science and technology, ministry of science and technology, 100 building, na haidiew village, chanthabouly district, vientiane capital, lao pdr. 2 royal botanic garden edinburgh, 20a inverleith row, edinburgh eh3 5lr, scotland, uk. * corresponding author: oudomphone_ins@most.gov.la or oudomphone.insisiengmay@mnhn.fr 2 email: mnewman@rbge.org.uk 3 email: thomas.haevermans@mnhn.fr abstract. two new species of kaempferia l. (zingiberaceae), kaempferia nemoralis insis. sp. nov. and kaempferia pascuorum insis. sp. nov., from cambodia and lao pdr are described and illustrated. morphological similarities to their closely related taxa are discussed. kaempferia nemoralis insis. sp. nov. is compared with kaempferia larsenii sirirugsa in its vegetative parts, but distinguished by the following characters: whole plant taller, leaf sheath and young shoot apex green, petiole absent. it differs from kaempferia rotunda l. in its floral parts by the following characters: presence of peduncle, floral tube longer, labellum purple with white line at centre, anther crest obovate, bifid, apex irregularly rounded and ovary glabrous. kaempferia pascuorum insis. sp. nov. is compared with kaempferia larsenii sirirugsa. proposed iucn conservation assessments are also given: kaempferia nemoralis insis. sp. nov. occurs in disturbed, open forest and is assessed as cr, whereas kaempferia pascuorum insis. sp. nov. occurs in short grassland and is assessed as en. keywords. kaempferia nemoralis insis. sp. nov., kaempferia pascuorum insis. sp. nov., cambodia, lao pdr. insisiengmay o., newman m.f. & haevermans t. 2020. two new species of kaempferia l. (zingiberaceae) from cambodia and lao pdr. european journal of taxonomy 712: 1–15. https://doi.org/10.5852/ejt.2020.712 introduction kaempferia l. (zingiberaceae, ginger family) is a genus of perennial herbs. around 40 species are currently accepted (mabberley 2017), although nearly 140 names are listed in the international plant names index (ipni 2020). the genus is distributed from india to southern china and peninsular malaysia. several species have medicinal properties, are used in spiritual rituals, or have culinary uses (e.g., kaempferia galanga l., known as kencur) and are widely cultivated, making it difficult to https://doi.org/10.5852/ejt.2020.712 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ mailto:oudomphone_ins%40most.gov.la?subject= mailto:oudomphone.insisiengmay%40mnhn.fr?subject= mailto:mnewman%40rbge.org.uk?subject= mailto:thomas.haevermans%40mnhn.fr?subject= https://doi.org/10.5852/ejt.2020.712 european journal of taxonomy 712: 1–15 (2020) 2 know their natural ranges. the centre of diversity is undoubtedly in the monsoonal parts of se asia, particularly thailand and its immediate neighbours. south of thailand, the diversity drops sharply with four species reported from peninsular malaysia, only one or two of them native (holttum 1950). to the north, only five species of kaempferia extend to the tropical provinces of china, and are probably more often cultivated or naturalised than native ones (wu & larsen 2000). it has been more than a century since the publication of the most recent monograph of all species of kaempferia (schumann 1904), and the most recent revision of the genus in cambodia, laos and vietnam (gagnepain 1908). revisions of kaempferia for the flora of thailand (t. jenjittikul et al., in prep.) and in cambodia, laos and vietnam (o. insisiengmay et al., in prep.) are under way, and have already revealed considerably more species than were known to gagnepain (1908) who recognised 13 species in cambodia, laos and vietnam. in more recent accounts, nine species were recorded in lao pdr (newman et al. 2007), but an additional six have been described in that country in the last 12 years (koonterm 2008; picheansoonthon & koonterm 2008, 2009b; picheansoonthon 2009; phokham et al. 2013), and more are expected from cambodia and vietnam. twenty-seven species are recognised in the current draft of the flora of thailand revision (ruchisansakul, pers. comm.), including 11 described in the last 9 years (nopporncharoenkul & jenjittikul 2017, 2018; nopporncharoenkul et al. 2020; phokham et al. 2013; picheansoonthon & koonterm 2009a; picheansoonthon 2010, 2011; wongsuwan et al. 2015) and at least three more are to be described. many species in the genus are very poorly known taxonomically, mainly because it is so difficult to make informative specimens from these plants, which usually bear only one, extremely delicate and short-lived flower per plant each day of the flowering period. furthermore, the inflorescence is often held between the leaves and is a complex structure of bracts, bracteoles and a number of flowers at different stages. several species bloom at night and must be collected in darkness. others bloom early in the rainy season, before the leaves appear, so to obtain all parts of the plant, two collections must be made a few weeks apart. therefore, herbarium collections are often of relatively little use to the taxonomist and must be supplemented by living material. morphologically, kaempferia is distinct from other genera of zingiberaceae. it is classified in the subfamily zingiberoideae tribe zingibereae which comprises genera with large, petaloid lateral staminodes, about the same size as the lip. the species of kaempferia are small herbaceous perennial plants, some with two leaves appressed to the ground, others with erect leaves to about 70 cm tall. the strongly zygomorphic flowers consist of the usual parts found in zingiberaceae, an inferior ovary, tubular calyx with three lobes, floral tube with three corolla lobes, a labellum, two lateral staminodes and a single fertile anther with two thecae which hold the style so that the stigma is presented just above the anther. the diagnostic character of the flowers of kaempferia is the deeply bilobed labellum and large lateral staminodes which are often held in a single plane so that the open flower resembles, at first glance, a tetramerous, salverform flower. a thorough account of the history of infrageneric classification of kaempferia is given by kam (1980), and the correct generic name for the african species by burtt (1982). the first classification of the species of kaempferia into infrageneric taxa was made by horaninow (1862) who described two taxa, kaempferia [unranked] soncorus and kaempferia [unranked] protanthium. the first of these was described as ‘flores centrales’ and the second as ‘flores praecoces, ante folia e caudice projecti’ which is to say that species in the soncorus group produce their inflorescences terminally on the leafy shoots while those in the protanthium group produce them early in the growing season, before the leaves, arising from the rhizome. horaninow placed kaempferia galanga l. with seven other species in his soncorus group and k. rotunda l. with two other species in his protanthium group. insisiengmay o. et al., two new species of kaempferia 3 bentham & hooker (1883) gave horaninow’s unranked infrageneric taxa the rank of section and added a third, kaempferia section stachyanthesis benth. & hook.f., in which they placed kaempferia scaposa (nimmo) benth. (= curcuma scaposa (nimmo) škorničk. & m.sabu) and kaempferia rosea schweinf. ex baker (= siphonochilus kirkii (hook.f.) b.l.burtt). baker (1890) raised these sections to subgenera and added a fourth, kaempferia subgenus monolophus (wall.) baker which is now treated as the genus monolophus delafosse, guill. & je.kuhn. finally, schumann (1904) recognised 5 subgenera, adding subgenus cienkowskia k.schum. to contain the african species now placed in siphonochilus. since k. galanga is the type species of the genus, kaempferia [unranked] soncorus is illegitimate; it must be called kaempferia [unranked] kaempferia (turland et al. 2018, art. 22.2). insisiengmay et al. (2018) were only able to locate a single element of original material of k. rotunda, and it did not match the written description in the protologue, so a proposal to conserve the name with a conserved type was made. this proposal, number 2581, has been recommended by the nomenclature committee for vascular plants (applequist 2020). during fieldwork in cambodia, laos and vietnam in 2016 and 2017, directed towards a revision of kaempferia in these three countries, we found additional undescribed taxa. here, we describe two new species, kaempferia nemoralis insis. sp. nov. and kaempferia pascuorum insis. sp. nov., the first known only in cambodia, the second distributed in southern lao pdr and cambodia. we compare the morphology of these new species to closely related taxa and we also propose a conservation status of each one using iucn criteria. material and methods a revision of kaempferia in cambodia, laos and vietnam formed part of the phd thesis of the first author (insisiengmay 2019). all names applied to species of kaempferia in these and surrounding countries were examined. protologues were gathered and specimens, including types at bk, bkf, bm, e, hnl, k, p, qbg, sing and vnm were studied. herbarium codes follow index herbariorum (thiers, continuously updated). specimens were collected during field expeditions in cambodia, laos and vietnam from april to august 2016 and may to september 2017. while most collections could be determined to an existing taxon, a small number of collections did not match any herbarium material examined. among these collections, were the specimens described as two new species below. each collection included: flowers and inflorescences in ca 70% alcohol, dried herbarium specimens, rhizomes for cultivation and leaf material in silica gel for molecular systematic study. a complete set of herbarium vouchers and the living collections were deposited at the national herbarium, muséum national d’histoire naturelle (p). the species descriptions are based on the spirit material because dried herbarium specimens lack three-dimensional structures and undergo changes in dimensions of parts of the plants which are very important for species description. the botanical terminology follows the kew plant glossary (beentje 2016). iucn conservation assessments of both species have been made using the guidelines for using the iucn red list categories and criteria ver. 3.1 (2012) and ver. 13 (2017). european journal of taxonomy 712: 1–15 (2020) 4 results description of new species order zingiberales griseb. family zingiberaceae martinov subfamily zingiberoideae tribe zingibereae meisn. genus kaempferia l. subgenus kaempferia l. (= soncorus horan.) kaempferia nemoralis insis. sp. nov. urn:lsid:ipni.org:names:77211164-1 figs 1–3, tables 1–2 diagnosis a species of kaempferia subg. kaempferia. vegetative parts similar to k. larsenii by the shape of the leaf blade which is glabrous, parallel-veined and erect but may be distinguished by the following characters: whole plant taller, leaf sheath and young shoot apex green, petiole absent, the flower of kaempferia nemoralis sp. nov. is completely different from that of k. larsenii in orientation, colour and size. fig. 1. kaempferia nemoralis insis. sp. nov. in natural habitat. photograph: oudomphone insisiengmay. http://www.ipni.org/urn:lsid:ipni.org:names:77211164-1 insisiengmay o. et al., two new species of kaempferia 5 inflorescences and flowers similar to those of k. rotunda (kaempferia subg. protanthium) by the presence of white staminodes and size of labellum but distinguished by the following characters: presence of peduncle, floral tube longer, 95 mm long (vs 50–55 mm long in k. rotunda), labellum purple with white line at centre, anther crest obovate, bifid, apex irregularly rounded and ovary glabrous, the vegetative parts differ by the shape and indumentum of the leaf blade, and absence of a petiole. etymology the specific epithet is from the latin ‘nemoralis’, meaning ‘of woods’. the only known locality is in open forest near a small stream. material examined type cambodia • prov. kratié, dist. snuol, sre roneam village, khseum commune; 12°17′ n, 106°25′ e; alt. 74 m; o. insisiengmay et al. oi 234; 9 jul. 2017; holotype: p (dried and spirit coll.); isotype: e (spirit coll. only). characters kaempferia larsenii kaempferia nemoralis sp. nov. leaves whole plant height 8–10 cm tall 8–20 cm tall leafless sheath colour red green young shoot colour red green longest leaf blade 90 × 10 mm 200 × 20 mm petiole present absent table 1. comparison of the vegetative parts of kaempferia larsenii sirirugsa and kaempferia nemoralis insis. sp. nov. characters kaempferia rotunda kaempferia nemoralis sp. nov. inflorescence peduncle shortly pedunculate 5 mm long floral tube 50–55 mm long 95 mm long staminodes white, purple-tinted, apex acute, pure white, apex rounded, 20 × 16 mm 40 × 12 mm labellum (whole) light purple, 40 × 20–25 mm purple, 47 × 22 mm crest oblong, 3-lobed, apex acute, obovate, bifid, apex irregularly 9–12 × 3–4 mm rounded, 10 × 6 mm stamen 3 mm long 10 mm long ovary villose glabrous table 2. comparison of the floral parts of kaempferia rotunda l. and kaempferia nemoralis insis. sp. nov. european journal of taxonomy 712: 1–15 (2020) 6 fig. 2. kaempferia nemoralis insis. sp. nov. a. whole plant. b. flower dissected. c. bract and bracteoles. d. calyx. e. calyx apex (back view). f. calyx apex (front view) (dorsal corolla lobe side). g. floral tube with stamen attached. h. anther. i. ovary and epigynous glands. j. ovary cross section. k. ligule. drawing from the type material by agathe haevermans (o. insisiengmay et al. oi 234 ). insisiengmay o. et al., two new species of kaempferia 7 description perennial herb, 8–20 cm tall. rhizome short, horizontal; roots of two kinds; tuberous, 10–15 × 3–5 mm, and filamentous. most individuals with one or two flowering shoots. leaves 2 with 2 leafless sheaths, green; ligule a very small rim at junction of sheaths and blade, < 1 mm long, glabrous. longest leaf blade 200 × 20 mm, oblong, erect, base attenuate, apex attenuate, glabrous, petiole absent. inflorescence terminal, peduncle 5 mm long, flowers 1–3. bracts narrowly elliptic to ensiform, villose, 35 × 7 mm, hyaline, subtending one flower; bracteoles 2 per flower, opposite, narrowly elliptic to ensiform, largest one 27 × 5 mm, diminishing to 25 × 2.2 mm, hyaline, villose. calyx tubular, 50 × 3 mm (not flattened), apex with two longer teeth dorsally, greenish and translucent, glabrous and ciliate at apex; floral tube 95 × 3 mm (not flattened), glabrous, white; dorsal corolla lobe linear-acuminate, 45 × 5 mm, white, glabrous, apex with 5 mm long mucro; lateral corolla lobes linear-acuminate, 40 × 4 mm, white, glabrous, apex acute; lateral staminodes oblong, 40 × 12 mm, white, glabrous; labellum obcordate, 47 × 22 mm, purple with white line at the centre, apex bifid, incision 15 mm, lobes narrowly oblong, emarginate; stamen: filament 10 mm long, thecae 5 mm long, dehiscing by longitudinal slits throughout their length, crest obovate, 10 × 6 mm, bifid, incision 6 mm deep, apex of sublobes irregularly rounded, white, glabrous; epigynous glands 2, colourless, subulate, 12–13 mm long; ovary ovoid, 8 × 3.5 mm, glabrous, trilocular with axile placentation, ovules 3–10 per locule, 1 × 0.5 mm; stigma 1.5 mm long, obcuneiform, curved longitudinally, ostiole ciliate. fruit unknown. distribution and habitat only known from the type locality where the plants were found growing in dry dipterocarp forest, near a stream, in moist, sandy soil at low altitude, 74 m. fig. 3. distribution of kaempferia nemoralis insis. sp. nov. (●) and kaempferia pascuorum insis. sp. nov. (▲). type locality of kaempferia pascuorum insis. sp. nov. (▲). european journal of taxonomy 712: 1–15 (2020) 8 conservation status proposed iucn conservation status: cr. aoo = 4 km2, this species is known only from the type locality which is near a path at the edge of a village where tractors and herds of cattle pass frequently. the area is not protected by law. the number of mature individuals is less than 20. notes other species of kaempferia, not yet determined, were also collected at the type locality of kaempferia nemoralis sp. nov. these species clearly differ from kaempferia nemoralis sp. nov. by having their leaves flat on the ground and by a number of characters of the floral parts. a high-resolution image of the type specimen will be deposited at rupp, the national herbarium of cambodia. normally, a type specimen would be deposited in the country of origin, but there is very little type material of kaempferia nemoralis sp. nov. and no paratypes, so it has been agreed with the curator of rupp that it is better to keep the types at e and p where the conditions for long-term conservation, especially of spirit material, are much better. the collections at the herbaria listed in material and methods were searched thoroughly, but no material of kaempferia nemoralis sp. nov. was discovered. kaempferia pascuorum insis. sp. nov. urn:lsid:ipni.org:names:77211165-1 figs 4–5, table 3 diagnosis belonging to kaempferia subg. kaempferia and most similar to k. larsenii by its habit, size around 6–10 cm tall, leaf blade erect, similar in shape, parallel-veined and glabrous but distinguished by the following characters: leaf sheath and young shoot apex green, staminodes white, labellum white with purple patch, crest flabellate, apex bifid, irregularly rounded, white. etymology this species epithet is derived from the latin ‘pascuorum’ (of pastures), referring to their habitat. material examined type lao pdr • prov. champassak, dist. khong, cambodian-laotian border; 13°56′ n, 106°1′ e; alt. 84 m; o. insisiengmay et al. oi. 116; 16 jun. 2016; holotype: hnl (dried coll. only); isotypes: e (dried coll. only), p (dried and spirit coll.), rupp (dried coll. only). additional material cambodia • prov. stung treng, dist. siem pang, siem pang village, sekong commune; 14°6′ n, 106°22′ e; alt. 71 m; o. insisiengmay et al. oi. 237; 10 jul. 2017; p (dried and spirit coll.), rupp (dried coll. only). lao pdr • champassak province, khong district; 14°5′ n, 105°52′ e; alt. 97 m; o. insisiengmay et al. oi. 112; 17 jun. 2016; p (spirit coll. only) • mounlapamok district, nong nga village, thong nong phue; 14°22′ n, 105°30′ e; alt. 107 m; v. lamxay et al. vl1881; 10 jun. 2009; e, natl. univ. laos, fac. science, sing, vnm. http://www.ipni.org/urn:lsid:ipni.org:names:77211165-1 insisiengmay o. et al., two new species of kaempferia 9 fig. 4. kaempferia pascuorum insis. sp. nov. in natural habitat. photograph: oudomphone insisiengmay. characters kaempferia larsenii kaempferia pascuorum sp. nov. 1. leaves leafless sheath colour red green young shoot colour red green longest leaf blade 90 × 10 mm 200 × 25 mm petiole present absent 2. inflorescence peduncle absent 5–18 mm long flowers 8 1–6 floral tube 55 mm long 75–90 mm staminode colour pink pure white labellum colour pink with white patch at middle white with purple patch at middle anther crest obovate, apex rounded, entire or flabellate, bifid, apex irregularly, slightly crenate, with pink patch white filament absent 2 mm long table 3. morphological comparison of kaempferia larsenii sirirugsa and kaempferia pascuorum insis. sp. nov. european journal of taxonomy 712: 1–15 (2020) 10 fig. 5. kaempferia pascuorum insis. sp. nov. a. whole plant. b. leaves. c. ligule. d. flower. e. flower dissected. f. floral tube with anther attached. g. anther crest from above. h. stamen and stigma. i. calyx. j–k. apex of calyx from two different views. l. bract. m–n. bracteoles. o. ovary and epigynous glands. drawing from the type material by agathe haevermans (o. insisiengmay et al. oi. 116 ). insisiengmay o. et al., two new species of kaempferia 11 description perennial herb, 6–10 cm tall. rhizome short, horizontal; roots of two kinds, tuberous, ca 6–10 × 3–5 mm, and filamentous. most individuals with one flowering shoot. leaves two with two leafless sheaths, 10–50 × 5–20 mm, green; ligule a very small rim at junction of sheath and blade, < 1 mm long, sparsely ciliate; longest leaf blade 200 × 25 mm, narrowly elliptic to narrowly ovate, erect, glabrous, base attenuate, apex attenuate; petiole absent. inflorescence terminal, peduncle 5–18 mm long, flowers 1–6. bracts narrowly elliptic, glabrous, 35 × 4 mm, hyaline, subtending a single flower; bracteoles 2 per flower, opposite, narrowly triangular to subulate, largest one 27 × 1.5 mm, diminishing to 25 × 1 mm, hyaline, glabrous. calyx tubular, 45 × 4 mm (not flattened), glabrous, apex 3-dentate, greenish and translucent; floral tube 75–90 × 4 mm (not flattened), glabrous, white; dorsal corolla lobe linear-acuminate, 40 × 6 mm, white, glabrous, apex with 5 mm long mucro; lateral corolla lobes linear-acuminate, 35 × 5 mm, white, glabrous, apex acute; lateral staminodes oblong, 30 × 10 mm, white, glabrous; labellum obcordate, 35–40 × 15–20 mm, white with purple patch at centre, glabrous, apex bifid, divided to 15–20 mm, lobes emarginate; stamen attached at mouth of floral tube, filament 2 mm long, thecae 4 mm long, dehiscing by longitudinal slits, crest flabellate, 3 × 5 mm to 12 × 6 mm, bifid, irregularly divided to 1–3 mm, white, glabrous; epigynous glands two, subulate, 6–8 mm long; ovary cylindrical, 5 × 3 mm, glabrous, trilocular with axile placentation, ovules 2–6 per locule, 1 × 0.5 mm; stigma 1 mm long, obcuneiform, curved longitudinally, ostiole ciliate. fruit dehiscent irregularly, cylindrical, obovate-oblong, ca 10–15 × 4–7 mm, calyx persistent; mature seeds not seen. distribution and habitat southern lao pdr and cambodia, paddy fields or in very open areas, in sandy soil. conservation status proposed iucn status en b1, b2, a, b(iii). eoo = 480 km2, aoo = 12 km2. this species is only known at three locations near the cambodian-lao border, none of which is protected in law. the main threat in lao pdr comes from agriculture, particularly the creation of pathways to and between fields. the cambodian location is within a built-up area in siem pang town. it may be developed in future. the number of mature individuals found at each location is less than 20. key to species of kaempferia in cambodia and lao pdr 1. flowers appearing before leaves (subgenus protanthium) ............................................................... 2 – flowers and leaves appearing at the same time (subgenus kaempferia) ......................................... 3 2. leaves 2–4; blade oblong, green or purple, pubescent; petiole 10–20 mm long ......... k. rotunda l. – leaves 5–7; blade elliptic, upper surface glabrous, lower pubescent; petiole absent ........................ .................................................................................... k. xiengkhouangensis picheans. & phokham 3. ligule present, > 2 mm long and clearly visible; flowers flat, directed upward ............................... 4 – ligules very small, ≤ 2 mm long or absent; flowers cup-shaped, directed upward or sideward, or flat, directed upward ................................................................................................................................ 8 4. leaves erect, blade oblong .....................................................k. sawanensis picheans. & koonterm – leaves flat on the ground, blade oblong or ovate to orbicular ......................................................... 5 5. lateral staminodes and labellum pink to purple; anther crest pink, apex non bifid ........................... ............................................................................................................k. marginata carey ex roscoe – lateral staminodes white; labellum white with violet patch; anther crest white, apex bifid ............ 6 european journal of taxonomy 712: 1–15 (2020) 12 6. anther crest rectangular, bifid, divided by 1 mm, apex rounded-emarginate ....... k. laotica gagnep. – anther crest elliptic or cuneate-flabellate, bifid, divided to the base ................................................ 7 7. anther crest elliptic, 4 × 4 mm, lobes rounded .............................................................. k. galanga l. – anther crest cuneate-flabellate, 6 × 7 mm, lobes emarginate-acute ........... k. harmandiana gagnep. 8. nyctanthous (night-flowering) .......................................................................................................... 9 – hemeranthous (day-flowering) ....................................................................................................... 10 9. ligule present, < 1 mm long; blade ensiform, coriaceous; lateral staminodes spathulate; stamen sessile ........................................................................................................................ k. fissa gagnep. – ligule absent; blade subulate, soft; lateral staminodes obovate; stamen attached at mouth of floral tube, filament 2 mm long .................................................................................... k. filifolia k.larsen 10. bracteoles two per flower, fused at base, outside of floral tube puberulous ....................................... ............................................................................................. k. gigantiphylla picheans. & koonterm – bracteoles two per flower, opposite, outside of floral tube glabrous ...............................................11 11. ovary puberulous ................................................................................... k. parviflora wall. ex baker – ovary glabrous ................................................................................................................................ 12 12. leaf one, blade flat on the ground, orbicular .................................................. k. siamensis sirirugsa – leaf one to three, blade flat on the ground to erect, oblong to ovate ............................................. 13 13. floral tube ≤ 55 mm long; lateral staminodes obovate ................................................................... 14 – floral tube ≥ 55 mm long; lateral staminodes oblong or obovate or spatulate ............................... 15 14. rhizome short, vertical; lateral staminode and labellum pink, anther crest entire or slightly crenate with pink patch ................................................................................................... k. larsenii sirirugsa – rhizome short horizontal and very slender; lateral staminodes and labellum pure white; anther crest bifid and pure white ....................................................... k. champasakensis picheans. & koonterm 15. floral orientation sideward; floral tube > 90 mm long; staminodes oblong, 40 × 12 mm; anther crest obovate, 10 × 6 mm; ovary 8 × 3.5 mm diameter ..................................... k. nemoralis insis. sp. nov. – floral orientation upward; floral tube ≤ 90 mm long; staminodes oblong or spatulate; anther crest flabellate or narrowly ovate to ovate; ovary 5 × 0.5–3 mm diameter .............................................. 16 16. floral tube 75-90 mm long; staminodes oblong, 30 × 10 mm; anther crest flabellate, 3 × 5 mm to 12 × 6 mm, white; ovary 5 × 3 mm diameter ......................................... k. pascuorum insis. sp. nov. – floral tube 55-62 mm long; staminodes spatulate, 20 × 12–16 mm; anther crest narrowly ovate to ovate, 4–6 × 3 mm; ovary 5 × 0.5 mm diameter .....................k. attapeuensis picheans. & koonterm discussion this paper describes and illustrates two new species of kaempferia, recently discovered in cambodia and lao pdr, adding to the overall biodiversity of these countries. a key is provided to distinguish the two new species from the other species known in these countries. the vietnamese species of kaempferia are not all included in this key because of the number of taxonomic uncertainties which remain. further work is under way to produce a revision of kaempferia in cambodia, lao pdr and vietnam, including a molecular phylogenetic survey of the genus. more field collections are needed to complete our studies of poorly known species and to discover new taxa for which we have not yet obtained complete material. an assessment of the variation of species across their ranges is also required. in conclusion, our taxonomic insisiengmay o. et al., two new species of kaempferia 13 and field work have already enriched botanical knowledge and collections, but more work is urgently needed in cambodia, lao pdr and vietnam which are experiencing a high rate of deforestation and land conversion (lang 2001). acknowledgements the authors wish to thank the staff of the ministry of science and technology, lao pdr, especially mrs keophayvanh douansavanh, general secretary of the cabinet of the lao academy of science and technology and mr phouthasack vichitra, director of the cabinet. also, the staff in science and technology at provincial and district level for all their support and help during our field work in lao pdr. we are grateful to assist. prof. dr vichith lamxay, mr sengmany bouta and miss kobkeo phethsomphou for their help. mrs youleang peou, mr sovanrith nheb and miss kunthea chieb of the department of biology, royal university of phnom penh, and miss sunisa sangvirotjanapat, chulalongkorn university, bangkok assisted us greatly during our cambodian field work. mrs agathe haevermans made the beautiful drawings. the authors wish to thank the franklinia foundation who contributed to this fieldwork through a grant to the muséum national d’histoire naturelle for the flora of cambodia, laos and vietnam. this paper is part of a project (aap3-96) supported by the sud expert plantes developpement durable programme. the royal botanic garden edinburgh (rbge) is supported by the scottish government’s rural and environmental science and analytical services division. references applequist w.a. 2020. report of the nomenclature committee for vascular plants: 71. taxon 69: 391– 397. https://doi.org/10.1002/tax.12217 baker j.g. 1890. scitamineae. in: hooker j.d. 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(eds) flora of china 24: 322–377. science press, beijing & missouri botanic garden press, st. louis. manuscript received: 5 december 2018 manuscript accepted: 10 june 2020 published on: 4 september 2020 topic editor: frederik leliaert desk editor: connie baak printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. european journal of taxonomy 176: 1–14 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2016.176 www.europeanjournaloftaxonomy.eu 2016 · yu d. et al. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:c94c7774-5176-460c-804d-a38520284d4b 1 tomoceridae (collembola, entomobryomorpha) from the southern annamitic cordillera: redescription of tomocerus ocreatus denis, 1948 and description of a new species of tomocerina yosii, 1955 daoyuan yu 1,*, le cong man 2 & louis deharveng 3 1 soil ecology lab, college of resources and environmental sciences, nanjing agricultural university, nanjing 210095, china. 2 department of biology, university of natural sciences, ho chi minh city national university, vietnam. 3 institut de systématique, evolution, biodiversité, isyeb umr 7205 cnrs, mnhn, upmc, ephe, muséum national d’histoire naturelle, sorbonne universités, 57 rue cuvier, cp 50, 75005 paris, france. * corresponding author. dyyu1987@hotmail.com; yudy@njau.edu.cn 2 email: tranletamlinh@yahoo.com.vn 3 email: deharven@mnhn.fr 1 urn:lsid:zoobank.org:author:07321441-2070-4db2-bc59-d3f1c8300bc3 2 urn:lsid:zoobank.org:author:0d7a5067-f6ef-456f-8b32-d1b61dcc4cdd 3 urn:lsid:zoobank.org:author:d8f5c679-c30c-442c-8621-d3b8edb17ef7 abstract. two species of tomoceridae were found near dalat, southern vietnam. tomocerus ocreatus denis, 1948 is redescribed based on a neotype specimen. previous records of tomocerus ocreatus in non-type localities are reevaluated. a new species tomocerina annamitica sp. nov. is described. the new species is mainly characterized by its small body size, pointed tenent hair, compound dental spines and the absence of intermediate teeth on mucro. key words. vietnam, southeast asia, tomocerinae, taxonomy, neotype. yu d., man l.c. & deharveng l. 2016. tomoceridae (collembola, entomobryomorpha) from the southern annamitic cordillera: redescription of tomocerus ocreatus denis, 1948 and description of a new species of tomocerina yosii, 1955. european journal of taxonomy 176: 1–14. http://dx.doi.org/10.5852/ejt.2016.176 introduction the family tomoceridae contains at least 169 known species (bellinger et al. 1996–2015) distributed worldwide. in asia tomoceridae are abundant in the north but have only a few reports in the south. three species were previously described from vietnam: tomocerus ocreatus denis, 1948 from dalat, lam dong province; tomocerus nodentalis nguyen, 1995 from moc chau, son la province; and tomocerus postantennalis yu, zhang & deharveng, 2014 from na hang, tuyen quang province. tomocerus ocreatus is characterized by the shape and arrangement of dental spines. tomocerus nodentalis is very http://dx.doi.org/10.5852/ejt.2016.176 https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:c94c7774-5176-460c-804d-a38520284d4b http://zoobank.org/urn:lsid:zoobank.org:author:07321441-2070-4db2-bc59-d3f1c8300bc3 http://zoobank.org/urn:lsid:zoobank.org:author:0d7a5067-f6ef-456f-8b32-d1b61dcc4cdd http://zoobank.org/urn:lsid:zoobank.org:author:d8f5c679-c30c-442c-8621-d3b8edb17ef7 http://dx.doi.org/10.5852/ejt.2016.176 european journal of taxonomy 176: 1–14 (2016) 2 peculiar for the absence of dental spines, which character is only observed in very early instars of other species (goto 1956; uchida & chiba 1958). tomocerus postantennalis is a cave species with developed postantennal organs in adult stage. to the present knowledge, tomocerus nodentalis and tomocerus postantennalis are both endemic species, whereas tomocerus ocreatus is known to be the most widely distributed asian species of tomocerus nicolet, 1842. besides its type locality, tomocerus ocreatus has records from himalaya (yosii 1966), japan (yosii 1956, 1967, 1969, 1977; chiba 1968), korea (yosii & lee 1963; lee 1974, 1975), china (stach 1964; liu et al. 1998), northern vietnam (stach 1965), eastern russia (sakhalin island) (martynova 1977), etc. but intraspecific differences were often reported by descriptions of non-type tomocerus ocreatus. for instance, chiba (1968) distinguished three varieties from japanese tomocerus ocreatus according to the variation in the body size and relative length of body parts. a molecular study (zhang et al. 2014) also revealed “cryptic” diversity among chinese specimens previously identified as tomocerus ocreatus. to clarify the true status and distribution of tomocerus ocreatus, a redescription of its type specimen is required, but the holotype and only specimen of the species has been lost in the muséum national d’histoire naturelle (paris) where the denis’ collection is currently stored. in a recent sample collected from dalat (lam dong province) in 2008, we have found a specimen that matches well the original description in morphology. we designate this specimen as neotype of tomocerus ocreatus. a new tomocerid species was found in the same sample as tomocerus ocreatus. we assign the new species to tomocerina yosii, 1955 because it has no toothlet on the outer basal mucronal tooth. to our knowledge, tomocerina annamitica sp. nov. is so far the southernmost record of the genus. materials and methods specimens were collected with aspirators or berlese funnels. photographs were taken under a jenoptik progres c10+ camera mounted on a leica mz 16 stereomicroscope. specimens were cleared in lactic acid and mounted in hoyer’s solution (krantz 1978). for some specimens the head, the furca and the legs were cut off from the trunk and mounted separately for detailed observation. the slide-mounted specimens were studied using a leica dmlb microscope or a nikon 80i microscope. we follow fjellberg (2007) for maxillary lamellae numbering, yu et al. (2014) for the pattern of cephalic dorsal chaetotaxy and christiansen (1964) for body macrochaetotaxy. the description of the body chaetotaxy refers to one side only since in most case it is symmetric. the exact morphology of each chaeta was uncertain due to shedding. the dental spines formula follows that of folsom (1913), in which the dental spines are arranged from basal to distal, with a slash indicating the separation between basal and medial subsegments and the roman numerals referring to spines that are noticeably larger. if not mentioned specially, all descriptions are based on fully developed individuals. abbreviations ant. = antennal segment pao = postantennal organ th. = thoracic segment abd. = abdominal segment institutional acronyms: hcmcu = ho chi minh city national university, vietnam mnhn = muséum national d’histoire naturelle, paris, france njau = nanjing agricultural university, nanjing, china yu d. et al., tomoceridae from southern annamitic cordillera 3 results class collembola lubbock, 1873 order entomobryomorpha börner, 1913 superfamily tomoceroidea szeptycki, 1979 family tomoceridae börner, 1913 subfamily tomocerinae schäffer, 1896 genus tomocerus nicolet, 1842 diagnosis moderate to large sized tomocerinae, usually longer than 3 mm; body colour pale to dark, some species with distinct colour pattern; eyes at most 6+6; trochantero-femoral organ reduced to 1, 1 chaetae; dens of furca basally without outer strong chaetae or inner large differentiated scales; shape of dental spines from simple to compound among different species; mucro with two dorsal lamellae and two basal teeth, outer basal tooth with corner toothlet. commonest group of tomocerinae in eurasia continent, especially abundant in east asia. tomocerus ocreatus denis, 1948 figs 1a, 2, 3 diagnosis typical tomocerus species with pale body colour, moderately long antennae and full set of 6+6 eyes. head without distinct pao; th. ii with relatively reduced number of macrochaetae; tenent hair clavate, moderately developed; unguis with 5–6 teeth; tenaculum unscaled, with numerous chaetae; manubrium without dorsal scales and blunt prominent chaetae; dental spines compound with numerous moderate sized denticles; no small dental spine between two distal large spines; mucro with 9–10 intermediate teeth. neotype vietnam: ♀, on slide. collected in hon giao, bi doup massif, northeast of dalat, lam dong province, 108°42’53”e, 12°11’11’’n, alt. 1630 m, 12 jun. 2008, by louis deharveng & anne bedos (sample code vn08-150). deposited in mnhn (specimen vn08-150_to1). description body length 3.6 mm. ground colour uniformly yellowish white. ant. iii+iv, antero-ventral part of head, coxae and tibiotarsi with dark pigment. eye patches black. scales brown (fig. 1a). body densely clothed by scales and various types of chaetae. scales of typical morphology of tomocerinae, with continuous longitudinal ridges on surface (lubbock 1873). ordinary chaetae of different sizes. microchaetae smooth and pointed. macrochaetae and mesochaetae from slightly to strongly ciliated, some slightly ciliated mesochaetae appearing to be smooth under optical microscope. most macrochaetae straight, rod-like and subcylindrical, some macrochaetae on posterior abdominal segments long, curved and acuminate. mesochaetae acuminate, shorter and thinner than macrochaetae. s-chaetae subcylindrical, more hyaline than ordinary chaetae, as small as microchaetae except long ones on abd. iv. dorso-inner chaetae on dens modified as strong pointed spines. pseudopores as small circular structures similar to chaetae sockets, distributed at least on th. ii to abd. iv, coxae, and manubrium. pao not seen. eyes 6+6. antennae nearly as long as body. antenna length ratio as i:ii:iii+iv= 1.0:1.7:15.4. ant. i and ant. ii dorsally scaled, ant. iii+iv unscaled. prelabral and labral chaetae (labral european journal of taxonomy 176: 1–14 (2016) 4 formula) 4/5, 5, 4, the distal 4 chaetae stronger. distal edge of labrum with four curved spine-like papillae (fig. 2a), and a well developed ventro-distal brush. mandibular head asymmetrical, the left one with 4 teeth (including a basal rounded one) and the right one with 5, left molar plate distally with a tapered tooth (fig. 2b). basal teeth of maxillary lamella 5 slightly longer than apical ones, without beard-like appendage (fig. 2c). maxillary outer lobe with trifurcate palp, one basal chaeta and 4 sublobal hairs (fig. 2d). both dorsal and ventral sides of head scaled. cephalic dorsal macrochaetotaxy: anterior area: 2, 2; interocular area: 2, 6, central uneven macrochaeta absent; postocular area: 2+2; posterior area: 2. posterior margin of head with about 20+20 small chaetae (fig. 2e). mentum with 5 chaetae, submentum with numerous chaetae. pattern of body chaetotaxy as in fig. 2f. bothriotricha 2, 1/0, 0, 1, 2, 0, 0 on th. ii–abd. vi, respectively, as typical in tomocerinae. macrochaetae densely arranged along anterior margin of th. ii (not shown in figure). th. ii with a row of macrochaetae behind anterior margin. number of macrochaetae or large mesochaetae in the posterior row as 3, 3/3, 3, 4, 2, 4 (3 dorsal+1 lateral) from th. ii to abd. v. on th. ii no macrochaeta near the pseudopore contrary to most other tomocerids; on abd. iii s-microchaeta and accompanying microchaeta posterior to lateral macrochaeta; abd. iv with one lateral macrochaeta and numerous long s-chaetae; on abd. v inner macrochaeta smaller than others in posterior row; abd. vi with numerous chaetae of moderate size. most mesochaetae laterally and posteriorly on terga. pseudopores near the axis of terga, 1, 1/1, 1, 1, 1, 0, 0 from th. ii to abd. vi. legs with numerous ordinary chaetae. trochantero-femoral organ with 1, 1 slender chaetae. front, middle and hind tibiotarsus dorsally with 1, 1, 3 long prominent chaetae, ventrally with 4–5, 5, 6 blunt spine-like chaetae (fig. 2g). each tibiotarsus with a distal whorl of 11 chaetae, ventral 6 as ordinary chaetae, dorsal 5 modified: tenent hair clavate, as long as inner edge of unguis; 2 accessory chaetae extremely minute; 2 guard chaetae thin and long, slightly shorter than tenent hair (fig. 3a, b). unguis fig. 1. appearance of tomocerus ocreatus denis, 1948 and tomocerina annamitica sp. nov. in alcohol. a. tomocerus ocreatus (lateral view). b. tomocerina annamitica sp. nov. (lateral view). scale bar: 1 mm. yu d. et al., tomoceridae from southern annamitic cordillera 5 slender, with baso-internal ridging visible in lateral view; lateral teeth pointed, of moderate size. inner edge of unguis with 5–6 teeth, the basal tooth smaller, the other subequal in size. unguiculus about half as long as unguis, its inner edge with 1 tooth. pretarsal chaetae 1+1, much larger than tenent-hair accessory chaetae (fig. 3b). fig. 2. tomocerus ocreatus denis, 1948. a. labrum (dorsal view). b. mandibular head (dorsal view). c. lamella 5 of maxillary head (dorsal view). d. maxillary outer lobe (ventral view). e. cephalic dorsal chaetotaxy (circles = sockets of chaetae, same as below; 1 = anterior area; 2 = interocular area; 3 = postocular area; 4 = posterior area). f. dorsal chaetotaxy of th. ii–abd. vi (circle with a slash = pseudopore, same as below). g. right tibiotarsi (lateral view). scale bars: a, b, g = 100 μm; c = 10 μm; d = 50 μm; e, f = 500 μm. european journal of taxonomy 176: 1–14 (2016) 6 ventral tube with scales on both anterior and posterior faces, lateral flap unscaled, anterior face with about 15 chaetae on each side, posterior face with about 55 chaetae, each lateral flap with 45–47 chaetae. rami of tenaculum with 4+4 teeth, anterior face with 13 small chaetae and without scale (fig. 3c). furca ratio manubrium: dens: mucro=2.9–3.1:4.1–4.4:1.0. manubrium ventrally scaled, without chaetae, laterally with large round scales and 11 chaetae, proximal 2 chaetae small and slender, distal 9 distinctly stronger and serrated; dorsal scales absent; each dorsal chaetal stripe with about 90 chaetae in different sizes, without blunt chaetae; pseudopores 11–12 on each side (fig. 3d); external corner chaeta as large as small mesochaetae in chaetal stripe (fig. 3e). dens basally without inner modified scale or outer fig. 3. tomocerus ocreatus denis, 1948. a. diagram of tibiotarsal distal chaetae (distal view; t = tenent hair; a = accessory chaeta; g = guard chaeta; p = pretarsal chaeta, same as below). b. hind claw (lateral view). c. tenaculum (anterior view). d. left side of manubrium (dorsal view). e. disto-external corner of manubrium (dorsal view). f. basal and middle subsegments of dens with dental spines (dorsal view). g. mucro (dorso-inner view). scale bars: b, c, e, g = 50 μm; d, f = 100 μm. yu d. et al., tomoceridae from southern annamitic cordillera 7 strong chaetae. dental spines formula as 3/4, ii, distal spine strongest, about 0.1 times as long as dens; all spines with numerous denticles of moderate size (fig. 3f). dens dorsally with ordinary chaetae and feather-like chaetae as typical in tomocerinae, ventrally with only scales. mucro elongated, bearing numerous smooth chaetae with elongated sockets; both basal teeth with proximal lamellae, the outer tooth with a toothlet; apical and subapical tooth subequal; structure of dorsal lamellae of tomocerus type, two dorsal lamellae running from subapical tooth, outer lamella ending in inner basal tooth, inner lamella ending freely at base of mucro; outer lamella with 9–10 subequal intermediate teeth (fig. 3g). remarks denis (1948) provided accurate description for tomocerus ocreatus. for instance, he described clearly the dental spines as “covered with secondary spines, more similar to scales than denticles”, and the figure showed that those scale-like denticles were of moderate size and covered at least basal half of each spine, on which account we rejected some records of non-type tomocerus ocreatus (see below). but naturally, some later revealed characters were not mentioned by him. for instance, chaetotaxy had not been used for taxonomy in tomocerinae before yosii (1956). the limit of original description is a main reason for the misidentification of tomocerus ocreatus in some subsequent records, and finally turned the species into a complex including a number of closely related forms from vietnam to eastern russia. to overcome the deep confusion about this species, a redescription of type specimen is necessary. because the holotype and unique type specimen of tomocerus ocreatus was lost, we propose to set up a neotype for the species from specimens of the type locality. denis (1948) described tomocerus ocreatus on one specimen, from “plateau du lang biang, 2400 m. alt., forêt tropicale”. it is not the lang bian peak itself that reaches 2400 m but the chu yang sin massif north of the lang bian (= dalat) plateau, the lang bian peak being only 2197 m in altitude. the chu yang sin massif was and still is of very difficult access, and was certainly not the place where the collector, dawydoff, operated. there are two main patches of subtropical forests around dalat: a small one on the lang bian peak, and a much larger one 25 km northeast on the bi doup massif, which reaches 2287 m and is less easy to access. in any case, it is most likely on the slopes of one of these massifs that tomocerus ocreatus was collected. on this account, we propose to designate a specimen that we obtained from litter sample in the bi doup mountain as the neotype. the neotype is highly identical with the original description in almost all described characters, including the body colour, the length of antennae, the structure of claws and the morphology and arrangement of dental spines. the only difference is that in the neotype specimen the numbers of ungual teeth, dental spines and mucronal teeth are slightly larger than in the original one. it could happen in tomoceridae that within the same species larger individuals have a few more teeth and dental spines than smaller ones, so the slight difference mentioned above can be treated as intraspecific considering the neotype specimen is slightly larger than the lost holotype (3.6 mm versus 3.3 mm). regarding previous non-type records of tomocerus ocreatus, stach (1964, 1965) redescribed two different forms of tomocerus ocreatus from southeastern china and northern vietnam, respectively, and also claimed that the japanese record of “tomocerus minor” by uchida (1953) was actually tomocerus ocreatus; yosii described a japanese species tomocerus kawamurai (yosii, 1954) and then synonymized it with tomocerus ocreatus (yosii 1956, 1967); chiba (1968) recognized three forms in japanese tomocerus ocreatus; lee (1975) made descriptive notes on korean specimens; martynova (1977) recorded tomocerus ocreatus in sakhalin and made some descriptive notes on it. these redescriptions had subtle to considerable differences to each other, and were never identical to the original description. the chinese “ocreatus” recorded in hangzhou (“hangchow”), zhejiang province by stach (1964) has brownish body colour and distinctly short antenna about half the length of body; the northern vietnamese record from lao cai province by the same author (stach 1965) bears a small dental spine between two large distal spines, and the denticles european journal of taxonomy 176: 1–14 (2016) 8 on spines are finer than those in the original description, therefore it is more similar to tomocerus folsomi denis, 1929 from yunnan province, china; the japanese “ocreatus” described by yosii (1967) has dorsal scales and 2+2 blunt principal chaetae on manubrium which are absent in the true tomocerus ocreatus; other records (chiba 1968; lee 1975; martynova 1977) all have dental spines with one or two whorls of crownlike denticles near the base which is not the morphology of the true ocreatus form. but these remarkable differences were treated as intraspecific variations when other characters showed high similarity. inferred from the present morphological review and the molecular study on chinese ocreatus complex (zhang et al. 2014), the aforementioned non-type “ocreatus” may each represent an independent species in the complex, and so far the true tomocerus ocreatus is known as only endemic to southern annamitic range in vietnam. genus tomocerina yosii, 1955 diagnosis small to moderate sized tomocerinae, usually less than 2 mm in length; body colour pale to light grey, seldom dark; antennae usually much shorter than body; eyes at most 6+6; trochantero-femoral organ with 1, 1 or more chaetae; dens of furca basally without outer strong chaetae or inner large differentiated scales; shape of dental spines from simple to compound; mucro with two dorsal lamellae and two basal teeth, outer basal tooth without corner toothlet. widely distributed in northern hemisphere. tomocerina annamitica sp. nov. urn:lsid:zoobank.org:act:9ad947ec-5a05-4385-9a92-8b0dc590a94d figs 1b, 4, 5 diagnosis tomocerina species with typical body size and pigmentation; antennae about 0.8 times as long as body, relatively long for tomocerina; th. ii with only one bothriotrichum; tibiotarsi with multiple strong chaetae; tenet hair small and pointed; unguis with 1–2 inner teeth; dental spines compound with denticles; only one distal spine distinctly larger; mucro without intermediate tooth. etymology specific name derived from its type locality: the southern annamitic range. type material holotype vietnam: ♀, on slide. collected in hon giao, bi doup massif, northeast of dalat, lam dong province, 108°42’53”e, 12°11’11’’n, alt. 1630 m, mixed forest litter, 12 jun. 2008, by louis deharveng & anne bedos (sample code vn08-150), deposited in mnhn (specimen vn08-150_ta1). paratypes vietnam: 2 ♀♀, on slides, same data as holotype, 1 in hcmcu (specimen vn08-150_ta2) and 1 in njau (specimen vn08-150_ta3). other material examined vietnam: 5 specimens on slides and 28 specimens in alcohol, collected in the hon ba nature reserve, kanh hoa province, located 25 km east-southeast of hon giao and 25 km south-southwest of nha trang, 108°56’55’’–108°58’51’’e, 12°06’35’’–12°07’09’’n, mixed forest litter, nov. 2013, by louis deharveng & anne bedos: alt. 850 m, vn13-072 (3 in alcohol); 890 m, vn13-126 (2 in alcohol), vn13135 (4 in alcohol); 1050 m, vn13-054 (2 ♀♀, on slides), vn13-057 (1 ♂ and 1 ♀, on sildes), vn13-114 (1 on slide, sex undetermined), vn13-120 (8 in alcohol); 1350 m, vn13-039 (5 in alcohol); 1500 m, vn13-231 (6 in alcohol). http://zoobank.org/urn:lsid:zoobank.org:act:9ad947ec-5a05-4385-9a92-8b0dc590a94d yu d. et al., tomoceridae from southern annamitic cordillera 9 description body length 1.6–1.8 mm. body colour light grey, antennae light purple, eye patches black, scales brown (fig. 1b). clothing of tomocerinae type, similar to that of tomocerus ocreatus. pao not seen. eyes 6+6. antenna about 0.75–0.90 times as long as body. antenna length ratio i:ii:iii: iv=1.0:1.3–1.5:5.5–6.0:2.2–2.5, ant. i and ant. ii dorsally scaled, ant. iii and ant. iv unscaled. labral formula 4/5, 5, 4, the distal 4 chaetae stronger. distal edge of labrum with four curved spine-like papillae, ventro-distal brush well developed. mandibular head asymmetrical, the left one with 4 teeth (including a fig. 4. tomocerina annamitica sp. nov. a. apical half of mandibular head (dorsal view). b. lamella 5 of maxillary head (dorsal view). c. maxillary outer lobe (ventral view). d. cephalic dorsal chaetotaxy. e. dorsal chaetotaxy of th. ii–abd. vi. f. left tibiotarsi (lateral view, b = blunt inner chaeta). scale bars: a, c = 50 μm; b = 10 μm; d, e = 250 μm; f = 100 μm. european journal of taxonomy 176: 1–14 (2016) 10 basal rounded one) and the right one with 5, left molar plate distally with tapered tooth (fig. 4a). basal teeth of maxillary lamella 5 elongated without beard-like appendage (fig. 4b). maxillary outer lobe with trifurcate pulp, basal chaeta and 4 sublobal hairs (fig. 4c). cephalic dorsal chaetotaxy: anterior area: 2, 4; interocular area: 2, 6, central chaeta absent; postocular area: 2+2; posterior area: 2; posterior margin: about 20+20 small chaetae of different sizes (fig. 4d). both dorsal and ventral side of head scaled. mentum with 5 chaetae, submentum with numerous chaetae. pattern of body chaetotaxy as fig. 4e. 1, 1/0, 0, 1, 2, 0, 0 bothriotricha on th. ii–abd. vi. macrochaetae densely along anterior margin of th. ii (not shown in figure). th. ii with a row of macrochaetae behind anterior margin. number of macroor large mesochaetae in the posterior row as 3, 3, 3, 3, 3, 5, 4 (3 dorsal+1 lateral) from th. ii to abd. v. on th. ii pseudopore near posterior macrochaeta of the 3 central ones; th. iii with anterior macrochaeta; on abd. iii s-microchaeta posterior to lateral macrochaetae; abd. iv with antero-lateral macrochaeta and numerous long s-chaetae, 4 inner mesochaetae in posterior fig. 5. tomocerina annamitica sp. nov. a. diagram of tibiotarsal distal chaetae (distal view). b. tibiotarsal distal chaetae (dorsal view). c. hind claw (lateral view). d. tenaculum (anterior view). e. left side of manubrium (dorsal view). f. disto-external corner of manubrium (dorsal view). g. basal and middle subsegments of dens with dental spines (dorsal view, 1 = ventral side of spine, showing only distal serrations; 2 = dorsal side of spine, showing fine longitudinal ribs; s = small inner scale). h. mucro (inner view). scale bars: b–d = 20 μm; e–h = 50 μm. yu d. et al., tomoceridae from southern annamitic cordillera 11 row prominent but smaller than outer macrochaeta; on abd. vi dorsal flap with 2+2 and lateral flap with 3 macrochaetae. most mesochaetae laterally and posteriorly on terga. pseudopores near the axis of terga, 1, 1/1, 1, 1, 1, 0, 0 from th. ii–abd. vi. legs with numerous ordinary chaetae. trochantero-femoral organ with 1, 1 chaetae. tibiotarsi dorsally with 1, 1, 3 prominent chaetae from front to hind leg, ventrally with 5–7, 4, 5–6 strong chaetae, only 2, 1, 2 of them blunt (fig. 4f). each tibiotarsus with a distal whorl of 11 chaetae, the dorsal 5 modified: tenent hair pointed, short and thin; 2 accessory chaetae about as long as but thicker than tenent hair, and longer than pretarsal chaetae; 2 outer guard chaetae strong, as long as inner edge of unguis (fig. 5a–c). scales absent from front and mid tibiotarsi but present on hind tibiotarsus. unguis slender, lateral teeth of moderate size, internal ridging present. inner edge of unguis with a small basal and 0–1 larger more distal teeth. unguiculus about half as long as unguis, without tooth on inner or outer edge. pretarsus with 1+1 chaetae (fig. 5c). ventral tube anteriorly with a few small scales, each side with 20–25 chaetae; posteriorly unscaled, with about 40 chaetae; lateral flap unscaled, each side with about 25 chaetae. tenaculum with 4+4 teeth, unscaled, anterior face with 1 chaeta about as long as rami (fig. 5d). ratio manubrium: dens: mucro= 2.0–2.2:2.9–3.2:1.0. manubrium ventrally with only scales, laterally with scales and 8 chaetae, proximal 2 chaetae small and slender, distal 6 chaetae distinctly stronger; dorsal scales absent; each dorsal chaetal stripe with about 70 chaetae in different sizes, without blunt chaetae; pseudopores 5–6 on each side (fig. 5e); external corner chaeta as large as small chaetae in chaetal stripe (fig. 5f). dental spines formula 4/3, i, a small pointed scale present ventrally to basal spines; all spines with several large denticles near base, superficial texture different between dorsal and ventral sides: dorsally with only fine longitudinal ribs, ventrally with longitudinal ribs and distal tiny serrations (fig. 5g). the largest spine about 0.1 times as long as dens. dens dorsally with feather-like chaetae between ordinary chaetae. stripe of feather-like chaetae starting from centre of middle subsegment of dens, ending a short distance to the apex of dens. ventral side of dens covered by scales, several chaetae present apically. mucro elongated and multisetaceous, with two dorsal lamellae; both basal teeth with proximal lamellae, outer tooth without toothlet; apical tooth elongated, stronger than subapical tooth; intermediate teeth absent (fig. 5h). remarks among tomocerina, the new species is similar to those of the minuta group in its small body size, grey body colour and shape of mucro, but is different from them mainly in the chaetotaxy and the shape of dental spines. it is similar to tomocerina purpurithora liu, hou & li, 1999 from sichuan province, china in the shape of dental spines, but is different from the latter mainly in the cephalic chaetotaxy, the number of tibiotarsal strong inner chaetae, the number of chaetae on tenaculum, the dental spines formula and the number of teeth on mucro (table 1). tomocerina annamitica sp. nov. is the southernmost distributed species of tomocerina, a genus mostly diversified in northern temperate to cold temperate regions. however, so far tomocerina is poorly defined by only a minute character: the absence of corner toothlet on outer basal tooth of mucro. in this respect, the presence of only 1+1 bothriotricha on th. ii of tomocerina annamitica sp. nov. might provide an interesting character, while most other species of tomocerinae have 2+2. investigation on this character in different groups of tomocerinae is in progress to assess its phylogenetic value. table 1. discrimination between tomocerina annamitica sp. nov. and tomocerina purpurithora liu, hou & li, 1999. species number of tibiotarsal strong inner chaetae number of chaetae on tenaculum dental spines formula number of intermediate teeth on mucro t. annamitica sp. nov. 5–7, 4, 5–6 1 4/3, i 0 t. purpurithora 0, 0, 1 5 5/7, 1 5–7 european journal of taxonomy 176: 1–14 (2016) 12 discussion tomoceridae are mostly distributed in the northern temperate areas, with a diversity peak in eastern palearctic: japan (yosii 1967), korea (lee 1975), russian far-east (martynova 1977) and china (ma 2004). the diversity rapidly decreases towards southern regions, but this family is still present in the caves or at high altitude of the subtropics (southern china and vietnam, denis 1929; yu et al. 2014; yu & deharveng 2015; nepal, yosii 1966), or even at high altitude in tropical regions (sumatra, oudemans 1891; vietnam, denis 1948). before this study, all three species of tomoceridae known from vietnam belong to tomocerus. the fourth species added here, tomocerina annamitica sp. nov., belongs to another genus which had so far a single species distributed south of 30°n (tomocerina simplex yosii, 1966 from nepal). unexpectedly, the new species was frequent in litter and soil as low as 1000 m of elevation. besides tomocerus ocreatus and tomocerina annamitica sp. nov., at least one other species is probably present in our collections from southern annamitic cordillera, confirming its status of southern extension for palaearctic fauna of springtails (deharveng & bedos 2000). acknowledgements we thank dr. anne bedos from mnhn paris who helped to collect specimens in lang bian, bi doup and hon ba, truong quang tam and nguyen tran vy from institute of tropical biology of ho chi minh city who organized the field trip and the staff of the bi doup-nui ba national park. field work in hon ba was carried out in the frame of a project with the nature reserve, and supported by a french grant of the agence nationale de la recherche under the labex anr-10-labx-0003-bcdiv. laboratory work in njau was supported by the national natural science foundation of china (41501056), the fundamental research funds for the central universities (kjqn201668) and the scientific grant for post-doctors of jiangsu province (1402054c). references bellinger p.f., christiansen k.a. & janssens f. 1996–2015. checklist of the collembola of the world [online]. available from http://www.collembola.org [accessed 20 mar. 2015] chiba s. 1968. collembola of the mt. hakkoda area. i. family tomoceridae. the science reports of the hirosaki university 15: 24–26. christiansen k. 1964. a revision of the nearctic members of the genus tomocerus (collembola: entomobryidae). revue d’écologie et de biologie du sol 1: 668–675. deharveng l. & bedos a. 2000. vietnura caerulea new genus, new species, from vietnam: first record of the palaearctic tribe neanurini in tropical asia (collembola: neanuridae). the raffles bulletin of zoology 48 (2): 209–214. denis j.r. 1929. notes sur les collemboles récoltés dans ses voyages par le prof. f. silvestri. bollettino del laboratorio di zoologia generale e agraria della r. scuola superiore d’agricoltura in portici 22: 166–180. denis j.r. 1948. collemboles d’indochine. notes d’entomologie chinoise 12: 183–311. fjellberg a. 2007. the collembola of fennoscandia and denmark. part ii. entomobryomorpha and symphypleona. fauna entomologica scandinavica 42, brill, leiden. http://dx.doi.org/10.1163/ ej.9789004157705.i-265 folsom j.w. 1913. north american spring-tails of the sub-family tomocerinae. proceedings of the united states national museum 46: 451–472. http://dx.doi.org/10.5479/si.00963801.46-2037.451 goto h.e. 1956. architomocerura denis, 1931 (collemb., isotomidae) an immature stage of tomocerus nicolet, 1841 (collemb., tomoceridae). entomologist’s monthly magazine 42: 49–51. http://dx.doi.org/10.1163/ej.9789004157705.i-265 http://dx.doi.org/10.1163/ej.9789004157705.i-265 yu d. et al., tomoceridae from southern annamitic cordillera 13 krantz g.w. 1978. a manual of acarology. second edition. oregon state university bookstore, inc. corvallis. lee b.h. 1974. étude de la faune coréenne des insectes collemboles. v. inventaire des grottes de corée et étude sur les tomoceridae cavernicoles avec la description d’une nouvelle espèce. annales de spéléologie 29: 403–418. lee b.h. 1975. étude de la faune coréenne des insectes collemboles. vi. sur la famille des tomoceridae, édaphiques, avec la description de quatre nouvelles espèces et d’une nouvelle sous-espèce. bulletin du muséum national d’histoire naturelle, série 3 317: 945–961. liu y.q., hou d.b. & li z.c. 1998. a checklist of collembola species from china. journal of southwest agricultural university 20: 125–131. lubbock j. 1873. monograph of the collembola and thysanura. ray society, london. http://dx.doi. org/10.5962/bhl.title.11583 ma y.t. 2004. taxonomic study on the genera dicranocentrus and entomobrya and the family tomoceridae from china (insecta: apterygota). phd thesis, nanjing university, china. martynova e.f. 1977. springtails of the family tomoceridae (collembola) from the fauna of the far east. insect fauna of the far east 46: 3–16. oudemans j.t. 1891. apterygota des indischen archipels. zoologische ergebnisse einer reisen in niederländische ost-indien 1: 73–91. stach j. 1964. materials to the knowledge of chinese collembolan fauna. acta zoologica cracoviensia 9: 1–26. stach j. 1965. on some collembola of north vietnam. acta zoologica cracoviensia 10: 346–372. uchida h. 1953. the insect fauna of mt. ishizuchi and omogo valley, iyo, japan. the collembola. transactions of the shikoku entomological society 4: 1–6. uchida h. & chiba s. 1958. studies on the development of tomocerus minutus tullberg (insecta: collembola). i. on the postembryonic development. zoological magazine 67: 242–248. yosii r. 1956. monographie zur höhlencollembolen japans. contributions from the biological laboratory kyoto university 3: 1–109. yosii r. & lee c.e. 1963. on some collembola of korea, with notes on the genus ptenothrix. contributions from the biological laboratory kyoto university 15: 1–37. yosii r. 1966. collembola of himalaya. journal of the college of arts and sciences, chiba university 4: 461–531. yosii r. 1967. studies on the collembolan family tomoceridae, with special reference to japanese forms. contributions from the biological laboratory kyoto university 20: 1–54. yosii r. 1969. collembola-arthropleona of the ibp-station in the shiga heights, central japan, i. bulletin of the national science museum tokyo 12: 531–556. yosii r. 1977. critical check list of the japanese species of collembola. contributions from the biological laboratory kyoto university 25: 142–170. yu d.y., zhang f. & deharveng l. 2014. a peculiar cave species of tomocerus (collembola, tomoceridae, tomocerinae) from vietnam, with a discussion of the postantennal organ and prelabral chaetae in tomocerinae. zookeys 408: 61–70. http://dx.doi.org/10.3897/zookeys.408.7030 http://dx.doi.org/10.5962/bhl.title.11583 http://dx.doi.org/10.5962/bhl.title.11583 european journal of taxonomy 176: 1–14 (2016) 14 yu d.y. & deharveng l. 2015. the first eyeless species of tomocerus from china (collembola, tomoceridae) with notes on genera tomocerus and pogonognathellus. zootaxa 3914 (2): 175–184. http://dx.doi.org/10.11646/zootaxa.3914.2.7 zhang f., yu d.y., luo y.z., ho s.y.w., wang b.x. & zhu c.d. 2014. cryptic diversity, diversification and vicariance in two species complexes of tomocerus (collembola, tomoceridae) from china. zoologica scripta 43 (4): 393–404. http://dx.doi.org/10.1111/zsc.12056 manuscript received: 17 october 2015 manuscript accepted: 6 november 2015 published on: 19 february 2016 topic editor: koen martens desk editor: charlotte thionois printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark. european journal of taxonomy 778: 86–89 issn 2118-9773 https://doi.org/10.5852/ejt.2021.778.1569 www.europeanjournaloftaxonomy.eu 2021 · gómez s. et al. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:2b639eaf-bed8-4223-b5d0-f1e030c677da 86 proposal of new genera and species of the subfamily diosaccinae (copepoda: harpacticoida: miraciidae) – corrigendum samuel gómez 1,*, paulo henrique costa corgosinho 2 & karen i. rivera-sánchez 3 1 universidad nacional autónoma de méxico, instituto de ciencias del mar y limnología, unidad académica mazatlán, mazatlán, sinaloa, méxico. 2 dep. biologia geral. universidade estadual de montes claros (unimontes), montes claros, mg, brazil. 3 posgrado en ciencias del mar y limnología, unidad académica mazatlán, mazatlán, sinaloa, méxico. * corresponding author: samuelgomez@ola.icmyl.unam.mx 2 email: pcorgo@gmail.com 3 email: kitzel.rivera10@gmail.com 1 urn:lsid:zoobank.org:author:7bbfab07-962f-4d10-9be0-87b936993fbd 2 urn:lsid:zoobank.org:author:2b499408-5c00-4a01-8a8e-18fa969123cf 3 urn:lsid:zoobank.org:author:ab43ce7f-6c8a-45af-b2d1-c3bd92644b18 gómez s., corgosinho p.h.c. & rivera-sánchez k.i. 2021. proposal of new genera and species of the subfamily diosaccinae (copepoda: harpacticoida: miraciidae) – corrigendum. european journal of taxonomy 778: 86–89. https://doi.org/10.5852/ejt.2021.778.1569 the present corrigendum corrects errors that occurred in gómez et al. (2021). junior homonym the genus dinetia gómez, corgosinho & rivera-sánchez, 2021 was proposed to accommodate dinetia minuta (dinet, 1971), previously included within bulbamphiascus by dinet (1971). we inadvertently overlooked that the genus name was preoccupied by dinetia decraemer & gourbault, 1997, proposed to accommodate the draconematid nematod dinetia nycterobia decraemer & gourbault, 1997 (decraemer & gourbault 1997). following the principle of homonymy (iczn 1999; art. 52.2), only the senior homonym may be used as a valid name. therefore, in adherence to art. 60.3 (icnz 1999), dinetocaris nom. nov. (gender: feminine), alluding to alain dinet (for his contribution to the taxonomy of harpacticoid copepods) followed by the suffix ‘caris’ (meaning ‘shrimp’), is proposed as the replacement name for dinetia gómez, corgosinho & rivera-sánchez, 2021. the genus dinetocaris nom. nov. includes dinetocaris minuta (dinet, 1971) comb. nov. as its type and only valid species, and it has been registered in zoobank (urn:lsid:zoobank.org:act:59e0ed74-c8d3-43bb-aa0c-4775840820a9). https://doi.org/10.5852/ejt.2021.778.1569 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:2b639eaf-bed8-4223-b5d0-f1e030c677da https://orcid.org/0000-0002-8597-8846 https://orcid.org/0000-0002-2851-6557 mailto:samuelgomez%40ola.icmyl.unam.mx?subject= mailto:pcorgo%40gmail.com?subject= mailto:kitzel.rivera10%40gmail.com?subject= http://zoobank.org/urn:lsid:zoobank.org:author:7bbfab07-962f-4d10-9be0-87b936993fbd http://zoobank.org/urn:lsid:zoobank.org:author:2b499408-5c00-4a01-8a8e-18fa969123cf http://zoobank.org/urn:lsid:zoobank.org:author:ab43ce7f-6c8a-45af-b2d1-c3bd92644b18 https://doi.org/10.5852/ejt.2021.778.1569 http://zoobank.org/urn:lsid:zoobank.org:act:59e0ed74-c8d3-43bb-aa0c-4775840820a9 gómez s. et al., on new species and genera of harpacticoida – corrigendum 87 the case of dineticola gen. nov. the name dineticola gen. nov. appears in table 4 (pages 10 and 11) and on fig. 19 (page 40) in gómez et al. (2021). dineticola gen. nov. was a slip of the pen and, therefore, it is a nomen nudum; it is unavailable because it was not accompanied by a description or a bibliographic reference, failing to conform to art. 13 (iczn 1999). the genus name dineticola gen. nov. should be changed to dinetocaris gen. nov. list of errors page 1, abstract, line 14: “and dinetia gen. nov.” should be changed to “and dinetocaris gen. nov.” page 2, introduction, line 30: “we propose dinetia gen. nov.” should be changed to “we propose dinetocaris gen. nov.” page 10, table 4, terminals: “dineticola gen. nov.” should be changed to “dinetocaris gen. nov.” page 11, table 4, terminals: “dineticola gen. nov.” should be changed to “dinetocaris gen. nov.” page 43, line 31: “dinetia gen. nov.” should be changed to “dinetocaris gen. nov.” page 43, line 35: “(= dinetia minuta (dinet, 1971)” should be changed to “(= dinetocaris minuta (dinet, 1971)” page 44, line 23: “apomorphies for dinetia gen. nov.” should be changed to “apomorphies for dinetocaris gen. nov.” page 44, lines 25, 30 and 37: “dinetia gen. nov.” should be changed to “dinetocaris gen. nov.” page 45, lines 12 and 17: “dinetia gen. nov.” should be changed to “dinetocaris gen. nov.” page 53, line 18: “phylogenetic position of dinetia gen. nov.,…” should be changed to “phylogenetic position of dinetocaris gen. nov.,…” page 53, lines 32–33: “…a clade composed of dinetia gen. nov.,…” should be changed to “…a clade composed of dinetocaris gen. nov.,…” page 55, lines 18, 19, 21, 25 and 29: “dinetia gen. nov.” should be changed to “dinetocaris gen. nov.” page 55, line 27: “…the dinetia–rhyncholagena clade…” should be changed to “…the dinetocaris–rhyncholagena clade…” page 56, line 18: “dinetia gen. nov.” should be changed to “dinetocaris gen. nov.” european journal of taxonomy 778: 86–89 (2021) 88 page 40, fig. 19 is in very low resolution. a new figure, with dineticola corrected to dinetocaris gen. nov., is provided here: fig. 19. strict consensus cladogram of the generic relationships within the diosaccinae. references decraemer w. & gourbault n. 1997. deep-sea nematodes (nemata, prochaetosomatinae): new taxa from hydrothermal vents and a polymetallic nodule formation of the pacific (east rise; north fiji and lau basins; clarion-clipperton fracture zone). zoologica scripta 26: 1–12. https://doi.org/10.1111/j.1463-6409.1997.tb00405.x dinet a. 1971. copépodes harpacticoïdes d’un sable fin organogène des environs de marseille. tethys 2: 747–762. gómez s., corgosinho, p.h.c. & rivera-sánchez k.i. 2021. proposal of new genera and species of the subfamily diosaccinae (copepoda: harpacticoida: miraciidae). european journal of taxonomy 759: 1–62. https://doi.org/10.5852/ejt.2021.759.1433 https://doi.org/10.1111/j.1463-6409.1997.tb00405.x https://doi.org/10.5852/ejt.2021.759.1433 gómez s. et al., on new species and genera of harpacticoida – corrigendum 89 international commission on zoological nomenclature (iczn). 1999. international code of zoological nomenclature. international trust for zoological nomenclature c/o the natural history museum, london. published on: 15 november 2021 printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d′histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. european journal of taxonomy 726: 24–37 issn 2118-9773 https://doi.org/10.5852/ejt.2020.726.1171 www.europeanjournaloftaxonomy.eu 2020 · ahmad z. et al. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:0fd044c0-6ecf-432e-875b-48013dfcf84e 24 parasitoids of the genus pholetesor mason, 1981 (hymenoptera: braconidae: microgastrinae) from the leafminers lepidoptera, with the description of three new species from india zubair ahmad 1, *, hamed a. ghramh 2 & kavita pandey 3 1, 2 research centre for advanced materials science (rcams), king khalid university, p.o. box 9004, abha 61413, saudi arabia. 1, 2 unit of bee research and honey production, faculty of science, king khalid university, p.o. box 9004, abha 61413, saudi arabia. 1 biology department, faculty of arts & sciences, king khalid university, dhahran al junoub, saudi arabia. 2 biology department, faculty of science, king khalid university, p.o. box 9004, abha 61413, saudi arabia. 3 department of zoology, aligarh muslim university, aligarh, 202002, up, india. * corresponding author: dzubair@gmail.com 2 email: hamedsa@hotmail.com 3 email: kavi_leaf@gmail.com 1 urn:lsid:zoobank.org:author:769585c9-7c8c-4381-9dfe-ed5c32efdcee 2 urn:lsid:zoobank.org:author:a2eaebaa-d809-4532-8b6a-eaf405530ea7 3 urn:lsid:zoobank.org:author:d976673b-19fe-4ca8-b0ff-e07cee18b048 abstract. pholetesor acrocercophagus sp. nov., p. camerariae sp. nov. and p. indicus sp. nov. (hymenoptera: braconidae: microgastrinae) are described as new to science. these three species were reared from acrocercops sp., acrocercops phaeospora meyrick, 1916 and cameraria virgulata meyrick, 1914 (lepidoptera: gracillariidae), respectively. characteristics of these new species and their affinities with related taxa are discussed. data on habitat, host records and host plant species for all the parasitoid species is provided. a key to the indian species of the genus pholetesor mason, 1981 reared from lepidopteran leafminers is also given. keywords. hymenoptera, braconidae, microgastrinae, pholetesor, parasitoids, lepidoptera, leafminer, new species, india. ahmad z., ghramh h.a. & pandey k. 2020. parasitoids of the genus pholetesor mason, 1981 (hymenoptera: braconidae: microgastrinae) from the leafminers lepidoptera, with the description of three new species from india. european journal of taxonomy 726: 24–37. https://doi.org/10.5852/ejt.2020.726.1171 https://doi.org/10.5852/ejt.2020.726.1171 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:0fd044c0-6ecf-432e-875b-48013dfcf84e mailto:dzubair%40gmail.com?subject= mailto:hamedsa%40hotmail.com?subject= mailto:kavi_leaf%40gmail.com?subject= http://zoobank.org/urn:lsid:zoobank.org:author:769585c9-7c8c-4381-9dfe-ed5c32efdcee http://zoobank.org/urn:lsid:zoobank.org:author:a2eaebaa-d809-4532-8b6a-eaf405530ea7 http://zoobank.org/urn:lsid:zoobank.org:author:d976673b-19fe-4ca8-b0ff-e07cee18b048 https://doi.org/10.5852/ejt.2020.726.1171 ahmad z. et al., genus pholetesor and description of three new species from india 25 introduction leaf-mining insects are considered serious pests which damage plant leaves in a variety of ways. if leaves are seriously attacked, crop can be reduced or seedling plants may be totally destroyed (spencer 1990). the majority of the leaf-mining larvae belong to the lepidoptera linnaeus, 1758, followed to a lesser degree by diptera linnaeus, 1758, coleoptera linnaeus, 1758 and hymenoptera linnaeus, 1758 (csoka 2003). among the lepidopteran leafminers, the family gracillariidae stainton, 1854 includes small-sized moths with white marks on the wings. several species are considered serious pests in several parts of the world (davis 1987). parasitoid insects play the most important role as natural enemies of leafminers, because in some cases, they can cause more than 90% mortality and consequently have great potential in biological pest control programs (hawkins et al. 1993). parasitoids of leafminers are exclusively from the order hymenoptera, with superfamilies chalcidoidea latreille, 1817, ichneumonidae latreille, 1802 and braconidae burmeister, 1829 being the most important groups associated with leafminers. mason (1981) erected the genus pholetesor mason, 1981 to accommodate nixon‘s (1965) ̒ circumscriptusgroupʼ and ʻbucculatricis-groupʼ of species of microgastrinae förster, 1862. this genus has 57 species recorded from the world, but only one species, p. hayati akhtar & ahmad, 2010, has been reported from india (akhtar et al. 2010; yu et al. 2016; fernandez-triana et al. 2020). known hosts of various species of pholetesor are almost exclusively leaf-mining lepidoptera (whitfield 2006). pholetesor is characterized by the following characters: 1) short and hairy and relatively dorsally attached ovipositor sheaths; 2) strong sub-lateral setiferous lobes of the metanotum; 3) propodeal areola (when present) strongly pentagonal rather than oval or diamond-shaped (whitfield 2006); 4) hypopygium medially folded but evenly sclerotized or only weakly translucent medially; 5) 2r-m of fore wing absent. as far as india is concerned, there is a lack of information on braconid parasitoids, especially reared from leaf-mining lepidopteran hosts. there were only four microgastrine parasitoid wasps described from india (wilkinson 1928; ahmad et al. 2019). in the present, work three new parasitoid species, viz, pholetesor acrocercophagus sp. nov., p. camerariae sp. nov. and p. indicus sp. nov. reared from acrocercops sp., acrocercops phaeospora meyrick, 1916 and cameraria virgulata meyrick, 1914 (lepidoptera: gracillariidae), respectively, are described and illustrated, and their affinities with related taxa are discussed. a key to the indian species of the genus pholetesor reared from lepidopteran leafminers is provided. this work was initially started in 2004 by first author za during his post-doctoral project on rearing braconid parasitoids from india. a part of this work is included in the phd thesis of the third author kp (pandey 2006). material and methods this study was conducted in the vicinity of western uttar pradesh (north india) in order to identify the parasitoids of leafminers along the roadside at amu university campus. the parasitoids were reared from acrocercops sp., a. phaeospora and c. virgulata on the tree leaves of achyranthes aspera l., millettia pinnata (l.) panigrahi and syzygium cuminii (l.) skeels, respectively, in the laboratory in glass jars. a complete data set such as the date of collection, locality and name of host plant was maintained. the emerged parasitoids were initially preserved in 75% alcohol with a few drops of glycerol. these specimens were later mounted on cards. the reared parasitoids were separated based on morphological characters. the pholetesor species keys of nixon (1973), papp (1983) and liu et al. (2016) were used for the identification. van achterberg (1993) is followed for the terminologies of various body parts and wing venation and eady (1968) for the terminology of micro-sculpture. the types of new species were deposited in the insect collection of the department of zoology, aligarh muslim university, aligarh, india (zdamu). european journal of taxonomy 726: 24–37 (2020) 26 abbreviations for morphological terms used in the text aol = anterior ocellar line (distance between the inner edges of an anterior ocellus and lateral ocellus) pol = posterior ocellar line (distance between the inner edges of lateral ocelli) ool = ocello-ocular line (distance from the outer edge of a lateral ocellus to the compound eye) od = ocellus diameter f = flagellomere t = metasomal tergite institutional abbreviations zdamu = department of zoology, aligarh muslim university, aligarh, india results taxonomic treatments class insecta linnaeus, 1758 order hymenoptera linnaeus, 1758 suborder apocrita latreille, 1810 superfamily ichneumonoidea latreille, 1802 family braconidae nees, 1811 subfamily microgastrinae förster 1862 genus pholetesor mason, 1981 pholetesor acrocercophagus sp. nov. urn:lsid:zoobank.org:act:6988b1d5-883f-422e-bf22-42750a323d49 fig. 1 diagnosis pholetesor acrocercophagus sp. nov. is closely related to p. circumscriptus (ness, 1834) in the european keys to the species of the ʻcircumscriptus-groupʼ (nixon 1973; papp 1983) on the basis of the following characters: thorax in profile less elongate; metanotum strongly retracted from scutellum anteriorly, exposing mesothoracic postphragma; pterostigma almost 3.0 × as wide as long; vein 1-r1 (metacarp) clearly longer than pterostigma, r and 2-sr slightly curved, meeting less angularly; propodeum largely smooth; hind femur yellowish; t1 largely smooth, polished and strongly narrowed apically; t2 + 3 smooth and polished. however, it differs from p. circumscriptus in the following characters: t2 subtriangular 2.0 × as wide as long posteriorly, hence more transverse (t2 strongly triangular 2.6 × as wide as long posteriorly in p. circumscriptus); scutellar sulcus narrow with fine foveation (scutellar sulcus narrow with fine groove in which there is no obvious foveation in p. circumscriptus); forewing vein r arising medially of the pterostigma (forewing vein r arising distally of the pterostigma in p. circumscriptus); f2 2.5 × as long as wide (f2 3.6 × as long as wide in p. circumscriptus); body comparatively small, about 1.5 mm (body larger, about 1.80–2.00 mm in p. circumscriptus). etymology the new species is named after its host insect. material examined holotype india • ♀; uttar pradesh, aligarh; 27°54′53.3″ n, 78°04′23.5″ e; 11 jul. 2005; z. ahmad leg.; ex. acrocercops sp. on achyranthes aspera l.; zdamu. http://zoobank.org/urn:lsid:zoobank.org:act:6988b1d5-883f-422e-bf22-42750a323d49 ahmad z. et al., genus pholetesor and description of three new species from india 27 paratype india • 1 ♀; same collection data as for holotype; zdamu. description female measurements. body lenght = 1.5 mm; length of fore wing = 1.6 mm; length of antenna = 1.5 mm. fig. 1. pholetesor acrocercophagus sp. nov., paratype, ♀ (zdamu). a. head, frontal view. b. mesosoma. c. metasoma. scale bars: 0.25 mm. european journal of taxonomy 726: 24–37 (2020) 28 head. almost 2 × as wide as long; eyes 1.6 × as long as temple in dorsal view; temple and vertex smooth, shiny, punctate with hairs; eyes setose, 1.4 × as long as wide, ool: pol: aol: od = 4: 3: 2: 2; frons concave, smooth and shiny, face medially convex, punctate with hairs; clypeus 3 × as wide as long, flattened, indistinctly punctate; malar space about as long as basal width of mandible; antennae about as long as body; scape 1.5 × as long as wide, pedicel slightly wider than long, f1 3.0 × as long as wide, f2 2.5 × as long as wide, flagellomeres gradually decreasing apically, apical flagellomere longer than f15 and pointed. mesosoma. about as long as wide and 1.5 × its height; mesoscutum distinctly punctate with hairs, punctations becomes obscured posteriorly, scutellar sulcus narrow with fine foveation; scutellum smooth and shiny, side of scutellum smooth and shiny; propodeum about 2.0 × as wide as long at longest point, largely smooth; posteriorly with series of ridges extending obliquely on either side from nucha; propleuron smooth and shiny; mesopleuron antero-dorsally smooth and shiny, postero-laterally punctate with hairs. legs. hind coxa punctate with hairs; length of femur, tibia and basitarsus of hind leg 3.0, 5.0 and 4.2 × their width, respectively. wings. fore wing 2.6 × as long as wide, pterostigma almost 3.0 × as wide as long, 1-r1 1.21 × as long as pterostigma, r and 2-sr slightly curved, meeting less angularly, r arising medially of pterostigma. metasoma. 2.4 × as long as wide; t1 about 3.0 × as long as its apical width, 1.8 × its basal width, strongly narrowed apically, smooth and shiny except few punctures; t2 2.0 × as wide as long, subtriangular, smooth and shiny, shorter than t3; t3 rectangular, 1.40 × as long as t2, unsculptured; hypopygium weakly sclerotized, transparent, medially folded, blunt at apex, 2.0 × as long as hind basitarsus; ovipositor sheaths elongate-fusiform, thick and hairy throughout their length, arising about half of valvifer, 1.2 × as long as hind basitarsus; ovipositor pointed at apex and curved downwards. colour. body black except for the following; hind femur, tibia and basitarsus infuscate; antennae, tegulae, latero-tergites dark brown; tip of hypopygium transparent; ovipositor reddish brown; tibial spurs, palpi creamish; fore leg except coxae, mid leg except coxae, hind tarsus except basitarsus and teleotarsus yellow. male unknown. distribution india: uttar pradesh. host acrocercops sp. remarks pholetesor acrocercophagus sp. nov. can be distinguished from all previously described species by the unique combination of characters as follows: metanotum strongly retracted from scutellum anteriorly, exposing mesothoracic postphragma; propodeum about 2.0 × as wide as long at longest point, largely smooth; posteriorly with a series of ridges extending obliquely on either side from nucha; t1 about 3.0 × as long as its apical width, 1.8 × its basal width, strongly narrowed apically, smooth and shiny except few punctures; t2 2.0 × as wide as long, subtriangular, smooth, shiny, shorter than t3; t3 rectangular, 1.40 × as long as t2, unsculptured; hypopygium weakly sclerotized, transparent, medially folded, blunt ahmad z. et al., genus pholetesor and description of three new species from india 29 at apex, 2.0 × as long as hind basitarsus; ovipositor sheaths elongate-fusiform, thick and hairy throughout their length, arising about half of valvifer; ovipositor pointed at apex and curved downwards. pholetesor camerariae sp. nov. urn:lsid:zoobank.org:act:a2b68eda-3dbf-4cd0-838f-1eff0b1e5da0 fig. 2 diagnosis pholetesor camerariae sp. nov. is closely related with indian species, viz, p. hayati. however, it differs from p. hayati in the following characters: hind coxae somewhat enlarged so that distal ends of hind coxae reach nearly to posterior end of t3 (hind coxae not reaching beyond t2 in p. hayati); ovipositor sheaths long, slender basally, then tapering broader to a bluntly bevelled tip, approximately as long as hind basitarsi, arising at mid length of valvifer (ovipositor sheath short, fusiform, 0.55 × as long as hind basitarsi, arising slightly below the mid length of valvifer in p. hayati); t2 smooth and shiny (t2 sculptured with a smooth raised median area p. hayati); t1 1.4 × as long as wide posteriorly (t1 1.1 × as long as wide posteriorly in p. hayati). pholetesor camerariae sp. nov. is also closely related with p. salalicus (mason, 1959) in the key to the chinese species of the genus pholetesor (liu et al. 2016). however, it differs from p. salalicus in the following characters: t2 smooth except few hairs, trapezoidal, 1.5 × as wide as long, (t2 rugose, subtriangular to trapezoidal, 1.7–2.0 × as wide as long in p. salalicus); posterior margin of t2 usually weakly concave (posterior margin of t2 straight in p. salalicus); propodeum about 2 × as wide as long, shallowly rugulose to rugose, smooth at antero-lateral corners (propodeum about 1.7 × as wide as long at longest point, punctate to weakly rugulose antero-laterally, smooth and depressed in postero-lateral corners). etymology the new species is named after its host insect. material examined holotype india • ♀; uttar pradesh, aligarh; 27°54′26.4″ n, 78°04′13.9″ e; 4 dec. 2005; z. ahmad leg.; ex. cameraria virgulata meyrick, 1914 on millettia pinnata (l.) panigrahi; zdamu. paratypes india • 17 ♀♀, 3 ♂♂; same collection data as for holotype; zdamu. description female measurements. body length = 1.8 mm; length of forewing = 2.0 mm; length of antenna = 2.0 mm. head. almost 2 × as wide as long in dorsal view; eyes as long as 2.0 × as temple in dorsal view; ool: pol: aol: od = 4: 3: 2: 2; temple smooth, shiny, punctate with hairs; vertex convex, smooth, shiny, indistinctly punctate with hairs; frons concave, smooth and shiny; face 1.3 × as wide as long, medially convex, punctate with hairs; clypeus convex, punctate with hairs; malar space about as long as basal width of mandible; antennae longer than body length, apical four flagellomeres without double rank of placodes, f1–f7 equal in length, f8–f15 gradually decreasing in length apically, apical flagellomere slightly longer than f15. http://zoobank.org/urn:lsid:zoobank.org:act:a2b68eda-3dbf-4cd0-838f-1eff0b1e5da0 european journal of taxonomy 726: 24–37 (2020) 30 mesosoma. 1.3 × as long as wide dorsally and just above the tegulae as wide as head; scutum shallowly punctate with hairs, becoming nearly impunctate at extreme posterior edge; scutellar sulcus almost straight and foveolate to crenulated; scutellum smooth with fine, shallow punctures; metanotum weakly retracted from scutellum; sublateral setiferous lobes nearly appressed to hind margin of scutellum; transverse carinae poorly developed; propodeum about 2 × as wide as long, shallowly rugulose, smooth at antero-lateral corners, with pair of ridges extending obliquely on either side from nucha; propleuron smooth and shiny; mesopleuron mostly smooth except few hairs anteriorly; metapleuron smooth and shiny except edges. wings. fore wing 2.5 × as long as wide; 1-r1 about as long as pterostigma; r slightly longer than 2-sr; r and 2-sr meeting angularly. hind wing with cu-a declivous; venal lobe evenly convex and hairy; pterostigma almost 3 × as long as wide. fig. 2. pholetesor camerariae sp. nov., paratype, ♀ (zdamu). a. head, frontal view. b. mesosoma, dorso-lateral view. c. t1 and t2, dorso-lateral view. d. metasoma, lateral view. scale bars: 0.25 mm. ahmad z. et al., genus pholetesor and description of three new species from india 31 legs. hind coxa smooth and shiny, almost reaching posterior end of t3; length of femur, tibia, and basitarsus of hind leg 4.0, 6.0 and 6 × their width, respectively, longer tibial spur 0.41 × as long as basitarsus (12:5). metasoma. about 3 × as long as wide; t1 2.5 × as long as its apical width, 1.66 × its basal width, parallel sided, only slightly narrowing apically, surface longitudinally aciculate anteriorly, posteriorly aciculorugose with less conspicuous longitudinal elements; t2 trapezoidal, 1.5 × as wide as long, smooth except few hairs, posterior margin of t2 usually straight and posteriorly marked by shallow groove; t3 rectangular, 1.30 × as long as t2, unsculptured; hypopygium weakly sclerotized basally, membranous apically, 1.5 × as long as hind basitarsus; ovipositor sheaths long, slender basally, then tapering broader to bluntly bevelled tip, hairy at apical half, approximately as long as hypopygium, arising at mid length of valvifer; ovipositor weakly decurved, approximately as long as hypopygium. colour. body black except for the following: ocelli testaceous; scape, pedicel, f5–f16, mandible brown; maxillary palpi, labial palpi, tibial spurs creamish; legs, tegulae and latero-tergites yellowish; wings hyaline; pterostigma, vein 1-r1, c + sc + r slightly pigmented and remaining veins colourless. male same as holotype. host cameraria virgulata. distribution india: uttar pradesh. remarks the new species bears strong resemblance to another indian species, p. hayati, in the colouration of the tegulae, wing veins, stigma and legs (except hind coxae), and the sculpturing of the propodeum and metasomal tergites (but not their exact shapes – p. hayati has a broader first tergite with rounded lateral margins and a somewhat longer second tergite). also showing some resemblance in the palearctic fauna is p. laetus (marshall, 1885) and p. salalicus, both of which shares a number of colour and tergite features with p. camerariae sp. nov., but both the palearctic species have more sculptured t2 than p. camerariae sp. nov., and scutellar sulcus reduced to a fine groove in which there is no obvious foveation, while p. camerariae sp. nov. has a distinct faveolated groove in scutellar sulcus. pholetesor indicus sp. nov. urn:lsid:zoobank.org:act:a0f294eb-cc47-4df7-a9fe-da2201becca2 fig. 3 diagnosis pholetesor indicus sp. nov. is closely related to p. bicolor (ness, 1874) in the european keys to the species of ̒ circumscriptus-groupʼ (nixon, 1973; papp 1983) on the basis of the following characters: metanotum strongly retracted from scutellum anteriorly, exposing mesothoracic postphragma; pterostigma almost 3.0 × as wide as long; vein 1-r1 (metacarp) clearly as long as or shorter than pterostigma, vein r and 2-sr meeting angularly; propodeum largely smooth; t1 largely smooth, polished and strongly narrowed apically; t2 + 3 smooth and polished, metasomal tergites and legs more often yellowish. however, it differs from p. bicolor in the following characters: mesoscutum punctations are prominent all along the mesoscutum surface (punctations becoming indistinct posteriorly near scutellum in p. bicolor); tegulae http://zoobank.org/urn:lsid:zoobank.org:act:a0f294eb-cc47-4df7-a9fe-da2201becca2 european journal of taxonomy 726: 24–37 (2020) 32 blackish brown (tegulae yellowish to pale yellowish in p. bicolor) pterostigma about 3.0 × as wide as long (pterostigma about 2.5 × as wide as long in p. bicolor); hind coxae black (hind coxae often yellowish in p. bicolor); ovipositor sheaths 0.9 × as long as hind basitarsi, small and thick (ovipositor sheaths 1.1–1.2 × as long as hind basitarsi, comparatively large with blunt apex in p. bicolor); f2 2.3 × as long as wide (f2 3.5 × as long as wide in p. bicolor); f5–f11 with double rank of placodes (f5–f6 with double rank of placodes in p. bicolor). the new species is also very similar to p. circumscriptus. however, it differs from p. circumscriptus in the following characters: vein 1-r1 (metacarp) as long as or shorter than pterostigma (vein 1-r1 (metacarp) clearly longer than pterostigma in p. circumscriptus); t2 subtriangular, 2.0 × as wide as fig. 3. pholetesor indicus sp. nov., paratype, ♀ (zdamu). a. mesosoma, dorsal view. b. propodeum, dorsal view. c. metasoma, dorsal view. scale bars: 0.25 mm. ahmad z. et al., genus pholetesor and description of three new species from india 33 long posteriorly, hence more transverse (t2 strongly triangular, 3 × as wide as long posteriorly in p. circumscriptus); forewing vein r and 2-sr meeting angularly (forewing vein r and 2-sr meeting each other less angularly in p. circumscriptus). apart from the similarities with p. bicolor, the new species also runs close to p. teresitergum liu & chen, 2016 in the key to the chinese species of the genus pholetesor (liu et al. 2016). however, it differs from p. teresitergum in the following characters: eyes 2.2 × as long as temple in dorsal view (eyes 1.6 × as long as temple in dorsal view in p. teresitergum); ocelli large (ocelli relatively small in p. teresitergum); scutellum highly polished (scutellum punctate with hairs in p. teresitergum); ovipositor sheaths small and thick, slender with a bluntly rounded tips, (ovipositor sheaths small and subfusiform in p. teresitergum); tegulae blackish brown (tegulae yellowish in p. teresitergum). material examined holotype india • ♀; uttar pradesh, aligarh; 27°54′51.0″ n, 78°04′24.7″ e; 27 oct. 2005; z. ahmad leg.; ex. acrocercops phaeospora meyrick, 1914 on syzygium cuminii (l.) skeels; zdamu. paratypes india • 5 ♀♀, 2 ♂♂; same collection data as for holotype; zdamu. etymology the new species is named after its type locality. description female measurements. body lenght = 1.7–1.8 mm; length of antenna = 2.0 mm; length of fore wing = 1.7 mm. head. transverse, almost 2 × as wide as long in dorsal view; eyes 2.2 × as long as temple in dorsal view; eyes 1.3 × as long as wide, setose; temple and vertex punctate with hairs; ool: pol: aol: od = 4: 3: 2: 2; frons concave, smooth, shiny, punctate with hairs; face medially convex, punctate with white pilosity, 1.3 × as wide as long; clypeus smooth, punctate with hairs; malar space 1.5 × as long as basal width of mandible; antennae longer than body, scape 1.2 × as long as wide, pedicel 0.66 × as long as wide, f2 2.3 × as long as wide, f14 1.5 × as long as wide , f5–f11 with double rank of placodes, apical segment pointed. mesosoma. 1.5 × as long as wide; mesoscutum as wide as head and strongly punctate with hairs, punctations prominent all along mesoscutum surface; scutellar sulcus narrow and crenulate; scutellum smooth and polished, side of scutellum smooth and shiny; propodeum almost 2 × as wide as long, largely smooth except antero-medially punctate with long hairs, posterior corners somewhat weakly rugulose; propleuron smooth and shiny; mesopleuron concave, smooth and shiny, antero-lateral, punctate with hairs; metapleuron smooth and shiny with few hairs. wings. fore wings hyaline, about 3.0 × as long as wide, pterostigma about 3.0 × as wide as long, 1-r1, about 0.9 × as long as length of pterostigma, r and 2-sr meeting angularly; venal lobe of hind wing convex. legs. hind coxae hairy, length of hind femur, tibia and basitarsus 2.6, 5.0 and 2.6 × their width, respectively, outer tibial spur 0.5 × as long as hind basitarsus. european journal of taxonomy 726: 24–37 (2020) 34 metasoma. 1.7 × as long as wide; t1 3.6 × as long as its apical width and 1.8 × its basal width, distinctly narrowed apically, smooth and shiny; t2 subtriangular, 3 × as wide as long, smooth except few punctations; hypopygium sclerotized, folded medially and blunt at apex; ovipositor sheaths small and thick, slender with bluntly rounded tips, 0.7 × as long as hypopygium and 0.9 × longer than hind basitarsi, hairy at apical one third, arising below at mid length of valvifer; ovipositor weakly curved downward. colour. body blackish brown except for the following: ocelli testaceous; palpi, labarum, fore leg, mid leg and hind leg yellowish except coxae; tegulae blackish brown; scape, pedicel, latero-tergites of t1 and t2 and ovipositor yellowish brown; antennae brown; ovipositor sheaths, fore, mid and hind coxae black; mid and hind femur infuscate; pterostigma, vein c + sc + r, 1-r1, 2-sr, cu-a, 1-cu1, 2-cu1 and 2-m brown; remaining veins colourless, wings hyaline. male same as holotype except antennae longer than body length, f1–f3 with double rank of placodes. host acrocercops phaeospora. distribution india: uttar pradesh. remarks pholetesor indicus sp. nov. can be distinguished from all previously described species by the unique combination of characters as follows: ocelli large; scutellum highly polished; propodeum almost 2 × as wide as long, largely smooth except antero-medially punctate with long hairs, posterior corners somewhat weakly rugulose; t1 3.6 × as long as its apical width and 1.8 × its basal width, distinctly narrowed apically, smooth and shiny; t2 subtriangular, 3 × as wide as long, smooth except few punctations; t3 rectangular, 1.40 × as long as t2, unsculptured; hypopygium sclerotized, folded medially and blunt at apex; ovipositor sheaths small and thick, slender with a bluntly rounded tips, 0.7 × as long as hypopygium, hairy at apical one third, arising below at mid length of valvifer; ovipositor weakly curved downward. key to the indian species of the genus pholetesor mason, 1981 reared from lepidopteran leafminers 1. t1 sculptured all along the surface, usually parallel sided, weakly narrowing only posteriorly; t2 trapezoidal; tegulae yellowish .......................................................................................................... 2 – t1 smooth except for few punctations, strongly narrowing posteriorly; t2 strongly triangular to subtriangular; tegulae dark brown to black ...................................................................................... 3 2. ovipositor sheaths short and fusiform in shape; t2 sculptured with a smooth raised median area; legs completely yellow ...................................................................p. hayati akhtar & ahmad, 2010 – ovipositor sheaths long, slender basally, then tapering broader to a bluntly bevelled tip; t2 completely smooth and shiny .............................................................................................p. camerariae sp. nov. 3. hypopygium heavily sclerotized; ovipositor sheaths small, nearly as long as hind basitarsi ............ ................................................................................................................................ p. indicus sp. nov. – hypopygium weakly sclerotized; ovipositor sheaths long, more than 1.2 × as long as hind basitarsi ................................................................................................................ p. acrocercophagus sp. nov. ahmad z. et al., genus pholetesor and description of three new species from india 35 discussion the genus pholetesor almost exclusively attacks leaf-mining lepidoptera (whitfield 2006). prior to the current study, only one species of pholetesor was described from india. however, within such a large expanse of land, covering more 3.2 million km2, many more species await discovery. approximately 21% of the country’s landmass is covered by forests (tree canopy density >10%), of which 12% comprises moderately or very dense forests (tree canopy density >40%) (cbd 2014). these include tropical rainforests of the andaman islands, the western ghats and northeast india; coniferous forests of himalaya; deciduous sal (shorea robusta gaertn.) forest of eastern india; the dry deciduous teak (tectona l.f. spp.) forest of central and southern india; and the babul (acacia mill.) dominated thorn forest of the central deccan and western gangetic plain (tritsch 2001). our findings of three species of the genus pholetesor reared from the leaf-mining lepidopteran larvae from the trees near the agrarian ecosystem provide important evidence that the wild area, as well as suburban areas (city parks) near the agrarian ecosystem could have more species fauna in the surrounding territories (owen et al. 1981; ahmad & ghramh 2018; banaszak-cibicka et al. 2018; ahmad et al. 2019; ghramh et al. 2019). the north indian landscape is covered mostly by the agrarian ecosystem, where lepidoptera are the key pests. the agrarian ecosystem is nevertheless rich in natural habitats, including small pockets of natural forests, and hosts a very diverse array of insects (cbd 2014). indeed, the presence of a natural wild reserve, near to suburban areas, can play a crucial role as a biodiversity reservoir from which beneficial species can spill over and colonize or decolonize perturbed areas. among the three new species two, p. acrocercophagus sp. nov. and p. indicus sp. nov., are readily placed in the ʻcircumscriptus-groupʼ of species of pholetesor. both the new species were differentiated from p. circumscriptus and p. bicolor, two species with a wide distribution range and demonstrating quite a large degree of variability in morphology according to their host and ecological preferences. owing to this morphological variation, the delimitation of the new species is quite problematic. shaw (2012) clearly addressed this problem and provides the detailed discussion about differentiating p. bicolor and p. circumscriptus. shaw (2012) has added strong evidence (biological and morphological data) that support p. bicolor and p. circumscriptus being considered as different species and also mentioned that these species are likely to be a complex of cryptic species. other two species, p. camerariae sp. nov. and p. hayati can be placed under ʻornigis-groupʼ of species of the genus pholetesor on the basis of the overall shape and sculpture patterns of propodeum and first metasoma tergite. acknowledgements the first author (za) extends his gratitude to the deanship of scientific research at king khalid university for funding through research program (rcams)-04/20. the first two authors (za and hag) also extend their appreciation to the honey bee research centre, king khalid university, abha, ksa. we also thank the editor, associate editor, and anonymous reviewers for their valuable and helpful comments to improve the quality of the manuscript. references ahmad z. & ghramh h.a. 2018. a new species of chelonus (areselonus) (hymenoptera, braconidae) from india reared from acrocercops lysibathra (meyrick) on cordia latifolia roxb. zookeys 737: 75– 80. https://doi.org/10.3897/zookeys.737.20835 ahmad z., al ghramh h.a., pandey k. & khan f. 2019. two new species of dolichogenidea (hymenoptera: braconidae: microgastrinae) parasitoids of leafminer lepidoptera from india. biologia 74: 1014–1018. https://doi.org/10.2478/s11756-019-00351-7 https://doi.org/10.3897/zookeys.737.20835 https://doi.org/10.2478/s11756-019-00351-7 european journal of taxonomy 726: 24–37 (2020) 36 akhtar m.s., ahmad z. & ramamurthy v.v. 2010. description of two new species of microgastrini (hymenoptera: braconidae) from india. zootaxa 2608: 57–62. https://doi.org/10.11646/zootaxa.2608.1.4 banaszak-cibicka w., twerd l. fliszkiewicz m. giejdasz k. & langowska a. 2018. city parks vs. natural areas – is it possible to preserve a natural level of bee richness and abundance in a city park? urban ecosystems 21 (4): 599–613. https://doi.org/10.1007/s11252-018-0756-8 cbd. 2014. india’s fifth national report to the convention on biological diversity. ministry of environment and forests, government of india, new delhi. csoka g. 2003. levélaknák és levélaknázók [leaf mines and leaf miners]. agro in form kiadóés nyomda, budapest. 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meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. https://doi.org/10.1007/978-94-009-1874-0 https://doi.org/10.11646/zootaxa.1144.1.1 https://doi.org/10.1017/s0007485300020393 another puzzle piece in the systematics of the chewing louse genus myrsidea, with a description of a new genus apomyrsidea stanislav kolencik 1,*, oldřich sychra 2 & julie m. allen 3 1,3 department of biology, university of nevada, reno, nv 89557, usa. 2 department of biology and wildlife diseases, faculty of veterinary hygiene and ecology, university of veterinary sciences brno, palackeho tr. 1946/1, 612 42 brno, czech republic. * corresponding author: skolencik@unr.edu 2 email: sychrao@vfu.cz 3 email: jallen23@unr.edu 1 urn:lsid:zoobank.org:author:af343a9b-3d7d-45d7-9fc5-eee51f990ba4 2 urn:lsid:zoobank.org:author:d28ceafb-0f34-4937-a66e-6ac8ba90e325 3 urn:lsid:zoobank.org:author:2bb2810c-80d1-41c7-8ddf-1f8e0230e94e abstract. a new avian chewing louse genus apomyrsidea gen. nov. is described based on species parasitizing birds in the family formicariidae. diagnostic characteristics and phylogenetic analyses were used to evaluate and confirm the generic status and merit its recognition as unique and different from myrsidea waterston, 1915. three species previously belonging to the genus myrsidea are placed in the new genus apomyrsidea gen. nov. and are discussed: apomyrsidea circumsternata (valim & weckstein, 2013) gen. et comb. nov., apomyrsidea isacantha (valim & weckstein, 2013) gen. et comb. nov. and apomyrsidea klimesi (sychra in sychra et al., 2006) gen. et comb. nov. keywords. phthiraptera, apomyrsidea, myrsidea, morphology, phylogeny. kolencik s., sychra o. & allen j.m. 2021. another puzzle in the systematics of the chewing louse genus myrsidea, with a description of a new genus apomyrsidea. european journal of taxonomy 748: 36–50. https://doi.org/10.5852/ejt.2021.748.1339 introduction chewing lice (psocodea hennig, 1966: phthiraptera haeckel, 1896) are obligate permanent ectoparasites, meaning they live their entire life cycle on their hosts (clayton et al. 2015). many are microhabitat specialists, often occupying specific body parts on the host (johnson et al. 2012). the species parasitizing birds feed on feathers, dead skin, skin secretions, and in some cases blood (price et al. 2003). at high prevalence, they can cause significant harm to feathers, skin irritation and even possible secondary infections that decrease the fitness of the bird hosts (møller et al. 1990; mullen & durden 2002). in the past, the taxonomy of chewing lice was based on morphology (clay 1966; price et al. 2003); however, in the last two decades, molecular and phylogenetic studies have helped to resolve the european journal of taxonomy 748: 36–50 issn 2118-9773 https://doi.org/10.5852/ejt.2021.748.1339 www.europeanjournaloftaxonomy.eu 2021 · kolencik s. et al. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:dafb8dde-a471-4274-b437-4606924be482 36 mailto:skolencik%40unr.edu?subject= mailto:sychrao%40vfu.cz?subject= mailto:jallen23%40unr.edu?subject= http://zoobank.org/urn:lsid:zoobank.org:author:af343a9b-3d7d-45d7-9fc5-eee51f990ba4 http://zoobank.org/urn:lsid:zoobank.org:author:d28ceafb-0f34-4937-a66e-6ac8ba90e325 http://zoobank.org/urn:lsid:zoobank.org:author:2bb2810c-80d1-41c7-8ddf-1f8e0230e94e https://doi.org/10.5852/ejt.2021.748.1339 https://doi.org/10.5852/ejt.2021.748.1339 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:dafb8dde-a471-4274-b437-4606924be482 systematics of this diverse group (johnson et al. 2004; smith et al. 2011). the chewing lice genus myrsidea waterston, 1915 is among the most diverse and host specific genera within the parasitic lice (phthiraptera), with more than 380 species described worldwide (kolencik & sychra, unpublished data). while there are 49 species of myrsidea described from neotropical suboscine birds (kolencik et al. 2018), only three have been described from birds of the family formicariidae gray, 1840 (sychra et al. 2006; valim & weckstein 2013). these three species show a set of characteristics that are unique among all species of myrsidea to the extent that sychra et al. (2006) and valim & weckstein (2013) suggested that they may be placed outside of myrsidea. here, we evaluate both morphological and molecular data of these three species along with representatives of the major clades in myrsidea. we find that the combination of differences found from both morphological and molecular data is strong enough to merit their recognition in a separate genus here named as apomyrsidea gen. nov. material and methods morphology all the morphological descriptions and characters as well as the terminology of chaetotaxy were taken from or follow those from clay (1961, 1962, 1966, 1969), sychra et al. (2006) and valim & weckstein (2013). taxonomy and nomenclature of the birds follows ioc bird world list ver. 10.1 (gill et al. 2020). phylogenetic reconstruction for the phylogenetic reconstruction we used a fragment of the mitochondrial gene cytochrome oxidase i (coi, 379bp). we amplified and sequenced this fragment from apomyrsidea klimesi (sychra in sychra et al., 2006) (3 nymphs ex formicarius analis (d’orbigny & lafresnaye, 1837) from zona protectora las tablas on the pacific slope of the cordillera de talamanca, 8°54′ n, 82°47′ w; 1300 m a.s.l.; identical sequences; genbank accession number: mw381016) using the technique described by martinu et al. (2015). all other sequences were downloaded from genbank (table 1) and aligned in seaview ver. 4.7 (gouy et al. 2010) using clustal omega (sievers et al. 2011) and checked visually. in total, we aligned 50 sequences from seven chewing lice genera from the suborder amblycera kellogg, 1896 and two sequences from the suborder ischnocera kellogg, 1896 as the outgroups. to determine the best model fit to our data we used partitionfinder ver. 2.1.1 (pf; lanfear et al. 2016) and compared all models using the corrected akaike information criterion (aicc). we selected two different approaches for a phylogenetic reconstruction to examine the similarity among the methods. first, the randomly accelerated maximum likelihood (raxml ver. 8.2.12; stamatakis 2014) method was used with three partitions and the model gtr + i + g for each and estimated 1000 bootstrap trees. second, mrbayes ver. 3.2 (ronquist & huelsenbeck 2003) was used for bayesian analysis with gtr + i + g model for two partitions and hky + g for the third. we conducted two parallel runs for 20 000 000 generations with four markov chains (huelsenbeck & bollback 2001), which were sampled every 1000 generations with a total of 20 000 parameter points estimates. markov chains were examined in tracer ver. 1.7.1 (rambaut et al. 2018) to determine that the chains had reached stationarity and 10% of trees were removed as burnin. a 50% majority rule consensus tree with posterior probabilities was generated. the outgroup ischnoceran genus philopterus nitzsch, 1818 was used for rooting in both trees. institutional abbreviations fmnh = field museum of natural history, chicago, usa inbio = instituto nacional de biodiversidad, costa rica mmbc = moravian museum, brno, czech republic mzusp = museum of zoology of the university of são paulo, brazil nsf = national science foundation, usa kolencik s. et al., a new louse genus apomyrsidea gen. nov. (phthiraptera) 37 lice species host species genbank number actornithophilus ceruleus anous tenuirostris af545666 actornithophilus erinaceus rostratula benghalensis af545667 actornithophilus piceus larus sp. af545668 apomyrsidea circumsternata* formicarius colma kf048105 apomyrsidea isacantha* chamaeza nobilis kf048115 apomyrsidea klimesi† formicarius analis mw381016 colpocephalum fregili corvus albus mf443951 colpocephalum indi ictinia mississippiensis mf443945 colpocephalum napiforme buteo lagopus mf443947 colpocephalum polybori caracara cheriway mf443950 colpocephalum subzerafae falco berigora mf443946 colpocephalum turbinatum circus approximans mf443944 dennyus bartoni aerodramus mearnsi dq139300 dennyus carljonesi carljonesi aerodramus v. vestitus dq139294 dennyus carljonesi forresteri aerodramus elaphrus dq139297 dennyus carljonesi forresteri aerodramus francicus dq139296 dennyus kristinae aerodramus s. spodiopygius dq139305 dennyus mimirogerorum aerodramus papuensis dq139301 dennyus singhi aerodramus assimilis dq139307 menacanthus alaudae calamonastes fasciolatus mg682392 menacanthus alaudae plocepasser mahali mg682398 menacanthus camelinus lanius collaris mg682399 menacanthus eurysternus curruca subcoerulea mg682400 menacanthus eurysternus prinia flavicans mg682401 menacanthus eurysternus turdus libonyana mg682393 menacanthus eurysternus melaenornis silens mg682384 menacanthus eurysternus turdus merula kj730692 menacanthus sp. lagonosticta rara dq887248 menacanthus sp. pogoniulus chrysoconus mg682396 menacanthus takayamai locustella luscinioides kj730535 myrsidea alexanderi pheugopedius maculipectus mf563536 myrsidea argentina spinus magellanicus ky113129 myrsidea cecilae ramphastos culminatus × ariel kf048126 myrsidea cf. bubalornithis bubalornis niger mg682394 myrsidea cf. textoris ploceus ocularis mg682397 myrsidea cinnamomei attila citriniventris kf048110 myrsidea contopi leptopogon amaurocephalus mf563537 myrsidea cruickshanki chlorothraupis carmioli gq454449 myrsidea icterocephalae tangara icterocephala kf048103 myrsidea incerta catharus ustulatus fj171268 myrsidea seminuda thraupis palmarum kf048116 myrsidea valimi euphonia anneae gq454450 table 1 (continued on next page). list of the louse species included in our phylogenetic analyses, with their hosts and genbank accession numbers. * = in genbank as species of myrsidea; † = a new sequence. european journal of taxonomy 748: 36–50 (2021) 38 results here, we find that species of apomyrsidea gen. nov. are unique and morphologically distinct from all species of myrsidea. specifically, the simple combination of the presence of the posterior dorsal head seta (dhs) 23 and the dorso-central pronotal setae (dps) 2 (fig. 3), sternite i lying inside the wide notch of sternite ii (fig. 5) and different type of fusion in the terminal sternites – male abdominal sternite viii and female sternite vii both at least partially but conspicuously separated from the subgenital plate, which is formed by a single sternite ix (in the male; fig. 2) or fusion of sternites viii and ix (in the female; fig. 1), make these species distinct. these characters were further defined in clay (1966), sychra et al. (2006) and valim & weckstein (2013). this separation is also supported with the present phylogenetic analyses of 379bp fragment of coi from 50 amblyceran species across 6 genera: actornithophilus ferris, 1916, colpocephalum nitzsch, 1818, dennyus neumann, 1906, myrsidea (including all three species of myrsidea occurring on formicariid hosts that are here placed in apomyrsidea gen. nov.), menacanthus neumann, 1912 and ricinus de geer, 1778, and with two ischnoceran lice, traihoriella laticeps (piaget, 1888) and philopterus solus (tendeiro, 1962), as outgroups (fig. 10). both bayesian analysis and maximum likelihood analysis (appendix) resulted in trees with a similar topology, differing in some intra-generic relationships. however, both trees coincide in topology of our main focus, ‘myrsidea from formicariidae’ – apomyrsidea gen. nov., and place it sister to all other species of myrsidea. class insecta linnaeus, 1758 superorder psocodea hennig, 1966 order phthiraptera haeckel, 1896 suborder amblycera kellogg, 1896 family menoponidae mjöberg, 1910 genus apomyrsidea gen. nov. urn:lsid:zoobank.org:act:5a86df15-251d-4262-9bdf-92a3e1be9c6c myrsidea waterston, 1915: 12 (in partim). lice species host species genbank number myrsidea violaceae euphonia hirundinacea ky113137 ricinus mugimaki cossypha dichroa kf768816 ricinus sp. atila spadiceus af545762 ricinus sp. cyanocompsa parellina af545763 ricinus sp. ficedula hyperythra af545764 ricinus sp. fringillaria tahapisi mg682387 ricinus sp. platysteira laticincta mk032013 ricinus sp. terpsiphone batesi ku187311 ricinus sp. terpsiphone viridis ky359404 outgroups philopterus solus rhinopomastus cyanomelas mg682416 traihoriella laticeps andigena nigrirostris ay149398 table 1 (continued). list of the louse species included in our phylogenetic analyses, with their hosts and genbank accession numbers. * = in genbank as species of myrsidea; † = a new sequence. kolencik s. et al., a new louse genus apomyrsidea gen. nov. (phthiraptera) 39 http://zoobank.org/urn:lsid:zoobank.org:act:5a86df15-251d-4262-9bdf-92a3e1be9c6c type species apomyrsidea klimesi (sychra in sychra et al., 2006: 55) gen. et comb. nov. diagnosis apomyrsidea can be characterized and distinguished from all other menoponid chewing lice genera with the combination of following characteristics: head rounded anteriorly, lacking lateral slit or notch, without sclerotized processes (oral spines) arising near the base of maxillary palpi (figs 1–3); figs 1–2. dorso-ventral view of apomyrsidea klimesi (sychra in sychra et al., 2006) gen. et comb. nov. (cr15). 1. ♀. 2. ♂. european journal of taxonomy 748: 36–50 (2021) 40 alveoli of dorsal head setae (dhs; marginal temporal setae by clay 1969) 26 and 27 not closely associated (fig. 3); dhs 18 (outer mid-dorsal head seta by clay (1966) or dorsal head seta „d“ by clay (1962) is missing; dhs 22 (outer occipital seta by valim & weckstein 2013 or posterior dorsal head seta „f“ by clay 1962) approximately as long as dhs 21 (inner occipital seta by valim & weckstein 2013), both surpassing pronotal carina; dhs 23 (posterior dorsal head seta “e” in clay 1962) present and anteriorly to line of bases of dhs 21 and 22 (fig. 3); head sensilla 3–5 sensu clay (1961) or c–e sensu clay (1969) absent; characteristic gular plate with the greater length and thickness of the posterior pair of setae compared to the rest ones (figs 1–2). figs 3–9. 3–6. apomyrsidea klimesi (sychra in sychra et al., 2006) gen. et comb. nov. (cr15). 3. dorsal view of head, prothorax and mesothorax of female. 4. prosternal plate of female. 5. metasternal plate and sternites i–iii of female. 6. male genitalia. – 7–9. male genital sac sclerites. 7. a. klimesi gen. et comb. nov. 8. a. circumsternata (valim & weckstein, 2013) gen. et comb. nov. 9. a. isacantha (valim & weckstein, 2013) gen. et comb. nov. abbreviations: a–b = head sensilla; dps = dorso-central pronotal setae; mps = marginal prothoracic setae; 8–30 = dorsal head setae. figs 7–9 are drawn to the same scale. kolencik s. et al., a new louse genus apomyrsidea gen. nov. (phthiraptera) 41 fig. 10. bayesian phylogenetic tree of selected amblyceran genera and species based on partial mitochondrial gene coi. tree is rooted with outgroup species philopterus solus (tendeiro, 1962). posterior probability values are shown above the nodes (values < 50% are not shown). blue colour indicates species of myrsidea waterston, 1915 (m); green colour indicates species of apomyrsidea gen. nov. (a); * symbol indicates a node between myrsidea and apomyrsidea. european journal of taxonomy 748: 36–50 (2021) 42 thorax prosternal plate well developed with straight anterior margin and two anterior setae (fig. 4); pronotum with one pair of minute dorso-central pronotal setae lying near the transverse carina (dps 2 by clay 1962) (fig. 3); pronotum with anterolateral pronotal setae (marginal prothoracic setae 1–3, mps 1–3 in clay 1962) in following arrangement: mps 1 and mps 3 spine-like, mps 2 fine and long; mps 1 and mps 2 located on each lateral corner of pronotum, with mps 3 posteriorly to mps 2 on pronotal margin (fig. 3); mesonotum well defined with only two anterior setae; mesonotum without median division, but with a y-shaped line just below postnotum, not forming a suture or even splitting mesonotum (fig. 3). this y-shaped line slightly less evident, but also discernible in good specimens from other host families; strongly sclerotized ring-like mesothorax – mesothorax with sternum, pleura and tergum fused to form strongly sclerotized ring round the body; femur iii without combs of spine-like setae but with thick or sparse brushes of setae. abdomen sternite i mostly surrounded by sternite ii (it lies inside the wide notch of sternite ii) (fig. 5); sternite ii enlarged with a clutch of heavy spine-like setae at each posterior-lateral margin called aster; male genitalia as in fig. 6. male genital sac sclerite with two roughly serrated spiculated lateral arms (figs 7–9); female vulva with smooth posterior margin (fig. 1); female ventral anal margin without lateral seta-bearing processes (see clay 1969); sternite vii fused with viii + ix + x, forming female subgenital plate, although with a distinct transverse fenestra distinctly enclosed at lateral sides of subgenital plate where seventh and eighth segments fused (in a. circumsternata and a. isacantha; see valim & weckstein 2013: fig. 11); in the case of a. klimesi – male abdominal sternite viii and female sternite vii both separated from the subgenital plate, which is formed by a single sternite ix (in the male) or fusion of sternites viii and ix (in the female) (figs 1–2). etymology the generic name apomyrsidea is formed by a combination of greek word ‘apo’ = ‘from’ and myrsidea, referring that it is separated from the genus myrsidea, where it was originally placed. the gender is feminine. included species three species are included in the apomyrsidea gen. nov., all are restricted to formicariid hosts: apomyrsidea circumsternata (valim & weckstein, 2013) gen. et comb. nov. apomyrsidea isacantha (valim & weckstein, 2013) gen. et comb. nov. apomyrsidea klimesi (sychra in sychra et al., 2006) gen. et comb. nov. descriptions of all three species are well presented in the original papers (sychra et al. 2006; valim & weckstein 2013). valim & weckstein (2013) also presented a key to their identification. apomyrsidea circumsternata (valim & weckstein, 2013) gen. et comb. nov. figs 8, 10 myrsidea circumsternata valim & weckstein, 2013: 383, figs 3–4, 13–15, 17, 19, 22 (type host: formicarius colma boddaert, 1783). kolencik s. et al., a new louse genus apomyrsidea gen. nov. (phthiraptera) 43 material examined holotype brazil • ♀; rio acanauí, município japurá, amazonas; 2°01′38″ s; 66°40′28″ w; 20 jul. 2007; weckstein leg.; ex formicarius colma boddaert, 1783; mzusp 2314. paratypes (2 ♀♀, 3 ♂♂) brazil • 1 ♂; same collection data as for holotype; fmnh-ins 94002 • 1 ♀; same collection data as for holotype; dna voucher mysp.foco.1.4.2011.3; fmnh-ins 94003 • 2 ♂♂; same collection data as for holotype; mzusp 2316, mzusp 2317 • 1 ♀; same collection data as for holotype; mzusp 2315. apomyrsidea isacantha (valim & weckstein, 2013) gen. et comb. nov. figs 9–10 myrsidea isacantha valim & weckstein, 2013: 381, figs 1–2, 11–12, 16, 18, 20–21 (type host: chamaeza nobilis gould, 1855). material examined holotype brazil • ♀; rio acanauí, município japurá, amazonas; 2°01′38″ s, 66°40′28″ w; 18 jul. 2007; weckstein leg.; ex chamaeza nobilis gould, 1855; mzusp 2310. paratypes (2 ♀♀, 3 ♂♂) brazil • 1 ♂; same collection data as for holotype; fmnh-ins 94000 • 1 ♀; same collection data as for holotype; dna voucher mysp.chno.1.4.2011.4; fmnh-ins 94001 • 2 ♂♂; same collection data as for holotype; mzusp 2312, mzusp 2313 • 1 ♀; same collection data as for holotype; mzusp 2311. apomyrsidea klimesi (sychra in sychra et al., 2006) gen. et comb. nov. figs 1–7, 10 myrsidea klimesi sychra in sychra et al., 2006: 55, figs 10–11, 14–15 (type host: formicarius analis (d’orbigny & lafresnaye, 1837)). material examined holotype costa rica • ♀; hitoy cerere br, provincia limón; 9°40′ n, 85°27′ w; 100 m a.s.l.; 27 aug. 2004; literak, capek and havlicek leg.; ex formicarius analis (d’orbigny & lafresnaye, 1837); inbio o.sychra cr15. allotype costa rica • ♂; same collection data as for holotype; inbio o.sychra cr15. paratypes (1 ♀, 1 ♂) costa rica • 1 ♀, 1 ♂; same collection data as for holotype; 27 and 31 aug. 2004; inbio o.sychra cr14, cr16. other material costa rica • 1 ♂; zona protectora las tablas on the pacific slope of the cordillera de talamanca; 8°54′ n, 82°47′ w; 1300 m a.s.l.; 21. aug. 2010; sychra and literak leg.; ex formicarius analis; mmbc o.sychra cr226. european journal of taxonomy 748: 36–50 (2021) 44 discussion in the key to the genera of the menoponidae mjöberg, 1910 of clay (1969), apomyrsidea gen. nov. would key to couplet 33 being closest to myrsidea and oculomenopon price & hellenthal, 2005. all three genera share unique diagnostic characters including: characteristic gular plate with the greater length and thickness of the posterior pair of setae compared to the other ones; missing dhs 18; well-developed prosternal plate with two anterior setae; strongly sclerotized ring-like mesothorax – mesothorax with sternum, pleura and tergum fused to form strongly sclerotized ring round the body; and general shape of male genitalia. moreover, enlarged sternite ii with aster of heavy setae at each posterior corner is a common feature of apomyrsidea gen. nov. and myrsidea, while presence of dhs 23 and the similar size of dhs 21 and dhs 22 is shared between apomyrsidea gen. nov. and oculomenopon (price & hellenthal 2005). the most important key characters useful for separating apomyrsidea gen. nov. from all species of myrsidea and oculomenopon are: the presence of dps 2, sternite i lying inside the wide notch of sternite ii and different type of fusion in the terminal sternites – male abdominal sternite viii and female sternite vii both at least partially but conspicuously separated from the subgenital plate, which is formed by a single sternite ix (in the male) or fusion of sternites viii and ix (in the female). here, we provide a key for these three genera that modifies and extends the first part of couplet 33 of the clay’s (1969) key as follows: 33. prosternum with 2 central setae ....................................................................................................33a 33a. dps 2 present; sternite i mostly surrounded by sternite ii (it lies inside the wide notch on anterior margin of sternite ii) ..................................................................................... apomyrsidea gen. nov. – both dps absent; sternite i lies above the sternite ii that has straight anterior margin without notch .......................................................................................................................................... 33b 33b. dhs 23 absent ..........................................................................................myrsidea waterston, 1915 – dhs 23 present ................................................................. oculomenopon price & hellenthal, 2005 valim & weckstein (2013) also discussed the presence of dps 2 and stated that, according to clay (1966), the absence of dorso-central pronotal setae (dps 1 and dps 2) is one of defining characteristics of the genus myrsidea. they wrongly interpreted clay (1969), when writing: “clay (1969) presented data showing that some species of myrsidea could have at least one pair of dorso-central pronotal setae (see couplet 32)”. when we checked couplet 32 in the key by clay (1969), there is a note “not more than one pair of dorso-central pronotal setae”. moreover, clay (1969: 11) wrote: “there are usually two pairs of dorso-central pronotal setae (dps) lying on or near the transverse carina, but in some species-groups (clay 1962: 237) or genera (myrsidea) they are reduced to one pair or absent.” we have examined many slides across groups and species of myrsidea and can confirm that dps are absent apart from the newly erected apomyrsidea. another interesting character is the presence of a cluster of heavy setae on each side of sternite iii (fig. 5). these setae are not as long as the aster, but they are heavier than spine-like setae on the lateral sides of other sternites and they are more conspicuous on females than on males. despite this, it cannot be used as a basic character of the genus, because it is present only in two species – a. circumsternata gen. et comb. nov. and a. klimesi gen. et comb. nov. an unusual cluster of three spine-like setae situated on one side of sternite iii was described by klockenhoff (1984) from only one female of m. serini (séguy, 1944). to our knowledge, no other author referred to such a structure on sternite iii and we have not found it in any other examined myrsidea. kolencik s. et al., a new louse genus apomyrsidea gen. nov. (phthiraptera) 45 a partial division of terminal sternites in females is known for example in some myrsidea from bulbuls (pycnonotidae g.r. gray, 1840; hellenthal & price 2003), or myrsidea abbreviata eichler, 1951 from ramphastos dicolorus linnaeus, 1766 (piciformes meyer & wolf, 1810: ramphastidae vigors, 1825; price et al. 2004). however, this partial division is noticeable only at the lateral margins of the sternites of these species. a medial division or totally divided terminal sternites are only found in species of apomyrsidea. according to clay (1969), this is not necessarily a generic character, but in our opinion, it can be useful for separating apomyrsidea gen. nov. from myrsidea. another interesting character is the length of dhs 21 and dhs 22. while clay (1966) considered the short length of dhs 22 compared with the long dhs 21 as one of the determining characteristics for the genus myrsidea, apomyrsidea gen. nov. shows dhs 22 as long as dhs 21 and both exceed the pronotal carina. interestingly, the enigmatic genus ramphasticola carriker, 1949, which has been difficult to place, with different authors suggesting it should be nested within myrsidea (hopkins & clay 1952; price et al. 2003), has both species with long and short dhs 22, further suggesting a molecular phylogenetic analysis is needed to validate the placement of this genus. the presence of an aster of heavy setae at each posterior corner of sternite ii in apomyrsidea gen. nov. and myrsidea may bring some new challenges to determinations. the aster is one of the most visible features of many species of myrsidea, and easily seen even under a stereoscope. however, in some species of myrsidea it can be reduced (e.g., some species of ramphastidae), while in species of apomyrsidea gen. nov. it is always present. this changes the defining characteristic of the aster as the determining morphological feature for myrsidea and broadens the use of this characteristic to a second genus. henceforth, if no aster is present in a louse, this does not necessarily mean that it is not a myrsidea, but the presence of asters will narrow the selection to two genera: myrsidea and apomyrsidea gen. nov. this further supports this character as an important one for morphological identifications. beside unique morphological characteristics, valim & weckstein (2013) referred to these species as “myrsidea from formicariidae” and suggested they would be phylogenetically distinct from nearly all other species of myrsidea. therein, instead of describing the new genus, the authors decided to broaden the set of diagnostic characters used to define the genus myrsidea. however, in their phylogenetic tree myrsidea sp. (genbank kf048123) from myiarchus panamensis lawrence, 1861 (the avian family tyrannidae vigors, 1825), appears to be in the same clade with both species of apomyrsidea gen. nov. used in the analysis (valim & weckstein 2013). this may be a misidentification of the sample and needs validation. further, when this sequence was compared to all other coi sequences (a 379bp fragment of mitochondrial gene cytochrome oxidase subunit i) in genbank, the highest match was menacanthus sp. (genbank af545726) from the tyrannid attila spadiceus (gmelin j.f., 1789), with a p-distance of only 0.79%, and menacanthus sp. (genbank kj730539) from the parid poecile sp. with a p-distance of 1.31%. a comparison between these two species of menacanthus showed the p-distance to be 1.05%, which is likely low enough to confirm it as the same species. on the basis of these facts, we believe that sequence kf048123 is potentially an untrustful sequence and we excluded it from our analysis. when we used a broader range of genera in our study, apomyrsidea gen. nov. shows a similar pattern creating a well separated clade from all other species of myrsidea (fig. 10). the familiar issue with lower support values (mostly in basal nodes) is often present when using only a single gene fragment (e.g., kolencik et al. 2017). unfortunately, the sequences, and their quality, for amblyceran species submitted to the genbank database are limited and mostly only for a 379bp fragment of coi and/ or a 347bp fragment of nuclear ef-1a (elongation factor 1 alpha) gene. besides that, in many cases both sequences for the same specimen did not overlap; moreover, all three species of apomyrsidea gen. nov. have only data for coi. thus, it leaves a 379bp coi’s fragment as the only one eligible for this study. the coi topology shows that a. klimesi gen. et comb. nov. is sister to a. circumsternata gen. et european journal of taxonomy 748: 36–50 (2021) 46 comb. nov., in accordance with their morphology, which was also used in the key for their identification by valim & weckstein (2013). in conclusion, the combination of unique morphological characteristics and the well-separated clade in the phylogenetic tree are strong enough to confirm apomyrsidea gen. nov. as a new genus, and a sister taxon to myrsidea. acknowledgements this study was supported by nsf grant deb-1925312. we are indebted to michel valim (biotério da universidade iguaçu, nova iguaçu, rio de janeiro, brazil), for his long-term cooperation and years of discussion about the taxonomy of myrsidea and chewing lice in general. we are also very indebted to our colleagues jana martinu and ivo papousek for their help in amplifying and processing the sequences from apomyrsidea klimesi gen. et comb. nov. references clay t. 1961. a new genus and species of the menoponidae (mallophaga, insecta) from apteryx. annals and magazine of natural history (series 13) 3: 571–576. https://doi.org/10.1080/00222936008651059 clay t. 1962. a key to the species of actornithophilus ferris with notes and descriptions of new species. bulletin of the british museum (natural history) entomology 11: 189–244. clay t. 1966. contributions towards a revision of myrsidea waterston. i. 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(phthiraptera) 49 https://doi.org/10.11646/zootaxa.1206.1.3 https://doi.org/10.14411/fp.2013.040 https://doi.org/10.5962/bhl.part.7786 appendix maximum likelihood phylogenetic tree of selected amblyceran genera and species on the basis of partial mitochondrial gene coi, with gtr + i + g model of molecular evolution. bootstraps values are next to the nodes (values below 50% are not shown). m = species of myrsidea; a = species of apomyrsidea gen. nov.; * = a node between myrsidea and apomyrsidea gen. nov. european journal of taxonomy 748: 36–50 (2021) 50 1 european journal of taxonomy 760: 1–15 issn 2118-9773 https://doi.org/10.5852/ejt.2021.760.1435 www.europeanjournaloftaxonomy.eu 2021 · vivallo f. & zanella f.c.v. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:bc4ec862-87d2-4578-b9c0-113f68f7fe48 relicthemisia, a new subgenus of the oil-collecting bee genus centris fabricius, 1804 with notes on distribution and host plants of c. xanthomelaena moure & castro, 2001 (hymenoptera: apidae) felipe vivallo 1 & fernando césar vieira zanella 2,* 1 hymn, laboratório de hymenoptera, departamento de entomologia, museu nacional, universidade federal do rio de janeiro, quinta da boa vista, são cristóvão 20940‒040, rio de janeiro, rj, brazil. 2 pós-graduação em biodiversidade neotropical, universidade federal da integração latino-americana, av. tarquinio joslin dos santos, 1000, jardim universitário, 85867-000 foz do iguaçu, pr, brazil. * corresponding author: fcvzanella@gmail.com 1 email: fvivallo@yahoo.com 1 urn:lsid:zoobank.org:author:ac109712-1474-4b5d-897b-1ee51459e792 2 urn:lsid:zoobank.org:author:8b9287c1-56a8-4dbc-964e-1022268c8beb abstract. centris xanthomelaena moure & castro, 2001 is a relict species, endemic to northeastern brazil and broadly recorded within the semiarid region of caatinga xerophilous open vegetation. it was originally included in the subgenus paracentris cameron, 1903 but posteriorly interpreted as remotely related to it or to the subgenus centris s. str. fabricius, 1804. in this paper it is proposed to recognize this species as the single member of the monotypic relicthemisia, a new subgenus which belongs to the ‘centris group’, one of the main internal lineages of the genus. the proposition of this new subgenus is based on both, morphological and molecular data which indicate its long history as a distinct lineage. distribution records, floral hosts as well as photographs of both sexes of c. xanthomelaena are also provided. keywords. caatinga, centridini, distribution, endemism, systematics. vivallo f. & zanella f.c.v. 2021. relicthemisia, a new subgenus of the oil-collecting bee genus centris fabricius, 1804 with notes on distribution and host plants of c. xanthomelaena moure & castro, 2001 (hymenoptera: apidae). european journal of taxonomy 760: 1–15. https://doi.org/10.5852/ejt.2021.760.1435 introduction centris fabricius, 1804 is one of the most abundant and diverse genus of solitary bees in the neotropical region. the taxonomy of this group is quite complex, mainly due to the lack of updated revisions, as well as the large number of described species. as a way to recognize apparently natural groups, several subgenera https://doi.org/10.5852/ejt.2021.760.1435 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:bc4ec862-87d2-4578-b9c0-113f68f7fe48 https://orcid.org/0000-0002-4487-0804 https://orcid.org/0000-0002-4817-1373 mailto:fcvzanella%40gmail.com?subject= mailto:fvivallo%40yahoo.com?subject= http://zoobank.org/urn:lsid:zoobank.org:author:ac109712-1474-4b5d-897b-1ee51459e792 http://zoobank.org/urn:lsid:zoobank.org:author:8b9287c1-56a8-4dbc-964e-1022268c8beb https://doi.org/10.5852/ejt.2021.760.1435 european journal of taxonomy 760: 1–15 (2021) 2 have been described in it, with paracentris cameron, 1903 being one of the most diverse, both in number of species and morphology (vivallo 2020). according to the distribution pattern of the species of this subgenus, sensu zanella (2002), it corresponds to a group with amphitropical distribution, with a large number of species occurring in north and south america, being practically absent in central america (michener 1979). the complete taxonomic revision of the species of paracentris was published by vivallo (2020), following the interpretation of zanella (2002) who carried out a phylogenetic analysis using morphological data of a large part of the taxa present in south america. before that taxonomic revision, some species were included in paracentris intuitively or for practical reasons. an example of this is the species of the ‘hyptidis group’ vivallo & melo, 2009, whose species have been cited in the subgenera ptilocentris snelling, 1984; centris. s. str.; ptilotopus klug, 1810; wagenknechtia moure, 1950; as well as in paracentris (moure et al. 2009), with the species of this group recently located in a new subgenus anisoctenodes vivallo, 2020. a similar case was also observed for penthemisia moure, 1950, synonymized with paracentris by snelling (1966) and still kept as a junior synonym by michener (2007), which was later recognized as a distinct lineage exclusive from southern south america and reinstated as a distinct subgenus of centris (zanella 2002). another similar case is observed in a species apparently endemic to northeastern brazil described as c. (paracentris) xanthomelaena moure & castro, 2001 and whose phylogenetic affiliation has remained uncertain until now. moure et al. (2007) cited this species in the subgenus paracentris, although the results obtained by zanella (2002) indicated that it is not closely related to that subgenus, but it would correspond to a relict and undescribed lineage close to centris s. str. the phylogenetic relationships of c. xanthomelaena with other species of the genus were newly reconstructed by martins & melo (2015) in a study using molecular characters. according to those authors, this species would correspond to a lineage not closely related to centris s. str., but remotely related to the south american species of paracentris. despite this inconsistency, in both cases c. xanthomelaena appears as a relict and distinct lineage, which is part of a larger clade known as the ‘centris group’ (zanella 2002). considering the results obtained by zanella (2002) and martins & melo (2015), we formally propose a new monotypic subgenus, relicthemisia subgen. nov., in this paper, containing c. xanthomelaena as type species and recognizing it as another of the major internal lineages of centris. both morphological and molecular evidence cited above support and justify the proposition of this new taxon. material and methods general morphological terminology follows michener (2007). specimen labels were transcribed under the section ‘material examined’. the backward slash (\) indicates different labels on the pin of the specimen. specimens marked with a cross ‘[†]’ were lost in the fire of the museu nacional of rio de janeiro on september 2nd, 2018. the literature cited below the name of the species and above its diagnosis corresponds to an update of the information presented by moure et al. (2007) in the catalogue of bees in the neotropical region. photographs were taken using a leica dfc 450 camera attached to a leica m205c stereo microscope and using extended-focus software leica application suite ver. 4.8.0. all images were prepared using combinezp ver. 7.0.0.1 software, and then enhanced with adobe photoshop® (ver. 7.0) without distorting the morphological characters of the specimens. the distribution map was created using arcview software (ver. 3.2 gis) and prepared from locality records taken from specimen labels and from records available in literature listed in the section ‘references’. the biogeographical provinces are according to morrone (2014) and were implemented vivallo f. & zanella f.c.v., relicthemisia, new subgenus of centris bees 3 with the shape file provided by löwenberg-neto (2014). this was also done for the compilation of the floral records (table 1). plant names were checked and updated according to the international plant names index (ipni.org). new distribution and/or floral records were marked with an asterisk (*). institutional abbreviations dzup = coleção de entomologia p.j.s. moure, departamento de zoologia, universidade federal do paraná, curitiba, paraná, brazil labe/ebda = laboratório de abelhas da empresa bahiana de desenvolvimento agrícola, salvador, bahia, brazil mnrj = museu nacional, universidade federal do rio de janeiro, rio de janeiro, brazil rpsp = coleção entomológica ‘prof. j.m.f. camargo’, universidade de são paulo, campus ribeirão preto, são paulo, brazil unila = coleção entomológica universidade federal da integração latino-americana, foz do iguaçu, paraná, brazil results systematics class insecta linnaeus, 1758 order hymenoptera linnaeus, 1758 family apidae latreille, 1802 tribe centridini cockerell & cockerell, 1901 genus centris fabricius, 1804 relicthemisia subgen. nov. urn:lsid:zoobank.org:act:361e0ae3-2711-4089-974a-da742a130e52 figs 1–4 type species centris xanthomelaena moure & castro, 2001 diagnosis integument dark brown to black, clypeus and labrum coriaceous with coarse and dense punctation, but the clypeus with an unpunctated area as a median longitudinal band on upper half (fig. 1a). female inner orbits of compound eyes converging downward (fig. 1a). mandible with four apically acute teeth, the fourth tooth slightly larger than the third (fig. 1a). basitibial plate elliptical, with s-like secondary plate (fig. 2a). elaiospathes normally developed. male clypeus, except lateral areas, and labrum yellow (fig. 1c). yellow spots on paraocular and supraclypeal areas (fig. 1c). apical margin of t7 with strong emargination (fig. 2b). s7 without emargination on the basal border (fig. 2b). s8, apical projection clearly defined, larger at middle and with rounded apex (fig. 2c). genital capsule with long dorsoapical projection of gonocoxite, ca 2/3 lengths of gonostylus (fig. 2e). etymology from latin ‘relictus’ (a survivor from a previous age) plus ‘hemisia’ (a junior synonym of centris) due to the antiquity and isolation of this lineage. https://ipni.org http://zoobank.org/urn:lsid:zoobank.org:act:361e0ae3-2711-4089-974a-da742a130e52 european journal of taxonomy 760: 1–15 (2021) 4 remarks centris xanthomelaena, the only species of the new subgenus relicthemisia, was recognized as a distinct lineage with no close relationship to other species, based on morphological (zanella 2002) and molecular data (martins & melo 2015). depending on the study, the phylogenetic position of this lineage was different, either as sister group of centris s. str. or paracentris cameron, 1903 respectively. nevertheless, it was always recovered as a distinct and relatively old lineage within the ‘centris group’. according to martins & melo (2015), c. xanthomelaena diverged from a south american clade formed by paracentris around 18 million years ago, at about the same time when the major lineages within the ‘centris group’ diverged from each other. the hypothetical relationship of this species with centris s. str. was based on the interpretation of two morphological characters: the strong emargination on the apical margin of t7 (fig. 2b; character 25: 0 in zanella 2002) and the short and wide translucent laminar projection on the dorsodistal region of the gonocoxite at the base of the long, giant bristles (fig. 2e–f; character 44: 1 in zanella 2002), but the states present in c. xanthomelaena are clearly unique and cannot be homologous to those present in species of centris s. str. despite the fact that new phylogenetic analyses using a higher number of terminals of centris s. str. and paracentris can provide new information regarding the history and relatedness of the lineage of c. xanthomelaena, its distinctness and old history are well supported (see martins & melo 2015). besides the uniqueness of the intense yellow slightly greenish pilosity covering the head, mesosoma (except the ventral surface) and on the anterior half of t1 that allow to easily recognize c. xanthomelaena from other species of the genus (fig. 1a–d), this monotypic subgenus presents a unique combination of characters of the male’s genitalia that distinguishes it from the other members of the ‘centris group’: an emargination on the apical margin of t7 (fig. 2b); a long dorsoapical projection of gonocoxite, ca 2/3 lengths of gonostylus (fig. 2e); the dorsomedial projections of the genital capsule (fig. 2e), as well as the s-like lower margin of the female’s secondary basitibial plate (fig. 2a). centris (relicthemisia) xanthomelaena moure & castro, 2001 figs 1–4 centris ‘xanthomelaena’ vogel & machado, 1991: 163–175, figs 6a–b, 9a, h (distribution, floral records, pollination). nomen nudum. centris (paracentris) xanthomelaena – moure & castro 2001: 330–332, figs 1–4 (original description). — silveira et al. 2002: 98, 253 (distribution, list). — urban 2003: 24, 43 (taxonomic note, cited). — azevedo & silveira 2005: 45 (cited). — batalha filho et al. 2007: 25 (distribution record). — moure et al. 2007: 120 (catalogue). — azevedo et al. 2008: 143 (distribution record). — machado & sazima 2008: 488 (floral record). — rodarte et al. 2008: 307 (distribution record). — pigozzo & viana 2010: 105 (distribution record). — vivallo & zanella 2012: 4, 6, 8–9, 13– 14, figs 37–38, 77–78 (distribution, key). — giannini et al. 2013: 78 (list). — silva 2014: 188 (distribution record). — martins et al. 2018: 770, figs 1–2 (bionomy, nesting behavior, distribution record). — barenbaum 2019: 222 (bionomy, sleeping behavior). — carneiro et al. 2019: 219 (distribution and floral records). centris xanthomelaena – zanella 2002: 438, 444, 447, 451, 453, 455, 457, 459, 483, 485, 486, figs 26, 164–170 (diagnosis, male description, distribution map, morphological characters, phylogenetic relationships). — aguiar 2003a: 42, 43 (distribution and floral records); 2003b: 464 (floral record). — zanella 2003: 234 (list). — aguiar et al. 2005: 249 (distribution record). — aguiar & zanella vivallo f. & zanella f.c.v., relicthemisia, new subgenus of centris bees 5 2005: 17, 19 (distribution record). — azevedo & silveira 2005: 47 (cited). — prevedello & carvalho 2006: 45 (cited). — zanella & vivallo 2009: 68 (cited). — martins & melo 2015: 7–8, 10 (phylogenetic relationships). — martins et al. 2018: 771 , 772 (sleeping behavior). — carneiro et al. 2019: 216, 220 (distribution and floral records, pollinator). diagnosis female integument dark brown to black, except flagellum dark brown and tegula yellowish brown (fig. 1a–b). wings brown with veins dark brown (fig. 1b). head, mesosoma (except the ventral surface) and anterior half of t1 with intense yellow slightly greenish pilosity, lighter on labrum and gena (fig. 1a–b). the rest of the body with blackish hairs, except posterior apex of the femur of forelegs with some yellowish hairs (fig. 1a–b). clypeus coriaceous with coarse and dense punctation (fig. 1a). clypeal disc with an unpunctated area on upper half, without smooth longitudinal band. labrum with the same punctation, but denser, without smooth basal margin. terga and sterna, except t6 and s6, with very narrow smooth distal margin, wider on t4. mandible with four apically acute teeth (fig. 1a). fourth teeth slightly larger fig. 1. centris (relicthemisia) xanthomelaena moure & castro, 2001. a –b. ♀. a. frontal view. b. habitus, lateral view. c–d. ♂. c. frontal view. d. habitus, lateral view. scale bars: a, c = 2 mm; b, d = 5 mm. european journal of taxonomy 760: 1–15 (2021) 6 than the third (fig. 1a). maxillary palpus 4-segmented. malar area very narrow (fig. 1a). labrum semicircular (fig. 1a). inner orbits of compound eyes converging downward (fig. 1a). elaiospathes normally developed. basitibial plate elliptical, with s-like secondary plate (fig. 2a). s2–s4 projected in the middle. apex of primary pygidial plate slightly rounded with the apex of the secondary plate open and slightly projected towards the distal edge of the primary plate (fig. 2d). male similar to the female, except for the following characters: integument dark brown to black, except basal segments of the flagellum brown and apical segments slightly orange (fig. 1c–d). supraclypeal area, discs of clypeus and labrum yellow (fig. 1c–d). tegula yellowish brown. clypeal disc with small unpunctated area on upper half, without smooth longitudinal band. terga and sterna, except t7 and s6, with relatively broad light brown smooth distal margin, wider on t4. mandible with three apically acute teeth (fig. 1c). distance between clypeus and compound eyes shorter than half of the shortest diameter of f1 (fig. 1c). apical margin of the hind tibia without tooth-like projection. apical half of fore and middle basitarsi without a row of long, erect, slightly spatulate and curved setae similar to an elaiospathe. pygidial plate absent. fig. 2. morphological characteristics of centris (relicthemisia) xanthomelaena moure & castro, 2001. a. female basitibial plate. b. male s7. c. male s8. d. female pygidial plate. e. genital capsule (dorsal view). f. genital capsule (ventral view). scale bars = 0.5 mm. vivallo f. & zanella f.c.v., relicthemisia, new subgenus of centris bees 7 type material holotype brazil – bahia state • ♀; milagres; 28 jan. 1998; 10h50; marina siqueira de castro leg.; chamaechrista amiciela (caesalpiniaceae)\1070; labe/ebda (not examined). paratypes brazil – bahia state • 1 ♀; milagres; 12º88.280´ s, 39º92.298´ w; 28 jan. 1998; 10h45; marina siqueira de castro leg.; chamaechrista amiciela (caesalpiniaceae)\1069; labe/ebda (not examined) • 1 ♀; milagres; 12º88.280´ s, 39º92.298´ w; 31 mar. 1997; 10h30; marina siqueira de castro leg.; in stimaphyllom auriculatum (malpighiaceae); dzup (not examined). material examined (specimens labeled as paratype or holotype below do not belong to the type series) brazil – alagoas state • 1 ♀; piranhas, poço da ingazeira; 9º50.594´ s, 37º88.113´ w; 28 oct. 2005; debora moura leg.; \ “centris xanthomelaena moure & castro 2001 schlindwein det.” \ hyap 4004; unila. – bahia state • 1 ♀; curaçá, faz. humaitá; 9º07´262˝ s, 39º42´859˝ w; 440 m a.s.l.; 4 may 2011; ppbio caatinga; f.c.v. zanella leg.; \ hyap 0582, unila • 1 ♀; monte santo; 10º43.958´ s, 39º33.566´ w; 3 feb. 2000; w.c.; malpighiaceae \ hyap 9706, unila • 2 ♀♀; camacari, dunas de tauá; 12º71.621´ s, 38º36.842´ w; 24 nov. 1993; j. becker leg. [†]; mnrj • 1 ♂; maracás; 13°26.467´ s, 40°25.850´ w; 20 jan. 1963 \ “coleção campos seabra” \ j. becker leg. \ “centris xanthemisia bicolor lepeletier” \ “c. paracentris xanthomelaena a.a. azevedo det.” [†]; mnrj • 1 ♀, 1 ♂; dias d’avila; 12º60.621´ s, 38º34.028´ w; 2 dec. 1951; luiz carlos leg. [†]; mnrj • 1 ♀, 2 ♂♂; dias d’avila; 12º60.621´ s, 38º34.028´ w; 16 dec. 1951; luiz carlos leg. [†]; mnrj • 2 ♀♀, 1 ♂; ipirá, santa quitéria; 12º15.642´ s, 39º76.319´ w; 6 jan. 2010; k. ramos and v. kanamura leg. [†]; mnrj. – minas gerais state • 1 ♀; conego marinho; 15º18˝ s, 44º25˝ w; 1 apr. 1988; nereu leg. 880824 \ “c. (paracentris) sp. nov. det. moure, 1992” \ “parátipo centris xanthomelaena sp. nov. f. zanella, 1999”; rpsp • 1 ♂; conego marinho; 15º18˝ s, 44º25˝ w; 29 mar. 1988; nereu leg. 880654\ “ex penthemisia moure, 1950 det. moure 1992” \ “c. (paracentris) sp. nov. det. moure, 1998” \ “parátipo centris xanthomelaena sp. nov. f. zanella, 1999”; rpsp. – paraíba state • 1 ♀; patos; 7º03.860´ s, 37º31.445´ w; 16 aug. 2002; gisllyana leg.; hyap 9300; unila. – pernambuco state • 1 ♀; alagoinha; 8º46.622´ s, 36º78.423´ w; 18 jun. 1987; i.c.s machado leg.; in ruellia sp. \ “c. (paracentris) sp. nov., det. moure 1992” \ “holótipo centris xanthomelaena sp. nov. f. zanella, 1999”; rpsp • 1 ♀; buique, vale do catimbau; 8º49.210´ s, 37º50.180´ w; 17 mar. 2005; r. pick leg. \ “centris xanthomelaena moure & castro 2001 schlindwein det.” \; hyap 4002; unila. – rio grande do norte state • 1 ♂; serra negra do norte, esec seridó; 6º66.572´ s, 37º40.515´ w; 12 aug. 1995; f. zanella leg.\ 0938 10b03\ hyap 9705; unila • 2 ♀♀; serra negra do norte, esec; 6º66.572´ s, 37º40.515´ w; 13 mar. 2005; f. zanella leg.; em pl. 1 roxa 10h\ hyap 4601; unila • 1 ♂; serra negra do norte, esec seridó; 6º66.572´ s, 37º40.515´ w; 5 jul. 2005; f. zanella leg.; pl. 27 chamaechrista 13h \ hyap 4598; unila • 1 ♀; serra negra do norte, esec seridó; 6º66.572´ s, 37º40.515´ w; 22 may 2005; f. zanella leg. \ hyap 4600; unila • 1 ♀; serra negra do norte, esec seridó; 6º66.572´ s, 37º40.515´ w; 12 jun. 2005; f. zanella leg.; krameria sp. 14h \ hyap 4853; unila • 1 ♂; serra negra do norte, esec seridó; 6º66.572´ s, 37º40.515´ w; 8 apr. 2005; f. zanella leg.; voando sobre arbusto 17h [†]; mnrj • 1 ♀; santana do seridó; 6º46˝ s, 36º44˝ w; 10 aug. 2007; j.m.f. camargo leg.; 07.1024; rpsp. type locality brazil: bahia state: milagres. european journal of taxonomy 760: 1–15 (2021) 8 distribution endemic to northeastern brazil, being recorded mainly in the dry open caatinga vegetation (fig. 3). the only record in central brazil, mato grosso state, needs confirmation. brazil: rio grande do norte state: serra negra do norte (zanella 2002, 2003; aguiar et al. 2003; silva 2014), *santana do seridó. paraíba state: (silveira et al. 2002). *patos. pernambuco state: alagoinha (vogel & machado 1991; machado & sazima 2008). petrolina (xavier et al. 2016; martins et al. 2018). salgueiro (xavier et al. 2016). *buique. alagoas state: *piranhas. bahia state: canudos (pigozzo & viana 2010; silva 2014). ibiraba (rodarte et al. 2008). itatim (aguiar 2003a, 2003b; aguiar et al. 2003; aguiar & zanella 2005; silva 2014). jequié (batalha filho et al. 2007; silva 2014). juazeiro (coelho et al. 2018). milagres (moure & castro 2001; silva 2014). monte santo (zanella 2002; aguiar et al. 2003). xique-xique (carneiro et al. 2019). *camacari (dunas de tauá), *dias d’ávila, *ipirá (santa quitéria), *maracás. mato grosso state: cf. serra do roncador (zanella 2002). minas gerais state: botumirim (azevedo et al. 2008). cônego marinho (zanella 2002). fig. 3. distribution records of centris (relicthemisia) xanthomelaena moure & castro, 2001. the limits of biogeographical provinces are depicted in the map. most records of this species are found in the caatinga province in northeastern brazil and marginally in the cerrado province, both in the south american diagonal of dry open areas. vivallo f. & zanella f.c.v., relicthemisia, new subgenus of centris bees 9 table 1. floral host records for centris (relicthemisia) xanthomelaena moure & castro, 2001 and distribution of host plant species. plant family and species main resource (elaiophore type) vegetation and distribution of host plant species 1, 2 host plant record 1 acanthaceae *ruellia sp. nectar ? (h) sp.? nectar ? (i) fabaceae chamaecrista amiciella (h.s. irwin & barneby) h.s. irwin & barneby pollen caatinga in northeast brazil and in go state (a) (j) c. pascuorum (benth.) h.s. irwin & barneby pollen caatinga, ‘campo rupestre’ and anthropized area in northeast brazil and mg state (a) (i) chamaecrista sp. ? (h) krameriaceae krameria grandiflora a. st. hill. oil (epithelial) caatinga, ‘campo rupestre’, restinga, cerrado (lato sensu) anthropized area from pa and rn to ms and es states in brazil (b) (k) krameria sp. (fig. 4) oil (epithelial) ? (l, h) malpighiaceae mcvaughia bahiana w.r. anderson3 oil (epithelial) sandy caatinga in ba state (c) (i, m) stigmaphyllon auriculatum a. juss. oil (epithelial) caatinga and restinga in northeast brazil and es and rj states (d) (j) plantaginaceae angelonia campestris nees & mart. (= bisaccata benth. = hookeriana gardner ex benth.) oil (trichomatic) caatinga and ‘campo rupestre’ in northeast brazil (e) (i, l, m) a. salicariifolia bonpl. (= hirta cham.) oil (trichomatic) caatinga in northeast brazil and cerrado (lato sensu) in ms state (e), reaching argentina and central america (f) (i, l, m) sterculiaceae melochia tomentosa l. nectar anthropized area, cerrado (lato sensu), rocky outcrops and rain forest in northeast and midwest brazil (g) (i, n, o) 1 references for plant species distribution and host plant record: (a) souza & bortoluzzi 2015; (b) costa-lima 2020; (c) almeida et al. 2019; (d) mamede 2015; (e) souza et al. 2020; (f) martins & alves-dos-santos 2013; (g) esteves 2015. references for plant hosts: (h) this work; (i) aguiar et al. 2003; (j) moure & castro 2001; (k) carneiro et al. 2019; (l) machado 2004; (m) vogel & machado 1991; (n) aguiar 2003a, 2003b; (o) machado & sazima 2008. 2 brazilian states: ba: bahia, es: espírito santo, go: goiás, mg: minas gerais, ms: mato grosso do sul, pa: pará, rj: rio de janeiro, rn: rio grande do norte. 3 cited as macvaughia bahiensis, a name that does not appear in ipni.org. european journal of taxonomy 760: 1–15 (2021) 10 discussion the distinctiveness of centris xanthomelaena as compared with the previously described subgenera of centris was already characterized by zanella (2002), especially based on the analysis of male genitalia. in the key for subgenera of centris of michener (2007), this species fits in paracentris, considering the “female’s basitibial plate with defined secondary plate that lacks sharp projecting margin” and the “margin of secondary plate extending [...] near posterior margin of basitibial plate” (dilemma 13), while the male’s lateral ocellus is separated from the eyes by a distance similar to the ocellar diameter, and the t2–t4 covered with dark pubescence (dilemma 20). nevertheless, males of c. xanthomelaena are clearly distinguished from males of paracentris by the strong emargination on the apical margin of t7 (fig. 2b), by the absence of emargination on the basal border of s7 (fig. 2b), and by the presence of a long dorsoapical projection of gonocoxite, ca ⅔ lengths of gonostylus (fig. 2e–f). this latter feature is unique, being a somewhat intermediate condition between that observed in centris s. str. and paracentris. fig. 4. female of centris (relicthemisia) xanthomelaena moure & castro, 2001 visiting krameria sp. (krameriaceae) in estação ecológica do seridó, rio grande do norte state, brazil. vivallo f. & zanella f.c.v., relicthemisia, new subgenus of centris bees 11 previously to its description and naming, centris xanthomelaena was already recognized as a legitimate pollinator of angelonia campestris nees & mart. and a. salicariifolia bonpl. (plantaginaceae) in the caatinga (vogel & machado 1991). the two pollen sources chamaecrista moench species and oil sources krameria grandiflora a. st. hill., mcvaughia bahiana w.r. anderson, stigmaphyllon auriculatum a. juss. and a. campestris are endemic to northeastern brazil and specially recorded in xerophilous caatinga vegetation or at least in open vegetation areas. a similar context was noted by aguiar et al. (2003) in relation to centris hyptidis ducke, 1908, another species endemic from northeastern brazil and typical from caatinga vegetation. this species also has a more specialized relation with pollen and oil sources in the caatinga biome as compared to other species with wider distribution patterns. this can be interpreted as a result of a longer history with plants of this semiarid region. differing from the interpretation of giannini et al. (2013), c. hyptidis, along with c. hyptidoides roig-alsina, 2000 and c. thelyopsis vivallo & melo, 2009, does not belong to the same lineage of the subgenus wagenknechtia, but to c. (anisoctenodes), an old lineage distributed in the south american diagonal of open dry vegetation (werneck 2011; zanella 2011), probably with association to flowers of angelonia (plantaginaceae) (vivallo & melo 2009; martins & melo 2015). a similar old history and restricted distribution is found in centris (relicthemisia) xanthomelaena, in spite of similar oil-collecting apparatuses found in other species of centris that also occur in the caatinga (vogel & machado 1991). those structures are specialized in exploring plant species with epithelial elaiophores (giannini et al. 2013). it must also be noted that there is no record of this bee visiting the introduced west indian cherry (malpighia emarginata d.c.) even though the intense collection effort in this tree at sites where c. (relicthemisia) xanthomelaena occurs (coelho et al. 2018; siqueira et al. 2011). acknowledgments we thank eduardo a.b. almeida (rpsp), diego marinho (mnrj) and the anonymous reviewers for their help. financial support was provided to felipe vivallo by conselho nacional de desenvolvimento científico e tecnológico (cnpq, grant 444320/2014-8) and projeto de informatização da coleção entomológica do museu nacional/ufrj, sibbr/cnpq (proc. 405588/2015-1), brazil. support for studies in the caatinga region was provided to fernando zanella by programa de pesquisa em biodiversidade semiárido do ministério da ciência e tecnologia (ppbio semiárido), by fundação de apoio à pesquisa da paraíba (fapesq) and cnpq, on the project ‘diversidade, ecologia e conservação de himenópteros na região semiárido do nordeste do brasil, com ênfase nas abelhas’. this study was registered at sistema nacional de gestão do patrimônio genético e do conhecimento tradicional associado under number a6f40c4. this paper is part of the sigma project nº 21565 mn/ufrj and the contribution number 57 from the hymn. references aguiar c.m.l. 2003a. flower visits of centris bees (hymenoptera: apidae) in an area of caatinga (bahia, brazil). studies on neotropical fauna environment 38: 41−45. https://doi.org/10.1076/snfe.38.1.41.14029 aguiar c.m.l. 2003b. utilização de recursos florais por abelhas (hymenoptera, apoidea) em uma área de caatinga (itatim, bahia, brasil). revista brasileira de zoologia 20: 457−467. https://doi.org/10.1590/s0101-81752003000300015 aguiar c.m.l. & zanella f.c.v. 2005. estrutura da comunidade de abelhas (hymenoptera: apoidea: apiformes) de uma área na margem do domínio da caatinga (itatim, ba). neotropical entomology 34: 15‒24. https://doi.org/10.1590/s1519-566x2005000100003 https://doi.org/10.1076/snfe.38.1.41.14029 https://doi.org/10.1590/s0101-81752003000300015 https://doi.org/10.1590/s1519-566x2005000100003 european journal of taxonomy 760: 1–15 (2021) 12 aguiar c.m.l., zanella f.c.v., martins c.f. & carvalho c.a.l. 2003. plantas visitadas por centris spp. (hymenoptera: apidae) na caatinga para obtenção de recursos florais. neotropical entomology 32: 247‒259. https://doi.org/10.1590/s1519-566x2003000200009 aguiar c.m.l., gimenes m. & rebouças p.l.o. 2005. abelhas (hymenoptera, apoidea). in: juncá f.a., funch l. & rocha w. 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(eds) biogeografia da américa do sul. padrões e processos: 198– 220. editora roca, são paulo, sp. available from https://www.researchgate.net/publication/350499172 [accessed 16 jun. 2021]. zanella f.c.v. & vivallo f. 2009. a new species of the bee genus centris from peru (hymenoptera: apidae). zootaxa 2175: 66–68. manuscript received: 22 june 2020 manuscript accepted: 13 april 2021 published on: 14 july 2021 topic editor: gavin broad desk editor: charlotte gérard printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. https://doi.org/10.1590/s0085-56262002000400001 https://www.researchgate.net/publication/235007025 https://www.researchgate.net/publication/350499172 european journal of taxonomy 748: 138–154 issn 2118-9773 https://doi.org/10.5852/ejt.2021.748.1347 www.europeanjournaloftaxonomy.eu 2021 · leduc d. & zhao z.q. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:e814bab1-3b3d-4458-9f94-e1ef89191881 138 litinium gludi sp. nov. (nematoda, oxystominidae) from kermadec trench, southwest pacific ocean daniel leduc 1,* & zeng qi zhao 2 1 national institute of water and atmospheric research (niwa) ltd, wellington, new zealand. 2 landcare research, private bag 92170, auckland mail centre, auckland 1142, new zealand. * corresponding author: daniel.leduc@niwa.co.nz 2 email: zhaoz@landcareresearch.co.nz 1 urn:lsid:zoobank.org:author:9393949f-3426-4ee2-8bde-defface3d9bc 2 urn:lsid:zoobank.org:author:f5bee9fd-801a-49c6-9585-c765ee77a123 abstract. recent work on the taxonomy of nematodes in southwest pacific ocean trenches has led to the discovery of taxa which so far appear to be restricted to the oceans’ deepest environments. here, litinium gludi sp. nov. is described based on specimens obtained from a deep basin within the kermadec trench at 9540 m water depth. the new species differs from other species of the genus in having a conico-cylindrical tail, papillose labial sensilla, and heartor leaf-shaped amphideal fovea. both ssu and lsu phylogenetic analyses provide strong support for the placement of the new species within a clade containing both thalassoalaimus and litinum, and within oxystomininae, which is consistent with the structure of the female reproductive system with only one posterior ovary in this subfamily. our molecular analyses also indicate that the new species is most closely related to thalassoalaimus despite lacking a caudal capsule, the main trait characterizing the latter genus, and despite being most morphologically similar to litinium, particularly in the unusual shape of the amphideal fovea. however, given the changing definitions of the closely-related genera thalassoalaimus and litinium in recent years, available genbank sequences may have been misidentified, which makes the interpretation of molecular phylogenetic analyses problematic. given the current morphological definitions of litinium and thalassoalaimus, we choose to place the new species within litinium, despite the apparently contradictory findings of molecular phylogenetic analyses. the placement of cricohalalaimus in a clade with thalassoalaimus and litinium in both ssu and lsu analyses indicates that this genus should be placed within the oxystomininae and not the halalaiminae as in the current classification. this new proposed grouping is consistent with variation in the structure of the female reproductive system, a feature which appears more taxonomically informative than amphid shape for subfamily-level classification. keywords. hadal, oxystomininae, cricohalalaimus, small subunit 18s rdna, d2-d3 region of large subunit 28s rdna. leduc d. & zhao z.q. 2021. litinium gludi sp. nov. (nematoda, oxystominidae) from kermadec trench, southwest pacific ocean. european journal of taxonomy 748: 138–154. https://doi.org/10.5852/ejt.2021.748.1347 https://doi.org/10.5852/ejt.2021.748.1347 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:e814bab1-3b3d-4458-9f94-e1ef89191881 mailto:daniel.leduc%40niwa.co.nz?subject= mailto:zhaoz%40landcareresearch.co.nz?subject= http://zoobank.org/urn:lsid:zoobank.org:author:9393949f-3426-4ee2-8bde-defface3d9bc http://zoobank.org/urn:lsid:zoobank.org:author:f5bee9fd-801a-49c6-9585-c765ee77a123 https://doi.org/10.5852/ejt.2021.748.1347 leduc d. & zhao z.q., new nematode species from kermadec trench 139 introduction the oxystominidae chitwood, 1935 (order enoplida filipjev, 1929) is among the most common nematode families in deep-sea environments (gambi et al. 2003; bik et al. 2010a; miljutin et al. 2010). this largely marine group is characterized by a thin elongated body, the outer labial and cephalic sensilla in separate circles, and the buccal cavity without teeth (lorenzen 1981). the family includes three subfamilies: the oxystomininae chitwood, 1935, halalaiminae de coninck, 1965 and paroxystomininae de coninck, 1965, which are differentiated based on the amphid shape, the structure of the female reproductive system and the presence/structure of the precloacal supplements. the oxystomininae comprises five genera, all of which except oxystomina chitwood, 1935 are exclusively marine; the presence of only the posterior ovary is considered a holapomorphy establishing the holophyly of the oxystomininae (lorenzen 1981). the halalaiminae originally comprised a single genus, halalaimus de man, 1888, which is characterized by an elongated longitudinal amphideal aperture, an holapomorphic trait establishing the holophyly of the subfamily. the genus cricohalalaimus bussau, 1993 was later included in the subfamily by smol et al. (2014) based on the similar structure of the amphideal aperture; this genus, however, is also characterized by the presence of a single posterior ovary in females, a feature of the oxystomininae. there is no holapomorphy for the paroxystominae, which comprises paroxystomina micoletzky, 1924 and maldivea gerlach, 1962 (lorenzen 1981). phylogenetic analyses based on ssu sequences do not provide support for the monophyly of the oxystominae or halalaminae (bik et al. 2010a), and no molecular sequences are yet available for the paroxystomininae. the majority of marine free-living nematode genera have wide water depth distributions spanning both coastal and deep-sea environments (> 200 m water depth), but some are mostly found in the deep sea (e.g., acantholaimus allgén, 1933, metasphaerolaimus gourbault & boucher, 1981), while others are found exclusively in continental slope and/or deeper environments (e.g., bathyeurystomina lambshead & platt, 1979, thelonema bussau, 1993, manganonema bussau, 1993). phylogenetic analyses based on ssu data suggest repeated and recent interchanges between the deep sea and shallow water environments, with habitat transitions thought to be frequent for free-living nematodes (bik et al. 2010b). in halalaimus and oxystomina, shallow water species appear to have evolved from deep water ancestors (bik et al. 2010b); however, within the oxystominidae, cricohalalaimus bussau, 1993 is the only genus with a distribution strictly restricted to the deep sea. limited core sampling has been conducted in hadal trenches to date (> 6000 m water depth), resulting in scant knowledge of nematode systematics in the deepest parts of the world’s oceans. taxonomic work on tonga and kermadec trench material has led to the discovery of new species and genera which so far appear to be restricted to hadal environments (leduc 2015, 2016). more recently, sampling within the deep kermadec trench axis provided core samples from the trench’s deepest basins, and the examination of specimens from a site located at 9540 m depth led to the discovery of a species of nematode, litinium gludi sp. nov., which is described here. phylogenetic relationships of this new species within the oxystominidae are investigated using ssu and d2-d3 of lsu rdna sequences. material and methods study area and sampling the kermadec trench extends from approximately from 26 to 36°s near the northeastern tip of new zealand’s north island, southwest pacific ocean. sediment samples were obtained from the deep axis of kermadec trench at 9540 m depth during rv tangaroa cruise tan1711 in december 2017. sediments were obtained using a usnel-type box corer (dimensions: 0.5 × 0.5 × 0.5 m, 0.125 m3 capacity). subsamples were obtained using a cut-off syringe with 29 mm internal diameter to a depth of 10 cm. subsamples for morphological analyses were sliced into 1 cm layers and fixed in 10% buffered formalin. nematodes were extracted from the remaining sediments by ludox flotation and transferred to pure european journal of taxonomy 748: 138–154 (2021) 140 glycerol (somerfield & warwick 1996). species descriptions were made from glycerol mounts using differential interference contrast microscopy and drawings were made with the aid of a camera lucida. measurements were obtained using an olympus bx53 compound microscope with cellsens standard software for digital image analysis. subsamples for molecular analyses were obtained as described above but the samples were frozen at -80°c instead of being fixed in formalin. in the laboratory, frozen sediment samples were thawed overnight, then sieved trough a 20 µm mesh to retain nematodes. nematodes were extracted using the ludox flotation method (somerfield & warwick 1996) and sorted under a dissecting microscope. one juvenile litinium gludi sp. nov. specimen was isolated and transferred to a temporary slide to confirm its identity. this specimen was identified based on the characteristic shape of the cephalic region, cephalic sensilla, amphids, and tail. all measurements are in µm (unless otherwise stated), and all curved structures are measured along the arc. type specimens are held in the niwa invertebrate collection, wellington, new zealand. list of abbreviations a = body length/maximum body diameter b = body length/pharynx length c = body length/tail length c’ = tail length/anal or cloacal body diameter cbd = corresponding body diameter l = total body length; n, number of specimens v = vulva distance from anterior end of body %v = v/total body length × 100 dna extraction, pcr and sequencing following observation under a compound microscope, the specimen was transferred to lysis buffer and kept frozen at -80°c prior to molecular analyses. dna was extracted by the method of zheng et al. (2002) with minor modifications (i.e., the nematode was not cut prior to being transferred to the lysis buffer solution). the dna extract was stored at -20°c until used as pcr template. primers for lsu amplification were forward primer d2a (5’ acaagtaccgtgagggaaagt 3’) and reverse primer d3b (5’ tgcgaaggaaccagctacta 3’) (nunn 1992). primers for the rdna small subunit (ssu) were the forward primer 1096f, 5’-ggtaattctggagctaatac-3’ and reverse primer 1912r, 5’-tttacggtcagaactaggg-3’ (first fragment), and the forward primer 1813f, 5’-ctgcgtgagaggtgaaat-3’ and reverse 2646r, 5’-gctaccttgttacgactttt-3’ (second fragment) (holterman et al. 2006). for both ssu and lsu, the 20 µl pcr contained 10 µl redextractn-amp pcr readymix (sigma-aldrich, usa), 1 µl (5 µm) each of forward and reverse primers, and 2 µl of dna template. the thermal cycling program was as follows: denaturation at 95°c for 3 min, followed by 40 cycles of denaturation at 94°c for 15 s, annealing at 55°c for 30 s, and extension at 72°c for 30 s. a final extension was performed at 72°c for 7 min. the amplicons were electrophoresed on 1% tae-agarose gel stained with sybr® safe, observed under uv illumination using the gel-doc system (biorad, hercules, ca, usa), and images processed using the image lab ver. 5.1 analysis software (biorad). the pcr products were sequenced bi-directionally using the amplification primers by ecogene (auckland, new zealand). sequences were obtained with a 3500xl genetic analyzer (applied biosystems, usa) and assembled and edited with sequencher ver. 4.10.1 (gene codes corp.). leduc d. & zhao z.q., new nematode species from kermadec trench 141 sequence alignment and phylogenetic inference the ribosomal dna ssu and lsu d2-d3 sequences were deposited in genbank under accession numbers mw209715 and mw209714, respectively. the placement of the new ssu and d2-d3 of lsu sequences was investigated through a phylogenetic analysis of the representative genera of the oxystominidae, as well as the rhaptothyreidae hope & murphy, 1969 and oncholaimoidea filipjev, 1916, which have been shown to be closely related (bik et al. 2010a; leduc et al. 2018), and rooted using ironidae de man, 1876 sequences (all ssu sequences > 1300 bp except four ca 800 bp, and d2-d3 of lsu sequences > 600 bp). dna sequences were aligned using muscle (edgar 2004a, 2004b) with default parameters. phylogenies were built in geneious ver. 10.2.6 (http://www.geneious.com, kearse et al. 2012). mrmodeltest ver. 2.3 (nylander 2004) in conjunction with paup* ver. 4.0b10 (swofford 2002) were used to select the best model using the akaike information criterion. the substitution model [gtr (general time-reversible) + i (proportion of invariable sites) + g (gamma distribution)] was selected as the best-fit model for ssu alignments (1522 bp) and lsu alignments (722 bp), respectively. the trees were run with chain length of 1 100 000, and burn-in length of 100 000. the perimeter files from multiple runs were inspected for chain convergence in tracer ver. 1.5 (rambaut & drummond 2007), and the trees were edited in figtree ver. 1.4.2 (http://tree.bio.ed.ac.uk/software/figtree) and powerpoint. these analyses were also conducted with phyml ver. 3.0 using the default settings in geneious ver. 10.2.6. the substitution model gtr, the nni (default, fast) topology search and 1000 bootstrap replicates (guindon et al. 2010) were selected for building the tree. results systematics phylum nematoda diesing, 1861 order enoplida filipjev, 1929 suborder ironina siddiqi, 1983 superfamily ironoidea de man, 1876 family oxystominidae chitwood, 1935 diagnosis (emended from smol et al. 2014) body elongated and very thin at anterior end. anterior sensilla in three separate circles, second and third circles clearly separated; inner labial sensilla papilliform or setiform, outer labial sensilla usually setiform (except in some species of litinium), cephalic sensilla setiform. buccal cavity narrow, tubular or funnelshaped and without teeth. sexual dimorphism in amphid shape sometimes present. only orthometanemes with very short caudal filaments present. pharynx inserts into body cuticle in region of buccal cavity; however, cephalic capsule not well developed. posterior section of pharynx with undulating outline. females didelphic-amphidelphic with antidromously reflexed ovaries or monodelphic-opisthodelphic. males diorchic with opposed testes or only one anterior testis. position of caudal glands variable. subfamily oxystomininae chitwood, 1935 diagnosis (modified from smol et al. 2014) only dorsolateral orthometanemes, ventral gland when present confined within pharyngeal region. amphideal fovea and aperture variable but amphideal aperture never elongated, precloacal supplements (when present) papilliform, in single ventral row. females monodelphic-opisthodelphic. http://www.geneious.com http://tree.bio.ed.ac.uk/software/figtree european journal of taxonomy 748: 138–154 (2021) 142 type genus oxystomina filipjev, 1918 other valid genera litinium cobb, 1920 nemanema cobb, 1920 thalassoalaimus de man, 1893 wieseria gerlach, 1956 litinium cobb, 1920 diagnosis (modified from tchesunov et al. 2014) circles of six inner and six outer labial papillae (< 2 µm) or setae (≥ 2 µm) situated close together on anterior end, subapically, with circle of four cephalic papillae or setae posterior to circles of inner and outer labial sensilla. amphideal fovea situated between circles of outer labial and cephalic sensilla. amphideal fovea varies in shape between species and may differ in males and females of same species: may be ovoid with anterior round aperture, horseshoe-like or crescent contoured or more complex. only one posterior antidromously reflexed ovary present; vulva shifted anteriorly. tail never clavate, more or less short, cylindrical with rounded tip, occasionally conical or conico-cylindrical with pointed tip; terminal caudal capsule (thick cuticular lining at tip of tail) absent or weakly developed. remarks the amphid of some species of litinium cobb, 1920 is described as being “horseshoe-shaped”. in the new species, our observations indicate that it is the superimposition of the amphideal aperture contour over the larger amphideal fovea contours that creates the appearance of a horseshoe-shaped structure. whether this is also the case in other species of litinium remains to be clarified. there has been some confusion regarding the definitions of litinium and thalassoalaimus. the diagnoses provided by smol et al. (2014) indicate that the two genera differ in the shape of the amphids (pocketshaped amphideal fovea and slit-like aperture in thalassoalaimus vs horseshoeor heart-shaped fovea and round aperture in litinium) and presence (thalassoalaimus) or absence (litinium) of a caudal capsule. however, we note that the pocket-shaped amphideal fovea and slit-like aperture in litinium subterraneum tchesunov, mokievsky & nguyen vu thanh, 2010 is identical to that of thalassoalaimus. on the other hand, litinium profundorum tchesunov, nguyen vu thanh & nguyen dinh tu, 2014 has a weak thickening of the inner cuticle layer of the tail tip resembling a caudal capsule but is characterised by a horseshoe-shaped amphideal fovea typical of the genus litinium. most recently, martelli et al. (2017) used the presence or absence of a caudal capsule as the sole distinguishing feature between the two genera, resulting in the transfer of several species of thalassoalaimus to litinium. some of the newly transferred species, namely l. aceratus (vitiello, 1970) martelli, lo russo, villares & pastor de ward, 2017 and l. longicaudatus (vitiello, 1970) martelli, lo russo, villares & pastor de ward, 2017, as well as l. gludi sp. nov., are characterised by conico-cylindrical tails, which require the genus diagnosis of tchesunov et al. (2014) to be emended. in addition, l. longicaudatus, l. qangi tchesunov, nguyen vu thanh & nguyen dinh tu, 2014, l. curticauda tchesunov, nguyen vu thanh & nguyen dinh tu, 2014 and l. gludi sp. nov. are characterised by papillose to short setose labial sensilla (0.5–2.5 µm long), unlike all other species of the genus which have exclusively setose (> 2 µm) labial sensilla. leduc d. & zhao z.q., new nematode species from kermadec trench 143 list of valid species l. abyssorum tchesunov, nguyen vu thanh & nguyen dinh tu, 2014 l. aceratus (vitiello, 1970) martelli, lo russo, villares & pastor de ward 2017 l. aequale cobb, 1920 l. australis martelli, lo russo, villares & pastor de ward, 2017 l. bananum gerlach, 1956 l. curticauda tchesunov, nguyen vu thanh & nguyen dinh tu, 2014 l. dispariseta yu & xu, 2018 l. egregius (steiner, 1916) martelli, lo russo, villares & pastor de ward 2017 l. gludi sp. nov. l. longicaudatus (vitiello, 1970) martelli, lo russo, villares & pastor de ward 2017 l. obtusilobus bussau, 1993 l. paramontemari (vitiello, 1970) martelli, lo russo, villares & pastor de ward 2017 l. parmatum wieser, 1954 l. pirum (lorenzen, 1969) martelli, lo russo, villares & pastor de ward 2017 l. profundorum tchesunov, nguyen vu thanh & nguyen dinh tu, 2014 l. quamgi tchesunov, nguyen vu thanh & nguyen dinh tu, 2014 l. setosus (timm, 1967) martelli, lo russo, villares & pastor de ward 2017 l. subterraneum tchesunov, mokievsky & nguyen vu thanh, 2010 l. volutum gerlach, 1962 litinium gludi sp. nov. urn:lsid:zoobank.org:act:fa79dfef-a389-48aa-af75-03984d69f202 figs 1–4, table 1 diagnosis litinium gludi sp. nov. is characterised by a slender body (a = 58–63), a body length of 895–1066 µm, papilliform labial sensilla, short cephalic sensilla 1.5–2.0 µm or ~0.2 cbd long, a large heartor leafshaped amphideal fovea 67–88% cbd wide, a smaller amphideal aperture with two central longitudinal ridges, males with dimorphic sperm, spicules 1.1–1.4 times cloacal body diameter, a short and simple gubernaculum, and two precloacal setae. differential diagnosis the new species resembles l. aceratus and l. longicaudatus, the only other species of the genus with conico-cylindrical tails, and is particularly similar to l. longicaudataus in having papillose (< 2 µm) labial sensilla. litinium gludi sp. nov. can be differentiated from l. longicaudatus by the shorter body (895–1066 vs 2180–2196 µm in l. longicaudatus), lower ‘a’ ratio (58–63 vs 75–86), and heartor leafshaped amphideal fovea (vs pocket-shaped). etymology the species is named after professor ronnie glud, leader of the hades-erc trench project and covoyage leader. material examined holotype pacific ocean • ♂; kermadec trench, voyage tan1711, station 25, site 3; 30.3815° s, 176.6417° w; 9540 m water depth, 0–2 cm sediment depth; 3 dec. 2017; niwa 139257. http://zoobank.org/urn:lsid:zoobank.org:act:fa79dfef-a389-48aa-af75-03984d69f202 european journal of taxonomy 748: 138–154 (2021) 144 males female paratype holotype paratypes specimen m1 m2 m3 f1 l 1027 895 1066 930 a 60 60 63 58 b 5 4 5 5 c 11 10 11 10 c’ 7.3 7.8 6.9 9.3 body diam. at outer labial sensilla 6 6 6 6 body diam. at cephalic setae 9 9 9 9 body diam. at amphids 9 8 9 9 length of cephalic sensilla 1.6–1.9 1.6–1.8 1.5–2.0 1.8 amphideal fovea height 10 8 9 9 amphideal fovea width 7 7 6 7 amphideal fovea width/cbd (%) 78 88 67 78 amphideal fovea from anterior end 4 5 4 5 amphideal aperture height 7 6 6 6 amphideal aperture width 3 3 3 3 nerve ring from anterior end 85 77 87 83 nerve ring cbd 17 15 17 16 pharynx length 208 200 211 195 pharyngeal diam. at base 12 9 11 10 pharynx cbd at base 15 12 15 13 max. body diam. 17 15 17 16 spicule length 18 15 16 – gubernacular apophyses length 3 2 3 – cloacal/anal body diam. 13 12 14 10 tail length 95 93 96 93 v – – – 329 %v – – – 35 vulval body diam. – – – 14 table 1. morphometrics (µm) of litinium gludi sp. nov. paratypes pacific ocean • 2 ♂♂, 1 ♀; kermadec trench, voyage tan1711, station 25, site 3; 30.3815° s, 176.6417° w; 9540 m water depth, 0–2 cm sediment depth; 3 dec. 2017; niwa 139258. description males body slender, cylindrical, widest at level of nerve ring, tapering slightly towards anterior extremity. cuticle smooth, without ornamentation; somatic setae absent, except for sparse sublateral setae on tail. cephalic region demarcated by slight constriction near posterior edge of amphideal fovea. inner labial sensilla not observed; six outer labial papillae, ~0.5 µm long, located slightly anterior to amphideal fovea and four short cephalic sensilla, 1.5–2.0 µm or ~0.2 cbd long, situated far posteriorly at level of posterior edge of amphideal fovea. amphideal fovea large, heartor drop-shaped, with lightly cuticularized outline; amphideal aperture smaller, ovalor drop shaped, with two lightly cuticularized longitudinal central ridges spanning long axis of amphideal fovea. buccal cavity narrow, cylindrical, with cuticularized portion 3–5 µm deep. pharynx muscular, cylindrical, surrounding posterior portion leduc d. & zhao z.q., new nematode species from kermadec trench 145 fig. 1. litinium gludi sp. nov. a. anterior body region of the male holotype (niwa 139257). b. cephalic region of the female paratype (niwa 139258). c. cephalic region of the male paratype (niwa 139258). d. posterior body region of the female paratype (niwa 139258). e. posterior body region of the male holotype (niwa 139257). scale bar: a = 65 µm, b–c = 15 µm, d–e = 25 µm. of buccal cavity, widening towards posterior extremity but not forming true bulb. nerve ring located slightly anterior to middle of pharynx. secretory-excretory system not observed. cardia 2–4 µm long, partially embedded in intestine. reproductive system with two opposed and outstretched testes; anterior testis located to the left or right of intestine, posterior testis located on opposite side or on same side. mature sperm in anterior testis large, elongated, 19–21 × 2–3 µm; mature sperm in posterior testis smaller, globular, 1.5 × 1.5–2.0 µm. spicules short, 1.1–1.4 times cloacal body diameter, curved distally, with ventrally bent capitulum. gubernaculum small, simple, block-shaped, without lateral pieces, with poorly developed apophyses. two precloacal papillae, 1.3–1.4 µm long; posterior-most papilla located 11–14 µm anterior to cloaca, european journal of taxonomy 748: 138–154 (2021) 146 fig. 2. litinium gludi sp. nov. a. entire female paratype (niwa 139258). b. entire male holotype (niwa 139257). scale bar = 200 µm. leduc d. & zhao z.q., new nematode species from kermadec trench 147 fig. 3. litinium gludi sp. nov., light micrographs of the male holotype (niwa 139257). a. surface view of cephalic region showing amphid and outer labial sensilla. b. optical cross-section of cephalic region showing narrow buccal cavity and pharynx. c. anterior testis showing elongated sperm cells (arrow). d. posterior testis showing globular sperm cells (arrow). scale bar = 10 µm. european journal of taxonomy 748: 138–154 (2021) 148 fig. 4. litinium gludi sp. nov., light micrograph of tail tip of the male holotype (niwa 139257), showing absence of caudal capsule and spinneret. scale bar = 2 µm. second precloacal papilla located 19–22 µm from posterior-most precloacal papilla. ejaculatory glands not observed. tail conico-cylindrical, with pointed tip, caudal capsule absent; three caudal glands present posterior to cloaca; spinneret not observed. female similar to males but without any somatic setae on tail. reproductive system with single posterior genital branch and reflexed ovary. oocyte up to 160 × 10 µm. vulva located far anteriorly at about ⅓ of body length from anterior extremity. vagina perpendicular to body surface; vaginal glands not observed. molecular phylogenetic relationships near full-length ssu (1516 bp) and d2-d3 of lsu sequences (756 bp) were obtained for litinium gludi sp. nov. the oxystominidae did not form a monophyletic clade in the consensus ssu tree due to the placement of rhaptothyerus typicus hope & murphy, 1969 and oncholaimoid sequences among the oxystominid sequences (fig. 5). neither the halalaiminae (represented by halalaimus and oxystomina) nor the oxystomininae (all other oxystominid genera) were recovered as monophyletic. litinium leduc d. & zhao z.q., new nematode species from kermadec trench 149 f ig . 5 . b ay es ia n tr ee i nf er re d fr om s su s eq ue nc es , a lig ne d us in g th e m u sc l e a lig nm en t al go ri th m u nd er t he g en er al t im ere ve rs ib le ( g t r ) + i (p ro po rt io n of in va ri ab le s ite s) + g am m a di st ri bu tio n (g ) m od el . t he n ew s eq ue nc e is s ho w n in b ol d fo nt o n a gr ey b ac kg ro un d. p os te ri or p ro ba bi lit ie s (l ef t) a nd b oo ts tr ap v al ue s (r ig ht ) g re at er th an o r e qu al to 5 0% a re g iv en o n ap pr op ri at e cl ad es . t he s ca le s ta nd s fo r s ub st itu tio ns p er s ite . : le ss th an 50 % b oo ts tr ap s up po rt ; # : s eq ue nc e fj 04 05 00 is la be lle d th al as so al ai m us p ir um in g en b an k bu t i s la be lle d li tin iu m p ir um h er e to r efl ec t t he n ew cl as si fic at io n of th is s pe ci es p ro po se d by m ar te lli e t a l. (2 01 7) . european journal of taxonomy 748: 138–154 (2021) 150 f ig . 6. b ay es ia n tr ee i nf er re d fr om d 2d 3 of l su s eq ue nc es , a lig ne d us in g th e m u sc l e a lig nm en t al go ri th m u nd er t he g en er al t im ere ve rs ib le (g t r ) + i (p ro po rt io n of in va ri ab le s ite s) + g am m a di st ri bu tio n (g ) m od el . p os te ri or p ro ba bi lit ie s (l ef t) a nd b oo ts tr ap v al ue s (r ig ht ) gr ea te r th an o r eq ua l t o 50 % a re g iv en o n ap pr op ri at e cl ad es . t he n ew s eq ue nc e is s ho w n in b ol d fo nt o n a gr ey b ac kg ro un d. t he s ca le s ta nd s fo r s ub st itu tio ns p er s ite . * : n o bo ot st ra p su pp or t; : le ss th an 5 0% b oo ts tr ap s up po rt . leduc d. & zhao z.q., new nematode species from kermadec trench 151 gludi sp. nov. was placed in a well-supported clade (100% posterior probability and 82% bootstrap support) with sequences of thalassoalaimus. the similarity between ssu sequences of litinium gludi sp. nov. and thalassoalaimus sp. was 94–95% with a difference of 69 in 1508 nucleotides with 4 gaps (mn250034), 83 in 1504 nucleotides with 1 gap (hm564634) and 43 in 798 nucleotides with 1 gap (mn250041). the litinium gludi sp. nov. + thalassoalaimus clade was part of a wider, strongly supported oxystominid clade including all sequences of litinium and cricohalalaimus (100% posterior probability and bootstrap support), although neither litinium nor thalassoalaimus were monophyletic. the similarity between ssu sequences of litinium gludi sp. nov. and litinium sp. was 87–94% with a difference of 84 in 1513 nucleotides with 5 gaps (fj040500), 116 in 1516 nucleotides with 3 gaps (hm564629), 113 in 1514 nucleotides with 2 gaps (hm564649), and 46 in 786 nucleotides with 1 gap (mk626773). sequences of halalaimus formed a strongly supported (100% posterior probability and bootstrap support) monophyletic sister group to the oncholaimoidea filipjev, 1916, the latter also forming a strongly supported monophyletic group (100% posterior probability and bootstrap support). the sequences of oxystomina formed a strongly supported group (100% posterior probability and bootstrap support), however this genus was not monophyletic due to the inclusion of a sequence of nemanema cobb, 1920 in the clade. the placement of another sequence of nemanema as a sister group to the halalaimus + oncholaimoidea clade was poorly supported (56% posterior probability and <50% bootstrap support). the topology of the d2-d3 of lsu consensus tree was similar to that of the ssu tree, with the oxystominidae again not recovered as monophyletic (fig. 6). sequences of halalaimus and oxystomina each formed strongly supported monophyletic clades (100% posterior probability and bootstrap support). as in the ssu tree, litinium, cricohalalaimus, and thalassohalaimus formed a strongly supported clade (100% posterior probability and bootstrap support), with litinium gludi sp. nov. most closely related to thalassoalaimus (100% posterior probability and 98% bootstrap support). the similarity between lsu sequences of litinium gludi sp. nov. and thalassoalaimus sp. was 92% with a difference of 118 in 664 nucleotides with 7 gaps (hm564880); the similarity between lsu sequences of litinium gludi sp. nov. and litinium sp. was 71–72% with a difference of 188 in 671 nucleotides with 13 gaps (hm564858), 184 in 671 nucleotides with 12 gaps (hm564874) and 183 in 671 nucleotides with 13 gaps (hm564875). discussion the topologies of our ssu and d2-d3 of lsu trees are very similar to the phylogeny of bik et al. (2010a), which recovered four sub-clades: oncholaimoidea, thalassoalaimus + cricohalalaimus + litinium, oxystomina, and halalaimus. bik et al. (2010a) noted that halalaimus was consistently recovered as a long-branch clade, which may have a destabilizing effect on the internal tree topology. the inclusion of rhaptothyreus typicus in the present study, which was also recovered as a long-branch clade, may have had a similar destabilizing effect, and the placement of this taxon in relation to the oxystominidae as well as wider enoplida remains uncertain (leduc et al. 2018). molecular phylogenetic analyses have not yet provided support for the monophyly of the oxystomininae or halalaiminae (bik et al. 2010a; present study). the morphology of litinium gludi sp. nov. is consistent with other species of litinium due to the shape of the amphids, the posterior position of the cephalic setae and the lack of a caudal capsule. however, both ssu and lsu trees indicate that the new genus is most closely related to thalassoalaimus. the latter genus is similar to litinium in the structure and position of cephalic sensilla, but differs from litinium in having a caudal capsule. it should be noted that most of the sequences of thalassoalaimus in genbank were not identified to the species level, and that given the recent changes in the definitions of litinium and thalassoalaimus (tchesunov et al. 2014; martelli et al. 2017), some or possibly all of the sequences of thalassoalaimus from genbank included in our phylogenetic analyses may in european journal of taxonomy 748: 138–154 (2021) 152 fact belong to litinium or vice versa. for example, the ssu sequence of thalssoalaimus pirum, the only sequence of thalassoalaimus identified to species level in genbank, has since been transferred to litinium by martelli et al. (2017) (see fig. 4). this uncertainty puts into question the apparent close relationship between the new species and thalassoalaimus in molecular analyses. therefore, given the current morphological definitions of litinium and thalassoalaimus, we choose to place the new species within litinium, despite the apparently contradictory findings of molecular analyses. our phylogenetic analyses provide strong support for the placement of the new genus within a clade containing both thalassoalaimus and litinum, and within the oxystomininae. interestingly, the placement of cricohalalaimus together with thalassoalaimus and litinium indicates that this genus, which is characterized by conflicting features used to define the halalaiminae (elongated amphideal aperture) and oxystomininae (female with single posterior ovary), should be placed within the oxystomininae and not in the halalaiminae as is currently the case (smol et al. 2014). an analogous change to the classification of the rhabdocoma cobb, 1920 (enoplida, trefusiidae gerlach, 1966 (de ley & blaxter 2004)) was suggested by shi & xu (2017), who found that ssu phylogenetic relationships supported the classification of this genus with the halononchinae wieser & hopper, 1967, a classification consistent with variation in the structure of the female reproductive system, and not in the trefusiinae gerlach, 1966, as previously suggested based on buccal morphology. similarly, in the case of cricohalalaimus, it appears that the shape of the amphideal aperture is not as informative as the structure of the female reproductive system for subfamily-level classification, and we therefore propose that cricohalalaimus be moved to the oxystomininae. acknowledgments we thank the co-voyage leaders ronnie n. glud and ashley a. rowden and science party of voyage tan1711, and the officers and crew of rv tangaroa for their contribution to sample collection. the voyage was funded by european research council advanced grant (erc adg 2014 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of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. https://doi.org/10.1007/s12526-017-0690-7 https://doi.org/10.11646/zootaxa.3872.1.5 european journal of taxonomy 595: 1–17 issn 2118-9773 https://doi.org/10.5852/ejt.2020.595 www.europeanjournaloftaxonomy.eu 2020 · gobert s. et al. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:f81a7282-a44b-4e70-9a44-fe8f67e5c1ea 1 schizorhynchia meixner, 1928 (platyhelminthes, rhabdocoela) of the iberian peninsula, with a description of four new species from portugal stefan gobert 1, marlies monnens 2,*, lise eerdekens 3, ernest schockaert 4, patrick reygel 5 & tom artois 6 1,2,3,4,5,6 hasselt university, centre for environmental sciences, research group zoology: biodiversity and toxicology, agoralaan gebouw d, b-3590 diepenbeek, belgium. * corresponding author: marlies.monnens@uhasselt.be 1 email: stefan.gobert@uhasselt.be 3 email: lise__eerdekens@hotmail.com 4 email: ernest.schockaert@uhasselt.be 5 email: patrick.reygel@uhasselt.be 6 email: tom.artois@uhasselt.be 1 urn:lsid:zoobank.org:author:5a55d3d7-b529-41fa-aa02-ee554f4a8cf9 2 urn:lsid:zoobank.org:author:782f71e0-ef84-48da-be72-8e205cb78eac 3 urn:lsid:zoobank.org:author:11c7606c-7677-4f9b-9295-604dabfc1dca 4 urn:lsid:zoobank.org:author:73da9dfc-69db-4168-88fa-b0ed54c88ddb 5 urn:lsid:zoobank.org:author:481991c8-ba09-457f-81ea-937c7a3dfd91 6 urn:lsid:zoobank.org:author:2edde35c-a2f0-4ca2-84aa-2a7893c40ac4 abstract. during several sampling campaigns in the regions of galicia and andalusia in spain and the algarve region in portugal, specimens of twelve species of schizorhynch rhabdocoels were collected. four of these are new to science: three species of proschizorhynchus (p. algarvensis sp. nov., p. arnautsae sp. nov. and p. troglodytus sp. nov.) and one species of schizochilus (s. coninxae sp. nov.). the new species of proschizorhynchus can be distinguished from their congeners by the curvature and length of their stylet, as well as the cirrus sheath and the organisation of the genital system. schizochilus coninxae sp. nov. has a distinct two-part stylet that is unique within the genus. in addition to these new species, new data are reported for the diascorhynchid diascorhynchus caligatus and the schizorhynchids carcharodorhynchus multidentatus, c. tenuis, c. flavidus, proschizorhynchus pectinatus and p. reniformis. finally, new records for c. flavidus from the hawaiian archipelago are presented. keywords. kalyptorhynchia, microturbellarians, meiobenthos, systematics, taxonomy. gobert s., monnens m., eerdekens l., schockaert e., reygel p. & artois t. 2020. schizorhynchia meixner, 1928 (platyhelminthes, rhabdocoela) of the iberian peninsula, with a description of four new species from portugal. european journal of taxonomy 595: 1–17. https://doi.org/10.5852/ejt.2020.595 https://doi.org/10.5852/2019.ejt.595 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:f81a7282-a44b-4e70-9a44-fe8f67e5c1ea mailto:marlies.monnens%40uhasselt.be?subject= mailto:stefan.gobert%40uhasselt.be?subject= mailto:lise__eerdekens%40hotmail.com?subject= mailto:ernest.schockaert%40uhasselt.be?subject= mailto:patrick.reygel%40uhasselt.be?subject= mailto:tom.artois%40uhasselt.be?subject= http://zoobank.org/urn:lsid:zoobank.org:author:5a55d3d7-b529-41fa-aa02-ee554f4a8cf9 http://zoobank.org/urn:lsid:zoobank.org:author:782f71e0-ef84-48da-be72-8e205cb78eac http://zoobank.org/urn:lsid:zoobank.org:author:11c7606c-7677-4f9b-9295-604dabfc1dca http://zoobank.org/urn:lsid:zoobank.org:author:73da9dfc-69db-4168-88fa-b0ed54c88ddb http://zoobank.org/urn:lsid:zoobank.org:author:481991c8-ba09-457f-81ea-937c7a3dfd91 http://zoobank.org/urn:lsid:zoobank.org:author:2edde35c-a2f0-4ca2-84aa-2a7893c40ac4 https://doi.org/10.5852/ejt.2020.595 european journal of taxonomy 595: 1–17 (2020) 2 introduction the species-rich platyhelminth taxon rhabdocoela ehrenberg, 1831 consists of two sister clades: kalyptorhynchia graff, 1905, easily recognised by the presence of a proboscis at the anterior body end, and dalytyphloplanida willems, wallberg, jondelius, littlewood, schockaert & artois, 2006, which lack such an organ (willems et al. 2006). within kalyptorhynchia, two major groups are distinguished: eukalyptorhynchia meixner, 1928, a paraphyletic assemblage of kalyptorhynchs with an undivided, conorhynch proboscis, and schizorhynchia meixner, 1928, a monophyletic group characterised by a split proboscis (tessens et al. 2014). so far, the study of microturbellarians in the iberian peninsula has focused primarily on non-marine environments (e.g., gieysztor 1931; farías et al. 1995; gamo & noreña-janssen 1998; noreña et al. 1999). the brackish-water habitats of galicia, asturias, cantabria and país vasco (noreña et al. 2007), the doñana national park in andalusia (van steenkiste et al. 2011) and the algarve (schockaert 2014; schockaert et al. 2017, 2019) have been investigated to some extent. in total, approximately 100 species of rhabdocoela have been reported from freshwater habitats in the iberian peninsula, the majority of which belong to dalytyphloplanida (tyler et al. 2006–2019a; van steenkiste et al. 2011). only 14 species of kalyptorhynchia are known from the iberian peninsula (noreña et al. 2007; schockaert et al. 2017). five of these belong to schizorhynchia: baltoplana magna karling, 1949, cheliplana cf. setosa evdonin, 1971, proschizorhynchus martinezi gobert, reygel & artois, 2017 and two records of unidentified species of karkinorhynchus meixner, 1928 (noreña et al. 2007; gobert et al. 2017). as part of an investigation into the systematics of schizorhynchia as a whole, sampling was undertaken in various locations across the iberian peninsula, including galicia (august 2008) and andalusia (march–april 2008) in spain, as well as the algarve region in portugal (september 2011, september 2012, may 2013, september 2014). in this paper, we provide an overview of the schizorhynch taxa found during these sampling campaigns. all twelve of the species found are new to the region and four are new to science. in addition, we report a number of new geographic records for some of these species outside of the iberian peninsula. material and methods specimens were collected from coastal regions throughout the iberian peninsula. sampling was performed at different time points in the summer and springtime of 2008, 2011, 2012, 2013 and 2014. sand samples were either retrieved from beaches or the subtidal zone. in the latter case, sediment was obtained manually or with a slurp gun by diving. conspecific specimens collected during previous sampling campaigns by members of the zoology group at hasselt university are also included in this work. this includes specimens from several locations in hawaii, usa. exact localities of these and the newly obtained material are listed for each species in the results section. meiofaunal interstitial kalyptorhynchs were extracted from the sediment using the mgcl2-decantation method (schockaert 1996). collected specimens were studied and drawn alive. of each species, at least one specimen was whole-mounted in lactophenol. whole-mounts were later studied in detail using nomarski interference contrast microscopy (leica dm 2500 and reichert-jung polyvar). measurements were taken axially unless indicated otherwise. the position of organs in the body is expressed as a percentage of the body length. all material, including holotypes, is deposited in the collection of hasselt university (hu numbers). abbreviations b = bursa cb = copulatory bulb gobert s. et al., schizorhynchia of the iberian peninsula 3 ci = cirrus gp = genital opening hg = haptic girdle ov = ovary p = proboscis pcs = post-cerebral septum pg = proboscis glands ph = pharynx pp = penis papilla sp = spines st = stylet t = testis vg = prostatic vesicle (‘vesicula granulorum’) vs = seminal vesicle. results class rhabditophora ehlers, 1985 subclass trepaxonemata ehlers, 1984 order rhabdocoela ehrenberg, 1831 suborder kalyptorhynchia graff, 1905 infraorder schizorhynchia meixner, 1928 family schizorhynchidae graff, 1905 genus schizochilus boaden, 1963 schizochilus coninxae sp. nov. urn:lsid:zoobank.org:act:001130a5-1784-4395-8619-0d5ad614007d fig. 1a–c etymology this species is named in honour of hasselt university researcher and close friend to the first author, laura coninx. material examined holotype portugal • 1 whole-mount; algarve region, lagoon near the ccmar ramalhete research station; 19 sep. 2012; e. schockaert leg.; high eulittoral zone, medium-fine sand among spartina von schreber, little detritus or silt, reduced layer more than 5 cm deep; hu 614. other material portugal • 3 whole-mounts (two of these of insufficient quality for taxonomic work); same collection data as for holotype; hu x.1.32 • 2 whole-mounts (photographs of live specimens available); olhos de água; 37°05′20″ n, 08°11′17″ w; 15 may 2013; b. tessens leg.; dry sand and algae mixture collected from eastern side of a rocky outcrop; hu x.1.33–x.1.34. description specimens approximately 2 mm long, measured on whole-mounts. eyes are absent. the anteriorlypositioned proboscis (p, fig. 1a) is relatively small (1⁄10–1/8 of the total body length). a pharynx rosulatus (ph, fig. 1a) is present in the caudal body half. a single, seemingly continuous haptic girdle is present at the caudal body end. http://zoobank.org/urn:lsid:zoobank.org:act:001130a5-1784-4395-8619-0d5ad614007d european journal of taxonomy 595: 1–17 (2020) 4 the common genital opening is situated in the posterior, at about 80%. a single row of four globular testes (t, fig. 1a) is present anterior to the pharynx. a pair of seminal vesicles (vs, fig. 1a) enter the proximal end of an elongate copulatory bulb separately. a large prostatic vesicle (vg, fig. 1a) occupies the proximal half of the copulatory bulb. the walls of the copulatory bulb are muscular, with several fig. 1. schizochilus coninxae sp. nov. a. habitus of live specimen collected at the type locality. b–c. distal end of the copulatory bulb, with stylet and penis papilla. b. holotype (hu 614). c. a specimen from the reference collection of hasselt university (hu x.1.32). scale bars: 20 µm. gobert s. et al., schizorhynchia of the iberian peninsula 5 marked constrictions present along the length of the bulb. the distal part contains the stylet (st, fig 1a– c). the stylet consists of a straight, tubular proximal part, 17–22 μm long (x̅ = 20 μm, n = 4) and with a diameter of 5 μm, and an irregular, folded ‘plate’or ‘sheet’-like distal part. the sheet-like distal part is a rather flexible extension that can be positioned anywhere from perpendicular to the tubular part of the stylet, to lying in its extension. in some live specimens, this part was twisted into a corkscrewlike shape. the entire stylet measures 58–63 μm (x̅ = 60 μm, n = 4) and lies within a thin-walled, unarmed cirrus. the distal part of the copulatory bulb forms a sclerotised penis papilla, which measures approximately 20 μm. remarks the unarmed, elongate, split proboscis unambiguously places the new species in the family schizorhynchidae. the combination of testes organisation in a single row, a pharynx occurring in the second body half and the sclerotised part of the copulatory organ consisting of (at least) a cirrus indicates that the specimens we retrieved belong to the genus schizochilus boaden, 1963. a tubular stylet is present within the cirrus, similar to the organisation of the copulatory organ in s. bruneti noldt & hoxhold, 1984, s. caecus l’hardy, 1963, s. choriurus boaden, 1963, s. hoxholdi karling, 1989, s. tubulatus brunet, 1970, s. parvulus brunet, 1970, s. santacruzensis noldt & hoxhold, 1984 and s. lanzarotensis gobert, reygel & artois, 2017. whereas in most of these species the cirrus is at least partly covered with spines, this is not the case in our specimens, nor in s. tubulatus or s. bruneti. our specimens differ from the latter two species in the sclerotisation of the cirrus: in s. tubulatus and s. bruneti, the cirrus is thick and characterised by longitudinal grooves, while it is only lightly sclerotised in our specimens. moreover, the distal sheet-like part we observed is unique for schizochilus and justifies the establishment of a new species: s. coninxae sp. nov. genus proschizorhynchus meixner, 1928 in the past, different terminologies have been used to describe the copulatory structures in proschizorhynchus (e.g., l’hardy 1965; noldt 1985). here, we apply the terminology put forward by noldt (1985), as this work comprises the most extensive overview of the genus to date. proschizorhynchus algarvensis sp. nov. urn:lsid:zoobank.org:act:b9703f24-3367-4329-bfb0-8a36051890b0 fig. 2a etymology the species epithet, ‘algarvensis’, refers to the algarve region of southern portugal, where the specimen was collected. material examined holotype portugal • 1 whole-mount (photographs of live specimen available); algarve region, sagres, east of mareta beach; 37°00′10″ n, 08°56′06″ w; 20 may 2013; b. tessens leg.; coarse sand with shell gravel; hu 615. description the living specimen is colourless. a pair of reniform eyes is present. a pair of small, round testes is situated in the anterior half of the body. the atrial genital organs are situated in the posterior quarter of the body, surrounding the common genital pore. a pair of seminal vesicles enters the proximal end of http://zoobank.org/urn:lsid:zoobank.org:act:b9703f24-3367-4329-bfb0-8a36051890b0 european journal of taxonomy 595: 1–17 (2020) 6 the elongate copulatory bulb, along with the gland necks of a number of prostatic glands. the distal part of the copulatory bulb contains the sclerotised parts of the copulatory apparatus (fig. 2a). these consist of a stylet (st, fig. 2a) enclosed by a sclerotised and grooved cirrus sheath (ci, fig. 2a) and surrounded by a penis papilla (pp, fig. 2a). the stylet measures 207 μm and is curved back upon itself. the distal edge of the penis papilla is armed with rod-like spines (sp, fig. 2a). the cirrus sheath surrounding the stylet measures 80 μm and has a distal opening of 73 μm. a small, sclerotised, tubular spermatic duct connects the bursa to the single ovary. remarks see taxonomic discussion following p. reniformis brunet, 1970. proschizorhynchus arnautsae sp. nov. urn:lsid:zoobank.org:act:e581df16-97c5-4f81-9702-bb79821ab620 fig. 2b–c etymology this species is named in honour of fellow hasselt university researcher and dear friend to the first author, natascha arnauts. material examined holotype portugal • 1 whole-mount (photographs of live specimen available); olhão, garganta canal; 37°01′34″ n, 07°50′56″ w; 13 sep. 2011; e. schockaert leg.; mid-eulittoral zone, fine, muddy sand; hu 616. other material portugal • 1 whole-mount (photographs of live specimen available); algarve region, faro, parque natural da rio formosa, culatra island; 37°00′00″ n, 07°49′16″ w; 17 may 2013; b. tessens leg.; fine, silty sand, with many bivalve shells and seagrass, from creek in a bay on the eastern shore of the island; hu x.1.35. • 1 whole-mount (photographs of live specimen available); algarve region, olhos de água; 37°05′20″ n, 08°11′16″ w; 15 may 2013; b. tessens leg.; coarse sand between rocks on the eastern side of a rocky outcrop; hu x.1.36 • 1 whole-mount (photographs of live specimen available); algarve region, olhos de água; 37°05′20″ n, 08°11′16″ w; 15 may 2013; b. tessens leg.; fine clean sand from intertidal at open beach, about 0.5 m deep, west of rocky outcrop; hu x.1.37. description live specimens measure approximately 1.6 mm in length and are colourless. eyes are absent. an elongate split proboscis, approximately 1⁄10 of the total body length (when relaxed), is situated at the anterior end of the body. connecting proximally to the muscular proboscis is a pair of small gland sacs. a rosulate pharynx is present in the anterior half of the body. two small, globular testes are positioned anterior to the pharynx. a pair of seminal vesicles enters an ovoid copulatory bulb, the proximal part of which is occupied by a prostatic bulb. distally, the copulatory bulb contains the sclerotised copulatory parts (fig. 2b–c), which consist of a very short, slender stylet, set in a cirrus sheath ornamented with parallel longitudinal grooves. the stylet is between 26 and 36 μm long (x̅ = 31 μm, n = 4). in most specimens, the stylet is completely enveloped by the cirrus sheath, meaning the tip does not protrude beyond the distal rim of the copulatory organ. however, in some instances it can protrude a little (as in fig. 2c), probably depending on the organ’s contraction or flattening of the specimen. the distal rim of the cirrus sheath does not bear spines. http://zoobank.org/urn:lsid:zoobank.org:act:e581df16-97c5-4f81-9702-bb79821ab620 gobert s. et al., schizorhynchia of the iberian peninsula 7 fig. 2. copulatory organ of species of proschizorhynchus meixner, 1928. a. proschizorhynchus algarvensis sp. nov., holotype (hu 615). b–c. p. arnautsae sp. nov. b. holotype (hu 616). c. a specimen from the reference collection of hasselt university (hu x.1.35). d–e. p. troglodytus sp. nov. d. holotype (hu 617). e. a specimen from the reference collection of hasselt university (hu x.1.40). f–g. p. pectinatus l’hardy, 1965 (hu x.1.43). f. copulatory organ. g. sclerotised spermatic duct. h. p. reniformis brunet, 1970 (hu x.1.45). scale bars: a, f = 50 µm; b–e, g–h = 20 µm. european journal of taxonomy 595: 1–17 (2020) 8 the female genital system could not be studied in detail. a uterus and bursa were observed to lie caudally with respect to the male copulatory apparatus. a single ovary was observed even more caudally. paired vitellaria run laterally along either side of the body. remarks see taxonomic discussion following p. reniformis. proschizorhynchus troglodytus sp. nov. urn:lsid:zoobank.org:act:c0548d31-d347-4b4d-8990-9d72f7fd8ced fig. 2d–e etymology the species epithet ‘troglodytus’, meaning ‘cave-dweller’, refers to the type locality, which is situated in a submerged cave. material examined holotype portugal • 1 whole-mount (photographs of live specimen available); algarve region, carvoeiro, beach at vale de covo; 37°05′34″ n, 08°27′33″ w; 27 may 2013; b. tessens leg.; fine, clean sand in rockpool near cave mouth at 1 m depth; hu 617. other material portugal • 1 whole-mount (photographs of live specimen available); algarve region, carvoeiro, beach below vale de covo; 37°05′32″ n, 08°27′33″ w; 27 may 2013; b. tessens leg.; inside a cave at about 0.5 m depth, fine to medium, clean sand, with fine shell debris; hu x.1.39 • 1 whole-mount (photographs of live specimen available); algarve region, carvoeiro, beach below vale de covo; 37°05′33″ n, 08°27′35″ w; 27 may 2013; b. tessens leg.; fine to medium, clean sand, with fine shell debris between rocks at beach in cave mouth, at low-water line; hu x.1.40. description specimens have a pair of pigmented eyes. the elongate, split proboscis is unarmed and lies at the anterior body end. the copulatory apparatus consists of an elongate bulb, with a pair of small seminal vesicles entering it proximally. the distal end of the copulatory bulb holds the sclerotised copulatory apparatus. this consists of a long stylet with a plate-like, widened base, set in a grooved cirrus sheath and surrounded by a sclerotised penis papilla (fig. 2d–e). the stylet is curved and measures 70–76 μm (x̅ = 74, n = 3) overall. approximately the distal half of the stylet protrudes beyond the rim of the penis papilla when the copulatory organ is in its relaxed state. the distal rim of the cirrus sheath is lined with 25-μm-long, curved spines. vitellaria are paired and occur laterally. a single ovary is situated near the caudal end of the specimen. the collected material did not allow for a more detailed study of the female system. remarks see taxonomic discussion following p. reniformis. http://zoobank.org/urn:lsid:zoobank.org:act:c0548d31-d347-4b4d-8990-9d72f7fd8ced gobert s. et al., schizorhynchia of the iberian peninsula 9 proschizorhynchus pectinatus l’hardy, 1965 fig. 2f–g material examined portugal • 1 whole-mount; algarve region, faro, ria formosa; 36°59′53″ n, 07°55′41″ w; 19 sep. 2012; e. schockaert leg.; low eulittoral zone, loose sand with some organic detritus and very little silt; hu x.1.42. spain • 2 whole-mounts; doñana national park; 6 apr. 2008; b. tessens and n. van steenkiste leg.; sand with arenicola marina (linnaeus, 1758) lamarck, 1801 collected from beach near the river mouth; hu x.1.43–x.1.44. previously known distribution english channel coast, france (l’hardy 1965); near marseille, france (brunet 1970); rhine-meusescheldt delta, the netherlands (schockaert et al. 1989). remarks specimens eyeless, measuring 2.5–2.7 mm in length (measured on two whole-mounts). the stylet (fig. 2f) is thin and strongly curved, with a widened proximal base. distally, the stylet shows a small, spindle-like piece that is separated from the rest of the stylet by a narrow constriction. the entire stylet has a length of around 93–102 μm (x̅ = 100 μm, n = 3). the stylet is surrounded by a 49 μm long curved cirrus sheath (cirrus in l’hardy 1965) with parallel grooves. the distal edge of the sheath is armed with spines, measuring 24–30 μm. these measurements are more or less in accordance with those provided by l’hardy (1965). a tubular, sclerotised spermatic duct with a wide (bulbous) region at the bursa-facing end is present (fig. 2g). proschizorhynchus reniformis brunet, 1970 fig. 2h material examined portugal • 1 whole-mount (photographs of live specimen available); algarve region, sagres, east of mareta beach; 37°00′10″ n, 08°56′06″ w; 20 may 2013; b. tessens leg.; coarse sand and shell gravel at 12 m depth; hu x.1.45. previously known distribution baie de cassis, provence-alpes-côte d’azur, france (brunet 1970). remarks the slender, curved stylet measures 65 μm in length and lies in a 30-μm-long, grooved cirrus sheath (fig. 2h). a sclerotised, plate-like structure, approximately 25 μm in length, extends from the stylet’s proximal base, perpendicular to the rim of the cirrus sheath, towards the distal end of the stylet. the stylet is slightly longer than the size range described by brunet (1970), who mentions stylet lengths of 40–45 μm. otherwise, our specimen conforms well to the description of p. reniformis. taxonomic discussion: proschizorhynchus proschizorhynchus is one of the most species-rich genera within schizorhynchidae (tyler et al. 2006– 2019b). the genus is relatively uniform in its general morphology and is characterised by a copulatory organ consisting of a grooved cirrus sheath, a stylet and a hardened penial papilla (see noldt 1985). european journal of taxonomy 595: 1–17 (2020) 10 the various species are distinguished from each other based largely on the structure of the copulatory organ. the highly characteristic copulatory apparatus, with a sheath showing longitudinal striation, places our three newly-described species in the genus proschizorhynchus. within schizorhynchia, only typhlorhynchus laidlaw, 1902 shares this feature of the male copulatory system. however, this genus is characterised by the lack of a proboscis, placing our specimens unambiguously within proschizorhynchus. the lack of spines or rods on the distal rim of the cirrus sheath in p. arnautsae sp. nov. is a feature shared with p. gullmarensis karling, 1950, p. reniformis brunet, 1970 and p. lunatus brunet, 1970. in p. gullmarensis and p. reniformis, however, the stylet is much longer, protruding far beyond the rim of the cirrus sheath and, in p. reniformis, curved back upon itself. furthermore, in both species, a bundle of spines is attached to the distal cirrus ridges (see noldt & hoxhold 1984), a feature not seen in p. arnautsae sp. nov. finally, both species have a larger plate-like extension at the base of the stylet that is not present either in p. arnautsae sp. nov. proschizorhynchus lunatus has a stylet comparable in length to that of p. arnautsae sp. nov. (30–35 µm and 26–36 µm long, respectively) and is generally similar in the proportions of the copulatory organ. however, in p. lunatus, the cirrus sheath forms the base of a curved, hardened distal penis, with a narrow distal opening (brunet 1970; noldt 1985). such a structure is unique within the genus and was not observed in any of the specimens of p. arnautsae sp. nov. either. proschizorhynchus troglodytus sp. nov. resembles p. pectinatus l’hardy, 1965 and p. arenarius de beauchamp, 1927 in its strongly curved and very long (with respect to the cirrus sheath) stylet. moreover, these three species all possess a plate-like extension at the base of the stylet, but this is shared with some other species within the genus (e.g., p. gullmarensis, see above). according to l’hardy (1965), p. pectinatus and p. arenarius are very difficult to distinguish. this is, at least in part, due to the inadequate description of p. arenarius (see noldt 1985). for example, it is unclear whether the continuation of the longitudinal ridges on the sheath of p. arenarius drawn and described by de beauchamp (1927) represent spines of the type present in p. pectinatus and most other representatives of the genus, or are the result of partial eversion of the cirrus (noldt 1985). both p. troglodytus sp. nov. and p. pectinatus have a sclerotised piece at the distal end of the stylet that is somewhat separated from the rest by a slight constriction. however, in p. pectinatus this piece is leaf-shaped, while in p. troglodytus sp. nov. it is cylindrical and robust. in fact, the entire stylet of p. troglodytus sp. nov. has a thicker and more robust appearance than that of p. pectinatus. furthermore, the stylet in p. troglodytus sp. nov. barely protrudes beyond the distal end of the spines on the cirrus sheath, while in p. pectinatus (and p. arenarius), the stylet is much longer relative to the cirrus sheath and spines. the copulatory apparatus of p. algarvensis sp. nov. also bears a great resemblance to that of p. pectinatus in the shape of the stylet and the presence of long spines on the distal rim of the penis papilla. however, the stylet of p. algarvensis sp. nov. is longer and much more strongly curved than that of p. pectinatus (207 μm in p. algarvensis sp. nov., approx. 100 μm in p. pectinatus). moreover, it appears to protrude less far beyond the spines of the cirrus sheath in p. pectinatus than it does in p. algarvensis sp. nov. in p. pectinatus the stylet also has a distal constriction, which is lacking in p. algarvensis sp. nov. furthermore, p. algarvensis sp. nov. has a pair of pigmented eyes, while p. pectinatus and p. arenarius are eyeless. gobert s. et al., schizorhynchia of the iberian peninsula 11 genus carcharodorhynchus meixner, 1938 carcharodorhynchus tenuis brunet, 1979 fig. 3a–d material examined portugal • 1 whole-mount (photographs of live specimen available); algarve region, sagres, eastward of mareta beach; 37°00′10″ n, 08°56′06″ w; 20 may 2013; b. tessens leg.; coarse sand and shell gravel, depth 12 m; hu x.1.46. previously known distribution bay of marseille, france (brunet 1979). remarks the specimen is eyeless. live specimens are colourless and transparent. a single, continuous adhesive girdle is present at the caudal body end. a rosulate pharynx is present at about 65%. the proboscis is highly asymmetrical: the ventral tongue measures approximately 44 μm, while the larger dorsal tongue measures 61 μm in length. the proboscis armature consists of rows of small, triangular teeth. on the ventral proboscis tongue, four to six teeth are set in a single row on either side of the tongue, with the teeth increasing in size distally. these teeth have a basal diameter of up to 4 μm and measure up to 3 μm in length. on the dorsal proboscis tongue, the teeth are arranged in multiple rows. in the distal part of the proboscis, the teeth are largest, with a diameter of 1–2 μm and a length of approximately 2 μm. these are more or less set in a single row. in the middle section of the proboscis tongue, the proboscis teeth are small and form a field of 3–4 rows. at the base of the dorsal tongue, the teeth are once again arranged in a single row. in the area connecting the tongues, the teeth form another wider field of about three rows. two very long seminal vesicles enter the proximal end of the copulatory bulb. the proximal ends of the seminal vesicles lie on either side of the pharynx. the male copulatory system holds a sclerotised, conical cirrus with a length of 15 μm, armed with very fine spines. in our specimen, the proximal spines have a diameter of 3–4 μm in the proximal region and 11–12 μm in the distal region of the cirrus. the male copulatory bulb connects to the male atrium, which is surrounded by circular to slightly diagonal muscles and leads into the common genital atrium. the common genital opening is situated at about 80%. a pair of ovaries is situated caudally, near the adhesive belt. no further details of the female reproductive system could be observed. the arrangement of the proboscis armature differs somewhat from the proboscis armature as described by brunet (1979). in particular, the fields of spines are wider (four to six rows of spines, instead of just three or four). carcharodorhynchus isolatus schilke, 1970 has a very similar organisation of the proboscis teeth. however, rather than two fields of spines, interrupted by a short section in which the spines are in a single row, the spines form a long stretch of three to four continuous rows. furthermore, c. isolatus has a substantially larger proboscis, even though the size of the proboscis teeth is approximately the same as those reported here. with a length of 15 µm, the cirrus of our portuguese specimen is slightly longer than what has been described for c. tenuis (9 µm). in addition, brunet (1979) mentions a tubular shape, while our specimen displays a more conically-shaped cirrus. however, this difference is likely the result of either the flattening of the specimen or the state of contraction of the copulatory bulb (or a combination of both). european journal of taxonomy 595: 1–17 (2020) 12 carcharodorhynchus flavidus brunet, 1967 fig. 3e–h material examined portugal • 1 whole-mount (photographs of live specimen available); algarve region, olhos de água; 37°05′20″ n, 08°11′17″ w; 15 may 2013; b. tessens leg.; dry sand and algae mixture, scraped from stones at eastern side of the rocky outcrop; hu x.1.47 • 1 whole-mount (photographs of live specimen available); algarve region, ilha de culatra; 36°59′10″ n, 07°50′20″ w; 17 may 2013; b. tessens leg.; exposed beach opposite village, at 30 cm depth, slightly above low-water line; hu x.1.48 • 1 wholemount; algarve region, carvoeiro, beach below vale de covo; 37°05′34″ n, 08°27′35″ w; 27 may 2013; b. tessens leg.; algae and sand scraped from rocks; hu x.1.49. usa – hawaii • 1 whole-mount (photographs of live specimen available); oahu, waimãnalo beach; 21°19′36″ n, 157°40′59″ w; 28 may 2010; n. van steenkiste leg; fine sand with organic material in sheltered habitats between coral at approximately 1.5 m depth; hu x.4.01 • 1 whole-mount (photographs of live specimen available); oahu, waimãnalo beach; 21°19′36″ n, 157°40′59″ w; 31 may 2010; n. van steenkiste leg; fine sand with organic material in between coral rocks; hu x.4.02 • 1 whole-mount (photographs of live specimen available); same collection data as for preceding; hu x.4.03. previously known distribution bay of marseille and nearby areas, mediterranean, france (brunet 1967); north carolina, usa (whitson et al. 2011); lanzarote, canary islands, spain (gobert et al. 2017); sardinia, italy (gobert et al. 2017). remarks live specimens are bright yellow. a single haptic girdle is present caudally. a large proboscis (fig. 3e–f) is present at the rostral end of the body. the proboscis is armed with triangular denticles on the inwardfacing surface. the fields of denticles are interrupted at the base of the proboscis tongues, producing a ‘basal gap’, typical for c. flavidus. a pair of seminal vesicles enter the copulatory bulb (fig. 3g–h) separately at its proximal end. prostatic glands occur proximally in the ovoid copulatory bulb. distally, it is provided with a straight cirrus, armed with small spines. the measurements of the portuguese specimens fall within the size ranges previously reported in the literature (brunet 1967; whitson et al. 2011; gobert et al. 2017). carcharodorhynchus flavidus has an extremely wide geographic range that includes the atlantic, the mediterranean and hawaii. this, along with the rather great variation in size this species exhibits, as well as some variation in proboscis dentition, has prompted some authors to suggest that c. flavidus as described by brunet (1967) may cover several species (whitson et al. 2011; gobert et al. 2017). carcharodorhynchus multidentatus brunet, 1979 fig. 3i–l material examined portugal • 1 whole-mount (photographs of live specimen available); algarve region, faro; 36°57′01″ n, 07°57′35″ w; 23 may 2013; b. tessens leg.; coarse sand with shell debris from the exposed part of the barrier island opposite faro at a depth of 26 m; hu x.1.50. spain • 1 whole-mount; andalusia, west of tarifa; 36°01′56″ n, 05°37′36″ w; 30 mar. 2008; b. tessens and n. van steenkiste leg.; fine sand from a long beach; hu x.2.01. gobert s. et al., schizorhynchia of the iberian peninsula 13 previously known distribution bay of marseille, france (brunet 1979). remarks our observations on the specimens collected in portugal and spain largely correspond to brunet’s (1979) original description. live specimens are transparent and measure 1.2–2.1 mm in length. a large proboscis, consisting of a pair of broad, flattened muscular tongues, is situated at the rostral body end (fig. 3i–j). the proboscis is more or less symmetrical, though the ventral tongue is more rounded than fig. 3. a–d. carcharodorhynchus tenuis brunet, 1979 (hu x.1.46). a–b. proboscis. c–d. copulatory bulb. e–h. c. flavidus brunet, 1967. e–f. proboscis (hu x.1.49). g–h. copulatory bulb (hu x.1.48– 49). i–l. c. multidentatus brunet, 1979 (hu x.1.50). i–j. proboscis. k–l. copulatory organ. scale bars: 20 µm. european journal of taxonomy 595: 1–17 (2020) 14 the dorsal tongue. two fields of numerous, very fine, sclerotised denticles run continuously across the inward-facing surface of both proboscis tongues. the denticles are arranged in 7–10 rows. the size of the denticles is largest on the rows closest to the lateral sides of the proboscis tongues and smallest in the rows facing the middle of the proboscis tongues. a rosulate pharynx is situated in the caudal body half, at approximately 65% of the body length. according to brunet (1979), paired rows of two to three testes follicles run laterally in the rostral body half. in the specimen from portugal, we observed only two large testes directly posterior to the proboscis. a pair of seminal vesicles empty into the proximal end of a short, ovoid-to-spherical copulatory bulb that is between 56 and 72 μm long (x̅ = 64 μm, n = 2). the proximal half to two-thirds of the copulatory bulb contains a prostatic vesicle. the cirrus occupies the distal third of the copulatory bulb, is 15 to 21 µm long and armed with minute spines, arranged in longitudinal rows. these measurements of the copulatory bulb are somewhat larger than what has been reported by brunet (1979) (cb: 53–57 μm, ci: 13–14 μm). a rather large size variation has also been reported in the copulatory organs of other species in the genus carcharodorhynchus, such as c. flavidus (see above). brunet (1979) did not describe the female system. in one live specimen, we observed a single ovary, situated to one side near the caudal adhesive belt. we were unable to ascertain whether a second ovary was present. the records mentioned here are the first published records of this species since its original description by brunet (1979) and are the first records outside of the french mediterranean. furthermore, two specimens present in the reference collection of hasselt university (hu viii.1.30 and hu viii.1.47), which were previously identified as c. flavidus (gobert et al. 2017), are now correctly identified as c. multidentatus. family diascorhynchidae meixner, 1929 genus diascorhynchus meixner, 1928 diascorhynchus caligatus ax, 1959 fig. 4 material examined portugal • 1 whole-mount (photographs of live specimen available); algarve region, olhos de água; 37°05′20″ n, 08°11′17″ w; 15 may 2013; b. tessens leg.; fine, clean sand from intertidal at open beach, west of rocky outcrop at depth of 0.5 m; hu x.2.06. fig. 4. diascorhynchus caligatus ax, 1959 (hu x.2.07). a. stylet. b. proboscis hooks. c. sclerotised spermatic duct. scale bars: 20 µm. gobert s. et al., schizorhynchia of the iberian peninsula 15 spain • 1 whole-mount; andalusia, doñana national park; 6 apr. 2008; b. tessens and n. van steenkiste leg.; beach at 5 km from river mouth, sandy sediment from the lower tidal mark; hu x.2.07. previously known distribution sile, black sea, turkey (ax 1959). remarks the overall habitus and internal organisation of the collected specimens correspond well to the description by ax (1959). the body length measures between 1.6 and 1.7 mm, which is at the lower end of the size range of 1.6–2.0 mm reported by ax (1959). the split proboscis bears a pair of large, sclerotised hooks (fig. 4b). the dorsal hook is more robust and measures 30–32 μm in length, while the ventral hook measures 31–34 μm. two elongate lateral gland sacs connect to the proximal end of the proboscis. the rosulate pharynx is situated slightly posterior to the body centre. six globular testes lie in a single, medial row anterior to the pharynx. the copulatory bulb varies in length between 61 μm and 86 μm and holds a sclerotised stylet (fig. 4a). the stylet measures 42 to 48 μm. these measurements are in line with the descriptions of ax (1959). a single ovary and a bursa occur near the caudal body end. in the specimen from doñana, we observed a sclerotised, spermatic duct, connecting the bursa to the ovary. similar to what was described by ax (1959), this duct is a straight tube, measuring around 26 μm in length and terminating at both ends in a small, sclerotised disk (fig. 4c). the spermatic duct was not observed in the portuguese specimen. vitellaria are paired and occur laterally, between the ovaries and the brain. discussion along with the newly-described species schizochilus coninxae sp. nov., proschizorhynchus algarvensis sp. nov., p. arnautsae sp. nov. and p. troglodytus sp. nov., all of the above-mentioned species are recorded from the iberian peninsula for the first time, bringing the total number of marine kalyptorhynch species in the area to 23, a near doubling of the number of species previously known from this region. for schizorhynchia in particular, the number of species has increased almost threefold with this contribution, going from only five previously-reported species to a total of 14. given the suggested ecological importance of schizorhynchia and other flatworms in the meiobenthic environment (see martens & schockaert 1986; boaden 1995), a thorough understanding of meiobenthic ecology and biodiversity in the iberian peninsula will require much more intensive sampling of this fauna. acknowledgements we thank the ministerio de educación y ciencia and the estación biológica de doñana (icts-rbd, spain) for the financial and logistical support of the fieldwork in doñana national park. collecting trips to the algarve were financed by the eu fp7 assemble grant agreement no. 227799. we would like to thank the centro de ciências do mar (centre of marine sciences) of the university of the algarve, portugal, for providing excellent working and sampling facilities, in particular dr ana amaral for the kind and efficient logistic support and dr jorge gonçalves for collecting the samples at the islands near sagres. special thanks also go to the captain of the zodiac for his help with sampling in the ria formosa. the research of s.g. was supported by a phd-grant of bof-uhasselt. m.m. is funded by a phd fellowship from the research foundation – flanders (fwo-vlaanderen). we would like to thank ria vanderspikken and natascha steffanie for their valuable assistance in the lab. dr bart tessens and dr niels van steenkiste are thanked for their involvement in collecting specimens from the algarve, portugal and doñana national park, spain. dave bosmans is cordially thanked for his help with processing the images included in this work european journal of taxonomy 595: 1–17 (2020) 16 references ax p. 1959. zür systematik, ökologie und tiergeographie der turbellarienfauna in den ponto-kaspischen brackwassergebieten. zoologische jahrbücher: abteilung für systematik, ökologie und geographie der tiere 87: 43–184. boaden p.j.s. 1995. where turbellaria? concerning knowledge and ignorance of marine turbellarian ecology. hydrobiologia 305: 91–99. https://doi.org/10.1007/bf00036368 brunet m. 1967. turbellariés schizorhynques de la région de marseille. sur carcharodorhynchus subterraneus et carcharodorhynchus flavidus nov. sp. bulletin de la société zoologique de france 92: 143–152. brunet m. 1970. turbellariés schizorhynchidae de sables infralittoraux de la côte marseillaise. cahiers de biologie marine 11: 279–306. brunet m. 1979. turbellaires calyptorhynques du golfe de marseille. revue de biologie et écologie méditerranéenne 6: 101–120. de beauchamp p. 1927. rhabdocoeles des sables à diatomées d’arcachon i. coup d’œil sur l’association schizorhynchidae. bulletin de la société zoologique de france 52: 351–386. farías f., gamo j. & noreña-janssen c. 1995. nuevas aportaciones al conocimiento de los microturbelarios de la península ibérica. graellsia 51: 93–100. https://doi.org/10.3989/graellsia.1995.v51.i0.399 gamo j. & noreña-janssen c. 1998. old and new records of turbellarians from the central areas of spain. hydrobiologia 383: 299–305. https://doi.org/10.1023/a:1003448029737 gieysztor m. 1931. contribution à la connaissance des turbellariés rhabdocoèles (turbellaria rhabdocoela) d’espagne. bulletin de l’académie polonaise des sciences et des lettres, classe des sciences mathematiques et naturelles b 2: 125–153. gobert s., reygel p. & artois t. 2017. schizorhynchia (platyhelminthes rhabdocoela) of lanzarote (canary islands), with the description of eight new species. marine biodiversity. https://doi.org/10.1007/s12526-017-0736-x l’hardy j.-p. 1965. turbellaries schizorhynchidae des sables de roscoff ii. le genre proschizorhynchus. cahiers de biologie marine 6: 135–161. martens p.m. & schockaert e.r. 1986. the importance of turbellarians in the marine meiobenthos: a review. hydrobiologia 132: 295–303. https://doi.org/10.1007/bf00046263 noldt u. 1985. typhlorhynchus syltensis n. sp. 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(turbellaria, kalyptorhynchia) from the coast of north carolina, usa. southeastern naturalist 10: 221–232. https://doi.org/10.1656/058.010.0203 willems w.r., wallberg a., jondelius u., littlewood d.t.j., backeljau t., schockaert e.r. & artois t.j. 2006. filling a gap in the phylogeny of flatworms: relationships within the rhabdocoela (platyhelminthes), inferred from 18s ribosomal dna sequences. zoologica scripta 35: 1–17. https://doi.org/10.1111/j.1463-6409.2005.00216.x manuscript received: 15 april 2019 manuscript accepted: 24 september 2019 published on: 21 january 2020 topic editor: rudy jocqué desk editor: pepe fernández printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. https://doi.org/10.11646/zootaxa.3872.5.8 http://dx.doi.org/10.11646/zootaxa.4227.1.3 http://dx.doi.org/10.11646/zootaxa.4603.1.4 https://doi.org/10.1111/zsc.12066 http://www.marinespecies.org/aphia.php?p=taxdetails&id=142393 http://www.marinespecies.org/aphia.php?p=taxdetails&id=853126 http://dx.doi.org/10.11646/zootaxa.2791.1.1 https://doi.org/10.1656/058.010.0203 https://doi.org/10.1111/j.1463-6409.2005.00216.x _goback european journal of taxonomy 613: 1–19 issn 2118-9773 https://doi.org/10.5852/ejt.2020.613 www.europeanjournaloftaxonomy.eu 2020 · marin i. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:66eb6c4d-0f36-46d7-871f-a1f988894e1d 1 a new species of the genus liljeborgia spence bate, 1862 (crustacea: amphipoda: liljeborgiidae) associated with the burrows of the spoon worm urechis unicinctus in the sea of japan ivan marin a.n. severtsov institute of ecology and evolution of ras, moscow, russia. email: coralliodecapoda@mail.ru, vanomarin@yahoo.com urn:lsid:zoobank.org:author:b26adaa5-5dbe-42b3-9784-3bc362540034 abstract. a new symbiotic species of liljeborgiid amphipods, liljeborgia associata sp. nov., is described from the burrows of the spoon worm urechis unicinctus (drasche, 1880) (annelida: polychaeta: echiura: urechidae) in the southern part of peter the great bay and posjeta bay in the sea of japan. the new species is mostly similar and probably related to liljeborgia geminata barnard, 1969, known from the californian coasts of the usa, and liljeborgia serratoides tzvetkova, 1967, described from posjeta bay in the sea of japan, but can be clearly distinguished from all congeners by morphological features of mouthparts, appendages and telson. the new species is only the fourth in the family liljeborgiidae to be described from the russian coast of the northwestern pacific and the first in association with spoon worms (echiura). keywords. diversity, association, northwestern pacific, boreal, barcoding. marin i. 2020. a new species of the genus liljeborgia spence bate, 1862 (crustacea: amphipoda: liljeborgiidae) associated with the burrows of the spoon worm urechis unicinctus in the sea of japan. european journal of taxonomy 613: 1–19. https://doi.org/10.5852/ejt.2020.613 introduction representatives of the family liljeborgiidae (crustacea: amphipoda) are relatively rare in the higher latitudes of the north pacific. only three genera, liljeborgia spence bate, 1862, idunella g.o. sars, 1894 (= listriella j.l. barnard, 1959) and sextonia chevreux, 1920, are known from the area. about 10+ species of the family are known from the ne pacific (eastern coasts of the bering sea; pacific coasts of north america), including four described species of the genus liljeborgia: liljeborgia cota barnard, 1962 known from the gulf of alaska to the northern baja california, mexico, at a depth of 366–2000 m; l. geminata barnard, 1969 (l. kinahani bate, 1862 is considered as a junior synonym), described from goleta to the northern baja california, at a depth of 3–70 m; l. marcinabrio barnard, 1979, known from bahia de los angeles, gulf of california, at a depth of 46 m; and l. pallida bate, 1857, having a wide area of distribution from ne atlantic to the central californian coasts, at a depth of 40–611 m (barnard 1959; barnard & karaman 1991; d’udekem d’acoz 2010; cadien 2015). moreover, two undescribed species are also presented in the ne pacific: one free-living, deep-water species and one commensal https://doi.org/10.5852/ejt.2020.613 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:66eb6c4d-0f36-46d7-871f-a1f988894e1d mailto:coralliodecapoda@mail.ru mailto:vanomarin@yahoo.com http://zoobank.org/urn:lsid:zoobank.org:author:b26adaa5-5dbe-42b3-9784-3bc362540034 https://doi.org/10.5852/ejt.2020.613 european journal of taxonomy 613: 1–19 (2020) 2 with pagurus hemphilli (j.e.benedict, 1892) (crustacea: decapoda: paguridae), living along the shores of central california (cadien 2015). the diversity of the family along the coasts of the nw pacific (western coasts of the bering sea; pacific coasts of russia and japan; sea of okhotsk; northern parts of the sea of japan and the yellow sea) is less studied and is obviously underestimated. only three species of the family have been reported from russian waters: liljeborgia serratoides tzvetkova, 1967 was described from the shallow waters of posjeta bay in the sea of japan (tzvetkova 1967), sextonia caecus labay, 2017 was recently described from the sea of okhotsk, at a depth of 109–309 m (labay 2017) and gurjanova (1951) reported northern atlantic and arctic lilljeborgia cf. fissicornis (м. sars, 1858) from the bering sea, representing the only known record of the species from the nw pacific (see d’udekem d’acoz & vader 2009). the geographically closest species of the family liljeborgiidae to the aforementioned species are known from the seto inland sea in southern japan, where liljeborgia japonica nagata, 1965 and l. serrata nagata, 1965, idunella chilkensis chilton, 1921 and i. curvidactyla nagata, 1965 were described by nagata (1965), and the yellow sea, where l. hwanghaensis kim & kim, 1990 and l. sinica ren, 1992 are also known (kim & kim 1990; ren 1992, 2007; ishimaru 1994). a variety of symbiotic communities associated with soft-bottom deep-burrowing invertebrates were investigated in the peter the great bay and posjeta bay in the sea of japan. the sampling of infaunal animals and symbiotic assemblage was carried out using a bait suction pump (yabby pump), which allowed us to study the diversity of burrowing crustaceans and their associates in russian waters (e.g., marin 2010, 2015, 2016, 2018a, 2018b; marin et al. 2011, 2013; marin & kornienko 2014) with the description of several new crustacean (marin 2013, 2017) and even a new phoronid species (temereva & chichvarkhin 2017). numerous specimens of liljeborgia sp. were also collected from the burrows of the spoon worm urechis unicinctus (drasche, 1880) (annelida: polychaeta: echiura: urechidae). previously, this species was identified as l. serratoides/geminata in accordance with its similar morphology and distinctive coloration, however, the presence of several specific morphological features has allowed its separation into a new species described herewithin. material and methods specimen sampling and imaging sampling was carried out in the estuary of volchanka river in vostok bay near the scientific station “vostok” (42°51′14.48″ n, 132°46′47.24″ e) and in astafieva bay (42°36′45.6″ n, 131°12′24.8″ e), both within peter the great bay, as well as troitza bay (42°38′60.0″ n, 131°07′27.8″ e) located within posjeta bay in sea of japan (see fig. 1). these bays are well known by large populations of the common spoon worm u. unicinctus and other burrowing animals. the hosts and symbiotic community were collected using a bait suction pump (yabby pump), which is actively used in the sampling of deep burrowing animals (e.g., eleftheriou & mcintyre 2005). unfortunately, in our study it was not possible to measure the length or volume of the host’s burrows. the collected specimens were photographed alive in situ using a canon g16 digital camera and then fixed with 90% ethanol solution. the drawings were made using camera a lucida attached to an olympus szx10 stereo microscope. molecular study to study the molecular genetic diversity, a fragment of the mitochondrial gene coding for cytochrome c oxidase subunit i (coi mtdna) gene marker was amplified, sequenced and compared. two female specimens (lemmi) from vostok bay were used for the moleculargenetic examination. total genomic dna was extracted from abdominal and pereiopod muscle tissue using the innuprep dna micro kit (analitikjena, germany) following the manufacturer’s protocol. the coi gene marker was amplified with the help of primers «m13polylco» marin i., a new species of liljeborgia from the sea of japan 3 (tgtaaaacgacggccagtgaytatwttcaacaaatcataaagatattgg) and «m13polyhco» (caggaaacagctatgactamacttcwgggtgaccaaaraatca) (carr et al. 2011); 16s with the help of +16sa (‘cgcctgtttatcaaaaacat’) and -16sh (‘ccggtctgaactcagatcacg’); 28s with the help of +c1 (‘acccgctgaatttaagcat’) and -d2 (‘tccgtgtttcaagacgg’). pcr products were performed on amplificator t100 (bio-rad, usa) under the following conditions: initial denaturation at 96°c for 1.5 min followed by 42 cycles of 95°c for 2 min, 49°c for 35 seconds, and 72°c for 1.5 min, followed by chain extension at 72°c for 7 min. the volume of 10 µl of reaction fig. 1. habitus of live female specimen of liljeborgia associata sp. nov. from vostok bay of the sea of japan with geographical indication of the collection sites in the peter the great bay and posjeta bay of the sea of japan. european journal of taxonomy 613: 1–19 (2020) 4 mixture contained 1 µl of total dna, 2 µl of 5 × pcr mix (dialat, russia) and 1 µl of each primer. the amplification products were separated by using gel electrophoresis of nucleic acids on a 1.5% agarose gel in 1 × tbe, and then stained and visualized with 0.003% etbr using imaging uv software. dna nucleotide sequences were determined using genetic analyzer abi 3500 (applied biosystems, usa) and bigdye 3.1 (applied biosystems, usa) with direct and reverse primers. unfortunately, the number of sequences of the representatives of the family liljeborgiidae deposited in genetic databases is still very small. as the phylogenetic relations of studied species are out of our interest, the obtained dna (coi mtdna) data are presented in the paper without any analysis. type material and morphological study the type material is deposited in the collection of zoological museum of moscow state university, moscow, russia (zmmu) and the laboratory of ecology and evolution of marine invertebrates (lemmi), a.n. severtzov institute of ecology and evolution, russian academy of sciences, moscow, russia. the body length (bl, mm), defined as the dorsal length from distal margin of head to the posterior margin of telson without the length of uropod iii and antennae, is used as a standard measurement. the terminology and the general model of the description are used in the study after d’udekem d’acoz & vader (2009) and d’udekem d’acoz (2010). abbreviations a1 = antenna 1 a2 = antenna 2 bl = body length ep1–3 = epimeral plates 1–3 gn1 = gnathopod 1 gn2 = gnathopod 2 md = mandible mx1 = maxilla 1 mx2 = maxilla 2 mxp = maxilliped p1–p7 = pereiopods 1–7 u1–u3 = uropods 1–3 results phylum arthropoda von siebold, 1848 class malacostraca latreille, 1802 order decapoda latreille, 1802 family liljeborgiidae stebbing, 1899 genus liljeborgia spence bate, 1862 liljeborgia associata sp. nov. urn:lsid:zoobank.org:act:cc2a771f-71b2-4eed-9c16-3ef17cd0655d figs 1–5 diagnosis a1 with stout articles 2 (about as long as wide) and 3 (about 1.5 times as long as wide), article 2 with dorsal projection produced into a rounded lobe; palp of md with long and slender articles, article 3 http://zoobank.org/urn:lsid:zoobank.org:act:cc2a771f-71b2-4eed-9c16-3ef17cd0655d marin i., a new species of liljeborgia from the sea of japan 5 almost equal to articles 1 and 2, about 5 times as long as wide; palp of mx1 with broad shovel-shaped article 1; stout articles 3 and 2 of a2, article 3 about as long as wide and article 2 about 3 times as long as wide; p1 and p2 with short and wide merus, about 3 times as long as wide; pp5–7 with slender propodal segments, about 6, 7 and 10 times as long as wide, respectively; posterodorsal area of pleonites 1 and 2 produced into 3 small teeth of which the median one is the longest; urosomites 1 and 2 with wellmarked dorsal crest; telson with long distal teeth, accompanied by 2 interdental long and slender spines. etymology the species is named after its symbiotic lifestyle. type material holotype pacific ocean • ♀ (bl 6.5 mm); sea of japan, primorye, peter the great bay, vostok bay, in front of the scientific station “vostok”; 42°54′35.8″ n, 132°44′08.7″ e; depth 1–1.5 m; 30–31 jul. 2017; i. marin leg.; shore in front of the laboratory, sandy-gravel bottom overgrown with sea grass; yabbypump; from burrows of spoon worm u. unicinctus; zmmu mb-1153. paratypes pacific ocean • 1 ♂; same data as for holotype; zmmu mb-1154 • 1 ♀; same data as for holotype; genbank: mn704855, mn704856; zmmu mb-1155. additional material pacific ocean • 8 ♀♀, 1 ♂; same data as for holotype; lemmi • 5 ♀♀; sea of japan, primorye, posjeta bay, troitza bay; 42°38′60.0″ n, 131°07′27.8″ e; depth 1–1.5 m; jul. 2019; i. marin leg.; muddy sand; inside burrow of spoon worm u. unicinctus; yabby-pump; lemmi • 1 ♀; peter the great bay, astafieva bay; 42°36′52.2″ n, 131°12′01.1″ e; depth 1–1.5 m; jul. 2019; i. marin leg.; sand bottom; inside burrow of spoon worm u. unicinctus; yabby-pump; lemmi. description head (fig. 2c). rostrum turned downward, distally acute; eye large (fig. 1), with well-developed ommatidia, eye brightly white in alive specimens (fig. 1). antenna 1 (fig. 2d). article 3 stout, about 1.5 times as long as wide; article 2 about as long as wide, with dorsal projection produced into a rounded lobe; major flagellum with 15–17 articles; accessory flagellum with 10–11 articles. antenna 2 (fig. 2e). article 3 about as long as wide, unarmed; article 2 about 3 times as long as wide, with 4 small dorsal spines, with 1 long distodorsal and 2 long simple distoventral spines, with simple setae ventrally; article 1 about 4–4.5 times as long as wide, with small dorsal spines, with 1 distoventral long spine, unarmed ventrally; flagellum with 13–15 articles. labrum (fig. 2a). upper lip with labrum broader than long and smaller than epistome, apical margin sinuous. epistome (fig. 2b). with rounded lobes, protruding in lateral view, armed with small setae dorsally. mandible (fig. 2f). lacinia mobilis large, anterior margin armed with 5 small strong teeth; palp consists of 3 long slender articles, similar in size, covered with long simple setae, article 1 almost equal to article 2, about 5–6 times as long as wide; article 2 about 5–6 times as long as wide; article 3 almost equal to articles 1 and 2, about 5 times as long as wide. european journal of taxonomy 613: 1–19 (2020) 6 fig. 2. liljeborgia associata sp. nov., female (lemmi) from vostok bay of the sea of japan. a. upper lip. b. lower lip. c. head. d. antenna 1. e. antenna 2. f. mandible. g. maxilla 1. h. maxilla 2. i. maxilliped. j. pleopod 1. k. telson. marin i., a new species of liljeborgia from the sea of japan 7 maxilla 1 (fig. 2g). outer plate with 8 large slender spines, ventrally denticulated; inner plate with a single long plumose seta; palp consists of 4 articles, article 1 shovel-shaped, with broad median part, about 1.5–2 times as long as wide, with 8–9 sharp robust spines along anterior margin, and simple small setae along dorsal margin. maxilla 2 (fig. 2h). inner and outer plates distally rounded, robust, covered with numerous long simple setae along anterior and lateral margins. maxilliped (fig. 2i). palp consists of 4 articles: article 1 of palp unarmed, article 2 with a cluster of long setae distodorsally, small robust setae distally and long simple setae along the inner margin, outer margin unarmed; article 3 with a cluster of long simple setae along anterior border and distal part of inner margin, outer margin unarmed, article 4 (dactylus) slender, curved, about 0.8 times as long as article 3, unarmed; outer plate with 13–15 robust spines along medial border (distal spines are narrow and rather long); inner plate covered with small setae along anterior margin. gnathopod 1 (fig. 3a). coxa trapezoidal, with anterior medial setae only, posterior border weakly concave; basis slender, about 6–7 times as long as wide, with ventral projection in proximal part, with long simple setae; ischium about as long as wide, with long simple setae along distoventral border; merus about 1.5 times as long as wide, sharping distoventrally, with several groups of setae along ventral margin; carpus slender, with blunt distoventral process, reaching ⅓ of palm length, not reaching propodal group of strong spines, armed with several groups of setae; propodus about twice as long as wide both in male and females (fig. 3b), with convex ventral margin, armed with small simple setae along almost all its length, with a depression and 3 long simple setae in proximal part level with end of dactylus, dorsal margin straight and unarmed; dactylus with 5–7 triangular teeth. gnathopod 2 (fig. 3c). very similar to gn1 in shape and slightly larger in size; coxa triangulo-elliptic; basis slender, about 6 times as long as wide, with long simple setae; ischium about as long as wide, with long simple setae along distoventral border; merus about 1.5 times as long as wide, sharpening distoventrally, with several groups of setae along ventral margin; carpus slender, with blunt distoventral process, reaching ¼ of palm length, not quite reaching propodal group of strong spines, armed with several groups of setae; propodus about 2 times as long as wide in both male (fig. 5a) and females (fig. 3d), dorsal margin straight and unarmed, with convex ventral margin, armed with small simple setae along almost entire length, with a depression and 3 long simple setae in proximal part level with end of dactylus; dactylus with 10–11 triangular teeth. pereiopod 3 (fig. 4a). coxa elliptic and narrow; basis slender, about 8–9 times as long as wide, with straight margins, covered with long simple setae; ischium about as long as wide, with long simple setae along distoventral border; merus broad, about 3 times as long as wide, slightly longer than carpus and equal to propodus, dorsal margin produced forward, with long simple setae along dorsal and ventral margins; carpus about 3 times as long as wide, with straight margins, unarmed dorsally and with simple long setae along ventral margin; propodus about 4–4.5 times as long as wide, slightly curved, unarmed dorsally and ventrally, with several long simple setae at distodorsal angle; dactylus of normal length, slender, weakly curved, slightly shorter than propodus and equal to carpus in length. pereiopod 4 (fig. 4b). coxa wide, with parallel anterior and posterior border, ventral border with 3 well marked serrations; basis slender, about 7–7.5 times as long as wide, with straight margins, covered with long simple setae; ischium about as long as wide, with long simple setae along distoventral border; merus broad, about 3 times as long as wide, longer than carpus and slightly shorter than propodus, dorsal margin produced forward, with long simple setae along dorsal and ventral margins; carpus about 2.5–3 times as long as wide, with straight margins, unarmed dorsally and with long simple setae along ventral margin; propodus about 5–5.5 times as long as wide, slightly curved, unarmed dorsally, with several european journal of taxonomy 613: 1–19 (2020) 8 fig. 3. liljeborgia associata sp. nov., female (lemmi) from vostok bay of the sea of japan. a. gnathopod 1. b. chela of gn1. c. gnathopod 2. d. chela of gn2. marin i., a new species of liljeborgia from the sea of japan 9 long simple setae at distodorsal angle, with 5 tiny spines along ventral margin; dactylus of normal length, slender, weakly curved, slightly shorter than propodus and slightly longer than carpus. pereiopod 5 (fig. 4c). coxa subquadrate, with medially concave ventral margin, unarmed; basis broad, with anterior and posterior border convex; anterior border with 12–13 small spines accompanied by small setae, posterior border with well-marked serration; ischium about as long as wide, with large simple spine on anterodistal corner; merus about 4 times as long as wide, with 4 short anterior spines, 1 large distodorsal spine paired with a spinule, and 1 simple distoventral spine; carpus about 4 times as long as wide, slightly shorter than merus, unarmed posteriorly, with 3 small anterior spines, and with a distal pair of spines anteriorly and posteriorly; propodus (fig. 4d) about 6 times as long as proximal width, slightly tapering distally, with 10 anterior spines and series of long medial setae; dactylus distinctly curved and of normal stoutness, with tip entire, about ½ of the length of propodus. pereiopod 6 (fig. 4e). coxa subquadrate, with medially concave ventral margin, unarmed; basis broad, with anterior and posterior border convex; anterior border with 10 small spines accompanied with small setae, posterior border with well-marked serration; ischium about as long as wide, with large simple spine on anterodistal corner; merus about 4 times as long as wide, with 5 short anterior spines, a pair of large distodorsal spines and 1 simple distoventral spine paired with a spinule; carpus about 4.5–5 times as long as wide, slightly shorter than merus, unarmed posteriorly, with 4 small anterior spines, and a distal pair of spines anteriorly and posteriorly; propodus (fig. 4f) about 7 times as long as proximal width, tapering distally, with 4 long anterior spines and series of long medial setae; dactylus distinctly curved, sharp, about ⅓ of the length of propodus. pereiopod 7 (fig. 4g). coxa almost rectangular; basis greatly broad, almost round, with convex anterior border and strongly convex posterior border; anterior border with 12 small conical spines, posterior border serrated, distal border with strong long spine; ischium about as long as wide, unarmed; merus broad, about 2.5–3 times as long as wide, with 4 anterior small spines and pair of anterodistal simple spines, 3 posterior spines and a pair of long posterodistal long spines; carpus about 5.5 times as long as wide, slightly longer than merus, with single or paired anterior spines and long simple setae posteriorly; propodus about 10 times as long as wide, tapering distally, with 4 small anterior spines and long setae posteriorly; dactylus straight, very long and slender, entire, about ⅓ of the length of propodus. pleonite 1 (fig. 5b). posterodorsal area produced into 3 small teeth of which median is longest (fig. 5e); ep1 with normally developed posteroventral tooth, with posterior border weakly convex, covered with numerous simple setae. pleonite 2 (fig. 5b). posterodorsal area produced into 3 small teeth of which median is longest (fig. 5e); ep2 with normally developed posteroventral triangular tooth, with posterior border distinctly convex, covered with numerous simple setae. pleonite 3 (fig. 5b). posterodorsal area toothless (fig. 5e); ep3 with small posteroventral tooth and distinct rounded notch (fig. 5c), with posterior border convex. urosomite 1. with well-developed dorsal lamina and posterodorsal tooth (fig. 5d); peduncle of u1 with medial distal corner rounded, with 4 dorsolateral spines of which proximal is longest, with 1 dorsomedial spine and 1 ventromedial spine; outer ramus with 5 small outer spines and 4 small medial spines; inner ramus with 4 small outer spines and 7 small medial spines. urosomite 2. with well-developed dorsal lamina and posterodorsal tooth (fig. 5d); peduncle of u2 with 4 slender dorsolateral distal spines, with 1 dorsomedial spine and 1 ventromedial spine; outer ramus european journal of taxonomy 613: 1–19 (2020) 10 fig. 4. liljeborgia associata sp. nov., female (lemmi) from vostok bay of the sea of japan. a. pereopod 3. b. pereopod 4. c. pereopod 5. d. distal segments of p5. e. pereopod 6. f. distal segments of p6. g. pereopod 7. marin i., a new species of liljeborgia from the sea of japan 11 fig. 5. liljeborgia associata sp. nov., male (a) (lemmi) and female (b–i) (lemmi) from vostok bay of the sea of japan. a. chela of gn2. b. pleonites 1–3 and urosomites 1–3. c. ventroproximal margin of pleonite 3. d. urosomites 1–3 and telson. e. dorsoproximal margins of pleonites 1–3. f. uropod 1. g. uropod 2. h. uropod 3. i. peduncle of uropod 1, outer view. european journal of taxonomy 613: 1–19 (2020) 12 with 5 small outer spines and 4 small medial spines; inner ramus with 4 small outer spines and 4 small medial spines. urosomite 3. with sharp medial projection (fig. 5i), with 1–2 lateral spines, 1 long posterolateral tooth and 3 posterodorsal spines; outer ramus with 3–4 small outer spines and 4 small medial spines; inner ramus with 6 small outer spines and 4 small medial spines (fig. 5h). telson (fig. 2k). with cleft reaching 0.85–0.9 of telson length; distal teeth about ¼ of the length of telson, medial teeth longer than outer teeth, 2 interdental spines long and slender, overreaching outer teeth. coloration. general coloration of body and all appendages translucent white with large light and dark brown spots; some segments of appendages dark red or with dark red spots; eyes brightly white (fig. 1). body size. largest collected female = bl 6.5 mm; largest collected male mostly similar in size to largest female, with bl 6.5 mm. taxonomic remarks the new species is distinctly morphologically similar to l. geminata and l. serratoides. at the same time, the new species can be clearly distinguished by several morphological features not known from the latter species. from l. geminata (see description, presented by barnard (1969)), the new species can be distinguished by the following characters: smaller dorsal crest on urosomites 1 and 2 (fig. 5d); mx1 with wide, shovelshaped article 3 of palp (fig. 2h); shorter and wider merus of p1 and p2 (see fig. 4a–b); slender propodal segments of pp5–7 (see fig. 4c–g); absence of long proximal seta of peduncle of u1 (see fig. 5f); shape of medial teeth of telson (fig. 2k), which are significantly longer and accompanied by 2 long interdental spines. at the same time, l. geminata seems to be more closely related to the new species by its very similar body coloration, the presence of a small dorsal crest on urosomites 1 and 2, short articles of peduncle of a1, relatively stout distal segments of pp3–4 (especially merus), and other morphological features that clearly separate both species from their relative l. serratoides. from l. serratiodes (after tzvetkova 1967), the new species is easily separated by the following characters: presence of a dorsal crest on urosomites 1 and 2; stout articles of peduncle of a1 and a2 (fig. 2d–e); wide distal article of palp of mx1; stout and wider distal segments, especially merus, of pp1–2; slender propodal segment of pp5–7; length and shape of medial teeth of telson, which are significantly longer and accompanied by 2 long interdental spines. in addition, relatively stout and wide segments of pp3–4 are distinguishing features that separate the aforementioned species from tropical relatives, such as l. japonica and l. serrata (e.g., nagata 1965; azman & othman 2013). from l. japonica and l. serrata, known from the seto inland sea in southern japan (see nagata 1965), the new species can be distinguished by longer rostrum, mandibular palp with longer and slender articles, especially distal one; shorter and stouter articles of peduncle a1 and a2; stouter distal segments of pp3–4, especially stout and wide merus of p3; significantly shorted dactyli of pp5–7; slender rami of u3 and longer median teeth of telson. from liljeborgia hwanghaensis kim & kim, 1990, known from the korean coasts of the yellow sea (37°23′ n, 126°35′ e) (see kim & kim 1990), the new species can be clearly separated by the posterodorsal armature of pleonites 1 and 2 (3 teeth in the new species vs 5 teeth in l. hwanghaensis) and shorter median teeth of telson. marin i., a new species of liljeborgia from the sea of japan 13 habitat and ecology specimens of liljeborgia associata sp. nov. were collected inside the burrows of urechis unicinctus (drasche, 1880) (polychaeta: echiura: urechidae), representing the first case of an association between liljeborgiid amphipods and spoon worms (echiura). the spoon worm u. unicinctus lives in large u-shaped burrows constructed in muddy and sandy sediments in the interand subtidal zones in the sea of japan and the yellow sea (abe et al. 2014; pers. obs.). about 20% of the studied spoon worms burrows were inhabited by a single individual of the new species, which may indicate aggressive territorial behavior. distribution the specimens of liljeborgia associata sp. nov. were found in the southern part of the peter the great bay and the northern part of posjeta bay in the sea of japan (russian federation) (see fig. 1), where the present study was accomplished. the distribution of the new species is probably related with the distribution range of its host, u. unicinctus (see abe et al. 2014). key to the species of the family liljeborgiidae from the boreal waters of the nw pacific (with species of liljeborgia from the ne pacific) 1. gn1 significantly larger than gn2; u3 large, paddle-like; postero-dorsal margin of pleonites 2–3 armed with marked teeth; eyes absent. nw pacific: sakhalin, the sea of okhotsk, depth 109– 309 m ................................................................................................... sextonia caecus labay, 2017 – gn1 similar to gn2; u3 smaller than u1 and u2; postero-dorsal margin of pleonites 2–3 unarmed; eyes present (except l. cota) ............................................................................................................ 2 2. carpus of g1 and g2 with strongly produced slender ventral lobe extending along hind margin of propodus .......................................................................................3 (liljeborgia spence bate, 1862) – carpus of g1 and g2 lacking produced ventral lobe. known from the ne pacific only ................... ................................................................ idunella g.o. sars, 1894 (= listriella j.l. barnard, 1959) 3. posterodorsal area of pleonites 1 and 2 produced into 5 teeth of which the median one is the longest. known from the korean coasts of the yellow sea .... liljeborgia hwanghaensis kim & kim, 1990 – posterodorsal area of pleonites 1 and 2 produced into 3 teeth of which the median one is the longest ............................................................................................................................................... 4 4. telson cleft only ¼ to ⅓, lacking terminal spines on lobes of telson; basis of pp5–7 more than twice as long as wide; blind. ne pacific: from the gulf of alaska to the northern baja california, depth 366–2000 m ...................................................................................... liljeborgia cota barnard, 1962 – telson cleft nearly to base, lobes with imbedded terminal spine; basis of pp5–7 about 1–1.5 times as long as wide; with eyes ................................................................................................................ 5 5. epimeral plate 1 concave above postero-ventral tooth. from ne atlantic to the coasts of central california, depth 40–611 m ................................................................liljeborgia pallida bate, 1857 – epimeral plate 1 convex above postero-ventral tooth ...................................................................... 6 6. cusps of lobes of the telson longer medially than laterally; eyes reniform. ne pacific: from bahia de los angeles, gulf of california, depth 46 m ....................... liljeborgia marcinabrio barnard, 1979 – cusps of lobes of the telson subequal longer laterally than medially; eyes oval to subquadrate ..... 7 7. urosomites 1–2 with well-marked dorsal crest, peduncle of a1 with short and stout articles (article 3 about 1.5 times as long as wide; article 2 about as long as wide) ................................................. 8 european journal of taxonomy 613: 1–19 (2020) 14 – urosomites 1–2 without dorsal crest, peduncle of a1 with slender articles (article 3 about 2–2.5 times as long as wide; article 2 about 1.5 times as long as wide). nw pacific: posjeta bay in the sea of japan, intertidal ............................................................. liljeborgia serratoides tzvetkova, 1967 8. medial teeth of telson about 0.15 length of telson, less than 1.5 times longer than outer teeth and accompanied by 1 long interdental spine. ne pacific: from goleta to the northern baja california, depth 3–70 m ............................................................................ liljeborgia geminata barnard, 1969 – medial teeth of telson about 0.25 length of telson, about 2 times longer than outer teeth and accompanied by 2 long interdental spines. nw pacific: the peter the great bay and the northern part of posjeta bay in the sea of japan, intertidal ......................................liljeborgia associata sp. nov. some other species of the genus liljeborgia are known from the region and the sea of japan (e.g., ishimaru 1994; ren 2007). for example, ishimaru (1994) notes liljeborgia aequabilis stebbing, 1888 from the tropical part of the seto inland sea influenced by the kuroshio current. this species is actually recognized in australia, new zealand and the sulu sea as a common associate of hermit crabs (stebbing 1888; after hurley 1954; see also vader 1995) and by most morphological features, especially long and slender segments of pp3–4 and the form of telson (see above), belong to the tropical representatives of the genus, such as l. japonica and l. serrata, or possibly represent an undescribed species. these species are known from warm (tropical) waters and probably are not present in the boreal zone of the northern pacific ocean. discussion the symbiotic communities associated with deep-burrowing invertebrates are very diverse and still include many undescribed species. burrows of spoon worms of the genus urechis as well as almost all large echiuroid worms are inhabited by various symbiotic animals, such as shrimps, crabs, bivalves and even fishes (e.g., fisher & macginitie 1928; anker et al. 2005, 2015; itani et al. 2005; goto & kato 2012; marin 2014; goto et al. 2017). the symbiotic fauna associated with u. unicinctus in the peter the great bay has not yet been fully studied, whereas the discovered symbiotic assemblage includes unidentified acoela turbellarian worms (platyhelminthes: rhabditophora: polycladida), crabs pinnixa rathbuni sakai, 1934 (crustacea: decapoda: pinnotheridae) and sestrostoma balssi (shen, 1932) (crustacea: decapoda: varunidae), copepod goidelia cf. japonica embleton, 1901 (crustacea: copepoda: poecilostomatoida: catiniidae) and goby gymnogobius heptacanthus (hilgendorf, 1879) (pisces: gobiidae) (pers. obs.). liljeborgia associata sp. nov. is probably only associated with the burrows of the spoon worms, since no free-living specimens of the genus liljeborgia or any other liljeborgiid amphipods were sampled on the surface of muddy or sand-bottom substrates in the peter the great bay and posjeta bay in the sea of japan during the present or previous studies (e.g., gurjanova 1951; tzvetkova 1967). moreover, tzvetkova (1967) described a quite common l. serratoides from mussel communities, overgrowing rocks and hard substrates in posjeta bay of the sea of japan, while liljeborgia species were not recorded on muddy and soft-bottom substrates that dominate the bay. this fact indirectly proves that liljeborgia associata sp. nov. rarely emerges from the host’s burrows and does not inhabit the surrounding softbottom substrates as a free-living species. at the same time, representatives of the family liljeborgiidae have been observed in association with other marine invertebrates. often liljeborgia spp. are observed is association with hermit crabs (taylor 1979; vader 1995). liljeborgia aequabilis stebbing, 1888, associated with the hermit crab dardanus arrosor (herbst, 1796) (crustacea: decapoda: diogenidae), has a specific colorful body coloration (e.g., vader & tandberg 2015: fig. 4; taylor 1979: fig. 1), similar to other symbiotic species of the family (barnard 1959; mills 1962: fig. 2), while none of the free-living species are known to have bright pigmentation in life (e.g., d’udekem d’acoz & vader 2009; d’udekem d’acoz 2010; cadien 2015). liljeborgia psaltrica krapp-schickel, 1975 is recorded among associates of the orange coral marin i., a new species of liljeborgia from the sea of japan 15 astroides calycularis (pallas, 1766) (cnidaria: scleractinia: dendrophylliidae) in the mediterranean (terrón-siglera et al. 2016). idunella clymenellae (mills, 1962) was found and described inside the tube arrays of the polychaete worm clymenella torquata (leidy, 1855) (polychaeta: maldanidae) in the intertidal flats of barnstable harbor, massachusetts (mills 1962; sanders et al. 1962). these amphipods inhabit about 20–25% of tubes with live polychaetes (mills 1962) and show a positive response to polychaetes mucus (batcheler & mills 1965). also, idunella albina (j.l. barnard, 1959) was commonly collected from dense beds of echiuroid worms off californian coasts (barnard 1959), however, the direct relationship of this species with spoon worms is not shown. an undescribed species of the genus idunella is reported from the burrows of the mud shrimp upogebia affinis (say, 1818) (crustacea: decapoda: upogebiidae) (fox & bynum 1975), while individuals of the genus idunella (= listriella) are common inhabitants of maldanid and terebellid polychaete tubes (barnard 1969; bousfield 1973) and are reported with holothurians (fox & bynum 1975; vader 1978). it is probable that associations of these animals with marine invertebrates are much more common than is currently known. as mentioned above, according to morphological data, the new species is closely related to l. geminata. unfortunately, minimal genetic data on the family liljeborgiidae are presented in genbank (ncbi) database, and genetic data are not available for the relative congeners of the described species. at the same time, it can be assumed that liljeborgia associata sp. nov. and l. geminata represent a pair of closely related vicariant taxa known as boreal amphi-pacific species. such species had a wide distribution area in the northern part of the pacific ocean during the pliocene–middle pleistocene period, characterized by a higher water temperature and sea level, which allowed the boreal fauna to move from east to west and even through the arctic basin [as great trans-arctic biotic interchange (gtai)] (einarsson et al. 1967; durham & macneil 1967; briggs 2003). during the temperature decreasing in the sea of okhotsk and the bering sea in the late pleistocene (about 2 ma ago), the ranges were subsequently divided into the western and the eastern distribution ranges (zenkevitch 1963; briggs 2003; maggs et al. 2008; marin 2018a). further genetic data will allow a more detailed description of the phylogenetic relationships within the representatives of the family from the northern part of the pacific ocean. acknowledgements the author is very grateful to the staff of the biological scientific station “vostok” of nscmb feb ras (vladivostok, russia) and personally k. dudka and dr a. mayorova for their help during the field sampling. the study was financially supported by the russian foundation for basic research (rfbr) (grant 18-04-01093_а “large burrowing crustaceans (callianassidae and upogebiidae) and their symbionts: diversity and trophic interaction”) and the russian scientific foundation (rsf) (grant 19-74-10104). references abe h., sato-okoshi w., tanaka m., okoshi k., teramoto w., kondoh t., nishitani g. & endo y. 2014. swimming behavior of the spoon worm urechis unicinctus (annelida, echiura). zoology 117 (3): 216–223. https://doi.org/10.1016/j.zool.2013.12.001 anker a., murina g.v., lira c., vera caripe j.a., palmer a.r. & jeng m.s. 2005. macrofauna associated with echiuran burrows: a review with new observations of the innkeeper worm, ochetostoma erythrogrammon leuckart and rüppel, in venezuela. zoological studies 44: 157–190. anker a., komai t. & marin i. 2015. a new echiuran-associated snapping shrimp (crustacea: decapoda: alpheidae) from the indo-west pacific. zootaxa 3914 (4): 441–455. https://doi.org/10.11646/zootaxa.3914.4.4 https://doi.org/10.1016/j.zool.2013.12.001 https://doi.org/10.11646/zootaxa.3914.4.4 european journal of taxonomy 613: 1–19 (2020) 16 azman b.a.r. & othman b.h.r. 2013. shallow water marine gammaridean amphipods of pulau tioman, malaysia, with the description of a new species. zookeys 335: 1–31. https://doi.org/10.3897/zookeys.335.5567 barnard j.l. 1959. liljeborgiid amphipods of southern california coastal bottoms, with a revision of the family. pacific naturalist 1 (3–4): 12–28. barnard, j.l. 1969. gammaridean amphipoda of the rocky intertidal of california: monterey bay to la jolla. bulletin of the united states national museum 258: 1–230. https://doi.org/10.5479/si.03629236.258.1 barnard j.l. & karaman g.s. 1991. the families and genera of marine gammaridean amphipoda (except marine gammaroids). part 1. records of the australian museum 13 (supplement): 1–417. https://doi.org/10.3853/j.0812–7387.13.1991.91 batcheller r. & mills e.l. 1965. behavoral studies on the commensal amphipod crustacean, listriella clymenellae mills. biological bulletin 129 (2): 398. bousfield e.l. 1973. shallow-water gammaridean amphipoda of new england. comstock publishing associates/cornell university press, ithaca, new york. briggs j.c. 2003. marine centers of origin as evolutionary engines. journal of biogeography 30: 1–18. https://doi.org/10.1046/j.1365-2699.2003.00810.x cadien d.b. 2015. amphipoda of the northeast pacific (equator to aleutians, intertidal to abyss): xxviii. ampeliscoidea – an updated review. southern california association of marine invertebrate taxonomists (scamit). available from amphipoda of the northeast pacific. ampeliscoidea [accessed 20 feb. 2020]. durham j.w. & macneil f. 1967. cenozoic migrations of marine invertebrates through the bering strait region. in: hopkins d.m. 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published on: 11 march 2020 topic editor: rudy jocqué desk editor: pepe fernández printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. https://doi.org/10.1002/iroh.19650500115 redescription of psolus tessellatus koehler, 1896 (echinodermata, holothuroidea) with neotype designation claude massin royal belgian institute of natural sciences, department of invertebrate biology, 29 rue vautier, 1000 brussels (belgium), e-mail: claude.massin@naturalsciences.be urn:lsid:zoobank.org:author:896580a2-73e2-4562-afcd-6aa69e5125b4 abstract. in the bycatch of a scientific campaign in the bay of biscay (vital 2002) a third specimen of psolus tessellatus koehler, 1896 was collected very close to the type locality. this specimen is here fully illustrated and designated as the neotype. key words. psolidae, bay of biscay, redescription, first illustration, neotype. massin c. 2013. redescription of psolus tessellatus koehler, 1896 (echinodermata, holothuroidea) with neotype designation. european journal of taxonomy 38: 1-5. http://dx.doi.org/10.5852/ejt.2013.38 introduction in 1895, the university of lyon organized a campaign on board of the “caudan” in the bay of biscay. during that expedition, material was collected between 110 and 1.710 m depth (koehler 1896). four new holothurian species were described and housed in the collections of the university of lyon: holothuria roulei koehler, 1896 (= mesothuria roulei (koehler, 1896)), paroriza pallens koehler, 1896 (= stichopus pallens (koehler, 1896)), benthogone rosea koehler, 1896 and psolus tessellatus koehler, 1896. in 1985, jangoux revisited the types in the collections of lyon and only found the type material of holothuria roulei and benthogone rosea. the type material of paroriza pallens and psolus tessellatus was missing and must be considered as lost. moreover, p. tessellatus was not illustrated by koehler (1896). a second catch of p. tessellatus was mentioned between the azores and madeira at 1.100 m depth by fiege & janssen (1998), who first illustrated the species. in 2002, ifremer nantes organized a campaign (vital) in the bay of biscay for the evaluation of fish populations. in the bycatch of trawled material one specimen of psolus tessellatus was collected. it is the object of the present note. material and methods the specimen of psolus tessellatus was collected at station 14 of the vital campaign (47°36’ n – 8°25’ w, 1399-1442 m, collecting date 23 aug. 2002). it was fixed on a small stone (40 x 44 x 50 mm) and housed in the collection of the rbins under the catalogue number ig31016-hol 1304. european journal of taxonomy 38: 1-5 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2013.38 www.europeanjournaloftaxonomy.eu 2013 · claude massin this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:a5b10105-a73f-4ab1-bf59-2095d95638f3 1 mailto:claude.massin%40naturalsciences.be?subject=claude.massin%40naturalsciences.be http://zoobank.org/urn:lsid:zoobank.org:author:896580a2-73e2-4562-afcd-6aa69e5125b4 http://www.europeanjournaloftaxonomy.eu/index.php/ejt http://creativecommons.org/licenses/by/3.0/ http://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:a5b10105-a73f-4ab1-bf59-2095d95638f3 abbreviations biogas = biologie golf gascogne ifremer = institut français de recherche pour l’exploitation de la mer. rbins = royal belgian institute of natural sciences vital = victor rove talus continental results phylum echinodermata brugière, 1791 order denrochirotida grube, 1840 family psolidae forbes, 1841 genus psolus oken, 1815 psolus tessellatus koehler, 1896 figs 1a-d, 2a-g psolus tessellatus koehler, 1896: 119. psolus tessellatus – mortensen 1927: 414. — fiege & janssen 1998: 341, figs 2-3. psolus tesselatus – costelloe et al. 2001: 348. fig. 1. psolus tessellatus koehler, 1896. a. dorsal view. b. buccal cone. c. anal cone. d. ventral view. european journal of taxonomy 38: 1-5 (2013) 2 redescription small oval specimen, 17.4 mm long, 10.6 mm broad and 2.4 mm thick (fig. 1a). dorsal body wall covered by large scales (from 850 x 550 to 1350 x 1100 µm) (figs 1a-c, 2a). the scale consists of one layer at the edge but rapidly becomes multilayered when going to the centre (fig. 2b). no tube fig. 2. psolus tessellatus koehler, 1896. a. dorsal scale. b. detail of dorsal scale. c. plates of ventral sole. d. rods of large tube feet. e. end plate of large tube feet. f. end plate of small tube feet. g. perforated plates at the base of the large tube feet. a b c d e f g a b-g500 µm 100 µm massin c., psolus tessellatus, redescription, neotype 3 feet dorsally. ten scales between mouth and anus. anus surrounded by irregular small plates (fig. 1c). mouth surrounded by 10 scales: five large triangular scales alternating with 5 small, elongated scales (fig. 1a, b). tentacles fully retracted, not observed. ventral sole very thin, translucent, calcareous ring and digestive tract visible by transparency (fig. 1d). ossicles as perforated plates, 100-160 µm long with 1-8 holes (fig. 2c). ventral sole with 2 rows of tube feet at the edge (fig. 1d). inner row of tube feet large, with perforated rods, 125-280 µm long (fig. 2d) and large end plate in one piece (more or less 300 µm across) (fig. 2f). external row with very small tube feet with small end plates made of one piece (more or less 100 µm across) (fig. 2e). at the base of the large tube feet, perforated plates, 100-170 µm long, similar to those of the ventral sole, sometimes knobbed (fig. 2g). no mid ventral tube feet except 1-2 at the rear (fig. 1d). discussion the specimen at hand belongs without any doubt to the genus psolus: thick dorsal surface covered by scales, thin ventral sole, tube feet restricted to the ventral sole. only 7 psolus species are known from the northeast atlantic ocean, viz p. fabricii (duben & koren, 1846), p. hypsinosus heding, 1942, p. nummularis r. perrier, 1899, p. phantapus (strussenfeld, 1767), p. pourtalesi théel, 1886, p. squamatus (müller, 1776), and p. tessellatus koehler, 1896. p. tessellatus is readily separated from the other mentioned species by the valves surrounding the mouth: 5 large triangular ones alternating with 5 narrow ones. it is also the only psolus species mentioned here with 10 scales present between mouth and anus. for p. hypsinosus and p. fabricii (see heding 1942) there are only 5-7 scales between mouth and anus, whereas for p. pourtalesi, p. squamatus and p. nummularis (see théel 1886; deichmann 1930; madsen & hansen 1994; perrier 1902) there are over 15. p. tessellatus is also the only psolus species present in the bay of biscay. the species seems to have a very restricted distribution and bathymetry in the bay of biscay: 45°47’ n – 6°15’ w and 1.700 m for the holotype, and 47°36’ n – 8°25’w and 1.400 m for the present specimen. in deeper water (1900-4800 m) holothurians are abundant (34 species, sibuet 1977) but p. tessellatus was not mentioned in spite of intensive sampling by ifremer brest (biogas, sibuet 1977; sibuet & segonzac 1985). ifremer nantes has intensively sampled at depths of 0-500 m with a few samples at 1000 m (j. martin, personal communication; sea cruises inventory 2012), but no p. tessellatus were collected. the rarity of p. tessellatus could be an artefact linked to the low intensity of sampling between 1.000 and 1.700 m, a zone obviously favourable to the species (fiege & janssen 1998). since the holotype is lost and koehler (1896) did not give any illustrations of the holotype (general aspect and ossicles) and since the specimen studied here was collected very close to the type locality, it is here designated as the neotype of p. tessellatus (see international code of zoological nomenclature, articles 75, 75.3.4, 75.3.6.). acknowledgements it is a pleasure to thank dr. y. samyn (rbins) for comments on the first draft of this manuscript, mme j. martin (ifremer nantes) for allowing me to study material from the vital campaign and mr. a. van haelen for the photographs on plate 1. european journal of taxonomy 38: 1-5 (2013) 4 references costelloe m., emblow c. & white r. 2001. european register of marine species: a check-list of the marine species in europe and a bibliography of guides to their identification. patrimoines naturels 50, muséum national d’histoire naturelle, paris. deichmann e. 1930. the holothurians of the western part of the atlantic ocean. bulletin of the museum of comparative zoology at harvard college 71 (3): 41-226. fiege d. & janssen r. 1998. ein fall von parasitismus bei psolus tessellatus einer ungewöhnlichen holothurie. natur und museum 128 (10): 341-344 heding s.g. 1942. holothuroidea ii. the danish ingolf-expedition iv (13): 1-39. jangoux m. 1985. catalogue commenté des types d’echinodermes actuels conservés dans les collections lyonnaises. nouvelles archives du muséum d’histoire naturelle de lyon 23: 3-11. koehler r. 1896. echinodermes. résultats scientifiques de la campagne du «caudan» dans le golfe de gascogne 1: 33-127. madsen f.j. & hansen b. 1994. echinodermata holothuroidea. marine invertebrates of scandinavia 9, scandinavian university press, norway. mortensen t.h. 1927. handbook of the echinoderms of the british isles. oxford university press, london. perrier r. 1902. holothuries. in: perrier r. (ed.) expéditions scientifiques du «travailleur» et du «talisman»: 273-554. masson, paris. sea cruises inventory. 2012. http://www.ifremer.fr/sismer/uk/catal/campagne/indexorga2_armat. htql?corg=ifrem sibuet m. 1977. répartition et diversité des echinodermes (holothurides-astérides) en zone profonde dans le golfe de gascogne. deep-sea research 24: 549-563. sibuet m. & segonzac m. 1985. abondance et répartition de l’épifaune mégabenthique. in: laubier l. & moniot c. (eds) peuplements profonds du golfe de gascogne, campagne biogas: 143-156. ifremer, brest. théel h. 1886. reports on the results of dredging in the gulf of mexico (1877-78), in the caribbean sea (1879-80), and along the eastern coast of the united states during the summer of 1880 by the u. s. coast survey steamer “blake”. bulletin of the museum of comparative zoology at harvard college 13 (1): 1-21. manuscript received: 5 december 2012 manuscript accepted: 28 january 2013 published on: 7 february 2013 topic editor: rudy jocqué desk editor: kristiaan hoedemakers printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; national botanic garden of belgium, meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark. massin c., psolus tessellatus, redescription, neotype 5 http://www.ifremer.fr/sismer/uk/catal/campagne/indexorga2_armat.htql?corg=ifrem http://www.ifremer.fr/sismer/uk/catal/campagne/indexorga2_armat.htql?corg=ifrem ejt-530_carvalho_.indd european journal of taxonomy 530: 1–24 issn 2118-9773 https://doi.org/10.5852/ejt.2019.530 www.europeanjournaloftaxonomy.eu 2019 · carvalho de santana e.c. et al. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:2cd56144-79d7-4325-bcc6-353c48e374bd 1 taxonomic revision of the canthidium erichson, 1847 species of the gigas group (coleoptera, scarabaeidae, scarabaeinae) edrielly c. carvalho de santana 1,*, thaynara l. pacheco 2 & fernando z. vaz-de-mello 3 1 programa de pós-graduação em zoologia, instituto de biociências, universidade federal de mato grosso, av. fernando correa da costa, 2367, boa esperança, 78060-900, cuiabá, mt, brazil. 2 programa de pós-graduação em zoologia, instituto de biociências, universidade federal de mato grosso, av. fernando correa da costa, 2367, boa esperança, 78060-900, cuiabá, mt, brazil; currently at programa de pós-graduação em zoologia, museu nacional, universidade federal do rio de janeiro, quinta da boa vista, s/n, são cristóvão, 20940-040, rio de janeiro, rj, brazil. 3 departamento de biologia e zoologia, instituto de biociências, universidade federal de mato grosso, av. fernando correa da costa, 2367, boa esperança, 78060-900, cuiabá, mt, brazil. * corresponding author: edrielly.carol@gmail.com 2 email: thay.lpacheco@gmail.com 3 email: vazdemello@gmail.com 1 urn:lsid:zoobank.org:author:68da9f2a-a005-4f96-9467-03575fadf91f 2 urn:lsid:zoobank.org:author:146cf81b-1394-456c-a768-bffde3e213d0 3 urn:lsid:zoobank.org:author:82872f67-e82a-40c6-b2a3-61f262488f21 abstract. the gigas species group of the subgenus canthidium (neocanthidium) is defi ned and described. this species group is composed of three described species [c. gigas balthasar, 1939, brazilian atlantic forest, including intrusions into cerrado, c. bokermanni (martínez et al., 1964), chaco and western cerrado in brazil, bolivia, paraguay, and argentina, and c. kelleri (martínez et al., 1964), brazilian cerrado and neighbouring open areas] and three new species: canthidium stofeli sp. nov. from the western and southern regions of the brazilian amazon, canthidium feeri sp. nov. from french guiana, and canthidium ayri sp. nov. from the brazilian atlantic forest. we present descriptions and redescriptions, illustrations, an identifi cation key and comments on the distributions of the species of the gigas group. keywords. dung beetles, taxonomy, new species, south america, neotropical region. carvalho de santana e.c., pacheco t.l. & vaz-de-mello f.z. 2019. taxonomic revision of the canthidium erichson, 1847 species of the gigas group. european journal of taxonomy 530: 1–24. https://doi.org/10.5852/ejt.2019.530 european journal of taxonomy 530: 1–24 (2019) 2 introduction the genus canthidium erichson, 1847 is diagnosed by the presence of nine-articulated antennae with a pubescent club; cylindrical labial palpi; ventral clypeal process usually absent, if present in the form of one longitudinal carina; mesoventrite very short; metaventrite usually simply convex; and dilation of meso and metatibia with a curved inner margin and an more or less straight outer margin in ventral view (vaz-de-mello et al. 2011). canthidium is currently subdivided into two subgenera, canthidium s. str. and neocanthidium martínez & halffter, 1986, with some species not presently assigned to any subgenus (see cupello 2018). since its original description (erichson 1847), many species have been described in general revisions (e.g., harold 1867a, 1867b), synopses (boucomont 1928; balthasar 1939; martínez et al. 1964) and regional studies (howden & young 1981; solís & kohlmann 2004; kohlmann & solís 2006). currently, the genus includes 170 valid species-group names (cupello 2018; génier & cupello 2018). due to the large number of species included in many new world dung beetle genera, efforts towards delimiting species groups that facilitate taxonomic revisions are highly desirable. so far, formal species groups have been delimited in several scarabaeinae genera such as dichotomius hope, 1838 (luederwaldt 1929; nunes & vaz-de-mello 2013), onthophagus latreille, 1802 (zunino & halffter 1997), eurysternus dalman, 1824 (génier 2009), ontherus erichson, 1847 (génier 1996), and canthon hoffmannsegg, 1817 (halffter & martínez 1977). in a preliminary attempt to discriminate some species group within canthidium, we propose a readily distinguishable species group around canthidium gigas balthasar, 1939. characters defi ning this group include the large size of its species in relation to the other species of the genus, strongly convex body and uniform black colour. the gigas group includes three previously described species ‒ namely, canthidium (n.) gigas balthasar, 1939, c. (n.) bokermanni (martínez et al., 1964) and c. (n.) kelleri (martínez et al., 1964), ‒ and three new species described below. material and methods institutional abbreviations a total of 129 specimens were examined from the following collections: ambc = ayr de moura bello personal collection, rio de janeiro, brazil bmnh = british museum of natural history, london, united kingdom cemt = setor de entomologia da coleção zoológica da universidade federal de mato grosso, cuiabá, brazil cmnc = canadian museum of nature, ottawa, canada macn = museo argentino de ciencias naturales “bernardino rivadavia”, buenos aires, argentina mzsp = museu de zoologia, universidade de são paulo, são paulo, brazil nmpc = národní muzeum (national museum), prague, czech republic oumnh = oxford university museum of natural history, oxford, united kingdom semc = snow entomological museum, lawrence, kansas, united states of america tlc = trond larsen personal collection, conservation international, washington, d.c., united states of america other abbreviations eecfa = estação experimental de ciências florestais anhembi esalq/usp = escola superior de agricultura luiz de queiroz/universidade de são paulo carvalho de santana e.c. et al., taxonomic revision of species of canthidium erichson, 1847 3 fit = fl ight interception trap pnb = parque nacional de brasilia r.f. ibge = instituto brasileiro de geografi a e estatística unesp = universidade estadual paulista unilavras = centro universitario de lavras pictures were taken using a leica m205c stereo microscope and the images were stacked with the manufacturer’s software. maps were generated with diva-gis (ver. 7.5.0), using the shapefi le of löwenberg-neto (2014) of the biogeographical regionalization of morrone (2014). holotype label data were transcribed verbatim in quotation marks; text lines are separated by a slash mark. the nomenclature used for the geographical distributions is based on the biogeographical regionalization scheme of morrone (2014) . the nomenclature used for the microsculpture is based on harris (1979). specimen lengths were measured from the mid-anterior part of the pronotum to the elytral apex to eliminate potential errors caused by various inclinations of the head; therefore, the length provided here is just a ‘partial length’, not the total length of the specimens. results class hexapoda blainville, 1816 order coleoptera linnaeus, 1758 family scarabaeidae latreille, 1802 subfamily scarabaeinae latreille, 1802 genus canthidium erichson, 1847 subgenus neocanthidium martínez et al., 1964 the gigas species group diagnosis species of the gigas group are readily recognizable from other members of the genus by their large (8 to 15 mm), globose black body, which contrasts with the light-coloured antennae. in addition, the group shows coarse punctation on head and anterior surface of the pronotum; dorsal surface of the eyes posteriorly narrowed; interocular distance approximately seven times wider than the maximum dorsal eye width, and clypeus with two well-defi ned rounded teeth separated by an acute angle. description clypeal margin well defi ned internally, with two more or less rounded teeth medially, separated by an acute angle. clypeogenal suture distinct. hypomeron with coarse longitudinal microsculpture between punctuation and obliquely-oriented setae. prosternum with few long setae, with greater concentration at central region, the apex of each setae reaches mesoventrite. profemora with ventral surface with a longitudinal carina along posterior margin and with a distinct row of punctures along anterior margin, each puncture with a golden seta; surface with microstriae; tibiofemoral joint with a tuft of golden setae. protibiae widening towards apex, external edge with three teeth, external edge serrate between proximal tooth and base, dorsal surface with punctation along the central axis, anteriorly with long setae; venter with longitudinal carina and a single row of punctures parallel to lateral edge. protibial spur tapering towards apex and more or less angulate at apical third in females or wider apically and bent ventrally at median half in males. tarsi with fi ve tarsomeres; claws simple, strongly curved. mesofemora with a carina along posterior edge, ventral surface with some setae anteroapically, centre and apex with minute punctation. mesotibiae with narrow basal half, apical half abruptly dilated, with carina and european journal of taxonomy 530: 1–24 (2019) 4 punctation along anterior margin, near the carina with punctation; external edge with small teeth, internal edge smooth. first mesotarsomere as long as next two mesotarsomeres combined; second to fourth mesotarsomeres decreasing gradually in length and fi fth mesotarsomere longer than the fourth; claws simple and strongly curved; two apical spurs, one as long as the fi rst mesotarsomere and the other as long as the fi rst and second mesotarsomeres combined. elytral striae wide and distinct, with circular punctures slightly wider than the striae. interstriae with small, scattered punctures. pseudepipleuron narrowing gradually towards the apex, where it dilates slightly at the end of the fi fth striae; surface with sparse fi ne punctation. abdomen glabrous, fi nely punctate. pygidium convex, margined along posterior edge, distinctly punctate. sexual dimorphism males and females can be differentiated by the protibial spur, which has a narrow apex in females and is fl attened in males. key to the species of the gigas group 1. head without transverse carina, horn or tubercle along frontoclypeal suture. cerrado-atlantic forest transition in são paulo state (brazil) .............................canthidium (neocanthidium) ayri sp. nov. – head with a transverse carina, conical horn or tubercle along the frontoclypeal suture .................. 2 fig. 1. a. canthidium gigas balthasar, 1939, hypomerum with complete hypomeral carina. b. canthidium kelleri (martínez et al., 1964), hypomerum with incomplete hypomeral carina. c. disc of pronotum completely punctated. d. disc of pronotum with indefi nite punctation. not to scale. carvalho de santana e.c. et al., taxonomic revision of species of canthidium erichson, 1847 5 fig. 2. canthidium gigas balthasar, 1939. a. dorsal view. b. ventral view. c. dorsal view of aedeagus. d. lateral view of aedeagus. scale bars: 1 mm. european journal of taxonomy 530: 1–24 (2019) 6 2. head with conical horn. hypomeron with incomplete hypomeral carina (fig. 1b). brazilian cerrado and neighbouring open area ............... canthidium (neocanthidium) kelleri (martínez et al., 1964) – head with either short or long transverse carina, lacking a horn. hypomeron with complete hypomeral carina (fig. 1a). known from forested/shaded habitats ................................................................... 3 3. head with a long transverse frontoclypeal carina, length greater than one half interocular width. pronotum anteriorly with a shallow excavation near centre (absent in small specimens and smaller in females than in males); bordered anteriorly by a transverse bilobed carina (reduced to a pair of lobes in smaller individuals). amazonia ........................................................................................... 4 – head with a short transverse frontoclypeal carina, length less than one-third of interocular width. pronotum convex anteriorly, without evident lobes or excavations. chaco, cerrado and atlantic forest ................................................................................................................................................ 5 4. frontoclypeal carina with length less than three-fourths of interocular width. pronotum with anterior carina and lobes not evident, positioned just behind anterior border and separated from border by distance less than the border width; anterior region of disc punctate, posterior region with indefi nite punctation (fig. 1d). french guiana ............................canthidium (neocanthidium) feeri sp. nov. fig. 3. canthidium bokermanni (martínez et al., 1964). a. dorsal view. b. lateral view of aedeagus. c. dorsal view of aedeagus. scale bars: 1 mm. carvalho de santana e.c. et al., taxonomic revision of species of canthidium erichson, 1847 7 – frontoclypeal carina with length greater than four-fi fths of interocular width. pronotum with both anterior carina and two lobes sharply evident, separated from border by a distance equal or greater than the latter’s width; disc with surface completely punctate (fig. 1c). amazon rainforest south of the amazon river in brazil and peru ........................ canthidium (neocanthidium) stofeli sp. nov. 5. parameres, in dorsal view, with strong median angulation and concave apical excavation (fig. 2c); in lateral view, with dorsomedial angulation of 120° (fig. 2d). brazilian atlantic forest, including intrusions into cerrado ...................................canthidium (neocanthidium) gigas balthasar, 1939 – parameres, in dorsal view, with feeble median angulation and shallow apical excavation (fig. 3c); in lateral view, with dorsomedial angulation of 160° (fig. 3b). chaco and western cerrado in brazil, bolivia, paraguay and argentina ........................................................................................................ .................................................. canthidium (neocanthidium) bokermanni (martínez et al., 1964) species accounts canthidium (neocanthidium) ayri sp. nov. urn:lsid:zoobank.org:act:0209148b-40fb-43de-8566-3eeb43646298 figs 4, 8 diagnosis canthidium ayri sp. nov. is differentiated from the other species in the group by the unarmed head and pronotum, lacking projections. etymology named for ayr de moura bello, great collector and enthusiast of brazilian beetles, directly or indirectly responsible for the present increase of studies on brazilian coleoptera. type material holotype brazil • ♂; “cerqueira césar/ sp-brasil/ ii-1995/ col. j. carlos”; cemt, ex ambc. paratypes brazil – são paulo • 1 ♀; anhembi; eecfa, esalq/usp; light trap, eucalyptus urophylla; 22°40′ s, 48°10′ w; e.n.l. ferreira leg.; cemt. – paraná • 1 ♀; icaraima, estância felipe; same collection data as for preceding; 23 dec. 2018–5 jan. 2019; d.f. rodrigues leg.; cemt. – minas gerais • 1 ♂; monjolinho; 19°33′10″ s, 49°15′25″ e; nov. 1989; cemt • 3 ♂♂; 6 ♀♀; same locality as for preceding; dec. 1988; cemt • 1 ♂; 1 ♀; same locality as for preceding; dec. 1988; bmnh • 1 ♀; três marias; 18°12′23″ s, 45°13′53″ e; nov. 1989; cemt • 1 ♀; same locality as for preceding; nov. 1991; j.c. zanúncio leg.; cemt. description (holotype) measurements. length: 8 mm. head. surface with microsculpture obscurely variolate. without frontoclypeal carina. pronotum. regularly convex, strongly and completely punctate, without projections along anterior margin. hypomeron. hypomeral carina complete. european journal of taxonomy 530: 1–24 (2019) 8 metaventrite. with simple and shallow punctation, laterobasal region with stronger punctation, lateral region with microsculpture obscurely variolate, almost semicircular. mesepimeron and mesepisternum. with shallow punctation, stronger microsculpture obscurely variolate, almost semicircular. hind leg. metafemora with a carina along posterior edge, anterior edge with setal fringe, surface with fi ne punctation. metatibiae elongated and gradually dilated towards the apex; external margin serrated. aedeagus. in dorsal view, parameres with median angulation nearly absent and with a shallow apical cavity; in lateral view, with dorsal angulation at the basal one-third. variation and sexual dimorphism length ranging from 8.00 to 8.31 mm. males with ventrite vi wider medially, female ventrite vi with the same length throughout. males also have protibial spur fl attened, while in females the protibial spur is narrowed and pointed. geographical distribution the specimens examined were collected in the brazilian state of são paulo in the paraná and chacoan dominion. fig. 4. canthidium ayri sp. nov., holotype (cemt) a. dorsal view. b. lateral view of aedeagus. c. dorsal view of aedeagus. scale bars: 1 mm. carvalho de santana e.c. et al., taxonomic revision of species of canthidium erichson, 1847 9 canthidium (neocanthidium) kelleri (martínez et al., 1964) figs 1b, 5, 8 neocanthidium kelleri martínez et al., 1964: 172–177 (original description). dichotomius (luederwaldtinia) paraguayanus gandini & aguilar, 2009: 139, fi gs 1–8 (original description) (synonymized by nunes & vaz-de-mello 2013: 416). canthidium (canthidium) kelleri – martínez & halffter 1986: 28 (new combination). — vaz-de-mello 2000: 191 (checklist). — nunes & vaz-de-mello 2013: 416 (senior synonymy). canthidium (neocanthidium) kelleri – cupello 2018: 470 (new subgeneric assignment). diagnosis canthidium kelleri differs from other species of the gigas group by the presence of a conical projection on the head which varies in development according to the size of the specimen. type specimens examined neocanthidium kelleri holotype argentina • ♂; handwritten label; “argentina/ missiones/ loreto/ en mongas him./ coll. martínez// enc: 958”; “holotypus”; “neocanthidium/ kelleri sp. n. ♂/ m., h y p./ a. martínez det.1963”; “macn-en 1223”; macn. dichotomius paraguayanus holotype paraguay • ♂; “paraguay: concepcion, zanja moroti. 2.xi.2004”; originally in gandini’s collection, now deposited in cemt; cemt. paratypes paraguay • 1 ♀; cordillera, cerro naranjo; 25°30′ s, 57°04′59″ w; 15 may 2005; j. rivas leg.; cemt. other specimens examined brazil • 1 ♀; distrito federal, brasília; 15°47′39.22″ s, 47°52′55.81″ w; r.f. ibge; dec.1997; i. diniz leg.; cemt • 1 ♀; brasília, fazenda água limpa; 15°57′11.05″ s, 47°58′23.76″ w; 16 sep. 1979; amadur leg.; cemt • 1 ♂ [paratype of n. kelleri]; goiás, rio verde; 17°47′33.79″ s, 50°55′10.77″ e; dr. nick leg.; cemt • 1 ♀; mato grosso, cuiabá, fazenda [“faz.”] mutuca, eucalipto-5; 15°18′51″ s, 55°58′18″ w; 6 mar. 2009; cemt • 1 ♀; same locality as preceding, cerrado-1; 3 feb. 2009; cemt • 1 ♂; chapada dos guimarães, cachoeira da geladeira; fit; 15°25′25″ s, 55°42′58″ w; 6,200 m a.s.l.; 9–30 jan. 2013; d. gimo leg.; cemt • 1 ♀; same locality as for preceding, comunidade joão carro; 15°02′01″ s, 55°43′00″ w; 300 m a.s.l., 3 may 2016; m. serrano leg.; cemt • 1 ♂; mato grosso do sul; selvíria, unesp farm, brachiaria decumbens pasture; black light trap; 20°25′6.82″ s, 51°20′51.87″ w; 21 sep. 2000; c.a.h. flechtmann leg.; cemt • 1 ♀; minas gerais, cordisburgo, fazenda [“faz”] pontinha; 19°5′42.30″ s, 44°17′3.58″ w; jan. 1994; f.z. vaz-de-mello leg.; cemt • 1 ♂; same locality as for preceding; jan. 1999; falqueto and vaz-de-mello leg.; bmnh • 1 ♀; same data as for preceding; cemt • 1 ♀; uberaba; 19°44′50.52″ s, 47°56′20.95″ w; cemt • 1 ♀; lavras; 21°18′50.15″ s, 44°59′36.95″ w; 20 jan. 2008; m.r. rocha and d.h.t. takahashi leg.; cemt • 1 ♀; arinos, rio urucuia; 16°14′34.15″ s, 45°32′44.46″ w; 20 apr. 1994; n. degallier leg.; bmnh • 1 ♂; buritis, fazenda [“faz”] querência; fit; 15°10′23,5″ s, 46°32′43,1″ w; 978 m a.s.l.; 10 jul. 2013–4 feb. 2014; c.m. oliveira leg.; cemt. european journal of taxonomy 530: 1–24 (2019) 10 redescription head. surface densely and distinctly punctated, especially on the clypeus. frontoclypeal suture with conical projection. pronotum. regularly convex, without projections, sometimes excavated anteriorly. anterior edge slightly curved, sides arched. hypomeron. hypomeral carina incomplete. metaventrite. with simple and shallow punctation, laterobasal region with stronger punctation, lateral region with microsculpture obscurely variolate, almost semicircular. mesepimeron and mesepisternum. with shallow punctation, stronger microsculpture obscurely variolate, almost semicircular. hind leg. metafemora with a carina along posterior edge, anterior edge with setal fringe, surface with fi ne punctuation. metatibiae elongated and gradually dilated towards the apex; external margin serrated. aedeagus. parameres, in lateral view, with dorsal angulation at basal two-fi fths. variation and sexual dimorphism length ranging from 8.95 to 11.54 mm. smaller specimens have the frontal horn reduced in size, appearing as a transverse gibbosity. pronotum with the anteromedial depression less visible and shallower in some fig. 5. canthidium kelleri (martínez et al., 1964). a. dorsal view. b. lateral view of aedeagus. c. dorsal view of aedeagus. scale bars: 1 mm. carvalho de santana e.c. et al., taxonomic revision of species of canthidium erichson, 1847 11 specimens. pygidium slightly less convex if compared with c. bokermanni. males present a small tooth in the upper ventral part of their metatibiae. males with ventrite vi wider medially, female ventrite vi with the same length throughout. males also have protibial spur fl attened, while in females the protibial spur is narrowed and pointed. geographical distribution the specimens examined were collected in paraguay (cordillera) and brazil (mato grosso, mato grosso do sul, goiás, distrito federal and minas gerais). it is present in the cerrado and the chaco biomes, which lie in the chacoan dominion. comments the holotype and one paratype originally identifi ed as male of d. paraguayanus were examined by us and were shown to be c. kelleri, thus confi rming the name’s synonymy with c. kelleri as fi rstly proposed by nunes & vaz-de-mello (2013). however, two paratypes originally said to be females by gandini & aguilar (2009) are actually males, and belong to c. bokermanni. see comments below. canthidium (neocanthidium) feeri sp. nov. urn:lsid:zoobank.org:act:eff5685c-71c4-434d-9a9f-9431b46b0b36 figs 1d, 6, 9 diagnosis canthidium feeri sp. nov. differs from other species of the group by the distinct anterior pronotal punctation, apparently lacking punctures on the centre of the disc. etymology named for françois feer, french scarabaeidologist, working at the muséum national d’histoire naturelle (paris, france) who collected half of the type series. type material holotype french guiana • ♂; “guyane française: st/ laurent du maroni, bélvédère/ de saul, 03°37′22″ s/ 53°12′57″ w, 17.i.2011”; “22363”; cemt. paratypes french guiana • 1 ♂; same data as for holotype; cemt • 1 ♂, 1 ♀; cayenne, nouragues nature reserve, primary forest, fl ight interception trap [“fit”]; 4°5′ n, 52°41′ w; 28 mar. 2002; f. feer leg.; cemt • 1 ♂; same location as for preceding; apr. 2003; f. feer leg.; cemt • 1 ♂; saül, mt. galbao; ex. fl ight int. trap; 3°37′18″ n 53°16′42″ n; 740 m a.s.l.; 5–7 jun. 1997; j. ashe and r. brooks leg.; fg1ab97; #154; cmnc. type locality bélvédère de saül, saint-laurent-du-maroni, french guiana, france. description (holotype) measurements. length: 9 mm. head. surface dense and distinctly punctated. frontoclypeal carina rectangular and with length less than three-fourths of interocular distance. european journal of taxonomy 530: 1–24 (2019) 12 pronotum. with anterior carina and lobes not evident, positioned just behind anterior border and separated from border by distance less than the border width; anterior region of disc punctated, posterior region with indefi nite punctation. hypomeron. hypomeral carina complete. metaventrite. with simple and shallow punctation, laterobasal and lateral region with microsculpture fi nely strigulate. mesepimeron and mesepisternum. with shallow punctation and microsculpture fi nely strigulate. hind leg. metafemora with a carina along posterior edge, anterior edge with setal fringe, surface with fi ne punctation. metatibiae elongated and gradually dilated towards the apex; external margin serrated. aedeagus. parameres, in lateral view, dorsally with angulation on basal third. variation and sexual dimorphism total length ranging from 9.00 to 10.67 mm. some males present a small tooth in the upper ventral part of their metatibiae and some have a protuberance on their basal half. males with ventrite vi wider fig. 6. canthidium feeri sp. nov., holotype (cemt) a. dorsal view. b. lateral view of aedeagus. c. dorsal view of aedeagus. scale bars: 1 mm. carvalho de santana e.c. et al., taxonomic revision of species of canthidium erichson, 1847 13 medially, female ventrite vi with the same length throughout. males also have protibial spur fl attened, while in females the protibial spur is narrowed and pointed. geographical distribution all the specimens examined for this work were collected in french guiana, which is located in the boreal brazilian dominion. canthidium (neocanthidium) stofeli sp. nov. urn:lsid:zoobank.org:act:1a3f5b17-8091-4acc-b7de-4311ada3400d figs 7, 9 diagnosis canthidium stofeli sp. nov. differs from the other species in the group by having two anterior medial projections, one on each side of the pronotum. etymology named after roberto stofel, for the fi eld support during collections in cotriguaçu. type material holotype brazil • ♂; “brasil: rondônia: guajará/ mirim: proprietário: fandinho/ 10°44′57.62″ s, 65°19′0.32″ w/ 138m. fl or. amaz. isca: baço bov/ 24.ii.2010 f.coletti”; “22366”; cemt. paratypes brazil • 1 ♂; acre, rio branco, fazenda [“faz”] catuaba; 9°57′19.77″ s, 65°51′55.72″ w; feb. 1997; f. vaz-de-mello leg.; cemt • 1 ♀; rondônia, guajará-mirim, sítio coletti; fl ight interception trap [“fit”]; 10°19′47.11″ s, 4°45′12.5″ w; 170 m a.s.l.; 28 jan. 2010; f. coletti leg.; cemt • 1 ♀; pará, tailândia, empresa agropalma; 9–16 jul. 2016; p. silva leg.; cemt • 1 ♀; belterra, forest; hum/ pig trap; 3°17′47″ s, 54°57′48″ w; 132 m a.s.l.; 24 mar. 2017; f. frança leg.; t2p3; cemt • 1 ♀; same locality as for preceding; t15p20; cemt • 1 ♀; mato grosso, querência, fazenda são luiz, amazon forest; fl ight interception trap [“fit”]; 12°39′3.21″ s, 52°21′9.37″ w; feb. 2009; r. andrade leg.; cemt • 2 ♂♂; cláudia, fazenda [“faz”] continental; fungus; 11°34′54″ s, 55°17′15″ w; 20 feb. 2010; m.f. souza leg.; cemt • 1 ♂; cotriguaçu, matinha; pitfall baited with fungus; 9°50′19″ s, 58°15′3″ w; 11–13 jul. 2009; f.z. vaz-de-mello leg.; bmnh • 1 ♂; same data as for preceding; cemt • 2 ♀♀; cotriguaçu, fazenda [“faz”] são nicolau; fl ight interception trap [“fit”]; 9°49′9″ s, 58°15′47″ w; 12–14 jul. 2009; f.z. vaz-de-mello leg.; cemt • 1 ♀; same data as for preceding; bmnh • 1 ♂; cotriguaçu, fazenda [“faz”] são nicolau; pitfall baited with fungus; 9°49′8″ s, 58°15′40″ w; 17–21 may 2012; f.z. vaz-de-mello leg.; cemt • 1 ♂; cotriguaçu, fazenda [“faz”] são nicolau, primary forest; fl ight interception trap [“fit”]; 9°50′24″ s, 58°15′10″ w; 250 m a.s.l.; 6 may 2009; vaz-demello leg.; cemt • 1 ♂, 1 ♀; fazenda são nicolau (site 6), primary forest; fl ight int. trap; 9°49′16″ s, 58°15′33″ w; 220 m a.s.l.; 1 dec. 2013; f. génier leg.; cmnc 2013-118 • 1 ♀; fazenda são nicolau (site 5), riparian primary forest; fl ight int. trap; 9°51′56″ s 58°13′09″ w; 220 m a.s.l.; 3 dec. 2013; f. génier leg.; cmnc 2013-129 • 1 ♀; fazenda são nicolau (site 6), primary forest; fl ight int. trap; 9°49′16″ s, 58°15′33″ w; 220 m a.s.l.; 3 dec. 2013; f. génier leg.; cmnc 2013-131. peru • 1 ♀; madre de dios, cicra los amigos b.s; 12°34′1″ s, 70°06′0″ w; 250 m a.s.l.; 2005; t. larsen leg.; tlc • 1 ♀; cicra field station, trail 6, research plot; blue pan trap; 12.55207° s, 70.10962° w; 295 m a.s.l.; 7–9 jun. 2011; chaboo team leg.; semc per-11-ptb-009 • 1 ♀; rio tambopata, jorge chavez; fl ight intercept trap; 12°38′59″ s, 69°06′24″ w, 230 m a.s.l.; 28–29 sep. 1999; european journal of taxonomy 530: 1–24 (2019) 14 t. larsen leg.; cmnc • 1 ♀; rio palma real grande, limon camp; fl ight intercept trap; 12°32′20″ s, 68°51′40″ w; 220 m a.s.l.; 9–10 oct. 1999; t. larsen leg.; cmnc • 1 ♀; rio palma real grande, limon camp; fl ight intercept trap; 12°32′20″ s, 68°51′40″ w; 220 m a.s.l.; 13–14 oct. 1999; t. larsen leg.; cmnc. type locality guajará-mirim, rondônia, brazil. description (holotype) measurements. length: 10 mm. head. surface densely and distinctly punctated; with rectangular frontoclypeal carina, length greater than four-fi fths of the interocular distance. pronotum. completely punctate, with dense and distinct punctation, becoming more accentuated towards the centre of the disc, regularly convex with two medial projections anteriorly, one on each side. hypomeron. hypomeral carina slightly curved toward edge. fig. 7. canthidium stofeli sp. nov., holotype (cemt) a. dorsal view. b. lateral view of aedeagus. c. dorsal view of aedeagus. scale bars: 1 mm. carvalho de santana e.c. et al., taxonomic revision of species of canthidium erichson, 1847 15 metaventrite. with simple and shallow punctation, laterobasal region with stronger punctation, lateral region with microsculpture obscurely variolate, almost semicircular. mesepimeron and mesepisternum. with shallow punctation, stronger microsculpture obscurely variolate, almost semicircular. hind leg. metafemora with a carina along posterior edge, anterior edge with setal fringe, surface with fi ne punctation. metatibiae elongated and gradually dilated towards the apex; external margin serrated. aedeagus. in dorsal view, parameres with a strong concave angulation and abruptly curved apex. variation and sexual dimorphism length ranging from 8.87 to 11.11 mm. the projections on the pronotal disc vary according to the size of the specimen, being allometrically developed as the size of the body increases. males with ventrite vi wider medially, female ventrite vi with the same length throughout. males also have protibial spur fl attened, while in females the protibial spur is narrowed and pointed. geographic distribution the specimens examined came from brazil (mato grosso, rondônia, pará) and peru (madre de dios). the species occurs in the southern region of the amazon biome, corresponding to the southeastern amazonian dominion and boreal brazilian dominion. canthidium (neocanthidium) gigas balthasar, 1939 figs 1.a, c, 2, 10 canthidium gigas balthasar, 1939: 127, 128 (original description) canthidium gigas – martínez 1947: 111 (checklist). canthidium (canthidium) gigas – martínez & halffter 1986: 27 (checklist). canthidium (neocanthidium) gigas – cupello 2018: 469 (new subgeneric assignment). neocanthidium gigas – vulcano & pereira 1964: 592 (identifi cation key). type material holotype brazil • ♀; handwritten label; “cayenne”; “coll c felsche/ kauf 20, 1918”; “c. gigas/ sp. m./ dr. v. balthasar det.”; “gigas/ m.”; “typus”; “holotype ♀”; nmpc. other specimens examined brazil • 1 ♀; espírito santo, domingos martins, parque estadual [“p.e.”] pedra azul; 20°23′19.93″ s, 41°0′55.49″ w; jan. 2000; lopes-andrade and f. vaz-de-mello leg.; cemt • 1 ♀; venda nova do imigrante; 20°19′39.02″ s, 41°8′9.36″ w; feb. 1998; falqueto and vaz-de-mello leg.; cemt • 1 ♀; rio de janeiro, itatiaia [“itatiaya”]; 22°29′29″ s, 44°33′33″ w; 12 dec. 1933; j.f. zikán leg.; cemt • 1 ♂; nova friburgo; 22°16′55″ s, 42°31′5″ w; dec. 1995; p. grossi and e. grossi leg.; cemt • 2 ♀♀, 1 ♂; nova friburgo; nov. 2001; e. grossi leg.; ambc • 1 ♀; nova friburgo, macaé de cima; 22°24′8.45″ s, 42°30′48.39″ w; jan. 1999; p. grossi leg.; cemt • 1 ♀; same locality as for preceding; mar. 2000; vazde-mello leg.; cemt • 1 ♀; same locality as for preceding; dec. 2001; e. and p. grossi leg.; cemt • 1 ♂; same locality as for preceding; jan. 2000; grossi leg.; cemt • 1 ♂; same data as for preceding; bmnh • 1 ♂; rio de janeiro, corcovado; dec. 1959; m. alvarenga leg.; mzsp • 1 ♀; são paulo, são luiz do paraitinga, parque estadual da [“pq. est.”] serra do mar, núcleo santa virgínia; nov. 2004; m. uehara leg.; ambc • 1 ♀; minas gerais, ingaí, reserva do boqueirão, unilavras, gallery forest; european journal of taxonomy 530: 1–24 (2019) 16 fl ight interception trap [“fit”]; 21°31′08.05″ s, 44°55′39.81″ e; 13 nov. 2002; silva and frieiro-costa leg.; cemt • 1 ♀, 1 ♂; viçosa, fit; 20°45′17.52″ s, 42°52′42.88″ w; nov. 1999; andrade and vaz-demello leg.; cemt • 1 ♀; same data as for preceding; bmnh • 1 ♀; same locality as for preceding; jan. 1999; vaz-de-mello leg.; cemt • 3 ♀♀, 1 ♂; same locality as for preceding; nov. 1998; vaz-de-mello leg.; cemt • 1 ♂; same locality as for preceding; universidade federal de viçosa; fit; 20°45′51.99″ s, 42°52′5.68″ w; mar. 1998; vaz-de-mello leg.; cemt • 1 ♂; viçosa, mata da biologia; 20°45′ s, 42°51′ w; nov. 2000; f.z. vaz-de-mello leg.; cemt • 2 ♀♀; same locality as for preceding; mata do paraíso; dec. 2000; vaz-de-mello leg.; cemt • 1 ♂; same locality as preceding; dec. 1998; falqueto and vaz-de-mello leg.; cemt • 1 ♂; same locality as preceding; dec. 1999, vaz-de-mello cemt • 1 ♀; same locality as for preceding, fit; 13 feb. 2015; s. aloquio, a. orsetti, c. lopes-andrade and m. bento leg.; cemt • 1 ♂; berizal, fazenda [‘faz”] veredão; 15°39′18″ s, 41°39′16″ w, 13–18 dec. 2012; p. grossi leg.; cemt • 1 ♀; distrito federal, brasília, fal-unb [“faz água limpa”], gallery forest; fit; 15°56′25.1″ s, 47°56′22.2″ w; 4 dec. 2013; t. bernardes leg.; cemt • 1 ♂; same locality as for preceding; pnb, gallery forest, fit; 14°44′49,5″ s, 48°00′32,3″ w; 11 nov. 2015; n.h.d.silva leg.; cemt • 1 ♂; same locality as for preceding; 21 dec. 2015; n.h.d. silva leg.; cemt • 1 ♀; same locality as for preceding; 27 nov. 2015; n.h.d. silva leg.; cemt • 1 ♀; same locality as for preceding; 12 jan. 2016; n.h.d.silva leg.; cemt • 1 ♀; paraná, campina grande do sul, estrada da mandaçaia; fit; 25°17′54″ s, 49°02′30″ w; 7–10 apr. 2014; f.w.t. leivas leg.; cemt. diagnosis canthidium gigas resembles c. bokermanni in the presence of a transverse carina on the head and distinct punctation on the pronotal disc. they differ only in the dorsal curvature of the parameres. in c. gigas, in dorsal view, the parameres have a strong angulation on their apical half and bear a concave medial excavation from the middle to the apex; in lateral view, the parameres have a dorsal angulation fig. 8. geographical distribution of canthidium kelleri (martínez et al., 1964) (circles) and canthidium ayri sp. nov. (triangles). carvalho de santana e.c. et al., taxonomic revision of species of canthidium erichson, 1847 17 of 120°. in c. bokermanni, in dorsal view, the apical angulation is almost absent and there is a shallow medial excavation at the apex; in lateral view, the parameres have a dorsal internal angulation of 160°. type locality brazilian atlantic forest. originally said to be cayenne, french guiana, but this is an error. see geographic distribution below. redescription head. surface with dense and distinct punctation, particularly on clypeus. with a transverse raised projection in the front. pronotum. without anteromedial depression, disc anteriorly convex, sometimes with an anterior sinuosity forming two broad and shallow depressions (both surrounding the medial line). surface with dense and distinct punctation, attenuating towards the centre of disc, where punctation is fi ne and sparse. hypomeron. with microstriae and setae in the basal part. hypomeral carina complete curved throughout its length. metaventrite. with simple and shallow punctation, laterobasal region with stronger punctation, lateral region with microsculpture obscurely variolate, almost semicircular. fig. 9. geographical distribution of canthidium stofeli sp. nov. (circles) and canthidium feeri sp. nov. (triangles).. european journal of taxonomy 530: 1–24 (2019) 18 mesepimeron and mesepisternum. with shallow punctation, stronger microsculpture obscurely variolate, almost semicircular. hind leg. metafemora with a carina along posterior edge, anterior edge with setal fringe, surface with fi ne punctation. metatibiae elongated and gradually dilated towards the apex presenting a sinuosity; external margin serrated. aedeagus. in dorsal view, parameres with a strong angulation in apical half and with a concave excavation from the middle to the apex; in lateral view, with internal medial angulation of 120° dorsally. variations and sexual dimorphism length ranging from 9.40 to 12.65 mm. transverse projection of head sometimes not well defi ned. the males have a tooth in the upper ventral part, more developed in some specimens than in others. males with ventrite vi wider medially, female ventrite vi of the same length throughout. males also have protibial spur fl attened, while in females the protibial spur is narrowed and pointed. geographical distribution the specimens examined were mostly collected in the brazilian states of minas gerais, são paulo, espírito santo and paraná in the atlantic rainforest biome, located in the paraná dominion. the record from distrito federal is surprising for the species. judging from the rest of the known distribution, its presence there may be due to gallery forest connecting it to the atlantic forest on the coast. although the holotype is labelled as coming from french guiana, we believe this represents a mislabelling, since fig. 10. geographical distribution of canthidium gigas balthasar, 1939 (circles) and canthidium bokermanni (martínez et al., 1964) (triangles).. carvalho de santana e.c. et al., taxonomic revision of species of canthidium erichson, 1847 19 no other specimens have been reported from that region. furthermore, the german entomologist carl felsche (who acquired the specimen) was known for buying insects, and this kind of mislabelling has been seen before with his specimens (fzvm, personal observation). canthidium (neocanthidium) bokermanni (martínez et al., 1964) figs 3, 10 neocanthidium bokermanni martínez et al., 1964: 168‒172 (original description). neocanthidium bokermanni – solís & kohlmann 2004: 5 (checklist). — kohlmann & solís 2006: 236 (checklist). — cupello 2018: 453, 465, 467 (checklist). canthidium (canthidium) bokermanni – martínez & halffter 1986: 21 (checklist). canthidium bokermanni – nunes & vaz-de-mello 2013: 416 (mention). canthidium (neocanthidium) bokermanni – cupello 2018: 467 (new subgeneric assignment). dichotomius (luederwaldtinia) paraguayanus gandini & aguilar, 2009 (parts): 139, fi gs 1–8 (2 paratypes). type material examined holotype argentina • ♂; handwritten label; “argentina, tucuman, dto burruyaçú/ km. 5 burruyaçu/ a san ramón/ coll. martínez// 28-iii-946”; “holotypus”; “neocanthidium/ bokermanni/ m., h. y p sp. n. ♂/ a. martínez det.1963”; “macn-en 903”; macn. paratypes bolivia • 1 ♀; santa cruz, santa cruz de la sierra; 17°52′00.00″ s, 63°00′00.00″ e; 10 nov. 1955; r. zischa leg.; cemt. dichotomius paraguayanus paratypes paraguay • 1 ♂; concepción, zanja morotí; 17 nov. 2001; martínez leg.; cemt • 1 ♂; boquerón, loma plata; 24 sep. 1995; drechsel leg.; cemt. other specimens examined argentina • 1 ♀; chaco, potrero; 26º08′35.8″ s, 61º41′49.5″ w; 31 oct. 2016; guerra alonso leg.; cemt. bolivia • 1 ♂; santa cruz, germán busch, puerto suárez, cerro de mutum; 18°58′00″ s, 57°47′53.00″ e; dec. 2012; p. wagner leg.; cemt • 1 ♀; tarija, between caiza and crevaux; blue light; 21°48′53″ s, 63°26′53″ w; 536 m a.s.l.; 5 jan. 2005; mann, hamel and herzog leg.; oumnh. brazil • 1 ♀; mato grosso, cuiabá; 8 mar. 2016; l.p. melo leg.; cemt • 1 ♀; chapada dos parecis, 30 km n uirapuru; fit; 14°17′ s, 59°15′ w; 1–15 dec. 2001; a. foucart leg.; cemt • 1 ♀; diamantino, vale da solidão, riparian forest; 14°21′50″ s, 56°7′23″ w; 18 feb. 2009; d.t.c. oliveira leg.; cemt • 1 ♂; same locality as for preceding; 26 jan. 2009; d.t.c. oliveira leg.; cemt • 1 ♀; same locality as for preceding; 14°21′52″ s, 56°7′23″ w; 23 dec. 2008–26 jan. 2009; d.t.c. oliveira leg.; cemt • 1 ♀; same locality as for preceding; 14°22′23″ s, 56°7′25″ w; 31 jan.–14 feb. 2009; d.t.c. oliveira leg.; cemt • 1 ♀; porto estrela, estação ecológica [“est. eco”] serra das araras; light trap; 15°39′14″ s, 57°13′1″ w; nov. 2003; l. kaminski leg.; cemt • 1 ♀; same locality as for preceding, boca do josé; fit; 15°39′14″ s, 57°13′1″ w; nov. 2003; r. nunes leg.; cemt • 1 ♂; tangará da serra; 27 mar. european journal of taxonomy 530: 1–24 (2019) 20 2016; g.a.a. arenhart leg.; bmnh • 2 ♀♀; tangará da serra, chácara asa branca; manual collection; 14°32′42″ s, 58°40′28″ w; nov. 2011; e. meurer leg.; cemt • 1 ♀; tangará da serra, campus unemat; light trap; 14º38′58″ s, 57º25′24″ w; 439 m a.s.l.; r.j. silva leg.; cemt • 1 ♀; chácara paraíso, semideciduous forest; fit; 14°39′4″ s, 57°24′55″ w; 460 m a.s.l.; 26 jan. 2011; cemt • 1 ♀; same data as for preceding; bmnh • 2 ♂♂; campos de júlio, chapada dos parecis, 30 km n of uirapuru; fit; 14°17′ s, 59°15′ w; dec. 2002; a. foucart leg.; cemt • 1 ♀; cuiabá; 8 mar. 2016; p.l. mello leg.; cemt • 1 ♀; cláudia, fazenda [“faz.”] iracema; 11º37′44″ s, 55º05′54″ w; 20 feb. 2011; m.f. souza leg.; cemt • 2 ♀♀; mato grosso do sul, corguinho, quinta do sol; fit; 19°49′57″ s, 54°49′45″ w; feb. 2011; l.o. bavutti leg.; cemt. diagnosis canthidium bokermanni resembles c. gigas in the presence of a transverse carina on the head and distinct punctation of the pronotal disc; it differs only in the dorsal angles of the parameres. see comments under c. gigas for more details. redescription head. surface with dense and distinctly punctation, especially on clypeus. with a transverse raised projection in the front. pronotum. without anteromedial depression, disc anteriorly convex, sometimes with an anterior sinuosity forming two broad and shallow depressions (both surrounding the medial line). hypomeron. with microstriae and setae in the basal part. hypomeral carina complete, curved throughout its length. metaventrite. with simple and shallow punctation, laterobasal region with stronger punctation, lateral region with microsculpture obscurely variolate, almost semicircular. mesepimeron and mesepisternum. with shallow punctation, stronger microsculpture obscurely variolate, semicircular. hind leg. metafemora with a carina along posterior edge, anterior edge with setal fringe, surface with fi ne punctation. metatibiae elongated and gradually dilated towards the apex; external margin serrated. aedeagus. in dorsal view, parameres lacking angulation apically and with a shallow excavation at apex; in lateral view, dorsally with internal median angulation of 160°. variation and sexual dimorphism total length ranges from 9 to 13 mm. the transverse carina of the head varies from rectangular to oval, regardless of the size and sex of the specimen. the males have a tooth in the upper ventral part, more developed in some specimens than in others. males have ventrite vi wider medially, female ventrite vi is of the same length throughout. males also have protibial spur fl attened, while in females the protibial spur is narrowed and pointed. geographical distribution the specimens examined were collected in argentina (chaco), brazil (mato grosso and mato grosso do sul), bolivia (santa cruz), and paraguay (concepción and boquerón). the species occurs in two different biogeographic dominions: the chacoan dominion and the south brazilian dominion. carvalho de santana e.c. et al., taxonomic revision of species of canthidium erichson, 1847 21 comments two paratypes of dichotomius paraguayanus, which were incorrectly interpreted as females by gandini & aguilar (2009), proved to be males and to belong to c. bokermanni. discussion as previously stated, species groups have so far been delimited, for practical purposes, in several scarabaeinae genera including dichotomius hope, 1838 (luederwaldt 1929; nunes & vaz-de-mello 2013), onthophagus latreille, 1802 (zunino & halffter 1997), eurysternus dalman, 1824 (génier 2009), ontherus erichson, 1847 (génier 1996) and canthon hoffmannsegg, 1817 (halffter & martínez 1977). we note that the members of the gigas group exhibit unique characters that distinguish them from their congeners. in fact, some authors have identifi ed specimens of the gigas group in other genera [such as dichotomius (nunes & vaz-de-mello 2013)], which shows how much they differ from other species of canthidium. in addition to the above described morphological characteristics that separate the species, the group is well established based on its geographical distribution. endemic to south america, the species occurs in different ecoregions. canthidium gigas and c. ayri sp. nov. are endemic to the atlantic forest, their occurrence being restricted respectively to the north and south of this biome. canthidium bokermanni and c. kelleri are found in cerrado vegetation; c. bokermanni in the west region of the biome, and c. kelleri in the east. finally, c. feeri sp. nov. is restricted to french guiana, north of the amazon biome, and c. stofeli sp. nov. occurs in the southern region of this biome. the group also has unique characteristics regarding feeding. most species of scarabaeinae are coprophagous, but there may be variations, such as c. kelleri and c. bokermanni feeding on the mycelia of fungi (halffter & matthews 1966). most recently, falqueto et al. (2005) reported c. gigas from fungus-baited pitfall traps. the two paratypes of c. ayri sp. nov. were collected with light traps and the other species often appear in fl ight interception traps, in addition to traps baited with fungi. more studies and fi eld observations are needed to confi rm if fungi are indeed a fundamental part of these species’ diet. conclusions this work dealt with the taxonomic revision of the canthidium species of the gigas group. we described three new species for the group, but we fi rmly believe that in order to broaden the distribution data more studies are required, including fi eld collecting and museum studies. acknowledgments we thank the curators ayr m. bello (ambc), sonia casari and carlos campaner (mzsp), darren mann (oumnh), jirí hajek (nmpc) and trond larsen (tlc) for loaning part of the material examined, and jirí hajek and martin fikacek, svata (in memoriam) and petr pokorný for hosting the third author during his visit to nmpc and prague. we thank olivier boilly and paolo gandini for making available and donating the type specimens of dichotomius paraguayanus. mario cupello is thanked for providing photographs of the holotypes of canthidium bokermanni and c. kelleri housed at macn. we are grateful also to mario cupello, renato machado, paul skelley, françois génier and anonymous reviewers for critically reading the manuscript and suggesting several improvements, we are grateful also to maxwell v.l. barclay for reviewing the english, and the funding agencies that made it possible to carry out this work: conselho nacional de desenvolvimento científi co e tecnológico (cnpq): 302997/20130; 484035/2013-4; 405697/2013-9; 440604/2015-0; 306745/2016-0; 142383/2017-1; 248299/2012-3; fundação de amparo à pesquisa do estado de mato grosso (fapemat): 568005/2014; 146138/2015; 0147956/2017; isyeb, umr 7205 cnr/mnhn; inct-cenbam / cnpq; comcerrado network / mma / cnpq; ufmt pibic 2014-15. european journal of taxonomy 530: 1–24 (2019) 22 references balthasar v. 1939. neue canthidium-arten. 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(coleoptera: scarabaeiodea) do brasil. in: martín-piera f., morrone j.j. & melic a. (eds) hacya un proyecto cyted para el inventario y estimación de la diversidad entomológica en iberoamérica: pribes-2000. m3m: monografi as tercer milenio. vol. 1: 183–195. sociedade entomológica aragonesa, zaragoza. vaz-de-mello f.z., edmonds w.d., ocampo f.c. & schoolmeesters p. 2011. a multilingual key to the genera and subgenera of the subfamily scarabaeinae of the new world (coleoptera: scarabaeidae). zootaxa 2854: 1–73. vulcano m.a. & pereira f.s. 1964. catalogue of the canthonini (col. scarab.) inhabiting the western hemisphere. entomologische arbeiten aus dem museum g. frey 15: 570–685. zunino m. & halffter g. 1997. sobre onthophagus latreille, 1802 americanos (coleoptera: scarabaeidae: scarabaeinae). elytron 11: 157–178. manuscript received: 24 december 2018 manuscript accepted: 16 april 2019 published on: 6 june 2019 topic editor: gavin broad desk editor: eva-maria levermann european journal of taxonomy 530: 1–24 (2019) 24 printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. description of the first palaearctic species of tineobius ashmead, 1896 with dna data, a checklist of world species, and nomenclatural changes in eupelmidae (hymenoptera, chalcidoidea) lucian fusu 1,* & antoni ribes 2 1 alexandru ioan cuza university, faculty of biology, bd. carol i no. 11, 700506, iasi, romania. 2 deceased 2 dec. 2014. former address: c/lleida 36, 25170 torres de segre, lleida, spain. * corresponding author: lucfusu@hotmail.com 1 urn:lsid:zoobank.org:author:894051b1-3121-4f10-90f5-a6e2284a7123 2 urn:lsid:zoobank.org:author:98b51d2c-3663-4382-bb10-c2dd8d1e80cc abstract. tineobius (tineobius) tamaricis ribes & fusu sp. nov. is newly described from parapodia sinaica (frauenfeld, 1859) (lepidoptera, gelechiidae) galls from catalonia in spain. this is the first record of the so far palaeotropical genus tineobius ashmead, 1896 in the palaearctic region. basic biological data and a dna barcode are provided for the new species. parapodia sinaica (the host of t. tamaricis sp. nov.) is reported for the first time to form galls on tamarix canariensis (willd). a checklist of described world tineobius species is provided, with nine species formally transferred to tineobius from anastatoidea gahan, 1927 and thirteen species newly assigned to t. (tineobius). metapelma seyrigi (risbec, 1952) is transferred to tineobius and the replacement name tineobius (tineobius) madagascariensis nom. nov. is proposed, as the name is preoccupied by tineobius (tineobius) seyrigi (ferrière, 1938) comb. nov.; tineobius (tineobius) albopalpalis (brues, 1907) comb. nov. is transferred from charitopus förster, 1856 (a genus in encyrtidae). one species is transferred from anastatoidea to eupelmus dalman, 1820 as eupelmus (episolindelia) ambatomangae (risbec, 1958) comb. nov. keywords. chalcidoidea, dna barcoding, tamarix, nomenclature, parapodia sinaica. fusu l. & ribes a. 2017. description of the first palaearctic species of tineobius ashmead, 1896 with dna data, a checklist of world species, and nomenclatural changes in eupelmidae (hymenoptera, chalcidoidea). european journal of taxonomy 263: 1–19. http://dx.doi.org/10.5852/ejt.2017.263 introduction tineobius ashmead, 1896 (chalcidoidea latreille, 1817; eupelmidae walker, 1833; eupelminae walker, 1833) is a palaeotropical genus known so far from the afrotropical, oriental, and australasian ecozones (gibson 1995), and never reported from the palaearctic region. tineobius is currently divided into three subgenera (gibson 1995), and known members are ectoparasitoids of lepidoptera and of their primary parasitoids: ichneumonidae (gahan 1927), braconidae (ferrière 1936, 1938) and tachinidae (bouček 1988). european journal of taxonomy 263: 1–19 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2017.263 www.europeanjournaloftaxonomy.eu 2017 · fusu l. & ribes a. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:8e4a56b0-ea35-4e3a-8d78-a11c934eaac6 1 mailto:lucfusu%40hotmail.com?subject= http://zoobank.org/urn:lsid:zoobank.org:author:894051b1-3121-4f10-90f5-a6e2284a7123 http://zoobank.org/urn:lsid:zoobank.org:author:98b51d2c-3663-4382-bb10-c2dd8d1e80cc http://dx.doi.org/10.5852/ejt.2017.263 http://dx.doi.org/10.5852/ejt.2017.263 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:8e4a56b0-ea35-4e3a-8d78-a11c934eaac6 most species of tineobius were revised by ferrière (1938) as anastatoidea gahan, 1927. after ferrière’s revision further species were described by risbec in anastatoidea, who also published a new key to the afrotropical species (risbec 1952, 1958), but out of four species described by this author two belong to eupelmus (episolindelia) girault, 1914 (fusu et al. 2015 and present paper). yoshimoto & ishii (1965) described two further species in anastatus motschulsky, 1859 (gibson et al. 2012) and bouček (1988) one species in duanellus bouček, 1988. the latter author synonymized anastatoidea with tineobius, transferring to tineobius all the australian species described by a.a. girault and some of the species described by c. ferrière (bouček 1988), but many of ferrière’s species were never formally transferred to tineobius. according to bouček (1988) and gibson (1995), more undescribed species are known, but all are palaeotropical. we report here on the unexpected discovery of the first species of tineobius in the palaearctic region. material and methods specimens of a species of tineobius (tineobius) were found in spain near lleida during a survey of the chalcidoidea of the region by ar. the first specimens emerged in 2008 from parapodia sinaica (frauenfeld, 1859) (lepidoptera, gelechiidae) galls on tamarix canariensis, in a small number compared to other parasitoids. further galls were collected from 2010 to 2013, stored indoor in polythene bags, controlled by ar for condensation and fungal growth, and checked periodically for the emerged chalcid wasps. other gall samples were dissected for further biological observations. emerged specimens were either killed with ethyl acetate or placed directly in ethanol. those in ethanol were dried using hmds (heraty & hawks 1998) before mounting on cards or points. the majority of the specimens were measured by ar with a stereomicroscope with a maximum magnification of 90 ×, fitted with a 10 mm ocular grid having 100 divisions. for paratype no. 10077 measurements were made by lf with an olympus szx9 stereomicroscope fitted with a 10 mm ocular grid having 100 divisions; to reduce glare, a double piece of tracing paper was inserted between the light source and the specimen. the habitus photograph of the holotype and acropleuron of paratype no. 13940 were taken by ar with a compact digital camera placed over the trinocular stereo microscope and the specimen illuminated with a 144-led ring as a light source. the serial images were combined using combinez5 software (alan hadley, http://www.bioinfo.org.uk/html/b148461.htm). all other photographs were taken by lf with a leica dfc 500 digital camera attached to a leica m205 a motorized stereomicroscope and the specimen illuminated within a cylinder made of white expanded polystyrene with an olympus kl1500 lcd and a kruss 150-watt light source. the serial images were combined with zerene stacker (zerene systems llc, http://www.zerenesystems.com/) and digitally retouched using adobe photoshop. specimens of the type series were labelled by lf using the data provided earlier by ar. the original labels contained only a catalogue number referring to an electronic database by ar that is currently inaccessible. these catalogue numbers are used throughout the text to uniquely designate a specimen and are provided in the material examined section after the name of the collector. the catalogue numbers were confidently correlated with collecting data in all cases except for paratypes 13940 and 22575 for which data could have been reversed. data from different labels of the holotype are separated in text by a slash. within the checklist of described world species of tineobius provided at the end of this paper, the species are listed alphabetically within the three subgenera. most morphological terms follow gibson (1995, 1997), wing folds terminology gibson (2004), and sculpture categories as defined in al khatib et al. (2014) and gibson & fusu (2016) except mesoscutellum is used instead of scutellum and metascutellum instead of dorsellum in accordance with the recommendations of the hymenoptera anatomy ontology portal (yoder et al. 2010). european journal of taxonomy 263: 1–19 (2017) 2 http://www.bioinfo.org.uk/html/b148461.htm http://www.zerenesystems.com/ abbreviations used in the descriptions fl1 = first flagellomere (anellus) fl2–fl8 = flagellomeres 2 –8 (funiculars 1–7) mpod = maximum posterior ocellus diameter mt1–mt8 = first to eighth metasomal tergites ool = occellocular line (distance between a posterior ocellus and inner orbit) pol = posterior ocellar line (distance between the posterior ocelli) for the molecular analyses dna was initially extracted by lf from non-type specimens 22591 and 22592 using a non-destructive protocol based on the dneasy® blood and tissue kit (qiagen, hilden, germany) as described in polaszek et al. (2014). because no pcr product was obtained, dna was subsequently extracted from the left mid and hind legs of the holotype using a chelex resin based protocol that was successfully used on other chalcid wasp taxa (gebiola et al. 2009; 2015). after dna extraction, the legs were dried with hmds and glued back to the specimen. we used an extraction protocol that avoids the boiling of the sample, as suggested by casquet et al. (2012). this result in double stranded dna that is much more stable during manipulations and for long term storage compared to the single stranded dna obtained with an extraction protocol that uses a final heating step at 95° or 100°c to inactivate the protease. briefly, the modified protocol is as follows. legs including coxae are placed in 25 µl of 5% chelex ® 100 (bio-rad laboratories) in a 1.5 ml microtube and proteinase k is added last (1.65 µl of 600–1000 u/ml, 14–22 mg/ml). if whole specimens are used, volumes are doubled. the tubes are incubated at 56°c for 5–6 hours. after the digestion, the microtubes are centrifuged briefly and the supernatant is directly used as template in a pcr. the standard barcode region (hebert et al. 2003) was amplified by pcr using the primer pair lco1490 / hco2198 (folmer et al. 1994). a standard 25 μl pcr was performed, containing 2.5 μl of 10 × pcr buffer, 1 μl of 50 mm mgcl2, 0.5 μl dntps solution (10 mm each), 1.25 μl of each primer (10 μm), 1.25 μl taq polymerase (1u/μl, red taq dna polymerase, rovalab), 4 μl dna extract, and pcr grade water to final volume. pcr conditions were: 94°c for 2 min, followed by 40 repeated cycles of 94°c for 30 s, 42°c for 60 s and 72°c for 35 s, a final extension at 72°c for 5 min, and incubation at 10°c. the pcr products were visualized on a 1% agarose gel. both dna strands were sequenced at macrogen europe (amsterdam) using the same primers used for the pcr. sequence verification was conducted by comparing forward and reverse chromatograms using pregap4 v1.5 and gap v4.10 in the staden package (bonfield et al. 1995). abbreviations used for repositories aicf = al. i. cuza university, iaşi, romania, lucian fusu collection anic = australian national insect collection, canberra, australia arpc = antoni ribes personal collection, lleida, spain [deposited in bmnh] bmnh = the natural history museum, london, u.k. bpbm = bernice pauahi bishop museum, honolulu, hawaii mncn = museo nacional de ciencias naturales, madrid, spain mnhn = muséum national d’histoire naturelle, paris, france mpmw = milwaukee public museum, milwaukee, wisconsin, u.s.a. qmba = queensland museum, brisbane, australia rrac = richard r. askew collection, beeston, cheshire, u.k. usnm = united states museum of natural history, washington d.c., u.s.a. fusu l. & ribes a., a new species of tineobius from spain 3 results class hexapoda blainville, 1816 order hymenoptera linnaeus, 1758 suborder apocrita latreille, 1810 superfamily chalcidoidea latreille, 1817 family eupelmidae walker, 1833 subfamily eupelminae walker, 1833 genus tineobius ashmead, 1896 diagnosis females can be recognized by the following combination of characters: vast majority of species with a conspicuously lengthened maxillary palpus with apical palpal segment elongate, distinctly curved or with inner margin angulate basally, prepectus with light-coloured and large frontal surface, propodeum with transverse plical depression and either with membranous white spot or line within depression (nominal subgenus and duanellus) or with whitish membranous bottom (in t. (progenitobius) gibson, 1995), mesotibia without oblique apical groove, gaster with deeply emarginate syntergum and very long and filamentous ovipositor sheaths (gibson 1995). excluding some t. (progenitobius), metatibia conspicuously compressed with laminate dorsal margin and white along lamina or at least dorsobasally, with the exception of some t. (duanellus). tineobius (tineobius) tamaricis ribes & fusu sp. nov. urn:lsid:zoobank.org:act:dfecb6fa-96d1-4eda-b742-85d7da57e85c fig. 1a–h etymology named from its associated plant, tamarix canariensis (willd.). a noun in the genitive case. type material examined holotype spain: ♀, torres de segre, pantà de camelis (lleida), 31t bg80, 160 m, leg. a. ribes, 22576 / parapodia sinaica galls on tamarix canariensis, col. 30.i.2013, em. 2013, a. ribes, 22576 / holotypus ♀ tineobius (tineobius) tamaricis ribes & fusu det. fusu l. 2015. original label: tineobius, tamarix, torres, ♀, 22576. dna voucher label: dna extracted, 15.vii.2015 fusu, tt.sp 03 (bmnh; bm hym type 5.4849, barcode number nhmuk010264039). condition: entire, uncontorted, glued by right side and wings to a card point. paratypes spain: 4 ♀♀, same data as holotype, except galls collected 17 oct. 2012, emerged 2013 (no. 22575?) (1 ♀ bmnh); galls collected 13 feb. 2010, emerged 2010 (no. 13940?) (1 ♀ mncn); 31t bf99, 140 m, galls collected on 20 oct. 2007, emerged in 2008 (no. 10077) (1♀ aicf); galls collected 13 feb. 2010, emerged 2010 (no. 22662) (1 ♀ with gaster missing, rrac). additional material examined spain: 2 ♀♀, same data as holotype, except galls collected 8 jan. 2011, emerged 22 aug. 2011 (no. 22591), and dna voucher label: dna extracted, 12.vi.2014 fusu, tt.sp 01 (1 ♀, damaged, head and metasoma chowed by psocids, arpc); galls collected 25 jan. 2012, inside galls 2013 (no. 22592), and dna voucher label: dna extracted, 15.vi.2014 fusu, tt.sp 02 (1 ♀, damaged and slightly mouldy, not emerged, antennae missing, with the head capsule of the host in a gelatine capsule on the same pin, arpc). european journal of taxonomy 263: 1–19 (2017) 4 http://zoobank.org/urn:lsid:zoobank.org:act:dfecb6fa-96d1-4eda-b742-85d7da57e85c description female (fig. 1a, c) length. 2.2–3.5 mm (about 3.2–6.3 mm including ovipositor); holotype 3.3 mm (about 6 mm including ovipositor). colour. head dark, with faint multicoloured metallic reflections on frontovertex and occiput changing colour and intensity depending on viewing angle and lighting conditions (cf. heads in figs 1a and 1c); more conspicuously metallic blue to greenish-blue variably extensively along eye orbit, in front of anterior ocellus and near posterior ocelli; interantennal region, scrobal depression and parascrobal region with green, bronze, coppery and violet reflections; lower face, gena and temple dark blue with some violet, greenish and bronze reflections. maxillary and labial palps dark brown. scape pale reddish to orangish brown, pedicel and flagellum dark brown. mesosoma including pronotum (fig. 1b, g) mainly similar in colour to frontovertex except as follows. mesoscutum dark blue to violet with green to coppery lustre more conspicuously on anteromedial convex lobe. axillae and about anterior half of mesoscutellum reddish to orangish brown with at most with faint metallic lustre, mesoscutellum progressively darker posteriorly with frenal area dark green with multicoloured metallic lustre. acropleuron (fig. 1a–c) reddish to orangish brown in posterior half to two-thirds and dark, mostly metallic bluish-green, anteriorly. tegula and prepectus dark reddish-brown, the frontal surface of prepectus dirty yellowish only in inferior third. metanotum and callar region of propodeum dark with blue reflections. legs reddish to orangish brown. front leg almost uniformly reddish-brown beyond coxa, except femur darkened on posterior surface. middle leg similar in colour to front leg, except femur and tibia somewhat darkened dorsally, anterodorsal angle of femur with a whitish spot, and mesotibial spur brown; mesotarsus with basal two tarsomeres dirty-white except yellowish ventrally and apically and subsequent tarsomeres gradually darker with apical tarsomere distinctly brown. mesotibial pegs reddish-brown, similar in colour to apex of tibia and mesotarsal pegs pale basally and brownish apically. hind leg with femur reddish to orangish brown dorsally and metallic brown ventrally to almost uniformly reddish-brown; tibia reddish-brown, with laminated dorsal margin yellowish-white; tarsus reddish-brown. gaster dark, with faint metallic lustre except basal tergite (mt2) dorsally with obvious bluish-green to bronze reflections subbasally. ovipositor sheaths dark brown, ventrally with a narrow, yellowish to orangish band along length, broadening subapically into an indistinctly delimited and dorsally incomplete pale ring. head. in dorsal view 1.8–1.95 × as wide as long, 1.1–1.15 × as wide as mesoscutum, and temples 0.22–0.35 × eye length; with interocular distance 0.31–0.34 × head width. ocelli in a right-angled triangle with pol 4.1–4.5 × ool and ool 0.4–0.43 × mpod. eyes 1.4–1.5 × as long as broad, with short sparse pilosity, inner eye orbits converging upwards. head in frontal view 1.15–1.2 × as broad as high, with dorsal margin of torulus in line with or slightly above lower orbit. malar space 0.5–0.55 × eye height, malar sulcus slightly out curved, mouth opening 1.4–1.6 × as broad as malar space. scrobal depression wide, about as broad as high, extending up to half eye height, carinately margined laterally at least in basal two thirds. vertex transversely reticulate-imbricate, frons conspicuously reticulate, with mesh size about as large as an ommatidium; scrobal depression including scrobes distinctly reticulate to transversely reticulate-rugulose. lower face, interantennal region and parascrobal regions with white, slightly lanceolate setae compared to thinner, more hair-like, dark setae on frontovertex. setae along outer orbit comparatively dense, white, slightly lanceolate and directed toward orbit, those along inner orbit and upper occiput long and semierect (fig. 1e, g). maxillary palpus enlarged, the last segment slightly curved and nearly as long as ventral length of mandible. antenna (fig. 1d) with scape 6–6.5 × as long as broad and 0.9–0.95 × eye length, not reaching to anterior ocellus. pedicel plus flagellum 1.43–1.5 × head width; pedicel in dorsal view 2.2–2.7 × as long as broad, 1.9–2.1 × as long as fl1 (anellus), and 0.45–0.55 × as long as combined length of fl1 and fl2. flagellum basally narrower than pedicel, the funiculars shorter and slightly wider distally; fl1 elongate, 1.4–1.7 × as long as broad; fl2 the fusu l. & ribes a., a new species of tineobius from spain 5 european journal of taxonomy 263: 1–19 (2017) 6 fig. 1. female of tineobius (tineobius) tamaricis ribes & fusu sp. nov. a. habitus of holotype before card mounting. b. mesosoma (paratype 13940). c–g. paratype 10077. c. habitus. d. antenna. e. head. f. fore wing. g. head and mesosoma, dorsal view. h. metanotum and propodeum (paratype 13940). scale bars: a, c = 1 mm; b, d–h = 0.2 mm. longest segment, 4.1–4.6 × as long as broad and 1.15–1.25 × as long as fl3; fl3 to fl8 2.85–3.5, 2.35–2.7, 1.75–2.15, 1.45–1.9, 1.3–1.5, and 1.15–1.25 × as long as broad respectively; clava 2.0–2.35 × as long as broad. mesosoma (fig. 1g). elongate, in dorsal view 2.0–2.05 × as long as broad. pronotum 0.30–0.35 × as long as mesoscutum, divided medially. anteromedial mesoscutal lobe 1–1.1 × as long as broad, lateral lobes markedly elevated; mesoscutum imbricate-reticulate with sculpture more superficial than on head, transversely imbricate-reticulate anteriorly, reticulate on depressed area behind anteromedial lobe, and with effaced sculpture posteriorly in front of mesoscutellum; with comparatively inconspicuous setae anteriorly and on lateral lobes, but with long, white conspicuous setae on bottom of median depressed area. mesoscutellar-axillar complex 0.9–1.05 × as long as broad, 0.53–0.6 × as long as mesoscutum, with isodiametric reticulate sculpture, coarser than on mesoscutum; frenal area differentiated by smoother, alutaceous-imbricate sculpture and sharply declivitous; mesoscutellum with sparse, erect black setae anterodorsally, and short suberect pale setae near frenal area. propodeum with deep plical depression with whitish membranous bottom and a differentiated whitish spot (fig. 1h) (exact structure often not visible, being overlaid by metascutellum, fig. 1g); callar region convex, outer half covered with dense, long white setae and inner half abruptly sloping from spiracle to plical depression, with a smaller tuft of long white setae; spiracle oval, situated on outer side of callus (fig. 1h). acropleuron (fig. 1b) reticulate in anterior third and ventrally, and in posterior third reticulate to striate with strongly elongated cells. mesotibia with 3–6 apical pegs in 1 or 2 rows; basitarsus ventrally with 10–15 pegs in row on each side, second tarsomere with 4–5, third with 3–5, and fourth with 3–4 pegs in row on each side. metatibia and metabasitarsus compressed, metatibia 4.9–5.7 × as long as broad, with laminate dorsal margin. fore wing (fig. 1f) 2.5–2.6 × as long as broad, broadly infuscate medially and very narrowly at extreme base, hyaline below submarginal vein and parastigma and apically beyond venation; with relatively long and thick, dark, hair-like setae in infuscate regions, and with very short and fine but dark setae in apical hyaline portion; basal hyaline region with a band of pale setae behind basal half of parastigma and apex of submarginal vein. basal cell variably extensively setose with pale setae, sometimes mostly bare but at least bare mesally and with setae sparser than on disk, open posteriorly because mediocubital fold, cubital and vanal areas bare. costal cell 8.5–9.5 × as long as broad, 1.3–1.35 × as long as marginal vein, dorsally near leading margin with row of dark setae in about basal half but bare in front of parastigma, ventrally with two rows of pale setae along length except more extensively setose basally. marginal vein 2.9–3.3 × as long as stigmal vein. stigmal vein nearly straight, stigma broadened with uncus slightly shorter than stigmal vein breadth. postmarginal vein 2.4–2.55 × as long as stigmal vein, and 0.75–0.85 × as long as marginal vein. gaster (excluding ovipositor sheaths) 0.85–0.9 × as long as mesosoma and 1.9–2.15 × as long as broad, with white setae mostly laterally, long and suberect on first tergite and shorter and more adpressed on rest. mt2 deeply incised, surface superficially coriaceous-imbricate to reticulate-imbricate; following tergites with similar but stronger sculpture, posterior margin of mt3 to mt5 sinuately emarginate, that of mt6 straight and mt7 slightly angulate posteriorly. mt7 of about same length as mt6 (concealed in various degrees under mt6 in all available specimens). ovipositor sheaths about 1.9–2 × length of gaster, 0.8–0.9 × body length, and 2.3–2.4 × length of metatibia, the pale portion about 1.5 × as long as the apical dark portion. male unknown. distribution spain. fusu l. & ribes a., a new species of tineobius from spain 7 remarks tineobius tamaricis sp. nov. is placed within tineobius s. str. because the female possess the following combination of characters: head in lateral view comparatively flat, with a band of differentiated reflective setae along outer orbit; fl2 not anelliform, much longer than fl1 or pedicel; metascutellum only slightly protuberant over propodeal plical depression; plical depression with a whitish membranous bottom and a differentiated anteromedian whitish spot; metatibia compressed with laminated dorsal margin yellowish-white; mt6 with posterior margin straight and mt7 transverse and of about the same length as mt6. females of t. (duanellus) share the same presumably derived state of the head, propodeum, metatibia and metasoma, but antennae are inserted near the centre of the face with fl2 anelliform and similar to fl1 (bouček 1988: figs 1011, 1013) and outer orbit lacks a band of spatulate or lanceolate setae (gibson 1995). the three known species of t. (progenitobius) of which one is undescribed (gibson 1995; fusu et al. 2015) all have a subglobose head (fusu et al. 2015: fig. 3c), mt6 with a broadly and deeply emarginate posterior margin similar to other tergites, and mt7 longer than mt6 and subtriangular. also, the two described species, both from madagascar, do not have a carinately compressed metatibia though basal part of dorsal margin is whitish (gibson 1995; fusu et al. 2015). however, one undescribed species from south africa is atypical for the subgenus in having a metatibia similar to that found in most species of tineobius s. str. (gibson 1995). the distribution of the character states within the genus and the potential paraphyly of t. (progenitobius) relative to tineobius s. str. + t. (duanellus) were discussed in detail by gibson (1995). using ferrière (1938) and risbec (1952) keys, the holotype of t. tamaricis sp. nov. runs near t. indicus (ferrière, 1938) and t. philippinensis (ferrière, 1938) comb. nov. because of the following combination of features: ovipositor sheaths with pale subapical band, fore wing with infuscate median area and sparse setation basally, mesosoma with some reddish to orangish brown parts in addition to the acropleuron, and metafemur mostly brown. it differs from these species by a different combination of colour characters, setation of wing disc, and proportion of antennal segments. tineobius tamaricis sp. nov. differs from t. indicus, a species from pakistan, in having reduced orangish brown areas (mesoscutum and gaster entirely dark, metallic), metatarsus dark brown, fore wing without spatulate setae, fl2 1.35–1.65 × as long as pedicel, and ovipositor sheaths almost as long as gaster plus mesosoma. tineobius indicus (holotype in bmnh examined) differs by the following: front leg including coxa, as well as prepectus, and acropleuron except anteriorly, brownish-orange; mesoscutum brownish-orange except posterior median depressed area and posterior half of median and lateral lobes dark with some metallic lustre; mesoscutellum and axillae orange; first and second gastral tergites pale, yellowish-brown; metatarsus with first and last tarsomeres brown, second paler, and third and fourth pale yellowish; fore wing with short, stout and spatulate setae below parastigma, marginal and stigmal veins; fl2 about as long as pedicel and ovipositor sheaths as long as gaster plus half of mesosoma. tineobius tamaricis sp. nov. differs from t. philippinensis (holotype in bmnh examined), a species from philippines, in having the pronotum dark metallic, a reddish to orangish brown scape, head without conspicuously modified setae, reticulate frontovertex, scrobal depression as broad as high, fore wing without lanceolate setae, and ovipositor sheaths 0.8–0.9 × as long as rest of body. tineobius philippinensis has the following: lateral panel of pronotum anteriorly and posteriorly, prepectus, posterior half of acropleuron, base of mesoscutellum and axillae reddish-brown; scape brown and only slightly paler than flagellum; head with a tuft of long, black, lanceolate setae at boundary of vertex and occiput and very long erect setae on frontovertex; frontovertex virtually polished, with effaced coriaceous sculpture; scrobal depression strongly transverse, п-shaped and widely separated from anterior ocellus; fore wing with lanceolate to spatulate setae below parastigma and marginal vein; and ovipositor sheaths 0.5 × as long as rest of body. one paratype of t. tamaricis sp. nov. has all the parts that have a noticeable reddish hue in other specimens (fig. 1a–b) more inconspicuously differentiated in colour, dark reddish-brown (fig. 1c, g), and therefore in available keys runs to the afrotropical species t. afer (ferrière, 1938) comb. nov. and european journal of taxonomy 263: 1–19 (2017) 8 t. montanus (ferrière, 1938) comb. nov. (type material in bmnh examined). specimens of t. tamaricis sp. nov. with this colour pattern differ most conspicuously from t. afer in having a slenderer antenna, with fl2 more than 4 × as long as broad and longer than pedicel, fore wing with hair-like setae behind marginal vein, and scrobal depression carinate laterally. in t. afer fl2 is 2 × as long as broad and slightly shorter than pedicel, fore wing with spatulate setae behind parastigma and basal ⅔ of marginal vein, and scrobal depression only slightly impressed, about as long as wide, and almost touching inner eye orbit. from t. montanus it differs in having a shorter fl1 and in the structure of fore wing setation and scrobal depression. in t. montanus fl1 is about 2 × longer than broad, and the fore wing and scrobal depression are as described for t. afer. in having only hair-like setae on fore wing, t. tamaricis sp. nov. is similar to t. lamborni (ferrière, 1938) comb. nov. (holotype in bmnh examined) described from sub-saharan africa (malawi), but this species differs in having the pedicel only 1.3 × as long as fl1 and about 0.5 × as long as fl2, a brown scape only slightly paler than flagellum, a narrow λ-shaped scrobal depression carinately margined laterally along entire length except dorsally below anterior ocellus and separated from anterior ocellus by only about one ocellar diameter; body mostly dull, black with very reduced metallic lustre; fore wing base without band of pale setae behind parastigma, and bare except about basal half of basal cell with scattered dark setae. dna barcoding a complete dna barcode sequence was obtained from the holotype and was deposited on genbank under the accession number kt962861. because this is the first dna barcode of a tineobius species, a blast search (altschul et al. 1990) in genbank revealed high similarities with cecidostiba thomson, 1878 (pteromalidae) and eupelmus dalman, 1820 (eupelmidae) sequences, but at least this attests that it is a genuine chalcidoidea sequence and not one from a contaminant or an endosymbiont. ecology tineobius tamaricis sp. nov. is a parasitoid of parapodia sinaica larvae that form galls on tamarix twigs. these galls are known so far from egypt (houard 1912), iran (rezaei et al. 2007), israel (gerling et al. 1976), and france (joannis 1912, as p. tamaricicola joannis, 1912; dauphin & aniotsbehere 1994). parapodia sinaica produces galls on several tamarix species including t. africana poir., t. gallica l., t. tetragyna ehrenb., t. nilotica (ehrenb.) bunge, and t. jordanis boiss. to the best of our knowledge, it was not recorded previously from spain or on t. canariensis (new host record). the galls are similar to those of amblypalpis olivierella (ragonot, 1886) reported by gerling et al. (1976), another gelechiidae moth galling several tamarix species, but differ in being smaller in size (12–13 × 6–8 mm), having a fusiform shape, and the cavities inside the gall leaving the central column of the vascular bundles intact. adult moths have the fore wings pale greyish brown, with a dark and pale spotted pattern. in the survey of parasitoids in tamarix galls in lleida, a number of parasitoids were found in the galls, the most common being a species of apanteles sp. (braconidae: microgastrinae), of which 186 specimens emerged, and occasionally other parasitoids of braconidae, ichneumonidae, eulophidae, eupelmidae and pteromalidae (a paper concerning this parasitoid complex is in preparation). a small number of t. tamaricis sp. nov. females emerged from the galls, with a total of six specimens over four years and one specimen (on one occasion two) per year. it therefore is not a common species, but with a stable population in this stand of tamarix in this locality. emergence dates of t. tamaricis sp. nov. adult females were not clearly established, but possibly between june and august, as they were found dry in the rearing bags during this period, after the usual emergence dates between may and june of the lepidopteran host and most of their parasitoids. from one sample of galls collected on 25 jan. 2012, and after the emergence period finished with no tineobius emerging, in one of the galls opened on 15 mar. 2013 an adult female was found inside as it was unable to emerge, and near the wasp there were the remains of a consumed p. sinaica larva, including its cephalic capsule. parapodia sinaica larvae fusu l. & ribes a., a new species of tineobius from spain 9 usually bore an exit hole in the wall of the woody gall before pupation, partly closed by the epidermis of the gall, and some other parasitoid species were also found dead inside galls, unable to complete the hole for exiting. we interpret this finding as evidence for the p. sinaica larva being the host. in this case, t. tamaricis sp. nov. was either a primary parasitoid or, more likely, a hyperparasitoid via apanteles sp., because other species of tineobius with known biology are reported as primary and often as secondary parasitoids of lepidoptera (gahan 1927; ferrière 1936, 1938). for example, the type series of tineobius seyrigi (= anastatoidea seyrigi) (mnhn) is accompanied by a large moth cocoon of about 33 by 16 mm with the label: “obtenu d’éclosion de prospilus / cohacarum m. (i. litt.) sur / borocera sp. – sur 500 cocons, / seuls ceux qui contenaient des / prospilus, étaient parasités par l’eupelmid” [obtained by rearing from prospilus cohacarum mihi (in litteris) on borocera sp. – out of 500 cocoons only those containing prospilus were parasitised by the eupelmid] (see also ferrière 1936). checklist of described world species of tineobius tineobius (duanellus) bouček, 1988 duanellus bouček, 1988: 566, figs 1011–1013. type species: anastatoidea jacobsoni ferrière, 1938 by original designation. synonymy by gibson (1995: 284). 1. tineobius (duanellus) facialis (bouček, 1988) duanellus facialis bouček, 1988: 566–567, figs 1012–1013. holotype ♀, by original designation (anic, not examined). type locality: australia, queensland, 14 km nw hope vale mission. tineobius (duanellus) facialis – gibson 1995: 286. 2. tineobius (duanellus) jacobsoni (ferrière, 1938) anastatoidea jacobsoni ferrière, 1938: 50–52, 57, fig. 7. holotype ♀, unique specimen (bmnh 5.942, examined by lf). type locality: sumatra, fort de kock. anastatoïdea jacobsoni – risbec 1952: 67 (catalogued). duanellus jacobsoni – bouček 1988: 566, fig. 1011. tineobius (duanellus) jacobsoni – gibson 1995: 286. tineobius (progenitobius) gibson, 1995 tineobius (progenitobius) gibson, 1995: 286–287, fig. 452. type species: anastatoidea elongata risbec, 1952 by monotypy and original designation. 3. tineobius (progenitobius) elongatus (risbec, 1952) anastatoïdea elongata risbec, 1952: 67, 69–71, fig. 17. lectotype ♀ designated by gibson (1995: 286) (mnhn, examined by lf). type locality: madagascar, bekily. anastatoidea elongata – risbec 1958: 109 (keyed). tineobius (progenitobius) elongatus – gibson 1995: 284, 287, fig. 452. 4. tineobius (progenitobius) mandrakae (risbec, 1952) cerambycobius mandrakae risbec, 1952: 138–141. lectotype ♀ designated by fusu et al. (2015: 468) (mnhn, examined by lf). type locality: madagascar, mandraka. european journal of taxonomy 263: 1–19 (2017) 10 eupelmus mandrakae – hedqvist 1970: 436 (catalogued). tineobius (progenitobius) mandrakae – fusu et al. 2015: 468–469, fig. 3a, c (type information). tineobius (tineobius) ashmead, 1896 tineobius ashmead, 1896: 14–15. type species: tineobius citri ashmead, 1896 by original designation. australeupelmus girault, 1921: 3. type species: eupelmus multicolor girault, 1915 by monotypy and original designation. synonymy by bouček (1988: 564). anastatoidea gahan, 1927: 122. type species: anastatoidea brachartonae gahan, 1927 by original designation. synonymy by bouček (1988: 564). anastatoïdea – risbec 1952: 61, 66–71 (misspelling). 5. tineobius (tineobius) adamsi (yoshimoto & ishii, 1965) anastatus adamsi yoshimoto & ishii, 1965: 157, 159. holotype ♀, by original designation (usnm 67527, not examined). type locality: caroline island group: ponape island, net point. tineobius (tineobius) adamsi – gibson et al. 2012: 55. 6. tineobius (tineobius) afer (ferrière, 1938) comb. nov. anastatoidea afra ferrière, 1938: 52–53, 58. syntypes ♀♀ (bmnh 5.943, examined by lf). type locality: south west africa [namibia], aus. anastatoïdea afra – risbec 1952: 67 (keyed). 7. tineobius (tineobius) albopalpalis (brues, 1907) comb. nov. charitopus albopalpalis brues, 1907: 47–48. holotype ♂, unique specimen (mpmw, not examined). type locality: south africa, cape colony, algoa bay. remark this new combination in tineobius is proposed based on a personal communication by g.a.p. gibson who examined the holotype. in the original description sex incorrectly stated as female. 8. tineobius (tineobius) beenleighi (girault, 1926) stat. nov. [new subgeneric status] metapelma beenleighi girault, 1926: 64. holotype ♀, unique specimen (qmba t.4210, not examined). type locality: australia, beenleigh. metapelma beenleighi – girault 1930a: 3 (repetition of original description). — dahms 1983: 118 (information on type). anastatoidea beenleighi – ferrière 1938: 45, 58 (keyed). — risbec 1952: 66 (keyed). tineobius beenleighi – bouček 1988: 565. 9. tineobius (tineobius) beharae (risbec, 1952) comb. nov. anastatoïdea beharae risbec, 1952: 67, 71–73. syntypes ♀♂ (mnhn, examined by lf). type locality: madagascar, behara. fusu l. & ribes a., a new species of tineobius from spain 11 10. tineobius (tineobius) brachartonae (gahan, 1927) stat. nov. [new subgeneric status] anastatoidea brachartonae gahan, 1927: 13–15. holotype ♀, by original designation (usnm 29444, not examined). type locality: central java. anastatoidea brachartonae – ferrière 1936: 195 (catalogued, distribution). — ferrière 1938: 48, 57 (keyed). — risbec 1952: 67 (catalogued). tineobius brachartonae – bouček 1988: 564. 11. tineobius (tineobius) capensis (ferrière, 1938) comb. nov. anastatoidea capensis ferrière, 1938: 53–54, 58. syntypes ♀ (bmnh 5.945, examined by lf). type locality: south africa: pondoland, fort st. john. anastatoïdea capensis – risbec 1952: 67, 68 (keyed, distribution). 12. tineobius (tineobius) citri ashmead, 1896 stat. nov. [new subgeneric status] tineobius citri ashmead, 1896: 15. holotype ♀, unique specimen (usnm 3465, not examined). type locality: australia, new south wales, paramatta. tineobius citri – bouček 1988: 565, figs 1004, 1005 (distribution, redescription). 13. tineobius (tineobius) columbi (girault, 1923) stat. nov. [new subgeneric status] metapelma columbi girault, 1923: 96–97. holotype ♀, unique specimen (qmba t.4215, not examined). type locality: australia, queensland, kuranda. metapelma columbi – dahms 1983: 189 (type information). anastatoidea columbi – gahan 1927: 15 (keyed). — ferrière 1936: 195 (catalogued). — ferrière 1938: 45, 57 (keyed, distribution). anastatoïdea colombi – risbec 1952: 67 (misspelling). tineobius columbi – bouček 1988: 565. 14. tineobius (tineobius) crassipes (yoshimoto & ishii,1965) anastatus crassipes yoshimoto & ishii, 1965: 156–158. holotype ♀, by original designation (bpbm 3581, not examined). type locality: caroline island group: palau island, koror. tineobius (tineobius) crassipes – gibson et al. 2012: 56. 15. tineobius (tineobius) decoratus (ferrière, 1938) stat. nov. [new subgeneric status] anastatoidea decorata ferrière, 1938: 45–46, 59, fig. 5. holotype ♀, by original designation (bmnh 5.940, examined by lf). type locality: fiji: lautoka. anastatoïdea decorata – risbec 1952: 67 (catalogued). tineobius decoratus – bouček 1988: 565 (distribution). european journal of taxonomy 263: 1–19 (2017) 12 16. tineobius (tineobius) gonometae (ferrière, 1938) comb. nov. anastatoidea gonometae ferrière, 1938: 55–56, 58. holotype ♀, unique specimen (bmnh 5.946, examined by lf). type locality: south africa: transvaal, louis trichardt. anastatoïdea gonometae – risbec 1952: 67 (keyed). 17. tineobius (tineobius) indicus (ferrière, 1938) stat. nov. [new subgeneric status] anastatoidea indica ferrière, 1938: 49–50, 58, fig. 6. holotype ♀, unique specimen (bmnh type no. 5.937, examined by lf). type locality: india [now pakistan]: punjab, lyallpur. anastatoïdea indica – risbec 1952: 66 (keyed). tineobius indicus – bouček 1988: 565. 18. tineobius (tineobius) lamborni (ferrière, 1938) comb. nov. anastatoidea lamborni ferrière, 1938: 56, 58. holotype ♀, unique specimen (bmnh type no. 5.947, examined by lf). type locality: east africa: nyasaland, kota-kotu. anastatoïdea lamborni – risbec 1952: 67 (keyed). 19. tineobius (tineobius) longfellowi (girault, 1923) stat. nov. [new subgeneric status] metapelma longfellowi girault, 1923: 99. holotype ♀, unique specimen (qmba t.4209, not examined). type locality: australia: kuranda. metapelma longfellowi – girault 1926: 64. — girault 1930a: 3 (compared with m. beenleighi). — dahms 1984: 764 (type information). anastatoidea longfellowi – gahan 1927: 15 (keyed). — ferrière 1938: 45, 57 (keyed). — risbec 1952: 67 (catalogued). tineobius longfellowi – bouček 1988: 565. 20. tineobius (tineobius) longicauda (ferrière, 1938) stat. nov. [new subgeneric status] anastatoidea longicauda ferrière, 1938: 49, 57. holotype ♀, unique specimen (bmnh 5.938, examined by lf). type locality: siam: bangkok. anastatoïdea longicauda – risbec 1952: 67 (catalogued). tineobius longicauda – bouček 1988: 565. 21. tineobius (tineobius) madagascariensis fusu & ribes nom. nov. metapelma seyrigi risbec, 1952: 63–66, fig. 16e, f. holotype ♀, unique specimen (mnhn, examined by lf). type locality: madagascar: ranomafana. secondary homonym of tineobius seyrigi (ferrière, 1938). 22. tineobius (tineobius) montanus (ferrière, 1938) comb. nov. anastatoidea montana ferrière, 1938: 53, 58. holotype ♀, unique specimen (bmnh 5.944, examined by lf). type locality: south africa: cape province, katberg, 1200 m. anastatoïdea montana – risbec 1952: 67 (keyed). fusu l. & ribes a., a new species of tineobius from spain 13 23. tineobius (tineobius) multicolor (girault, 1915) stat. nov. [new subgeneric status] eupelmus multicolor girault, 1915: 10. holotype ♀, unique specimen (qmba hy.2489, not examined). type locality: australia: queensland, gordonvale (cairns). eupelmus multicolor – dahms 1984: 835–836 (type information). australeupelmus multicolor – girault 1921: 3. tineobius multicolor – bouček 1988: 565 (distribution, stated as very close to tineobius citri). 24. tineobius (tineobius) nassaui (girault, 1933) stat. nov. [new subgeneric status] metapelma (tineobius) nassaui girault, 1933: 3. holotype ♀, unique specimen (qmba t.4211, not examined). type locality: e. australia [brisbane according to label data]. metapelma nassaui – girault 1930b: 3 (nomen nudum, discovered by dahms 1986: 321). metapelma nassaui – dahms 1986: 321 (type locality data). tineobius nassaui – bouček 1988: 565 (possible synonym of t. citri). 25. tineobius (tineobius) philippinensis (ferrière, 1938) comb. nov. anastatoidea philippinensis ferrière, 1938: 47–48, 58. holotype ♀, unique specimen (bmnh 5.939, examined by lf). type locality: philippines: mindanao, zamboanga. anastatoïdea philippinensis – risbec 1952: 66 (keyed). — baltazar 1966: 129 (catalogued). 26. tineobius (tineobius) rufescens (ferrière, 1938) comb. nov. anastatoidea rufescens ferrière, 1938: 54–55, 59. syntypes ♀♀ ( bmnh 5.948, examined by lf). type locality: south africa: pondoland, port st. john. anastatoïdea rufescens – risbec 1952: 67, 68 (keyed, distribution) 27. tineobius (tineobius) salomonis (ferrière, 1938) stat. nov. [new subgeneric status] anastatoidea salomonis ferrière, 1938: 46–47, 59. syntypes ♀♀ (bmnh 5.941, examined by lf). type locality: solomon island group: tulagi and guadalcanal. anastatoïdea salomonis – risbec 1952: 67 (catalogued). tineobius salomonis – bouček 1988: 565. 28. tineobius (tineobius) seyrigi (ferrière, 1936) comb. nov. anastatoidea seyrigi ferrière, 1936: 196–198, fig. 4. syntypes ♀♂ (bmnh 5.949, mnhn, examined by lf). type locality: madagascar, bekily. anastatoidea seyrigi – ferrière 1938: 52, 57 (keyed). — risbec 1952: 66, 68 (keyed). 29. tineobius (tineobius) sidneyi (girault, 1930) stat. nov. [new subgeneric status] metapelma sidneyi girault, 1930b: 3. syntypes ♀♀ (anic, qmba t.4214, not examined). type locality: australia, new south wales, sydney. metapelma sidneyi – dahms 1986: 532 (type information). — naumann et al. 1994: 72 (type information). tineobius sidneyi – bouček 1988: 565. european journal of taxonomy 263: 1–19 (2017) 14 30. tineobius (tineobius) superbus (dodd, 1917) stat. nov. [new subgeneric status] metapelma superba dodd, 1917: 357–358. holotype ♀, by original designation (qmba, not examined). type locality: australia: queensland, townsville. metapelma superba – girault 1923: 96, 99 (compared with t. columbi and t. longfellowi). anastatoidea superba – gahan 1927: 15 (keyed). — ferrière 1938: 45, 57 (keyed). — risbec 1952: 67 (catalogued). tineobius superbus – bouček 1988: 566. 31. tineobius (tineobius) tamaricis ribes & fusu sp. nov. species described in anastatoidea but excluded from tineobius 1. eupelmus (episolindelia) ambatomangae (risbec, 1958) comb. nov. anastatoidea ambatomangae risbec, 1958: 109, 112–114, fig. 2. holotype ♀, unique specimen (mnhn, examined by lf). type locality: madagascar, ambatomanga. 2. eupelmus (episolindelia) pennisetae (risbec, 1958) anastatoidea pennisetae risbec, 1958: 109–112, fig. 1. syntypes ♀♂ (mnhn, examined by lf). type locality: madagascar: terrain d’aviation de majunga. eupelmus (episolindelia) pennisetae – fusu et al. 2015: 460 (type information). discussion tineobius is a diverse genus known from the afrotropical, oriental, and australasian regions, including numerous undescribed species (gibson 1995), but previously not recorded from the palaearctic region. the newly described t. tamaricis sp. nov. from spain represents the first species of the genus for the palaearctic region. the most geographically close records of other tineobius species are from pakistan and the sub-saharan afrotropical region. the association of t. tamaricis sp. nov. with parapodia sinaica, which is probably distributed over the entire mediterranean area on several tamarix species, implies that t. tamaricis sp. nov. could also be distributed over a larger area. the distribution range of several tamarix species extends further into the oriental and afrotropical regions, mainly through near east where p. sinaica galls were found to attack three different tamarix species (gerling et al. 1976). via an iranian, ethiopian or saharan path, there might be a connection between the populations of t. tamaricis sp. nov. in spain with the tropical regions were tineobius is more diverse. on the other hand, the localised distribution of t. tamaricis sp. nov., the fact that it was not collected before despite the eupelmidae of spain being well investigated (askew & nieves-aldrey 2000, 2004, 2006) and this being the sole species of tineobius present in the palaearctic region, might indicate that it is in fact an introduced species and not a local endemic. also, the absence of any males reared over the four years might indicate it is a thelytokous species. if so, the likelihood of the species being introduced from elsewhere is further increased as there are examples of several eupelmid species in north america being introduced from other regions and with only females known (parthenogenesis favours the establishment of introduced species; hoffman et al. 2008). this is the case of balcha indica (mani & kaul, 1973) (gibson 2005), eupelmus pini taylor, 1927 (gibson 2011; gibson & fusu 2016), and eupelmus vesicularis (retzius, 1783) (s. lat., fusu 2010). as exotic species of tamarix are widely used as ornamental plants or for reforestation throughout europe, t. tamaricis sp. nov. might have been introduced together with infested plants from africa or asia. fusu l. & ribes a., a new species of tineobius from spain 15 acknowledgements the authors are grateful to g.a.p. gibson (cnc, ottawa, canada) for information on the type material of some tineobius species and to peter huemer (tiroler landesmuseen-betriebsges, innsbruck, austria) for the verification of the identification of parapodia sinaica. l. fusu is thankful to r.r. askew (beeston, cheshire, uk) for locating the unlabelled type series of t. tamaricis sp. nov. in arpc and mailing it to aicf for curation and to two anonymous referees for their valuable comments. this study was supported by the research grant gi-2015-02, internal grants 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botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands. fusu l. & ribes a., a new species of tineobius from spain 19 http://dx.doi.org/10.1080/00222933.2013.852264 http://dx.doi.org/10.1371/journal.pone.0015991 phylogeny of miliusa (magnoliales: annonaceae: malmeoideae: miliuseae), with descriptions of two new species from malesia – corrigendum tanawat chaowasku1, paul j.a. keßler2 & lars w. chatrou3 1 naturalis biodiversity center (section nhn), leiden university, p.o. box 9514, 2300 ra leiden, the netherlands e-mail: cyathostemma@yahoo.com (corresponding author) 2 hortus botanicus leiden, p.o. box 9516, 2300 ra leiden, the netherlands 3 wageningen university biosystematics group, droevendaalsesteeg 1, 6708 pb wageningen, the netherlands authorship of the new species remains unchanged. chaowasku t., keßler p.j.a. & chatrou l.w. 2013. phylogeny of miliusa (magnoliales: annonaceae: malmeoideae: miliuseae), with descriptions of two new species from malesia – corrigendum. european journal of taxonomy 64: 1. http://dx.doi.org/10.5852/ejt.2013.64 published on: 18 november 2013 printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; national botanic garden of belgium, meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark. european journal of taxonomy 64: 1 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2013.64 www.europeanjournaloftaxonomy.eu 2013 · tanawat chaowasku, paul j.a.keßler & lars w. chatrou this work is licensed under a creative commons attribution 3.0 license. c o r r i g e n d u m the third author has been added to paper no. 54 (http://dx.doi.org/10.5852/ejt.2013.54) 1 mailto:cyathostemma%40yahoo.com?subject=cyathostemma%40yahoo.com http://dx.doi.org/10.5852/ejt.2013.64 http://dx.doi.org/10.5852/ejt.2013.64 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index http://creativecommons.org/licenses/by/3.0/ http://dx.doi.org/10.5852/ejt.2013.54 1 european journal of taxonomy 729: 1–10 issn 2118-9773 https://doi.org/10.5852/ejt.2020.729.1185 www.europeanjournaloftaxonomy.eu 2020 · bitencourt c. et al. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e hemipogon trilobatus sp. nov. (apocynaceae: asclepiadoideae), a new microendemic from chapada dos veadeiros, central brazil cássia bitencourt 1,*, amanda p.b. santos 2, cristiane snak 3 & alessandro rapini 4 1,2,3,4 universidade estadual de feira de santana, programa de pós-graduação em botânica, av. transnordestina, s/n, novo horizonte, 44036-900, feira de santana, bahia, brazil. 3 universidade do estado de santa catarina, departamento de engenharia de pesca e ciências biológicas, rua cel. fernandes martins 270, progresso, 88790-000, laguna, santa catarina, brazil. * corresponding author: ca.biten@gmail.com 2 email: amanda.pricilla@hotmail.com 3 email: cristianesnak@gmail.com 4 email: rapinibot@yahoo.com.br abstract. hemipogon s. str. (apocynaceae: asclepiadoideae) currently consists of three species sharing an erect herbaceous habit, narrow leaves and corona-less flowers with urceolate, internally bearded corolla, that are mainly distributed in savannahs of the cerrado biodiversity hotspot, south america. here, we describe and illustrate a new species of hemipogon, h. trilobatus bitencourt & rapini sp. nov., from an open savannah in chapada dos veadeiros, central brazil. hemipogon trilobatus sp. nov. differs from the other species of the genus mainly by the presence of a reduced staminal corona with 3-lobed lobes, but also by opposite leaves and triangular anthers. distribution and habitat data, as well as a key and a comparative table to distinguish the four species currently accepted in hemipogon s. str., are provided. based on criteria b2ab(i,ii,iii,iv) of the international union for conservation of nature (iucn), the species is provisionally assessed as critically endangered. keywords. cerrado, metastelmatinae, neotropics, savannah, taxonomy. bitencourt c., santos a.p.b., snak c. & rapini a. 2020. hemipogon trilobatus sp. nov. (apocynaceae: asclepiadoideae), a new microendemic from chapada dos veadeiros, central brazil. european journal of taxonomy 729: 1–10. https://doi.org/10.5852/ejt.2020.729.1185 introduction hemipogon decne. (apocynaceae juss.: asclepiadoideae burnett) belongs to the neotropical subtribe metastelmatinae endl. ex meisn. and originally comprised erect perennial herbs with narrow leaves and urceolate, internally bearded, corona-less flowers (fournier 1885). the genus was enlarged with the inclusion of twining plants with corona-less flowers classified in the american astephanus r.br. and erect herbs with corona-bearing flowers segregated from metastelma r.br. (rapini et al. 2001; rapini 2002). phylogenetic studies (ribeiro et al. 2012, 2014; silva et al. 2012), however, showed that these two groups form a lineage predominantly distributed through the campos rupestres (rocky https://doi.org/10.5852/ejt.2020.729.1185 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ mailto:ca.biten%40gmail.com?subject= mailto:amanda.pricilla%40hotmail.com?subject= mailto:cristianesnak%40gmail.com?subject= mailto:rapinibot%40yahoo.com.br?subject= https://doi.org/10.5852/ejt.2020.729.1185 european journal of taxonomy 729: 1–10 (2020) 2 grasslands on quartzite outcrops) of the espinhaço range and are not closely related to hemipogon acerosus decne., the type of the genus. the phylogenetic inference indicates that the corona was lost several times within metastelmatinae and thus hemipogon should be treated under its narrow concept, comprising only three species, h. acerosus, h. setaceus decne. and h. irwinii fontella & paixão, or four, if h. acerosus var. platyphyllus hoehne would be recognised at the species level (bitencourt 2019). using this narrow circumscription, hemipogon is predominantly distributed in savannahs, and less often in campos rupestres of the cerrado domain, in central brazil, reaching bolivia, peru and paraguay. cerrado is a biodiversity hotspot (myers et al. 2000) and the world’s most diverse savannah, with around 6910 species of flowering plants, approximately 3815 of which are endemic (silva & bates 2002; flora do brasil 2020). despite its great importance, about half of the area originally occupied by cerrado has been lost due to the implantation of pasture and commercial crops, mainly soybean and corn, and only 10% of its original area is protected in conservation units (beuchle et al. 2015). chapada dos veadeiros in the brazilian cerrado houses savannahs in lowlands and patches of campos rupestres over mountaintops up to 1683 m a.s.l. (topographic map, https://www.arcgis.com/), and is considered a conservation priority area due to the high level of unique evolutionary diversity (carvalho et al. 2015; fenker et al. 2020). describing the biodiversity is an essential step to understand evolutionary and biogeographic patterns, and establish conservation strategies (brach & boufford 2011; ulloa et al. 2017). this task is especially relevant in areas under high pressure, such as the cerrado hotspot. herbaria are important source of new species (prance 2001; joppa et al. 2010; bebber et al. 2010) and most undescribed species are discovered a long time after a first specimen was collected (bebber et al. 2010). here, we describe a new species from chapada dos veadeiros first collected 28 years ago. it fits the narrow concept of hemipogon and is here provisionally assessed as critically endangered. material and methods as part of a large study which aims to propose a new circumscription for the genus hemipogon, specimens from different species of south american savannahs were examined in herbaria and collected in situ. morphological measurements for this study were taken from specimens in the herbaria bhcb, cen, hbr, hrb, huefs, ibge, lpb, mbm, mo, ny, r, rb, ub, ufg, ufmt and usz (acronyms according to the index herbariorum, thiers continuously updated: http://sweetgum.nybg.org/science/ih/). morphological terms follow beentje (2016) and endress et al. (2018). a provisional species conservation status was inferred based on iucn (2019), according to the categories, criteria and conditions outlined in the red list guidelines. we used the geocat tool (bachman et al. 2011) to calculate the area of occupancy (aoo) using a 2 × 2 km2 cell resolution. https://www.arcgis.com/ http://sweetgum.nybg.org/science/ih/ bitencourt c. et al., a new species of hemipogon 3 results class magnoliopsida brongn. order gentianales bercht. & j.presl family apocynaceae juss. subfamily asclepiadoideae burnett genus hemipogon decne. hemipogon trilobatus bitencourt & rapini sp. nov. urn:lsid:ipni.org:names:77213240-1 figs 1–2 diagnosis flowers with reduced, apically 3-lobed corona lobes. similar to h. acerosus due to the acicular leaves, sessile cymes and urceolate corolla, internally bearded along the lobe basal half, but can be distinguished by the opposite leaves (vs verticillate in h. acerosus) and presence of a corona (vs absence in h. acerosus). etymology the epithet ʻtrilobatusʼ refers to the apically 3-lobed corona lobes. type material brazil • goiás, alto paraíso de goiás, chapada dos veadeiros, 9 km de alto paraíso para vila são jorge, cachoeira são bento; 9 sep. 1994; m. aparecida da silva, t.s. filgueiras, r.c. mendonça, m.l.f. resende, f.c.a. oliveira and e. cardoso 2320; holotype: ibge[33546]!; isotype: rb[496900]!. paratype brazil • goiás, alto paraíso de goiás, chapada dos veadeiros; 14 aug. 1992; h.d. ferreira 2579; ufg[14114]!. description herbs to subshrub erect, 20–30 cm tall; stems 2 or 3, diverging at base, cylindrical, glabrescent; latex white. leaves opposite, obliquely erect, sessile, acicular to narrowly elliptic, 0.7–1 × 0.1–0.13 mm, apex acute, base cuneate, margins entire, coriaceous, glabrescent. cymes alternate, 2or 4-flowered, subaxillary; bracts lanceolate, ca 1.5 × 0.6 mm, glabrous, sessile. flowers with pedicel 1–1.5 mm long; sepals ovate, 1.5–1.6 × 0.5–0.7 mm, apex acute, glabrescent, axillary colleters not seen; corolla urceolate, probably cream or white, abaxially glabrescent, adaxially bearded from base to centre of lobes, tube 0.5– 0.7 × 0.8–1.1 mm, lobes triangular, 1–1.2 × 0.8–1 mm, apex acute and recurved; corona staminal, lobes short, at base of anthers, oblate, 0.3–0.45 × 0.7–0.76 mm, apex 3-lobed; anthers 0.45–0.5 × 0.6–0.7 mm, triangular, wings triangular, 0.3–0.4 × 0.15–0.2 mm, connective appendage oblong, 0.3–0.4 mm long, over gynostegium; corpusculum ellipsoid, 0.17–0.25 × 0.08–0.1 mm; caudicles horizontal, ca 0.05 mm long, subapically attached to pollinia; pollinia ellipsoid, 0.16–0.2 × 0.06–0.08 mm; style-head apically mammillate, included within corolla tube. follicles not seen. distribution, habitat and phenology hemipogon trilobatus sp. nov. is known from a small population nearby the são bento waterfall, in chapada dos veadeiros, goiás, brazil (fig. 2). it grows in open savannah and wet grasslands locally known as “várzea” and was collected with flowers in august and september. the species has not been found since a fire burned the type locality (m.a. da silva, pers. com., 2015). http://www.ipni.org/urn:lsid:ipni.org:names:77213240-1 european journal of taxonomy 729: 1–10 (2020) 4 fig. 1. hemipogon trilobatus bitencourt & rapini sp. nov. a. habitus. b. cyme with two flowers and a bud. c. flower. d. flower with calyx and two corolla lobes removed to show the gynostegium. e. corolla lobe, adaxial view. f. anther with a staminal corona lobe basally attached. g. corona lobe showing the 3-lobed apex, adaxial view. h. gynostegium, viewed from above. i. pollinarium. drawn by pétala gomes ribeiro from the holotype (ibge[33546]). bitencourt c. et al., a new species of hemipogon 5 fig. 2. habitat and distribution of hemipogon trilobatus bitencourt & rapini sp. nov. a. open savannah in chapada dos veadeiros, state of goiás, brazil. photo by cb. b. map showing the geographic distribution of h. trilobatus sp. nov. in the cerrado domain (shaded area at reference map in a) and its occurrence (black star) nearby the chapada dos veadeiros national park. images and distribution maps were built and exported using arcgis online (https://www.arcgis.com). © esri and its licensors, all rights reserved. https://www.arcgis.com european journal of taxonomy 729: 1–10 (2020) 6 provisional conservation status hemipogon trilobatus sp. nov. is known from only two collections, both near the são bento waterfall, in chapada dos veadeiros. specimen labels provide no information about population density. however, the species is known to occur in a tourist location, surrounded by farms and cattle raising, has an area of occupancy (aoo) estimated at 8 km2, probably under continuing decline in geographic range and habitat quality. during fieldwork in the type locality, we did not find any specimen of h. trilobatus sp. nov. and the new species has not been recollected since 1994. based on criteria b2ab(i,ii,iii,iv) of the international union for conservation of nature (iucn 2019) red list assessment, we provisionally classify the species as critically endangered (cr). taxonomic note hemipogon trilobatus sp. nov. has narrow leaves, urceolate corolla with bearded lobes and was found in a savannah of central brazil. together, these features support its classification in hemipogon s. str. it most closely resembles h. acerosus based on the acicular leaves and sessile cymes, but differs from all other taxa in the genus by the opposite leaves and flowers with reduced corona lobes, as presented in the key below and table 1. key to species of the genus hemipogon s. str. 1. leaves opposite; flowers with short, 3-lobed staminal corona lobes ................................................. ..........................................................................................h. trilobatus bitencourt & rapini sp. nov. – leaves verticillate or spirally arranged, flowers without corona ...................................................... 2 2. leaf blade filiform, spirally arranged ............................................................................................... 3 – leaf blade acicular or ovate to lanceolate, verticillate ..................................................................... 4 3. corolla lobes 8.5–10 mm long, twisted in the bud, adaxially barbate on the basal half, sericeous on the apical half ....................................................................................... h. irwinii fontella & paixão – corolla lobes 3.8–7 mm long, straight in the bud, adaxially barbate on the basal third, puberulous towards the apex .................................................................................................. h. setaceus decne. 4. leaves sessile, the blade acicular, ≤ 1.5 mm wide ....................... h. acerosus var. acerosus decne. – leaves petiolate, the blade ovate to lanceolate, ca 3 mm wide .......................................................... ............................................................................................... h. acerosus var. platyphyllus hoehne discussion asclepiadoideae is a cosmopolitan subfamily with approximately 180 genera and 3200 species (endress et al. 2018); around one third of this diversity is represented in the americas (good 1952; rapini 2012). the proportion of narrowly distributed species and genera in the subfamily is high (good 1952), and many species are known only from the type locality (endress et al. 2018). brazil harbours 35 native genera of asclepiadoideae, only five of them including more than 15 species. narrowly distributed species of asclepiadoideae are common in the country (rapini et al. 2001, 2002, 2009; rapini 2010) and several brazilian microendemics have been described in the last years (e.g., bitencourt et al. 2020). this contributed to the increase of the number of asclepiadoideae species in brazil from 376 (rapini et al. 2005) to 400 (flora do brazil 2020) in the last 15 years. some of the asclepiadoideae microendemics are poorly known due to the limited number of collections. hemipogon abietoides e.fourn., described in flora brasiliensis (fournier 1885) from a specimen collected by riedel during the legendary expedition headed by langsdorff through the espinhaço range of minas gerais, for instance, was recollected only 175 years after its original collection (rapini bitencourt c. et al., a new species of hemipogon 7 et al. 2010). molecular phylogenetic analyses (ribeiro et al. 2012, 2014; silva et al. 2012), however, showed that this species together with h. hemipogonoides (malme) rapini and h. hatschbachii (fontella & marquete) rapini, both erect herbs with narrow leaves and endemic to the campos rupestres (rocky grasslands on quartzite outcrops) of serra do cipó in the espinhaço range, belong to a lineage distant to hemipogon s. str. and sister to the clade of twining plants predominantly distributed in the espinhaço range and assigned to the recently described morilloa fontella, goes & s.cáceres (= the former american astephanus; fontella-pereira et al. 2014). unpublished plastome phylogenetic analyses confirmed that convergences in metastelmatinae are more common than previously supposed. they showed that two other species from the espinhaço range, also morphologically similar to hemipogon s. lat. but endemic to cangas (rocky grasslands on iron outcrops) of the iron quadrangle form a clade separate from both hemipogon s. str. and morilloa (bitencourt 2019). the two species were recently described in the non-monophyletic ditassa r.br., as d. cangae bitencourt & rapini and d. ferricola bitencourt & rapini (bitencourt et al. 2020). the erect habit, narrow leaves and urceolate corolla of hemipogon trilobatus sp. nov. may also represent another example of morphological convergence. in preliminary phylogenetic analyses of its rdna sequence data, h. acerosus and h. trilobatus sp. nov. appeared in a polytomy, but not closely related to morilloa (unpublished results). the geographical distribution of h. trilobatus sp. nov., restricted to an open savannah in central brazil rather than cangas or campos rupestres of espinhaço range, can be regarded as additional evidence for its placement in hemipogon until further phylogenetic studies may provide new insights towards a new generic circumscription in metastelmatinae and a more confident placement for h. trilobatus sp. nov. acknowledgements we thank betânia goes and maria aparecida silva for supporting cb’s visit to ibge reserve, marina resende (ibge) and rafaela campostrini forzza (rb) for specimen loans, moabe f. fernandes for help with fieldwork and suggestions on a first draft of the manuscript, three anonymous reviewers and the topical editor frederik leliaert for corrections and suggestions and pétala gomes ribeiro for the illustration. this work is part of the phd thesis of cb, developed at ppgbot-uefs with a fellowship from the coordenação de aperfeiçoamento de pessoal de nível superior – brasil (capes) – finance codes #1514632 (ds) and #88881.135731/2016-01 (pdse), and supported by the project universal (cnpq #485468/2013-1). cb acknowledges the international association of plant taxonomy (iapt, 2016) grant, neotropical grasslands conservancy memorial (ngc, 2016) and shirley a. graham (missouri botanical garden, 2018) fellowships. ar is supported by cnpq (productivity fellowship no. 307396/2019-3). characters h. trilobatus sp. nov. h. acerosus h. irwinii h. setaceus leaves opposite and acicular verticillate and acicular spirally arranged and filiform spirally arranged and filiform corolla probably cream or white cream white cream corona present absent absent absent anther triangular subtriangular subsagittate subtringular corpusculum ellipsoid ellipsoid oblong ellipsoid table 1. diagnostic characters to distinguish the species of hemipogon s. str. european journal of taxonomy 729: 1–10 (2020) 8 authors’ contributions ar 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meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. https://doi.org/10.1600/036364412x648733 http://sweetgum.nybg.org/science/ih/ https://doi.org/10.1126/science.aao0398 european journal of taxonomy 586: 1–22 issn 2118-9773 https://doi.org/10.5852/ejt.2019.586 www.europeanjournaloftaxonomy.eu 2019 · chester c. et al. this work is licensed under a creative commons attribution 3.0 license. o p i n i o n p a p e r 1 ejt editorial standard for the semantic enhancement of specimen data in taxonomy literature chloë chester 1,*, donat agosti 2, guido sautter 3, terry catapano 4, koen martens 5, isabelle gérard 6 & laurence bénichou 7 1,7 muséum national d’histoire naturelle, 57 rue cuvier, 75005 paris, france. 2,3,4 plazi, zinggstrasse 16, 3007 bern, switzerland. 5 royal belgian institute for natural sciences, vautierstraat 29, b-1000 brussels, belgium. 6 royal museum for central africa, 13 leuvensesteenweg, b-3080 tervuren, belgium. * corresponding author: chloe.chester@mnhn.fr 2 email: agosti@amnh.org 3 email: gsautter@gmail.com 4 email: catapano@plazi.org 5 email: koen.martens@naturalsciences.be 6 email: isabelle.gerard@africamuseum.be 7 email: laurence.benichou@mnhn.fr abstract. this paper describes a set of guidelines for the citation of zoological and botanical specimens in the european journal of taxonomy. the guidelines stipulate controlled vocabularies and precise formats for presenting the specimens examined within a taxonomic publication, which allow for the rich data associated with the primary research material to be harvested, distributed and interlinked online via international biodiversity data aggregators. herein we explain how the ejt editorial standard was defined and how this initiative fits into the journal’s project to semantically enhance its publications using the plazi taxpub dtd extension. by establishing a standardised format for the citation of taxonomic specimens, the journal intends to widen the distribution of and improve accessibility to the data it publishes. authors who conform to these guidelines will benefit from higher visibility and new ways of visualising their work. in a wider context, we hope that other taxonomy journals will adopt this approach to their publications, adapting their working methods to enable domain-specific text mining to take place. if specimen data can be efficiently cited, harvested and linked to wider resources, we propose that there is also the potential to develop alternative metrics for assessing impact and productivity within the natural sciences. keywords. xml, publishing standard, taxonomy, fair data, open science. chester c., agosti d., sautter g., catapano t., martens k., gérard i. & bénichou l. 2019. ejt editorial standard for the semantic enhancement of specimen data in taxonomy literature. european journal of taxonomy 586: 1–22. https://doi.org/10.5852/ejt.2019.586 european journal of taxonomy 586: 1–22 (2019) 2 table of contents introduction ............................................................................................................................................. 2 approach, tools and partners ................................................................................................................... 4 semantic enhancement of specimen citations .......................................................................................... 5 material citations formatting guidelines ................................................................................................. 8 general presentation ......................................................................................................................... 8 1. order ......................................................................................................................................... 8 2. punctuation ............................................................................................................................... 9 3. type material ............................................................................................................................ 9 4. repetitive data ........................................................................................................................ 10 5. ‘missing’ elements ................................................................................................................... 11 6. label citations .......................................................................................................................... 11 7. author interpretation ................................................................................................................ 11 data fields ....................................................................................................................................... 12 1. country/water body ................................................................................................................ 12 2. specimen count (zoology) ...................................................................................................... 12 3. locality ................................................................................................................................... 12 4. geographic coordinates .......................................................................................................... 13 5. altitude/elevation/depth .......................................................................................................... 13 6. collection date ......................................................................................................................... 13 7. collector (zoology) .................................................................................................................. 13 8. collector and collection number (botany) ............................................................................... 13 9. additional data ......................................................................................................................... 13 10. associated sequences ............................................................................................................ 14 11. repository data ...................................................................................................................... 14 discussion ............................................................................................................................................. 15 conclusion ............................................................................................................................................. 18 acknowledgements ................................................................................................................................ 18 references .............................................................................................................................................. 18 introduction taxonomy and the urgent need for integrated bioinformatics publications containing descriptive taxonomy and nomenclatural acts constitute a pillar for developing robust hypotheses on identity and relationships in the natural world. species names and the treatments associated with them are a prerequisite for reliable research across the natural sciences (wägele et al. 2011), which in turn plays a pivotal role in effective conservation management and sustainable development (e.g., groombridge 1992; heywood et al. 1995; mccook et al. 2010). at a time when there is growing concern across the planet for the cascading effects of climate change and biodiversity loss on agriculture, land use and human welfare (ipcc 2018), it is now urgent for taxon concepts to move beyond the description of isolated organisms and towards an integrative systems approach that is able to address species interactions, both with their environment and with other species (hardisty et al. 2013). despite its fundamental role in subsequent fields of research, alpha taxonomy is suffering from impediments that effect end-user accessibility, as well as the valorisation of its authors and their output (ebach et al. 2011). the establishment of best practices in the citation of collection specimens, taxon concepts and bibliographic works would contribute to the interlinked resource infrastructure that is called for by the community (hobern et al. 2019) and help to resolve some of the issues. however, the challenge therein is twofold, as we must overcome both sociological (e.g., habits and ‘traditional’ ways of working) and technical barriers in order to engineer change. this notion was a guiding vector throughout the european journal of taxonomy’s project for the semantic enhancement of its chester c. et al., semantic enhancement of specimen citations 3 publications. achieving dynamic data exchange was not the unique goal; as well as the chosen workflow (the technical framework), the applicability and relevance of the method (the sociological factor) were equally paramount to ensuring that authors embrace the increasingly technological research paradigm. the european journal of taxonomy: spearheading innovative publishing workflows the european journal of taxonomy (ejt) is a peer-reviewed international journal in descriptive taxonomy of eukaryotic organisms (zoology, entomology, botany, palaeontology). the journal was founded by a consortium of european natural history institutions to take advantage of the shift from paper to online publications, as supported by the recent modifications in the governing codes of nomenclature in zoology (iczn 2009, 2012) and botany (mcneill et al. 2012, turland et al. 2018). from the journal’s onset in 2011, ejt’s articles have been published directly online as individual pdfs in diamond open access (bénichou et al. 2011). however, in accordance with the journal’s founding principle to spearhead innovative publishing techniques, it had always been envisaged to provide publications in a machine-readable format, namely extensible mark-up language (xml). beyond the advantages of xml’s stability as an archiving format (morrissey et al. 2010; coalition s 2019; library of congress 2019), which would serve to guarantee the longevity of its publications, ejt understood that the way to increase the visibility of taxonomic research was to provide machinereadable and semantically enhanced text (agosti et al. 2007; agosti & egloff 2009; penev et al. 2010; penev et al. 2011a; penev et al. 2012; miller et al. 2012; miller et al. 2015). moreover, in light of recent academic movements such as the fair data principles (findable, accessible, interoperable, reusable) for open science (wilkinson et al. 2016) and international initiatives to mobilise biodiversity data – e.g., gbif (https://gbif.org), the catalogue of life (https://www.catalogueoflife.org), dissco (https://www.dissco.eu) – it became an increasing priority to spread the alpha taxonomy data contained within ejt articles throughout the ecosystem of dynamic, stable, free-to-use and interconnected platforms available on the web. a community-wide solution the promotion of taxonomy, systematics and collection-based research via scientific publishing is a vision that ejt shares with the consortium of european taxonomic facilities (cetaf), who officially endorsed the journal as its flagship title in 2016. ejt therefore aims to provide the taxonomic community with all of the modern interactive web-based facilities expected of a high-level, high-impact journal. an additional motivation behind the creation of an international, cross-institutional journal was to enable its members to collectively tackle the challenges of the digital transition. indeed, the unprecedented technological advances associated with 21st-century scientific publishing have given rise to complex strategic and technical issues related to the visibility, access, format and financial structure of academic journals, especially publicly funded titles (bénichou et al. 2011, 2012; côtez et al. 2018). xml has proven itself as an indispensable format. the availability of a “machine readable format (for example xml)” now features in the mandatory quality criteria for plan s-compliant journals (coalition s 2019: art. 9.2). however, this is a veritable technical barrier for many smaller open access publishers, especially those who do not impose author processing charges (apcs). given the fact that ejt is representative of typical natural history journals and their technical set-up, publishing on behalf of public-sector authors, the ejt-xml project reflects the operational and sociological constraints experienced by many other independent journals. thus, in line with ejt’s role as incubator on behalf of its members, the solution had to be transposable to other institutional journals managed within the consortium’s network, and even further afield into the wider community of taxonomic publishing. we here report on the process of defining a new postpublishing xml workflow and the associated formatting that enables optimised encoding of the published content. european journal of taxonomy 586: 1–22 (2019) 4 approach, tools and partners project analysis the ejt steering committee, consisting of the general directors of the members of the ejt consortium, asked for an assessment of the potential ways to achieve a semantically enhanced xml version of ejt’s publications. in 2015, the naturalis biodiversity centre (an ejt consortium member) was tasked with investigating the different methods of obtaining and utilizing an xml version of ejt content. the report submitted by naturalis in march 2017 concluded that, while the field of taxonomy provided exciting opportunities for interlinking data to the wider biodiversity community, elaborating an in-house workflow operated by the ejt production team was not advisable. given the journal’s independent position (i.e., entirely managed, produced and published directly by the consortium members), the development of a tailor-made xml production platform was considered too costly, and performing manual encoding would be too time-consuming and technical for the desk editors to undertake. the only existing platform that offered integrated services for xml encoding of taxonomy literature was the pensoft arpha journal publishing system (see penev et al. 2019). ejt felt that it was important to investigate an alternative to this unique solution which, although highly performant, represented issues of workflow rigidity, cost and commercial monopoly. plazi and goldengate based on its findings, the naturalis team carried out a proof-of-concept using goldengate to apply mark-up to ejt articles published in pdf (fig. 1). goldengate is a semi-automatic retro-conversion tool for encoding taxonomic literature (sautter et al. 2007). the open-source program was developed by the swiss ngo plazi (http://plazi.org) to support and promote the interoperability of taxonomic treatments with other relevant cyber infrastructure components (name servers, biodiversity resources, etc.). it was specifically conceived to process “born-digital” pdfs (as opposed to digitised paper publications) by recovering the text from the rendering instructions embedded in the file and discovering structural elements such as figures, tables and textual sub-sections e.g., taxonomic treatments. once identified, these elements are parsed using the tags defined in the taxpub extension to the jats dtd (catapano 2010), which in turn allows the text to be annotated with scalable links to external sources. the process can be highly automated by developing a journal-specific template, chaining the various steps in the conversion process and using batch processing. the ejt-plazi workflow the solution of a collaboration between ejt and plazi was chosen owing to the latter’s valuable expertise in domain-specific mark-up for taxonomic literature (agosti et al. 2019a), as well as for their established distribution network for the harvested content; both essential points for ejt. furthermore, plazi’s data warehouse treatment bank (https://treatmentbank.org), combined with the group’s active status as a trusted data provider for the global biodiversity infrastructure facility (gbif) and the biodiversity literature repository (blr; https://zenodo.org/communities/biosyslit/; agosti 2019b) offered further scope for the distribution of the taxonomic treatments, images, specimens and other data published by ejt. using goldengate and taxpub, the taxonomic treatments and specimen data featured within ejt articles are converted into darwin core (dwc) archives (wieczorek et al. 2012), the biodiversity informatics data standard developed by the biodiversity information standards (tdwg), which is the preferred format for publishing data to the gbif network. once converted, data relating to the treatments and specimens published in ejt is accessible via treatment bank and gbif within a few days of publication, at no extra cost or effort for the author. the figures and captions, along with a full copy of the article, are also available via the biodiversity literature repository. all data harvested from chester c. et al., semantic enhancement of specimen citations 5 an article is explicitly linked to the original publication using full original citations and a doi. this means that the sub-article elements are integrated into a much larger network of biodiversity data, which helps to ensure their preservation but also maximises the reach of the article (fig. 2). semantic enhancement of specimen citations a goldmine of data the details given by authors about the specimens studied in their research – especially for the type material used in nomenclatural acts – are extremely rich and highly structured. within the realm of biodiversity informatics, the physical specimens studied in a systematic account can be used as a valuable anchor to unambiguously identify taxa, analyse the associated collecting data and track any research based on these vectors. fig. 1. the goldengate interface. using pattern recognition and natural language processing techniques, the article structure is encoded allowing the extraction of the taxon treatment sections. furthermore, the taxpub extension allows for fine-grain mark-up of sub-article elements such as the scientific names (a) and specimen citations (b). a b european journal of taxonomy 586: 1–22 (2019) 6 fig. 2. the ejt-plazi workflow. plazi processes the pdf of an ejt publication to extract the subarticle elements and distribute them to biodiversity aggregators. taxonomic treatments, images, tables, scientific names and even the fine-grain specimen data are semantically enhanced and available on a variety of platforms. fig. 3. a taxonomic treatment extracted from the european journal of taxonomy and displayed on the plazi treatment bank. in the right-hand sidebar, the parsing performed on the specimen citations has been used to generate graphic charts of the material studied. in the inset, a map has been generated via googlemaps by plotting the extracted geocoordinates. chester c. et al., semantic enhancement of specimen citations 7 plazi demonstrated how the information relating to the collection and preservation of physical specimens can be parsed to a fine degree and used to generate darwin core archives of occurrence records (wieczorek et al. 2012). once available in a machine-readable format, these records can be used to search, cite and track specimens through a multitude of criteria (by locality, date, collector, repository etc.) (fig. 3), providing researchers and collection managers with sophisticated ways to query data sets and carry out their work (e.g., nicolson & tucker 2017; miller 2019). structuring text for automatic parsing during the first test phase, ejt submitted to plazi a sample of 30 articles representing the diversity of disciplines (entomology, zoology, palaeontology, botany) and article types (species descriptions, revisions, check-lists...) published by the journal. these publications were processed within goldengate using algorithms to automatically mark-up the article structure and the material sections. certain highly structured elements such as dates and geo-coordinates were correctly identified and encoded. however, the wide range of methods, punctuation and vocabularies used to describe the specimens examined resulted in a considerable amount of errors that were time-consuming to correct manually. difficulties mainly stemmed from matching occurrence data to the correct specimen and delimiting unstructured data, e.g., distinguishing a habitat description from a locality within a text string. thus it became clear that a standardised format for the presentation of specimen data was required in order to facilitate fine-grain harvesting of details related to the physical material (specimens) studied. if correctly formatted, these data could be harvested, converted into occurrence records, then integrated into the wider infrastructure of biodiversity informatics (dikow 2019). moreover, by establishing standardised formatting that different authors and journals could follow, we also hoped to enable text mining for other taxonomy journals who publish in pdf. defining the ejt format the specimen citations contained within the test set of articles, as well as those found in further articles published by ejt and other taxonomic journals (zoosystema, adansonia, geodiversitas, zookeys, zootaxa, the zoological journal of the linnean society, the botanical journal of the linnean society, phytokeys, phytotaxa), were analysed to establish the types of data most commonly presented. once the specimen citations had been broken down into a list of highly recurrent fields, these fields were mapped to dwc terms (http://rs.tdwg.org/dwc/terms/). this permitted the creation of a flexible template for specimen citations, designed to fit a broad array of specimen data, which was tested on a wide sample of citations and refined according to feedback from plazi, the ejt production staff and scientific editors, as well as several active ejt authors. the resulting template and formatting guidelines are described in detail hereafter as the material citations formatting guidelines. european journal of taxonomy 586: 1–22 (2019) 8 material citations formatting guidelines in accordance with the european journal of taxonomy’s fair data & open science policy (available from https://europeanjournaloftaxonomy.eu), the formatting guide for zoological and botanical specimen citations is presented below. authors are encouraged to prepare their manuscripts according to this model prior to submission, but they will also be given an opportunity to comply upon acceptance of the article. while ejt strongly recommends that authors adhere to the guidelines given below, the fine-grain formatting of the material citations is not compulsory; if an author decides not to comply or that the material is not appropriate, ejt will perform reduced formatting during production. in this case, the majority of the specimen data will not be tagged and converted into a machine-readable format; this means that the specimen-related information from the paper will not be included in major databases. only specimen data presented in the ‘materials examined’ section will be tagged and converted for distribution. at this time, any specimen data presented in a separate table or section of the paper cannot be linked back to the specimen citation to form a full occurrence record. general presentation 1. order each material citation is composed of diverse data fields (number of specimens, locality, date collected, etc.) that ejt identifies using darwin core (dwc) terms. to efficiently perform this, it is important to ensure that the different fields of a material citation are consistently presented in the same order throughout the article or, at the very least, within a taxon treatment. the preferred order for data fields differs for zoology and botany, and are as follows: country • specimen(s) (e.g., “1 ♂”); geographic / locality data (from largest to smallest); geographic coordinates; altitude / elevation / depth (using alt. / elev. / m a.s.l. etc.); date (format: dd mmm. yyyy, e.g., “16 jan. 1998”); collector (followed by “leg.”); other collecting data (e.g., micro habitat / host / method of collecting / “dna voucher specimen”/ “vial with detached elements”, etc.); barcodes/identifiers (e.g., “genbank: mg779236”); institution code and specimen code (e.g., “cbf 06023”). country • locality data (from least to most specific); geographic coordinates; altitude / elevation / depth (using alt. / elev. / m a.s.l. etc.); other collecting data (e.g., habitat / substrate / method of collecting / “herbarium specimen” / “dna voucher specimen”); date (format: dd mmm. yyyy, e.g., “16 jan. 1998”); phenology (e.g., fl / fr); collection number (collector’s name + number in italics); determinator (followed by “det.”); repository[identifier in the repository] (format: herbarium acronym[identifier]; e.g., “ny[133679], ny[3774223], ny[3774224], l[l.1264510]!, plp[plp-0012346], br[awh10018408]); genetic identifiers (e.g., “genbank no.: mg779236”). zoology botany details on how to format each data field are provided in the ‘data fields’ section. chester c. et al., semantic enhancement of specimen citations 9 2. punctuation a bullet point “•” (unicode: hex 2022, decimal 8226) is used to signify the beginning of a material citation. in microsoft word, the following keyboard shortcuts can be used to obtain a bullet point: • for mac: alt + 8 (qwerty keyboard) / alt + shift + full stop (azerty) • for windows: alt + 0149 on the numeric keypad within a citation, a semicolon “;” delimits each different field. semicolons should not be used elsewhere in a material citation. a single field can be composed of several details, which are separated by commas (e.g., the details region, area and town for the ‘locality’ field). in the following example, the ‘locality’ field is composed of two details: province (“eastern cape province”) and town (“cradock”): south africa • ♂; eastern cape province, cradock; 30°19′ s, 25°39′ e; aug.–oct. 1985; museum staff leg.; pitfall trap; nmba 1170. ohelopapa flexilis (setch.) f.rousseau, martin-lescanne, payri & l.le gall comb. nov. fig. 3, table 2 basionym laurencia flexilis setch., university of california publications in botany 12: 101, pl. 19, figs 1–6 (setchell 1926). type: france • french polynesia, tahiti; reef at tahara mountain; setchell w.a., setchell c.b. & parks h.e. 5246; holotype: jeps[uc261333]. original material is also preserved in sap fide masuda et al. (1999, 2006). afroriccardia comosa (steph.) reeb & gradst. comb. nov. figs 3–4 aneura comosa steph., botanical gazette 15 (11): 281 (stephani 1890). – riccardia comosa (steph.) e.w jones, transactions of the british bryological society 3: 74 (jones 1956, nom. inval.). – type: france • la réunion; 1889; rodriguez s.n.; holotype: g[g00045027]!; isotype: pc[pc0103522]! 3. type material zoology type material should be presented in separate paragraphs with relevant subheadings (holotype, paratypes, etc.). botany basionyms & synonyms in botanical articles, the type material of basionyms and homotypic synonyms is presented in the same paragraph as the relative scientific name and bibliographic reference (just under the treatment heading), preceded by the mention “type:” in bold. all heterotypic synonyms under the recognised name are cited accordingly with their basionyms. this presentation should be used regardless of whether the specimen has been examined (indicated by an exclamation mark in this context) or not. european journal of taxonomy 586: 1–22 (2019) 10 lectotypification example of material designated as lectotype in a previous publication vs material designated within the paper: type peru • region pasco, prov. oxapampa, dist. palcazu, parque nacional yanachaga-chemillén, sector paujil; 150 m from entrance to las cavernas on trail from paujil; 10°20′40″ s, 75°15′1″ w; alt. 432 m; 25 feb. 2016; moonlight & daza 318; holotype: mol; isotypes: e, mo, usm. new taxa type material for new taxa, which do not have an existing bibliographic reference (e.g., it is proposed for the first time in the publication), should be presented as follows, under the heading “type” or “type material”: paratypes south africa • 1 ♂; same data as for holotype • 1 ♀; limpopo province, “pietersburg” [now polokwane]; 23°54′ s, 27°23′ e; 4 nov. 1916; c.a. thompson leg.; tm 13375 (formerly 2217). if a material citation is identical to another with only minor differences, the exceptions should be listed after the mention of repeated data e.g.: south africa • 1 ♂; eastern cape province, middelburg; 31°49′ s, 25°00′ e; 8 sep. 1995; m. de jager leg.; pit traps; nca 95/394 • 1 ♂; same collection data as for preceding; 3 sep. 1995; nca 95/243 • 2 ♂♂; same collection data as for preceding; 8 may 1999; associated with termites; nca 91/1051. zoology botany india – arunachal-pradesh • dirang dzong; 8 aug. 1938; ward 14055; bm • senge dzong; 18 aug. 1938; ward 14091; bm. – mizoram • hmuifang; jul. 1926; parry 45 • ibid.; jul. 1926; parry 46; k • ibid.; jul. 1926; parry 47; k • lungleh; 1 sep. 1931; wenger 320; k. 4. repetitive data repetitive data can be indicated with terms such as “same data as for holotype”, “same data as for preceding”, “same locality”, “ibid.”, etc., as long as the method used is consistent throughout the paper. begonia acetosella craib figs 6–7 bulletin of miscellaneous information, kew 1912: 153 (craib 1912). – type: thailand • chiengmai, doi sootep; 21 mar. 1909; 18°50′ n, 98°54′ e; a.f.g. kerr 557; lectotype: k[k000761199], designated by tebbit 2003a, sheet 2 k[k000761201]; isolectotype: b. begonia tetragona irmsch., mitteilungen aus dem institut für allgemeine botanik in hamburg 10: 515 (irmscher 1939). – type: china • yunnan, mengtze; henry 10737a; lectotype: b[b100238046], here designated; isolectotype: e[e00315022]. chester c. et al., semantic enhancement of specimen citations 11 5. ‘missing’ elements zoology it is not necessary to include information such as “no date” or “no locality data”; list only the elements that are available. botany all commonly used abbreviations can be used where appropriate (“s.n.”/ “s.loc.”/ “s.d.”, “s.coll” etc.). type ivory coast • régio du bas-sassandra, km 41 sassandra-san pedro road; 16 nov. 1968; fl. bud, fr.; breteler 6052; holotype: wag; isotype: br, k, mo n.v., pre n.v., w n.v. basionym laurencia crustiformans mcdermaid, phycologia 28: 352, figs 2–8 (mcdermaid 1989). type: usa • hawaii, oahu, lualualei beach park; 3 jul. 1988; k.j. mcdermaid s.n.; holotype: bish[km 2050] (abbott 1999: 384). lectotype (here designated) montenegro • 1 specimen; “popovo höhle bei njegus” [popovo cave near njegus]; [43.5291° n, 19.2074° e]; 30 may 1903; sturany leg.; nhmw 38260a. 6. label citations double quotation marks (“ ”) are used to represent label citations that cannot be reliably interpreted and formatted as recommended in these guidelines. this data will simply be parsed as a verbatim citation. ejt recommends including photos of labels as figures if they contain data that cannot be standardised. only quotation marks should be used to present verbatim label data and they should not appear elsewhere in a material citation. 7. author interpretation use square brackets [ ] to distinguish data that has been interpreted from a label e.g., coordinates interpreted from a locality, or translations of foreign text: european journal of taxonomy 586: 1–22 (2019) 12 3. locality the locality data is listed from least to most specific, using commas to divide each detail. it is recommended to employ the english name in current usage where possible. if a different system is used, e.g., variant spellings or archaic names from label transcriptions, these should preferably be identified using quotes, with their current names given in square brackets. south africa • 1 ♂; eastern cape province, middelburg; 31°49′ s, 25°00′ e; 8 sep. 1995; m. de jager leg.; pit traps; nca 95/394 • 2 ♀♀; same collection data as for preceding; 3 sep. 1995; nca 95/243 • 3 ♂♂; same collection data as for preceding; 8 may 1991; associated with termites; nca 91/1051. 2. specimen count (zoology) this field can contain several indications about the specimen(s) cited: number, nature (e.g., specimen, juv., shell, exuviae), sex and type status. all subsequent data in the same citation will be applied to the specimen(s) presented. thailand • 3 shells, same data as for preceding; hnhm 97479 • 16 specimens (preserved in ethanol); same data as for preceding; uf 76457. data fields the different data fields of a material citation that ejt identifies for conversion and diffusion are explained below, along with the format required to achieve maximum output and accuracy. 1. country/ water body the citations must be listed by either country or water body (e.g., ocean/sea), using a separate paragraph for each new zone. the country or water body is presented in capital letters. if the material is organised by region, use the following format: madagascar – mahajanga province • betsiboka region, forêt de kasijy, kelifely; nov. 1974; fl; morat 4700; p, tan • boeny region, majunga; 30 mar. 1912; fr; afzelius 259; p • soalala, réserve naturelle intégrale de namoroka, ca 40 km s of soalala; 3 feb. 2000; fr; davis, rakotonasolo & wilkin 2529; br, k, tan • majunga; dunes; feb. 1915; fl, fr; perrier de la bâthie 3504; p • ambongo; 17 feb. 1841; fr; pervillé 680; p. china – guangdong province • 1 ♀; tianma, xinhui; 24 apr. 1956; z.b. zhou leg.; scau • 4 ♀♀; xinhui; nov. 1956; scau • 1 ♀; nanhai; 14 oct. 1955; l.b. huo leg.; scau. – guangxi province • 1 ♂; longsheng; 6 apr. 1974; y.l. luo leg.; scau • 1 ♂; huaan; 26 apr. 1982; y.q. tang leg.; scau. – hainan province • 1 ♂, 3 ♀♀; diaoluoshan; jul. 1995; z.q. peng leg.; scau • 1 ♂; diaoluoshan; 8 may 2005; x.m. wang leg.; scau • 1 ♂; dongfang; 27 nov. 1997; z.q. peng leg.; scau. myanmar • 1 shell, holotype of p. ponsonbyi (d = 17.8 mm); “burmah”, hlindet; nhmuk 1913.3.14.9. montenegro • “popovo höhle bei njegus” [popovo cave near njegus]; 43.5291° n, 19.2074° e; 30 may 1903; sturany leg.; nhmw 38260a. chester c. et al., semantic enhancement of specimen citations 13 4. geographic coordinates various formats are accepted but it is important to include the degree symbol (°) as well as the direction (n/e/s/w), which distinguishes the data as a geographic coordinate: • degrees minutes seconds: 40°26′46″ n, 79°58′56″ w • degrees decimal minutes: 40°26.767′ n, 79°58.933′ w • decimal degrees: 40.446° n, 79.982° w geographic coordinates should be presented to a maximum of 5 decimal places. latitude and longitude are separated with a comma. latitude is cited first, then longitude. 5. altitude/elevation/depth this type of measurement should be explicit in the material citations, e.g.: • altitude: alt. 489 m or 547 m a.s.l. • depth: depth 20 m 6. collection date format: d(d) mmm. yyyy date ranges should be shown with an n-dash, e.g., jan.–may 2018 / 5 feb.–6 apr. 2016 / 14 dec. 2008–3 feb. 2009 / 1950–1953. 7. collector (zoology) the name(s) of the collector(s) should always be followed by “leg.”; for expeditions, “exped.” can be used, e.g., “mnhn exped.” 8. collector and collection number (botany) the collector’s name and field number are cited together in italics. for botanical disciplines that do not catalogue specimens on sheets (e.g., algae, diatoms), we ask that authors use “collected by: x”, because the term “leg.” does not have the same signification across all botanical fields. 9. additional data ideally, the data fields identified above should be listed before any other collection data. if a different order is used, it is important to be as consistent as possible throughout the paper, or at least within a single treatment. semicolons may be used to separate any additional data into appropriate fields, e.g.: sweden • ♂; halland, halmstad; gårdshult, buskastycket; 56.41° n, 13.91° e; 3–25 may 2005; swedish malaise trap project leg.; trap 35; collecting event: 1786; malaise trap; hay meadow; nhrs cec1405 • ♂; öland, mörbylånga, llevi; 56.61° n, 16.60° e; 8 apr.–9 may 2016; m. & c. jaschhof leg.; malaise trap; herb-rich meadow near forest; sdei cec1429. european journal of taxonomy 586: 1–22 (2019) 14 holotype myanmar • 1 shell, holotype of p. dextrorse (d = 16 mm); tenasserium valley; nhmuk 1906.2.2.144. other material thailand • 2 ♂♂, 1 ♂ (gonopods lost); prachuap khiri khan province, kui buri district, hat kham subdistrict, ban yan sue; 12°03′12″ n, 99°37′52″ e; ca 147 m a.s.l.; 31 aug. 2007; asru members leg.; cumz. additional data can also be given in the appropriate field between brackets, e.g.: 10. associated sequences specimen accession numbers, barcodes and dna accessions should be identified as such, e.g., “genbank: u34853.1”, “accession no.: 5587453”. brazil • ♂; pernambuco, off recife, approximately 20 nautical miles from the coast; rec i, dredge 4; 8°08′51.5″ s, 34°34′08.0″ w; 65 m depth; 7 feb. 2018; agglomerations of sponges, rhodoliths and calcareous algae; genbank 16s gene: mk918616; moufpe 19470. 11. repository data the repository data field should be composed of an institution acronym followed by a specimen code/ catalogue number/barcode (where available). zoology institution acronym acronyms for repositories must feature in a distinct list in the materials and methods section, under a heading called “repositories”, “institutional acronyms” or “institutional abbreviations”. institution codes must follow grscicoll (https://gbif.org/grscicoll) where possible. specimen code where a specimen code is available, it should be explicit which specimen it refers to. this guarantees unambiguous interpretation, both by human readers and upon encoding. for example, in the citation below, we cannot distinguish which specimens are catalogued under which code: namibia • 2 ♂♂, 4 imm.; grootfontein, nosib cave; 8 feb. 1995; segl leg.; samc b7732, b8870. namibia • 2 ♂♂; grootfontein, nosib cave; 8 feb. 1995; segl leg.; samc b7732 • 4 imm.; same data as for preceding; samc b8870. this citation should be presented as follows: use the word “to” instead of a hyphen or an n-dash in order to show a range of specimen numbers. e.g., “nhmuk 213584 to 213595”. chester c. et al., semantic enhancement of specimen citations 15 in case of type material, use the same convention for all fields, except for repository[identifier in the repository]: introduce the field with the nature of the type: holotype, lectotype... for example: “lectotype: bm[bm000043986], here designated; isolectotypes: p, k, wag.” “holotype: br[br0000013468675]!” “holotype: ny[133679]!; isotypes: ny[3774223][3774224], l[l.1264510]!” “holotype: bm[slide bm001166601] illustrated in figs 3–62, the valve representing the holotype is illustrated in fig. 9; isotype: brm[slide zu10/14], friedrich hustedt diatom collection.” discussion first results by respecting the standardised format explained in this paper, the specimens cited in ejt publications are quickly and efficiently converted into dwc archives by plazi using a semi-automatic mark-up workflow developed within goldengate. since the project was launched in early 2018, ejt and plazi have contributed 2155 treatments and 7888 materials citations to major aggregators of biodiversity data, as well as 2288 figures and 7324 bibliographic references: all directly linked to the original publication. identifier in the repository each identifier (barcode and/or other catalogue numbers) should be cited exactly as it is registered in the repository. each individual code is presented within square brackets immediately after the herbarium acronym. e.g.: material and methods repositories acronyms of herbaria follow index herbariorum (http://sweetgum.nybg.org/science/ih/), except for the following repository: uant = university of antwerp, belgium botany repository acronyms of herbaria must follow index herbariorum (http://sweetgum.nybg.org/science/ih) and a phrase to this effect will be included in the ‘materials and methods’ section under the heading ‘repositories’. any acronyms used for repositories that do not feature in the index herbariorum must also be given here, e.g.: usa • hawaii, hawaii island, isaac hale beach park; alt. 250 m; jan. 2008; a. kurihara s.n. sish[ars03327][766726]. france – la réunion • “sur les mousses, source pétrifiante de hell-bourge”; g. de l’isle 220; pc[pc0716023] • “sous piton de la fournaise, le long gr2, réserve de mare longue”; 21°20′30″ s, 55°44′30″ e; 175–300 m a.s.l.; vojko 9435b; egr • g de l’isle 570bis; pc[pc0716024], g[g00264057] • rodriques s.n.; g[g-00264058] • “plaine des palmistes”; s.coll. 56; pc[pc0716026]. european journal of taxonomy 586: 1–22 (2019) 16 spot checks carried out on the occurrence records obtained from the encoded specimen citations reveal that the use of the standard formatting greatly improves the precision and quantity of harvested data. eventual discrepancies in the distributed data compared to the original data (stemming from misinterpretation by the parsing algorithms) can be easily reported (“contact” details available directly of the derived web pages) and corrected (updates performed by plazi and automatically propagated throughout the network). workflow evaluation the most evident drawback of the goldengate workflow is the approach of retro-conversion, whereby pattern recognition and natural language processing techniques are used to decipher the pdf postpublication. this method is seldom chosen by publishers setting up digital workflows owing to the difficulties connected to a computer interpreting the text correctly. however, for the ejt “born-digital” pdf publications, the text is recovered from the rendering instructions embedded in the pdf, which facilitates the task. in addition, a template has been built to automate the conversion process, right up to the fine granularity of material citations. using plazi’s expertise and the ejt standardised formatting as much as possible, we have not only improved the quality and quantity of the data harvested from the articles, but also the speed with which plazi can perform encoding. finding a consensus for best practices the process of establishing a standard for citing specimens within taxonomic publications exposed the variety of methods currently used and accepted by publishers. while it is important for researchers to benefit from flexibility in journals to present their work in a form that befits the nuances of their subject matter, it is clear that the taxonomic community needs to invest time and effort into reaching a consensus for standardising data and sharing best practices. reaching a consensus on working methods that call on community-wide standards, such as those suggested in the material citations formatting guidelines, could have a considerable impact on issues that contribute to the “taxonomic impediment” but the solutions – even those easily attainable by all – will only be effective through adequate buy-in and through our willingness to change. for example, the appropriate citation of taxonomic works, acts and authorities has long been evoked as a subject that could improve the recognition of taxonomy researchers (ebach et al. 2011), yet there is still much confusion about which course of action should be followed, resulting in anarchical practices across different journals and even from one article to another (bénichou et al. 2018). this is a purely sociological problem, which was directly addressed in the recent paper “consortium of european taxonomic facilities (cetaf) best practices in electronic publishing in taxonomy” (bénichou et al. 2018). an authoritative response is therefore available; the question now remains whether the community will assimilate these best practices. best practices and the interoperability of biodiversity data are recurrent themes in cetaf meetings, workgroups and publications (e.g., keklikoglou et al. 2019), and other leading publishers of taxonomy are equally concerned with these issues, e.g., pensoft (penev et al. 2019), who also recommends that authors submitting to the journal zookeys apply the format developed by ejt and described in this paper (see: https://zookeys.pensoft.net/about#materialsexaminedformattingguidelines). another pertinent example of the need for controlled vocabularies and well-maintained registers is the citation of digital bio-collections in taxonomical work. over the past decade, a huge effort has been made by natural history institutions to digitize their collections. this initiative demands serious resources, but also consensus on how to identify, retrieve and cite specimens once they are digitised. once again, the cetaf has published directives on recommended best practices (güntsch et al. 2017) for the implementation of globally unique identifiers (guids), but bio-collection managers, informatics chester c. et al., semantic enhancement of specimen citations 17 communities, researchers and publishers still need to agree on which type of identifier to use, when to assign it and how to cite it, in order for real progress to be made (guralnick et al. 2015). the citation of specimen guids has not been covered within the present guidelines for this very reason but will be addressed in a subsequent paper. producing well-formatted occurrence records and handling associated guids may take extra effort from researchers and from editors during manuscript preparation, but they constitute a solid foundation for subsequent study and will greatly contribute to the creation of an interlinked network of robust data sources. the standardised documentation of physical specimens and collection events would facilitate bio-collection management and, if stricter practices for the citation of material were consistently respected by researchers and publishers, improve interpretation by both humans and machines. using digital identifiers to assess impact the traceable nature of linked data could potentially offer alternative metrics for evaluating research impact (mcdade et al. 2011). however, in order to achieve metrics that are sufficiently representative in terms of quality and quantity to make reliable statements, it is urgent to reach a consensus in the use of controlled vocabularies and identifiers, starting with unique ids for physical specimens, collections and institutions. this would allow specimens to be tracked from the moment they are collected to their deposition in a repository and throughout any subsequent movements, permitting the identification and retrieval of a given physical specimen despite eventual changes in local collection management. with such a tagging system in place, it would also be possible to infer the number of times a specimen has been used in research papers by tracking its identifier through every publication that cites it, or to imagine using guids to gather statistics about the representation of museum collections across different fields of research (guralnick et al. 2015; nicolson et al. 2019). however, for as long as there is no standardised approach to the citation of biological specimens, any data obtained on a large scale will have a limited power. as an institutional publisher, a specific area of interest for ejt was the identification of institution codes within research articles, indicating where specimens are held. however, the lack of an agreed method for identifying a given institution, in fields other than botany, means that these results cannot currently be exploited. for example, within the test batch of converted ejt articles, the natural history museum of london was identified by several acronyms: nhm, nhmuk and bmnh. without a recognised register of official acronyms for zoological collections, as is the case with botanical collections and the institutional register index herbariorum, it would be difficult to make any statements about the collections represented by the journal’s publications. if we hope to align our efforts and build a truly seamless global infrastructure for biodiversity knowledge, the institutions dedicated to the curation, documentation and preservation of the primary research matter must take a central role in any proposed scheme. the recommendation in this paper is to follow index herbariorum (for botany) and for all other fields the global register of scientific collections (grscicoll – a compendium of earlier directories recently merged together and re-launched by gbif), but this resource will only be as strong as the will of its contributors, i.e., the community must make the effort to contribute, develop and utilise the registry. we therefore recommend that institutions take an active role in enriching and controlling this resource, which offers institutions and collections a foothold in the developing landscape of interlinked data, specimens, publications and institutions. while awaiting the community-curation functionality to be activated by gbif (intended for late 2019), management should consult the website (https://www.gbif.org/grscicoll) and reflect upon how their institution and collections should be presented. this includes consensus on and communication about a unique and unambiguous acronym to represent their institution (and collections) that should be consistently used as an identifier in all scholarly output. european journal of taxonomy 586: 1–22 (2019) 18 conclusion by contributing data to aggregators such as gbif and blr, the ejt-plazi workflow not only improves access to and citation of taxonomic literature, but also means that the specimens studied and their linked taxonomic information are rapidly made available to the wider scientific community. another huge benefit of this workflow is that the original data source is consistently cited and linked throughout all subsequent representations. the ejt-xml project was presented to the international community of biodiversity scientists at the biodiversity next meeting in leiden, the netherlands in october 2019 (bénichou et al. 2019), where it was received with enthusiasm. we hope that this paper will now serve as a reference for progress in the field of biodiversity informatics, as well as for ejt authors to prepare their manuscripts. the standard formatting for material citations presented in this paper will be adapted and implemented by four other titles produced by the muséum national d’histoire naturelle (paris), and additional journals produced by ejt member institutions are expected to follow suit. we hope that any taxonomic journal or author who would like to contribute to the dynamic exchange of biodiversity data might consider adopting a similar approach to formatting and inform plazi of their efforts. within the scope of this project, ejt has concentrated on standardising the presentation of specimen data to facilitate semantic enhancement and text mining. the next step will be to work on integrating persistent identifiers for sub-article elements (e.g., treatments), specimens and institutions, with the aim of further improving the accessibility of its publications and establishing alternative metrics for taxonomy. ejt intends to take an active role in addressing these issues (cetaf e-publishing workgroup; biodiversity next 2019; tdwg working groups) and will continue to collaborate closely with the cetaf to define best practices across european natural science institutions. acknowledgements the authors would like to thank daphné duin and jeremy miller (naturalis biodiversity centre) for the initial project analysis carried out on behalf of ejt, as well as natacha beau (meise botanic garden), alejandro quintanar sánchez (real jardín botánico de madrid), emmanuel côtez, anne mabille and marc pignal from the muséum national d’histoire naturelle, paris, for their valuable help in defining aspects of the material citations formatting guidelines. we are also grateful for the support from the team of scientific editors and desk editors at the european journal of taxonomy regarding the implementation of the project described in this paper. all projects carried out by the european journal of taxonomy are made possible by the funding granted to the journal by the consortium of member institutions: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. ejt would also like to thank the cetaf for their continued support and endorsement of the journal. the authors would like to thank lyubomir penov and michelle price for reviewing the present article and improving the original manuscript with their comments, suggestions and corrections. references agosti d. & egloff w. 2009. taxonomic information exchange and copyright: the plazi approach. bmc research notes 2: 53. https://doi.org/10.1186/1756-0500-2-53 chester c. et al., semantic enhancement of specimen citations 19 agosti d., klingenberg c., sautter g., johnson n., stephenson c. & catapano t. 2007. why not let the computer save you time by reading 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(asteraceae). an apterous viviparous female of uroleucon (lambersius) robinsoni sp. nov. from oregon and apterous and alate viviparous females of uroleucon (lambersius) grindeliae sp. nov. from colorado are described and illustrated. taxonomical notes of the new and other grindelia-feeding taxa of uroleucon are given and an updated key to the apterae of the grindelia-feeding species of uroleucon in the world is provided. keywords. macrosiphini, uroleucon, lambersius, new taxa, north america. barjadze s. & kanturski m. 2020. two new species of the aphid genus uroleucon (hemiptera: aphididae) living on grindelia in the usa. european journal of taxonomy 729: 42–53. https://doi.org/10.5852/ejt.2020.729.1189 introduction the genus uroleucon mordvilko, 1914, which comprises 239 aphid species in the world, is one of the most speciose genera in the tribe macrosiphini wilson, 1910 (favret 2020). the genus has six subgenera: belochilum börner, 1932 (one species), divium pashtshenko, 2000 (one species), lambersius olive, 1965 (53 species), satula olive, 1963 (one species), uroleucon mordvilko, 1914 (122 species) and uromelan mordvilko, 1914 (61 species) (favret 2020). all of the species live on herbaceous plants that belong to the families asteraceae (more than 160 species) and campanulaceae (11 species) without any host alternation; only nine species use host plants from other plant families (blackman 2010; blackman & eastop 2020). the coloration of species of uroleucon varies from green to blackish brown (heie 1995; blackman & eastop 2020). the genus is generally characterized by well-developed antennal tubercles with diverging inner sides, the presence of secondary rhinaria on the basal part of antennal segment iii in apterae, often a very long processus terminalis, ultimate rostral segments with blunt apices, long https://doi.org/10.5852/ejt.2020.729.1189 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:5d9a5938-4858-4f8d-a50d-6e2cf5623e69 mailto:mariusz.kanturski%40us.edu.pl?subject= mailto:shalva.barjadze%40yahoo.com?subject= http://zoobank.org/urn:lsid:zoobank.org:author:63abe1b2-8a56-42c8-bc34-1119d3a2ecbc http://zoobank.org/urn:lsid:zoobank.org:author:78c290a3-d07b-4af9-9358-ed8c05a702bf https://doi.org/10.5852/ejt.2020.729.1189 barjadze s. & kanturski m., new american uroleucon 43 and cylindrical reticulated siphunculi, a finger-shaped cauda with a pointed apex and five setae (a few species have three or four setae) on the first tarsal segments. in many species, the abdominal dorsal setae are located on pigmented scleroites (blackman & eastop 2020). during an examination of the aphid collection in the natural history museum of london (bmnh), slides with two undescribed species of lambersius, which had been sampled on grindelia hirsutula hook. & arn. and g. squarrosa (pursh) dunal (asteraceae) in oregon and colorado, respectively, were recognized. in blackman & eastop (2006: 496), these species were included in the key to apterae aphids on grindelia willd. as “uroleucon (lambersius) sp. (on g. stricta dc., oregon)” and “uroleucon (lambersius) sp. (on g. squarrosa, colorado)”. the aim of this paper is to describe these two grindeliafeeding species of lambersius from the usa. material and methods the specimens were examined using a leica dm 3000 led light microscope and photographed using a leica mc 190 hd camera. the measurements were taken according to ilharco & van harten (1987). the measurements are given in millimeters. the material examined for each species is given in the review of the species. the host plant names are given according to the most recent edition of the plant list (2013). abbreviations for morphological terms used in the text ant = antennae or their lengths ant i-vi = antennal segments i, ii, iii, iv, v and vi or their lengths (ratios between the antennal segments are simply given as, e.g., ‘vi: iii’) base = basal part of last antennal segment or its length bd iii = basal articular diameter of ant iii bl = body length (from the anterior border of the head to the end of the cauda) iii femur = hind femur or its length hw = greatest head width across the compound eyes ht ii = second segment of the hind tarsus or its length ls ant iii = length of longest setae of ant iii pt = processus terminalis of the last antennal segment or its length siph = siphunculus or its length siph w = maximum width of the siphunculus iii tibia = hind tibia or its length and urs the ultimate segments of the rostrum (iv + v) or their lengths abbreviations for depositories for the type material nhmuk = natural history museum, london, uk dzus = hemiptera collection of the department of zoology, university of silesia, katowice, poland european journal of taxonomy 729: 42–53 (2020) 44 results taxonomy class insecta linnaeus, 1758 order hemiptera linnaeus, 1758 family aphididae latreille, 1802 genus uroleucon mordvilko, 1914 uroleucon (lambersius) robinsoni sp. nov. urn:lsid:zoobank.org:act:c7c38dd0-01c5-4bb4-9938-1d3cd3d8a81a figs 1–2, table 1 diagnosis apterous viviparous females of grindelia-feeding lambersius have the following combinations of characters: pt 4.66–6.50 × base; ant iii with 12–21 secondary rhinaria (fig. 2a–b); urs 1.06– fig. 1. new species of uroleucon mordvilko, 1914 feeding on grindelia willd. a. uroleucon (lambersius) robinsoni sp. nov., holotype, apterous viviparous female (nhmuk 010121495) b. u. (l.) grindeliae sp. nov., holotype, apterous viviparous female (nhmuk 010121473) c. u. (l.) grindeliae sp. nov., paratype, alate viviparous female (nhmuk 010121477). http://zoobank.org/urn:lsid:zoobank.org:act:c7c38dd0-01c5-4bb4-9938-1d3cd3d8a81a barjadze s. & kanturski m., new american uroleucon 45 1.33 × base and 1.18–1.25 × ht ii with 7–9 short, fine, pointed accessory setae (fig. 2d); ht i with 5:5:5 setae; siph subcylindrical, slightly tapering without a flange (fig. 2e); its reticulated zone 0.17– 0.28 × siph. siph 2.30–2.68 × cauda and 0.31–0.41 × bl; scleroites on setal bases absent on abd i–v, while setae on abd vi–viii arise from barely visible, rounded scleroites; cauda narrow finger-shaped without constriction and with 7–9 setae (fig. 2f). etymology we are very pleased to name the new species in honor of a.g. robinson (university of manitoba, canada), who worked on the north american macrosiphini. material examined holotype usa • ♀ apterous viviparous; colorado, carson hole; 38°43′ n, 108°36′ w; alt. 2348 m; 16 sep. 1956; hottes and hille ris lambers leg.; grindelia squarrosa; nhmuk 010121495. fig. 2. uroleucon (lambersius) robinsoni sp. nov., holotype, apterous viviparous female (nhmuk 010121495). a. ant iii. b. secondary rhinaria on ant iii. c. ant vi. d. ultimate rostral (iii–v) segments. e. siphunculus. f. cauda. european journal of taxonomy 729: 42–53 (2020) 46 paratypes usa • 2 ♀♀ apterous viviparous; same collection data as for holotype; nhmuk 010121495 • 2 ♀♀ apterous viviparous; same collection data as for holotype; nhmuk 010121496 • 2 ♀♀ apterous viviparous; same collection data as for holotype; dzus 010121481 • 2 ♀♀ apterous viviparous; same collection data as for holotype; nhmuk 010121482. description apterous viviparous female (n = 9) color in life. unknown. pigmentation of cleared specimens on slide. body generally membranous, pale to yellow; ant brown with pale ant i–ii and basal part of ant iii; coxae and trochanters pale; femora yellow with light brown distal ends; tibiae brown with paler middle section or pale with only distal part darker; tarsi light brown; siph yellow with light brown to brown distal ends; cauda pale (fig. 1a). morphometric characters. hw 0.17–0.19 × ant. head with thick, rigid setae with blunt or narrowly capitate apices, 0.030–0.040 mm long. ant tubercles each with 2–3 setae. ant 0.83–1.04 × bl. ant iii with 12–21 rounded secondary rhinaria of different sizes, with sclerotized rims (fig. 2a–b), ant iv longer than ant v. pt 4.66–6.50 × base (fig. 2c). other antennal ratios: vi:iii 1.13–1.28, v:iii 0.61– 0.70, iv:iii 0.70–0.76, pt:iii 0.93–1.11, pt:iv 1.22–1.47 and pt:v 1.32–1.69. ant bearing medium length thick, rigid setae with blunt or narrowly capitate apices. ant iii setae 0.020–0.045 mm long, ls ant iii 0.84–1.13 × bd iii. ant i with 4–5, ant ii with 3–4, ant iii with 12–21, ant iv with 8–12, ant v with 7–9 setae. ant vi with 3 basal, 4 apical and 2–5 setae on pt. rostrum reaching middle coxae in some specimens to as far as hind coxae in other specimens. urs 0.21–0.24 × ant iii, 0.17–0.20 × ant vi, 0.20–0.24 × pt, 1.06–1.33 × base and 1.18–1.25 × ht ii with 7–9 short, fine and pointed accessory setae (fig. 2d). mesosternal furca fused, wide, t-shaped. hind femora bearing table 1. measurements of the known morphs of uroleucon (lambersius) robinsoni sp. nov. and u. (l.) grindeliae sp. nov. measurements of the holotype are given in parentheses after the ranges. character uroleucon robinsoni uroleucon grindeliae apterous viviparous female apterous viviparous female alate viviparous female bl 2.50–3.20 (2.90) 2.17–3.02 (2.67) 2.02–2.95 hw 0.46–0.50 (0.46) 0.47–0.52 (0.47) 0.46–0.50 ant 2.39–2.91(2.43) 1.99–2.48 (2.23) 2.25–2.54 ant iii 0.62–0.75 (0.65) 0.51–0.70 (0.61) 0.64–0.70 ant iv 0.45–0.57 (0.46) 0.37–0.47 (0.41) 0.42–0.50 ant v 0.40–0.53 (0.40) 0.32–0.42 (0.38) 0.41–0.46 ant vi 0.73–0.90 (0.74) 0.55–0.67 (0.62) 0.58–0.70 base 0.12–0.15 (0.12) 0.12–0.16 (0.14) 0.12–0.16 pt 0.61–0.78 (0.62) 0.43–0.52 (0.48) 0.46–0.54 urs 0.15–0.16 (0.15) 0.15–0.16 (0.16) 0.15–0.17 femora iii 0.80–0.97 (0.80) 0.84–1.05 (0.90) 0.84–0.97 tibiae iii 1.50–1.87 (1.52) 1.47–1.82 (1.70) 1.52–1.77 ht ii 0.12–01.3 (0.12) 0.12–0.13 (0.12) 0.11–0.12 siph 0.86–1.10 (0.92) 0.62–0.80 (0.66) 0.58–0.68 cauda 0.37–0.44 (0.40) 0.40–0.45 (0.40) 0.37–0.40 barjadze s. & kanturski m., new american uroleucon 47 medium length, thick, rigid setae with narrowly capitate or slightly pointed apices, 0.020–0.040 mm long. hind tibiae bearing thick, rigid setae, most slightly pointed or blunt, shorter than width of tibiae, 0.025–0.045 mm long. in some specimens, tibiae bearing few pseudosensoria. ht i with 5:5:5 setae, ht ii 0.18–0.19 × ant iii, 0.14–0.16 × ant vi, 0.16–0.20 × pt and 0.86–1.08 × base. siph subcylindrical, slightly tapering and curved externally with distinct zone of subapical reticulation and without flange (fig. 2e). reticulated zone 0.17–0.28 × siph. siph 2.30–2.68 × cauda, 0.31–0.41 × bl, and 1.38–1.57 × ant iii. abdomen membranous with medium length, thick rigid setae with narrowly capitate or blunt apices, 0.025–0.040 mm long on abd i–v and 0.030–0.040 mm long on abd vi– viii. abd viii with 4 setae. scleroites on setal bases absent on abd i–v, while setae on abd vi–viii arising from very poorly visible, rounded and pale scleroites. genital plate with two anterior setae longer than others, 7–10 posterior setae. cauda narrow finger-shaped without constriction, 2.05–2.75 × its width at base and 0.13–0.15 × bl with 7–9 fine setae (fig. 2f). remarks of the six grindelia-feeding aphids of uroleucon, only u. penderum robinson, 1986, u. robinsoni sp. nov. and u. grindeliae sp. nov. have shared characters: (1) ant iii has the majority of the secondary rhinaria on the basal half and (2) their urs/ht ii ratio is 1.15 or more and (3) the coxae and trochanters are pale (robinson 1986; blackman & eastop 2020). the apterous viviparous females of uroleucon robinsoni sp. nov. differ from the same morphs of u. penderum: (1) their siph/cauda l ratio is 2.30–2.68 in u. robinsoni sp. nov., while it is 1.65–1.95 in u. penderum, (2) their urs/ht ii ratio is 1.18–1.25 in u. robinsoni sp. nov., while it is 1.3–1.6 in u. penderum, (3) the pigmentation of the tibiae is dusky or dark for most of their length in uroleucon robinsoni, while their tibiae are pale except at the apices in u. penderum (blackman & eastop 2020). the apterous viviparous females of the two new species differ from each other in the following characters: (1) their ant pt/base ratio is 4.66–6.50 in u. robinsoni sp. nov., while it is 3.31–3.61 in u. grindeliae sp. nov., (2) their siph/cauda ratio is 2.30–2.68, while it is 1.37–1.77 in u. grindeliae sp. nov. and (3) their cauda is finger shaped with no constriction in u. robinsoni sp. nov. (fig. 2f), while it is tongue shaped with an evident constriction near the base in u. grindeliae sp. nov. (fig. 3f). biology and distribution the species was collected from grindelia squarrosa in colorado, usa. other morphs and the life cycle of the new species are unknown. uroleucon (lambersius) grindeliae sp. nov. urn:lsid:zoobank.org:act:714de06e-c8cb-4404-8fda-569c01524adc figs 1, 3, table 1 diagnosis apterous viviparous females of grindelia-feeding lambersius share the following combinations of characters: pt 3.31–3.61 × base; ant iii with 7–16 secondary rhinaria (fig. 3a–b); urs 1.03– 1.37 × base and 1.19–1.37 × ht ii with 6–9 short, fine, pointed accessory setae (fig. 3d); ht i with 5:5:5 setae; siph subcylindrical, slightly tapering without a flange (fig. 3e); its reticulated zone 0.27– 0.31 × siph. siph 1.37–1.77 × cauda and 0.24–0.30 × bl; scleroites on setal bases absent on abd i–v, while the setae on abd vi–viii arising from barely visible, rounded scleroites; cauda tongue-shaped with evident constriction near its base and with 9–12 setae (fig. 3f). http://zoobank.org/urn:lsid:zoobank.org:act:714de06e-c8cb-4404-8fda-569c01524adc european journal of taxonomy 729: 42–53 (2020) 48 etymology the name of the new species is derived from the host plant genus grindelia. material examined holotype usa • ♀ apterous viviparous; oregon, salt marsh, near newport; 44°39′ n, 124°1′ w; alt. 113 m; 19 apr. 1980; hille ris lambers leg.; grindelia stricta (= g. hirsutula); nhmuk 010121473. paratypes usa • 1 ♀ alate viviparous; same collection data as for holotype; nhmuk 010121473 • 2 ♀♀ apterous, 2 ♀♀ alate viviparous; same collection data as for holotype; nhmuk 010121465 • 1 ♀ apterous viviparous, 1 ♀ alate viviparous; same collection data as for holotype; nhmuk 010121476 • 1 ♀ apterous viviparous, 1 ♀ alate viviparous; same collection data as for holotype; nhmuk 010121478 • 1 ♀ apterous viviparous, 1 ♀ alate viviparous; same collection data as for holotype; nhmuk 010121474 • 1 ♀ apterous viviparous, 1 ♀ alate viviparous; same collection data as for holotype; fig. 3. uroleucon (lambersius) grindeliae sp. nov., paratype, apterous viviparous female (nhmuk 010121474). a. ant iii. b. secondary rhinaria on ant iii. c. ant vi. d. ultimate rostral (iii–v) segments. e. siphunculus. f. cauda. barjadze s. & kanturski m., new american uroleucon 49 nhmuk 010121475 • 1 ♀ apterous viviparous, 1 ♀ alate viviparous; same collection data as for holotype; nhmuk 010121477 • 3 ♀♀ apterous viviparous, 2 ♀♀ alate viviparous; same collection data as for holotype; dzus 010121479 • 1 ♀ apterous, 1 ♀ alate viviparous; same collection data as for holotype; nhmuk 010121480. description apterous viviparous female (n = 12) color in life. unknown. pigmentation of cleared specimens on slide. body generally yellow to pale brown; ant yellow with light brown ant iii (very basal part pale) or ant yellow with darker apices of ant iii–v; coxae and trochanters pale; femora of legs yellow with very distal part light brown or uniformly yellow; tibiae yellow to light brown with slightly darker distal ends; tarsi light brown; siph pale to yellow with darker distal half; cauda pale (fig. 1b). morphometric characters. hw 0.20–0.25 × ant. head with thick, rigid setae with flat or narrowly capitate apices, 0.035–0.049 mm long. ant tubercles each with 3 setae. ant 0.82–0.96 × bl. ant iii with 7–16 rounded secondary rhinaria of different sizes with sclerotized rims (fig. 3a–b), ant iv longer than ant v. pt 3.31–3.61 × base (fig. 3c). other antennal ratios: vi:iii 0.90–1.18, v:iii 0.57– 0.66, iv:iii 0.65–0.72, pt:iii 0.70–0.92, pt:iv 1.07–1.35 and pt:v 1.22–1.35. ant chaetotaxy: ant bearing thick, rigid setae with blunt or narrow capitate apices. ant iii setae 0.020–0.042 mm long, ls ant iii 1.00–1.20 × bd iii. ant i with 4–6, ant ii with 3–4, ant iii with 12–15, ant iv with 6–10 and ant v with 5–8 setae. ant vi with 3–4 basal, 3–4 apical and 3–5 setae on pt. rostrum reaching hind coxae. urs 0.22–0.31 × ant iii, 0.23–0.30 × ant vi, 0.29–0.38 × pt, 1.03–1.37 × base and 1.19–1.37 × ht ii with 6–9 short, fine, pointed accessory setae (fig. 3d). mesosternal furca fused, wide, t-shaped. hind femora bearing medium length, thick, rigid setae with narrow capitate or slightly pointed setae, 0.020–0.040 mm long. hind tibiae bearing thick, rigid setae with mostly slightly pointed or blunt, shorter than width of tibiae, 0.025–0.050 mm long. ht i with 5:5:5 setae, ht ii 0.18–0.23 × ant iii, 0.18–0.21 × ant vi, 0.23–0.27 × pt and 0.80–1.00 × base. siph subcylindrical, slightly tapering, straight with distinct zone of subapical reticulation and without a flange (fig. 3e). reticulated zone 0.27– 0.31 × siph. siph 1.37–1.77 × cauda, 0.24–0.30 × bl and 1.08–1.33 × ant iii. abdomen membranous with medium length, thick rigid setae with narrowly capitate or blunt apices, 0.035–0.045 mm long on abd i-v and 0.035–0.050 mm long on abd vi–viii. abd viii with 4 setae. scleroites on setal bases absent on abd i–v, while setae on abd vi–viii arising from very poorly visible, rounded and pale scleroites. genital plate with two anterior setae longer than others, 7–10 posterior setae. cauda tongueshaped with evident constriction near base, 1.55–2.52 × its width at base and 0.14–0.19 × bl, with 9–12 long and fine setae (fig. 3f). alate viviparous female (n = 11) color in life. unknown. pigmentation of cleared specimens on slide. head and thorax slightly sclerotized, pale to pale yellow; ant yellow with light brown ant iii (except base) and brown basal part of ant iv; coxae and trochanters pale; femora of legs yellow with very distal ends light; tibiae brown with paler middle section; tarsi yellow to light brown; siph yellow to brown with paler bases; cauda pale (fig. 1c). morphometric characters. hw 0.18–0.20 × ant. head with thick rigid setae with blunt or narrow capitate apices, 0.020–0.035 mm long. ant tubercles each with 2–4 setae. ant 0.81–1.13 × bl. ant iii with 20–26 rounded secondary rhinaria of different sizes with sclerotized rims, ant iv longer than ant v. pt 3.33–3.83 × base. other antennal ratios: vi:iii 0.89–1.09, v:iii 0.62–0.68, iv:iii 0.65– european journal of taxonomy 729: 42–53 (2020) 50 0.76, pt:iii 0.70–0.84, pt:iv 1.06–1.11 and pt:v 1.09–1.22. ant bearing medium length thick and rigid setae with blunt or narrowly capitate apices. ant iii setae 0.020–0.040 mm long, ls ant iii 0.94–1.33 × bd iii. ant i with 5–6, ant ii with 3–4, ant iii with 12–16, ant iv with 7–10, ant v with 7–8 setae. ant vi with 3–4 basal, 4–5 apical and 3–4 setae on pt. rostrum reaching mesosternum. urs 0.23–0.26 × ant iii, 0.23–0.28 × ant vi, 0.31–0.35 × pt, 1.03–1.37 × base and 1.29–1.41 × ht ii with 7–9 fine-pointed accessory setae. hind femora bearing medium length, thick rigid setae with narrowly capitate or slightly pointed setae, 0.020–0.037 mm long. hind tibiae bearing thick, rigid setae with most slightly pointed or blunt, shorter than width of tibiae, 0.025–0.050 mm long. ht i with 5:5:5 setae, ht ii 0.17–0.19 × ant iii, 0.18–0.20 × ant vi, 0.23–0.25 × pt and 0.78–0.97 × base. siph subcylindrical, slightly tapering, straight with distinct zone of subapical reticulation and without a flange. reticulated zone 0.32–0.40 × siph. siph 1.55–1.71 × cauda, 0.22–0.29 × bl and 0.89– 1.00 × ant iii. abdomen membranous with medium length thick, rigid setae with narrowly capitate or blunt apices, 0.025–0.045 mm long on abd i–v and 0.030–0.052 mm long on abd vi-viii. abd viii with 4 setae. scleroites on setal bases absent on abd i–v, while setae on abd vi–vii arising from very poorly visible, rounded scleroites. genital plate with two anterior setae longer than others, 8–10 posterior setae. cauda tongue-shaped with evident constriction near base, 1.95–2.92 × its width at base and 0.12–0.18 × bl with 9–12 long, fine setae. remarks of the six grindelia-feeding aphids of uroleucon, only u. penderum, u. robinsoni sp. nov. and u. grindeliae sp. nov. have shared characters: (1) ant iii has the majority of the secondary rhinaria on the basal half, (2) their urs/ht ii ratio is 1.15 or more and (3) the coxae and trochanters are pale (robinson 1986; blackman & eastop 2020). the apterous viviparous females of u. grindeliae sp. nov. differ from the same morphs of u. penderum and u. robinsoni sp. nov. in the following characters: (1) their ant pt/base ratio is 3.31–3.58 in u. grindeliae, while it is 3.6–6.5 in u. penderum and u. robinsoni and (2) their ant pt/cauda ratio is 1.00–1.20 in u. grindeliae, while it is 1.5–2.0 in u. penderum and u. robinsoni (blackman & eastop 2020). biology and distribution representatives of this species live on grindelia hirsutula in oregon, usa. other morphs and the life cycle of the new species are unknown. discussion the subgenus lambersius is a nearctic group of yellowish-green or green aphids that have a pale cauda and dark siphunculi with pale or almost pale bases or completely pale siphunculi (robinson 1986; heie 1995; nieto nafría et al. 2007). robinson (1986) prepared an annotated list of the species of lambersius that are known north of mexico in north america, provided a key for 29 species and described seven new taxa. at present, 26 species of the subgenus lambersius have been reported from the usa (blackman & eastop 2020). there are six species of uroleucon living on grindelia spp. in north and south america: uroleucon (uroleucon) brevisiphon de carvalho, 1998 on baccharis l. spp. and grindelia sp. in chile; uroleucon (uroleucon) chani robinson, 1985 on grindelia hirsutula and gnaphalium l. sp. in canada and mexico; oligophagous uroleucon (lambersius) erigeronense (thomas, 1878) on grindelia in the usa; uroleucon (lambersius) payuniense ortego & nieto nafría, 2007 on grindelia chiloensis (cornel.) cabrera in argentina; uroleucon (lambersius) penderum on grindelia integrifolia dc., grindelia sp. and heterotheca inuloides cass. in canada, the usa and mexico and uroleucon (lambersius) richardsi barjadze s. & kanturski m., new american uroleucon 51 (robinson, 1964) on grindelia spp. and viguiera kunth sp. in canada, the usa and mexico (robinson 1964, 1985, 1986; nieto nafría et al. 2007, 2011, 2019; jensen et al. 2010; blackman & eastop 2020). the apterous viviparous females of u. brevisiphon and u. chani have a completely dark siph and pale cauda and belong to the subgenus uroleucon, while the remaining four species have dark siphunculi with pale or almost pale bases or completely pale siphunculi, pale coxae and trochanters and belong to the subgenus lambersius. uroleucon brevisiphon is listed in square brackets in the list of grindeliaassociated aphids as a dubious record (blackman & eastop 2020). two grindelia-feeding species fit the diagnosis of the genus uroleucon well and are members of the genus lambersius based on the pigmentation of the siphunculi, coxae and trochanters. after our investigation, the number of species of lambersius increased from 53 to 55 species. it would be interesting to conduct combined morphological, morphometric and molecular (dna barcoding) investigations of species of the subgenus lambersius in the future. key to the apterae of uroleucon on grindelia spp. note: the key is modified from that of blackman & eastop (2020) 7. siph entirely dark ........................................................................ uroleucon chani robinson, 1985 – siph pale, at least on basal part (figs 2e, 3e) .................................................................................. 8 8. ant iii with 12–31 rhinaria extending to 0.80–0.98 from base with majority on distal half ........... ............................................................................................... uroleucon richardsi (robinson, 1964) – ant iii with 5–26 rhinaria extending to 0.4–0.8 from base with majority on the basal half (figs 2a, 3a) ..................................................................................................................................................... 9 9 r iv+v 0.9–1.2 × ht ii, with 6–10 accessory hairs. siph 1.85–2.5 × cauda. ant pt/base 3.2– 5.7 ................................................................................................................................................... 10 – r iv+v 1.15–1.5 × ht ii with 7–13 accessory hairs (figs 2d, 3d). siph 1.5–2.6 × cauda. ant pt/ base 3.0–5.2 ..................................................................................................................................11 10. siphunculi pale at base, dark over most of their length. distal caudal hairs usually short and blunt or capitate. dorsal cephalic hairs 18–35 μm long ................. uroleucon erigeronense (thomas, 1878) – siphunculi pale. distal caudal hairs pointed. dorsal cephalic hairs 45–55 μm long ......................... ......................................................................... uroleucon payuniense ortego & nieto nafría, 2007 11. ant pt/base 3.31–3.58. ant pt 1.0–1.2 × cauda .. uroleucon (lambersius) grindeliae sp. nov. – ant pt/base 3.6–5.2. ant pt 1.5–2.0 × cauda ......................................................................... 12 12. siph 2.30–2.68 × cauda. r iv+v 1.18–1.25 × ht ii. tibiae dusky/dark for most of their length (fig. 1a) .........................................................................uroleucon (lambersius) robinsoni sp. nov. – siph 1.65–1.95 × cauda. r iv+v 1.3–1.6 × ht ii. tibiae pale except at apices ................................ ................................................................................................ uroleucon penderum robinson, 1986 acknowledgments we are sincerely grateful to paul a. brown and david ouvrard (nhmuk, london, uk) for their kind assistance and support during our visits to the collection as well as for the loan of the material. special thanks go to eric maw (agriculture and agri-food canada, ottawa, canada) and andrew s. jensen (california academy of sciences, san francisco, ca, usa) for the linguistic assistance and comments on the manuscript. we would like to thank the editor and two reviewers for their valuable comments and suggestions, which helped the authors to improve the first version of the manuscript. mariusz kanturski european journal of taxonomy 729: 42–53 (2020) 52 gratefully acknowledges the scholarship for outstanding young scientists from the ministry of science and higher education of poland (1165/e-340/styp/12/17). references blackman r.l. & eastop v.f. 2006. aphids on the world’s herbaceous plants and shrubs. john wiley and sons, london. blackman r.l. 2010. aphids – aphidinae (macrosiphini). handbooks for the identification of british insects 2 (7). royal entomological society, st albans. blackman r.l. & eastop v.f. 2020. aphids of the world’s plants: an online identification and information guide. available from http://www.aphidsonworldsplants.info [accessed 15 mar. 2020]. favret c. 2020. aphid species file. version 5.0/5.0. available from http://aphid.speciesfile.org [accessed 15 mar. 2020]. heie o.e. 1995. the aphidoidea of fennoscandia and denmark vi. aphidinae. part 3 of macrosiphini and lachnidae. fauna entomologica scandinavica 31: 1–222. ilharco f.a. & van harten a. 1987. systematics. in: minks a.k. & harrewijn p (eds) aphids: their biology, natural enemies and control: 51–77. elsevier science publishers, amsterdam. jensen a.s., miller g.l. & carmichael a. 2010. host range and biology of uroleucon (lambersius) erigeronense (thomas, 1878), and its synonymy with uroleucon (lambersius) escalantii (knowlton) 1928 (hemiptera: aphididae). proceedings of the entomological society of washington 112 (2): 239– 245. https://doi.org/10.4289/0013-8797-112.2.239 nieto nafría j.m., mier durante m.p., ortego j. & seco fernández m.v. 2007. the genus uroleucon (hemiptera: aphididae: macrosiphini) in argentina, with descriptions of five new species. the canadian entomologist 139 (2): 154–178. https://doi.org/10.4039/n05-100 nieto nafría j.m., mier durante m.p. & pérez hidalgo n. 2011. mexican uroleucon (hemiptera: aphididae) from the collection of the muséum national d’histoire naturelle of paris with eleven new species. florida entomologist 94 (3): 622–648. https://doi.org/10.1653/024.094.0329 nieto nafría j.m., von dohlen c.d., moreno-gonzález v., ortego j. & mier durante m.p. 2019. the species of uroleucon (hemiptera: aphididae) living on adesmia (fabaceae) in argentina, with the description of a new species. zootaxa 4555 (4): 561–572. https://doi.org/10.11646/zootaxa.4555.4.7 robinson a.g. 1964. a new species of dactynotus rafinesque (homoptera: aphididae) from grindelia squarrosa (pursh) dunal. the canadian entomologist 96 (10): 1330–1332. https://doi.org/10.4039/ent961330-10 robinson a.g. 1985. annotated list of uroleucon (uroleucon, uromelan, satula) (homoptera: aphididae) of america north of mexico, with keys and descriptions of new species. the canadian entomologist 117 (8): 1029–1054. https://doi.org/10.4039/ent1171029-8 robinson a.g. 1986. annotated list of uroleucon (lambersius) (homoptera: aphididae) of america north of mexico, with a key and descriptions of new species. the canadian entomologist 118 (6): 559–576. https://doi.org/10.4039/ent118559-6 the plant list. 2013. version 1.1. available from http://www.theplantlist.org/ [accessed 1 mar. 2020)]. http://www.aphidsonworldsplants.info http://aphid.speciesfile.org https://doi.org/10.4289/0013-8797-112.2.239 https://doi.org/10.4039/n05-100 https://doi.org/10.1653/024.094.0329 https://doi.org/10.11646/zootaxa.4555.4.7 https://doi.org/10.4039/ent961330-10 https://doi.org/10.4039/ent1171029-8 https://doi.org/10.4039/ent118559-6 http://www.theplantlist.org/ barjadze s. & kanturski m., new american uroleucon 53 manuscript received: 29 april 2020 manuscript accepted: 14 october 2020 published on: 16 december 2020 topic editor: nesrine akkari section editor: anna namyatova desk editor: eva-maria levermann printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. 181 european journal of taxonomy 860: 181–182 issn 2118-9773 https://doi.org/10.5852/ejt.2023.860.2059 www.europeanjournaloftaxonomy.eu 2023 · zheng y. et al. this work is licensed under a creative commons attribution license (cc by 4.0). c o r r i g e n d u m urn:lsid:zoobank.org:pub:2ed9296a-d8e2-4ee3-99d7-2eecc970fc40 new fossil records of xyelidae (hymenoptera) from the middle jurassic of inner mongolia, china – corrigendum yan zheng 1,*, haiyan hu 2, dong chen 3, jun chen 4, haichun zhang 5 & alexandr p. rasnitsyn 6,* 1,4 institute of geology and paleontology, linyi university, shuangling rd., linyi 276000, china. 1,4,5 state key laboratory of palaeobiology and stratigraphy, nanjing institute of geology and palaeontology, east beijing road, nanjing 210008, china. 2 school of agronomy and environment, weifang university of science and techonology, jinguang road, shouguang, 262700, china. 3 school of environmental and municipal engineering, qingdao university of technology, qingdao 266033, china. 6 palaeontological institute, russian academy of sciences, moscow, 117647, russia. 6 natural history museum, london sw7 5bd, uk. * corresponding authors: zhengyan536@163.com, alex.rasnitsyn@gmail.com 2 email: huhaiyanzy@163.com 3 email: chendong_cau@163.com 4 email: rubiscada@sina.com 5 email: hczhang@nigpas.ac.cn 1 urn:lsid:zoobank.org:author:28eb8d72-5909-4435-b0f2-0a48a5174cf9 2 urn:lsid:zoobank.org:author:91b2fb61-31a9-449b-a949-7ae9efd69f56 3 urn:lsid:zoobank.org:author:51d01636-eb69-4100-b5f6-329235eb5c52 4 urn:lsid:zoobank.org:author:8bab244f-8248-45c6-b31e-6b9f48962055 5 urn:lsid:zoobank.org:author:18a0b9f9-537a-46ef-b745-3942f6a5ab58 6 urn:lsid:zoobank.org:author:e7277cab-3892-49d4-8a5d-647b4a342c13 zheng y., hu h., chen d., chen j., zhang h. & rasnitsyn a.p. 2023. new fossil records of xyelidae (hymenoptera) from the middle jurassic of inner mongolia, china – corrigendum. european journal of taxonomy 860: 181–182. https://doi.org/10.5852/ejt.2023.860.2059 the present corrigendum corrects errors that occurred in zhen et al. (2021). https://doi.org/10.5852/ejt.2021.733.1229 the authors regret to inform that the specimen numbers in our original paper (zhen et al. 2021) are not official: “nnd0176” should actually be “nigp201142” and “nnd0185” should actually be “nigp201143”. the authors would like to apologise for any inconvenience caused. https://doi.org/10.5852/ejt.2023.860.2059 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ https://zoobank.org/urn:lsid:zoobank.org:pub:2ed9296a-d8e2-4ee3-99d7-2eecc970fc40 mailto:zhengyan536%40163.com?subject= mailto:alex.rasnitsyn%40gmail.com?subject= mailto:huhaiyanzy%40163.com?subject= mailto:chendong_cau%40163.com?subject= mailto:rubiscada%40sina.com?subject= mailto:hczhang%40nigpas.ac.cn?subject= https://zoobank.org/urn:lsid:zoobank.org:author:28eb8d72-5909-4435-b0f2-0a48a5174cf9 https://zoobank.org/urn:lsid:zoobank.org:author:91b2fb61-31a9-449b-a949-7ae9efd69f56 https://zoobank.org/urn:lsid:zoobank.org:author:51d01636-eb69-4100-b5f6-329235eb5c52 https://zoobank.org/urn:lsid:zoobank.org:author:8bab244f-8248-45c6-b31e-6b9f48962055 https://zoobank.org/urn:lsid:zoobank.org:author:18a0b9f9-537a-46ef-b745-3942f6a5ab58 https://zoobank.org/urn:lsid:zoobank.org:author:e7277cab-3892-49d4-8a5d-647b4a342c13 https://doi.org/10.5852/ejt.2023.860.2059 https://doi.org/10.5852/ejt.2021.733.1229 european journal of taxonomy 860: 181–182 (2023) 182 references zheng y., hu h., chen d., chen j., zhang h. & rasnitsyn a.p. 2021. new fossil records of xyelidae (hymenoptera) from the middle jurassic of inner mongolia, china. european journal of taxonomy 733: 146–159. https://doi.org/10.5852/ejt.2021.733.1229 published on: 3 march 2023 printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; leibniz institute for the analysis of biodiversity change, bonn – hamburg, germany; national museum, prague, czech republic. https://doi.org/10.5852/ejt.2021.733.1229 european journal of taxonomy 819: 158–165 issn 2118-9773 https://doi.org/10.5852/ejt.2022.819.1787 www.europeanjournaloftaxonomy.eu 2022 · johanson k.a. et al. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:80de5f71-8a1c-472f-9205-f956639d0605 158 description of two new species of oecetis (trichoptera, leptoceridae) from borneo kjell arne johanson 1,*, tobias malm 2 & tin sjöberg 3 1,2,3 department of zoology, swedish museum of natural history, box 50007, 10405 stockholm, sweden. * corresponding author: kjell.arne.johanson@nrm.se 2 email: tobias.malm@nrm.se 3 email: tin.sjoberg@nrm.se 1 urn:lsid:zoobank.org:author:f2a38cf6-59eb-4f88-bfeb-761dbea7b01a 2 urn:lsid:zoobank.org:author:4cd5e640-d2f2-4123-913e-e854959424ea 3 urn:lsid:zoobank.org:author:f20b3a9c-7173-409d-8c89-92977b6d2b8d abstract. two new species of oecetis from maliau basin, malaysian borneo, are described for the first time, o. mesospina sp. nov. and o. apelqvisti sp. nov. these two new species bring the total number of oecetis found on the island of borneo up to 16. keywords. taxonomy, caddisflies, oriental, leptocerini, malaysia. johanson k.a., malm t. & sjöberg t. 2022. description of two new species of oecetis (trichoptera, leptoceridae) from borneo. european journal of taxonomy 819: 158–165. https://doi.org/10.5852/ejt.2022.819.1787 introduction with almost 2000 known species (morse 2020) the family leptoceridae leach, 1815 represents about 12% of the species diversity in the trichoptera and constitute one of the largest families in the order. within the family, the subfamily leptocerinae leach, 1815 holds the majority of the species and also has the widest distribution, with species recorded from all continents except antarctica. around 600 species are recognized in the leptocerine genus oecetis mclachlan, 1877 (morse 2020; johanson et al. 2020a, 2020b), the largest genus in the entire family. a few attempts based on morphological characteristics have been made to cluster its species into more heuristic groups. chen (1993) presented a phylogenetic hypothesis based on morphological characters and revealed four main clades; these were erected as subgenera of oecetis. the eurasian members of the genus were subsequently divided into species groups by malicky (2005), but without any critical examinations of the characters used of or phylogenetic context. about 270 species have been described from the oriental biogeographical region, and of these, 14 are recorded from the island of borneo (malicky 2010). of these, the following four species have been recorded from the malaysian states: from sabah, o. paris malicky, 2006 and o. lynkeus malicky, 2005; from sarawak, o. koyana kimmins, 1955 and o. maron malicky & chantaramongkol, 2005. the latter https://doi.org/10.5852/ejt.2022.819.1787 https://creativecommons.org/licenses/by/4.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:80de5f71-8a1c-472f-9205-f956639d0605 mailto:kjell.arne.johanson@nrm.se mailto:tobias.malm%40nrm.se?subject= mailto:tin.sjoberg%40nrm.se?subject= http://zoobank.org/urn:lsid:zoobank.org:author:f2a38cf6-59eb-4f88-bfeb-761dbea7b01a http://zoobank.org/urn:lsid:zoobank.org:author:4cd5e640-d2f2-4123-913e-e854959424ea http://zoobank.org/urn:lsid:zoobank.org:author:f20b3a9c-7173-409d-8c89-92977b6d2b8d https://doi.org/10.5852/ejt.2022.819.1787 johanson k.a. et al., new oecetis from borneo 159 is also known from indonesian borneo (kalimantan). in total, eight species are known from indonesian borneo, and three are recorded from brunei. here we describe two new species from sabah, bringing to 16 the total for the island. material and methods caddisfly adults were collected from 22w circular uv lamps and in malaise traps situated closely to the river during december 9–14 2007. the material was preserved in 80% ethanol. sorting and determinations were executed in the laboratory at the swedish museum of natural history. individual specimens were initially grouped based on genital morphology, after which the abdomen was separated from the rest of the body. extraction of dna was carried out on the abdomen with qiaamp® dna micro kit (qiagen), which also macerated the abdomen. the abdomen was dehydrated in absolute alcohol and temporarily suspended in euparal on a microscope slide. illustrations of genitalia were drawn using a laborlux compound microscope with a drawing tube. the illustrations were subsequently scanned and finalized in adobe® illustrator® 15.1 and photoshop® 12.1. after examination and illustration, the abdomens were transferred back to 80% ethanol together with the rest of the specimen. in situations where the description is based on a single specimen, species are confirmed to be distinct from other bornean and oriental species based on a combination of dna sequences and morphology. all material is deposited in the swedish museum of natural history (nhrs). the terminology applied to genitalia follows that of nielsen (1957). results phylum arthropoda von siebold, 1848 class insecta linneaus, 1758 order trichoptera kirby, 1813 family leptoceridae leach, 1815 genus oecetis mclachlan, 1877 oecetis mesospina sp. nov. urn:lsid:zoobank.org:act:2896e944-5e55-49fb-89d5-76df8d8e9ec2 figs 1–5, 11 diagnosis the species is distinguished from many of the oriental and palearctic species by the absence of modified tergites on segment vi–viii. it resembles the bornean oecetis peleus malicky, 2005 and oecetis halirrhotios malicky, 2005 in genitalic features, particularly in the shape of tergum x, the inferior appendages of the inferior appendages and the superior appendages. however, it is distinguished from both species by the presence of a row of short megasetae forming a comb along the mesal margin of the inferior appendages and by the phallus bearing a pair of sharply triangular lateral processes on the posterior part of the ventral side compared to rounded processes in o. peleus and o. halirrhotios. it is also distinguished from o. peleus by the more strongly curved tergum x in lateral view. etymology mesospina, referring to the row of spine-like megasetae along the mesal margin of the coxopodites. type material holotype malaysia • ♂ (in alcohol); sabah, tawau, maliau basin, nepenthes camp, crossing stream; 4°43′58.9″ n, 116°52′40.7″ e; 994 m a.s.l.; 9–14 dec. 2007; b. viklund and n. jönsson leg.; 6 m malaise trap, loc# vkbs-2007-27; dna voucher jl5; nhrs. http://zoobank.org/urn:lsid:zoobank.org:act:2896e944-5e55-49fb-89d5-76df8d8e9ec2 european journal of taxonomy 819: 158–165 (2022) 160 figs 1–5. oecetis mesospina sp. nov., holotype, ♂ (nhrs). 1. genitalia, lateral view. 2. genitalia, dorsal view. 3. genitalia, ventral view. 4. phallus, lateral view. 5. phallus, ventral. scale bar = 0.1 mm. johanson k.a. et al., new oecetis from borneo 161 paratype malaysia • 1 ♂ (in alcohol); same collection data as for holotype; dna voucher jo7; nhrs. description of male body. forewing: 5.6–6.4 mm (n = 2). tergites vi–viii without reticulate pattern. genitalia (figs 1–5). higher than long in lateral view. in lateral view segment ix with almost straight anterior margin, dorsal and ventral parts slightly produced anteriorly and posteriorly. acrotergite of segment ix well developed, forming single transverse wart-like aggregation of setal bases, more than three times wider than long. pair of dorsal processes of segment ix minute, wart-like. in lateral view lower posterolateral plate of segment ix short, posteriorly rounded, situated at same level as base of inferior appendages; upper posterolateral plate of segment ix short, sharply triangular, located medially between superior appendages and inferior appendages. longitudinal apodemes absent. tergum x reaching as far posteriorly as inferior appendages, about twice as long as superior appendages; divided into long central and two shorter lateral processes; in lateral view, central process longer than lateral processes, curving posteroventrally, with apical megasetae, narrow in dorsal view; lateral processes curving posteroventrally, each with single, apical megaseta; in dorsal view forming triangular plates. in lateral view, superior appendages almost circular, about half as long as tergum x; in dorsal view situated closely to each other, mesally fused, almost circular. in lateral view, each inferior appendage slender along their, slightly sigmoid, parallel-sided, except narrowing apically; four very long dorsally oriented setae located at mid-length of each inferior appendage; in ventral view inferior appendages well separated after basal plate and pair of small rounded processes; each mesal plate with comb of short megasetae along mesal margin. phallus about as long as tergum x. in lateral view almost as high as long; in ventral view anterior margin almost straight, widest anteriorly and weakly narrowing posteriorly; sharply triangular pair of lateral processes after mid-length; internal sclerite horse-shoe shaped. distribution the species is only known from the type locality (fig. 11). oecetis apelqvisti sp. nov. urn:lsid:zoobank.org:act:0463dede-cf3b-43ba-9b3d-af23699c165c figs 6–11 diagnosis the species is distinguished from many of the oriental and palearctic species by the absence of modified tergites on segment vi–viii. it resembles o. idas malicky, 2005 in genitalic features, particularly the long and slightly curving superior appendages, long and strongly ventrally curving tergum x, with right branch longer than left branch, and long pair of dorsal processes. oecetis apelqvisti sp. nov. can be distinguished from o. idas in lateral view by the clearly bifid inferior appendages, and the apical part of phallus with a single hook-shaped sclerotized process. etymology apelqvisti, named after one of the collectors, niklas apelqvist (formerly niklas jönsson). type material holotype malaysia • ♂ (in alcohol); sabah, tawau, maliau basin, nepenthes camp, 150 m from jalan babi; 4°43′56.1″ n, 116°52′52.0″ e; 1031 m a.s.l.; 13 dec. 2007; n. jönsson, t. malm and b. viklund; light trap, loc#vkbs-2007-47; dna voucher jq1; nhrs. http://zoobank.org/urn:lsid:zoobank.org:act:0463dede-cf3b-43ba-9b3d-af23699c165c european journal of taxonomy 819: 158–165 (2022) 162 figs 6–10. oecetis apelqvisti sp. nov., holotype, ♂ (nhrs). 6. genitalia, lateral view. 7. genitalia, dorsal view. 8. genitalia, ventral view. 9. phallus, lateral view. 10. phallus, ventral view. scale bar = 0.1 mm. johanson k.a. et al., new oecetis from borneo 163 description of male body. forewing: 6.7 mm (n = 1). tergites vi–viii unmodified. genitalia (figs 6–10). longer than high in lateral view. segment ix oriented almost vertically, anterior margin uniformly convex along its height; ventral part almost circularly expanded posteriorly; dorsal part reduced. acrotergite of segment ix well developed, forming single oval aggregation of setal bases about width of dorsal processes combined. pair of dorsal processes of segment ix elongate, tube-like; segment lacking membranous area. lower posterolateral plate of segment ix short, high, located from basis of inferior appendages to mid-height of segment; upper posterolateral plate of segment ix lacking. longitudinal apodeme absent. tergum x in lateral view twice length of superior appendages, curving ventrally along its length to pointed apex; in dorsal view bilobed from half length, both lobes tapering. superior appendages very long, tubular, originating at anterior margin of segment ix at junction between dorsal and ventral parts; oriented dorsally before curving posteriorly over segment ix. each inferior appendage high and wide, about as long as superior appendages, all setae shorter than length of inferior appendages; in lateral view 2-branched, basal two-thirds widening from base, sharply triangular dorsal branch, ventral margin expanded around mid-length; apical branch digitate, narrow, oriented posterodorsally; in ventral view, each inferior appendage with basimesal expansions, sharply narrowing after basal one-third, apical one-third narrow, digitate. phallus about as long as tergum x in lateral view, uniformly slender, oriented posteriorly before curving ventrally at mid-length, apex with single hook-shaped sclerotized process; in ventral view about length of inferior appendages, tubular, slightly widening apically, hook-shaped sclerotized process located on left side. distribution the species is only known from the type locality (fig. 11). fig. 11. map with the position of the two sampling sites in maliau basin shown as orange dots. the position of the maliau basin area on borneo is indicated with a blue rectangular in the upper-left map. european journal of taxonomy 819: 158–165 (2022) 164 discussion the study is based on only three specimens. both specimens of o. mesospina sp. nov. were collected in a malaise trap and the single specimen of o. apelqvisti sp. nov. was taken from a light trap. no specimens of previously described species were collected from the visited sites. this could indicate that species of oecetis are rarely collected in the area or that they are seasonal. it is possible that the diversity might be much higher than indicated by the known species number. with around 270 species described, the diversity of oecetis in the oriental region represents about 45% of the world fauna of oecetis. a recent paper by quinteiro & almeida (2021) on the neotropical diversity of oecetis gave the number of neotropical species as 73 and of these, 49 were assigned to six evolutionary groups based on phylogenetic analyses of morphological characters. to fully understand the evolutionary success of oecetis in the oriental region, a phylogenetic analysis involving dna characters is needed, involving as many species and representatives of different biogeographical regions as possible. acknowledgements we are very grateful to the collectors of these specimens, the late mr bert viklund (nhrs) and mr niklas apelqvist (formerly jönsson, nhrs). we also like to extend our thanks to the editor and the two referees for their help in improving the manuscript. references chen y.e. 1993. revision of the oecetis (trichoptera: leptoceridae) of the world. phd thesis, clemson university. johanson k.a., pham t.h., malm t. & sjöberg t. 2020a. description of six new species of oecetis (trichoptera, leptoceridae) from vietnam. zootaxa 4816 (3): 311–324. https://doi.org/10.11646/zootaxa.4816.3.2 johanson k.a., mary j.n., sjöberg t. & malm t. 2020b. eighteen new species of oecetis (trichoptera, leptoceridae) from new caledonia. zootaxa 4809 (2): 201–240. https://doi.org/10.11646/zootaxa.4809.2.1 malicky h. 2005. beiträge zur kenntnis asiatischer oecetis (trichoptera, leptoceridae). linzer biologische beiträge 27: 605–669. malicky h. 2010. atlas of southeast asian trichoptera. chiang mai university. morse j.c. 2020. trichoptera world checklist. available from https://entweb.sites.clemson.edu/database/trichopt/ [accessed 2 jan. 2020]. nielsen a. 1957. a comparative study of the genital segments and their appendages in male trichoptera. biologiske skrifter 8 (5): 1–159. quinteiro f.b. & almeida e.a.b. 2021. systematics of neotropical oecetis mclachlan, 1877 (trichoptera: leptoceridae): when the taxonomy and phylogeny meet. zoologischer anzeiger 293: 233–246. https://doi.org/10.1016/j.jcz.2021.06.005 manuscript received: 10 august 2021 manuscript accepted: 4 march 2022 published on: 18 may 2022 section editor: nesrine akkari section editor: helen m. barber-james desk editor: pepe fernández https://doi.org/10.11646/zootaxa.4816.3.2 https://doi.org/10.11646/zootaxa.4809.2.1 https://entweb.sites.clemson.edu/database/trichopt/ https://doi.org/10.1016/j.jcz.2021.06.005 johanson k.a. et al., new oecetis from borneo 165 printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. european journal of taxonomy 750: 140–155 issn 2118-9773 https://doi.org/10.5852/ejt.2021.750.1369 www.europeanjournaloftaxonomy.eu 2021 · delicado d. et al. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:ffdad9f7-5e6a-40da-9d2f-1713ce7aff4a 140 arganiella giusti & pezzoli, 1980 (caenogastropoda: truncatelloidea: hydrobiidae): a widespread genus or several narrow-range endemic genera? diana delicado 1,*, vladimir pešić 2 & marian a. ramos 3 1 justus liebig university giessen, department of animal ecology & systematics, heinrich-buff-ring 26-32 ifzd, 35392 giessen, germany. 2 university of montenegro, faculty of sciences, department of biology, cetinjski put b.b., 81000 podgorica, montenegro. 3 museo nacional de ciencias naturales (mncn-csic), josé gutiérrez abascal 2, 28006 madrid, spain. * corresponding author: didelicado@gmail.com 2 email: vladopesic@gmail.com 3 email: m.ramos@mncn.csic.es 1 urn:lsid:zoobank.org:author:ed8cd4c8-7310-40e3-8821-7721615a5992 2 urn:lsid:zoobank.org:author:2b46f869-1202-47a9-9359-0be04d691f6c 3 urn:lsid:zoobank.org:author:c108bacd-06a7-4c6d-8466-2a1cfe349deb abstract. most valvatiform genera of the gastropod family hydrobiidae are narrow-range taxa. one exception is the genus arganiella, which is comprised of three congeners: the type species a. pescei from the apennine peninsula, a. wolfi from the iberian peninsula and a. tabanensis from the balkans. the genus assignment of the latter two species was based on morphological similarities with a. pescei in the shell, operculum, radula and genitalia. given that the morphology of hydrobiids is sometimes susceptible to convergence, this study re-evaluates the taxonomic status of species of arganiella by analysing mitochondrial (mtcoi) and nuclear (18s rrna) sequences of topotypes or near topotypes to infer their phylogenetic position. our phylogenetic analyses depicted arganiella as a non-monophyletic group within hydrobiidae, and sequence divergence among the three species ranged from 14.5 to 16.7% for mtcoi and 2.0 to 3.8% for 18s. we also re-examined the extent of morphological variation among species of arganiella and found a few differences among them and other valvatiform genera. consequently, we propose two new genera for a. wolfi and a. tabanensis. our results conflict with the classification of valvatiform hydrobiid species solely based on traditional phenotypical methods and suggest further taxonomic evaluation within a molecular framework. keywords. springsnails, valvatiform hydrobiids, phylogeny, morphology, mediterranean region. delicado d., pešić v. & ramos m.a. 2021. arganiella giusti & pezzoli, 1980 (caenogastropoda: truncatelloidea: hydrobiidae): a widespread genus or several narrow-range endemic genera? european journal of taxonomy 750: 140–155. https://doi.org/10.5852/ejt.2021.750.1369 https://doi.org/10.5852/ejt.2021.750.1369 https://creativecommons.org/licenses/by/4.0/ urn:lsid:zoobank.org:pub:ffdad9f7-5e6a-40da-9d2f-1713ce7aff4a mailto:didelicado%40gmail.com?subject= mailto:vladopesic%40gmail.com?subject= mailto:m.ramos%40mncn.csic.es?subject= http://zoobank.org/urn:lsid:zoobank.org:author:ed8cd4c8-7310-40e3-8821-7721615a5992 http://zoobank.org/urn:lsid:zoobank.org:author:2b46f869-1202-47a9-9359-0be04d691f6c http://zoobank.org/urn:lsid:zoobank.org:author:c108bacd-06a7-4c6d-8466-2a1cfe349deb https://doi.org/10.5852/ejt.2021.750.1369 delicado d. et al., systematics of the hydrobiid genus arganiella 141 introduction the gastropod genus arganiella giusti & pezzoli, 1980 (family hydrobiidae stimpson, 1865) comprises three narrow-range endemic species, each living on a different mediterranean peninsula. these tiny springsnails (1.0–1.5 mm) have a depressed trochiform to valvatiform shell and simplified anatomical structures. the type species, a. pescei giusti & pezzoli, 1980, was described from several wells and springs located in the central-eastern apennine peninsula (giusti & pezzoli 1980). a proposed representative of the genus boetersiella arconada & ramos, 2001, b. wolfi boeters & glöer, 2007, which is endemic to the southwestern iberian peninsula, was transferred to arganiella by arconada & ramos (2007a, 2007b). based on morphological and geographical evidence, these authors also synonymized the species a. tartesica arconada & ramos, 2007 with a. wolfi (arconada & ramos 2007b). the third species, a. tabanensis boeters, glöer & pešić, 2014, was recently erected for a hydrobiid population living in a small rheocrene spring in montenegro, in the balkan peninsula (boeters et al. 2014). the assignment of a. wolfi and a. tabanensis to arganiella was based on morphological similarity with the type species in shell, penis and female distal genital features (arconada & ramos 2007a, 2007b; boeters et al. 2014). however, such characters are susceptible to convergent evolution in valvatiform hydrobiid genera (bodon et al. 2001) and, therefore, the systematics of the known species of arganiella needs to be re-evaluated through molecular analyses. to date, dna sequence data are available only for a. wolfi, and phylogenetic relationships inferred from a multilocus dataset of valvatiform hydrobiid genera from the iberian peninsula resolved a. wolfi as the sister taxon of the iberian genus iberhoratia arconada & ramos, 2007 (delicado et al. 2019). sequence data from the other two species of arganiella are needed to confidently assess the taxonomic composition of this wide-ranging genus. as might be expected in taxa with poor dispersal capabilities, we hypothesize that phylogenetic analyses would depict species of arganiella as three unrelated, narrow-range lineages rather than as a widely distributed monophyletic group. this result would challenge the genus assignment of these valvatiform hydrobiids, which was based on morphology, and limit the known geographic distribution of arganiella to the apennine peninsula. to test these assumptions, we analyse new mitochondrial and nuclear dna sequence data from the type species a. pescei and from a. tabanensis with previously published molecular datasets of valvatiform hydrobiids that also include sequences of a. wolfi (delicado et al. 2019) in order to infer the phylogenetic relationships of the species of arganiella and to quantify the degree of divergence among them. we also provide previously missing descriptions of the radular features of a. tabanensis for a morphological comparison. finally, the taxonomic status of species of arganiella is discussed in light of the evidence from the phylogenetic analyses and morphological comparisons of these taxa with other valvatiform hydrobiid genera recorded from the same mediterranean regions. material and methods we assessed the taxonomic status of the three recognized species of arganiella using dna sequence and morphological information from these and other valvatiform genera occurring in the apennine, balkan and iberian peninsulas. one individual of a. pescei was used for the dna assessment. this specimen was collected from susanna springs, in the region of the type locality (i.e., central-eastern apennine peninsula), and deposited in the collection of the university of giessen systematics and biodiversity (ugsb) (diehl et al. 2018) in germany (ugsb 10365). following an exhaustive morphological examination, bodon et al. (2001) had assigned the hydrobiid populations living in these springs to a. pescei. morphological characters from a. pescei were scored using the original description by giusti & pezzoli (1980) and the re-description by bodon et al. (2001). for the iberian a. wolfi, we used the morphological description by arconada & ramos (2007a) and boeters & glöer (2007), and the sequences of a topotype used by delicado et al. (2019) for the molecular analyses. the shell morphology and anatomy of the balkan a. tabanensis are as illustrated by boeters et al. (2014). additional data on european journal of taxonomy 750: 140–155 (2021) 142 radular and opercular features of this species, as well as partial sequences of the studied dna markers, were also collected for the present study. we collected ca 50 topotypes from the a. tabanensis type locality (taban spring, montenegro; 42.52795° n, 19.21921° e) in 2015 and preserved them in 80% ethanol in the field. shells and opercula were photographed using a keyence vhx 2000 3d digital microscope. six adults were dissected, and their buccal mass extracted, under an olympus szx12 stereo microscope. radulae were extracted, cleaned and prepared as described by delicado et al. (2016) for imaging on a field emission scanning electron microscope (fesem) dsm982 gemini (carl zeiss gmbh, germany). the collected specimens were then deposited in the ugsb collection (ugsb 18847). we isolated total dna from one individual per species (for a. pescei and a. tabanensis) following the ctab protocol performed by wilke et al. (2006). fragments of the mitochondrial cytochrome c oxidase subunit i (coi) and nuclear ribosomal 18s rrna (18s) were amplified and sequenced using the primer pairs lco1490 (folmer et al. 1994) and cor722b (davis et al. 1998) for coi and the universal primers for metazoan 18s (holland et al. 1991). amplification conditions for both gene fragments were as previously described by delicado et al. (2012). the annealing temperature used was 52°c. samples were sequenced in an abi 3730 xl sequencer (life technologies, carlsbad, ca, usa) using a bigdye terminator kit ver. 3.1 (life technologies). the new sequences were deposited in genbank (table 1). we assessed the taxonomy of the three species of arganiella by analysing the coi and 18s sequences of these species along with those of other european valvatiform and (closely related) non-valvatiform genera available from genbank (table 1). forward and reverse sequences were aligned and edited in sequencher ver. 5.4.6 (gene codes, ann arbor, mi). mega ver. 7.0.14 (kumar et al. 2016) was used to assemble the gene-partition datasets and to calculate genetic distances (uncorrected p-distances). the coi dataset was manually aligned also using mega. the rrna 18s fragment was aligned using mafft ver. 7.402 (katoh & standley 2013), with default settings for gap penalties. according to the corrected akaike’s information criterion (aicc; akaike 1974; sugiura 1978; hurvich & tsai 1989), jmodeltest ver. 2.1.7 (darriba et al. 2012) selected trn (tamura & nei 1993) + i (invariable sites) + g (rate variation among sites) and trnef (tamura-nei model with equal base frequencies; tamura & nei 1993) + i + g models of nucleotide evolution for the coi and 18s datasets, respectively. we used dambe ver. 7 (xia 2018) and the proportion of invariant sites (pinv = 0.47) obtained in jmodeltest to conduct the saturation test (xia et al. 2003; xia & lemey 2009) on the coi partition. the observed saturation (iss = 0.39) was significantly lower than the critical values (iss.c = 0.71; p < 0.001), suggesting little saturation in our coi dataset. phylogenetic analyses based on maximum likelihood (ml) methods were conducted using the raxml blackbox web-server [https://raxml-ng.vital-it.ch/#/; kozlov et al. 2019] with 10 random starting trees and the optimal substitution models for each gene partition selected in jmodeltest. bayesian inference (bi) analyses were run with mixed substitution models in mrbayes ver. 3.2.6 (ronquist et al. 2012) for 5 million generations with a sampling frequency of 1000. after verifying convergence of the bi analysis (standard deviation of split frequencies < 0.01), the first 10% of generations were discarded as burn-in. branch robustness was evaluated by rapid bootstrapping (bs) (stamatakis et al. 2008) with an automatic cut-off for ml and by bayesian posterior probability (bpp) for bi. inferred topologies and branch supports were visualized in figtree ver. 1.4.3 (rambaut 2010). for the morphological comparisons of species of arganiella with other valvatiform hydrobiid genera from the iberian, apennine and balkan peninsulas (table 2), we scored morphological character states, including features of the shell, radula, operculum and anatomy, according to the information provided in the following studies: arconada & ramos (2001) for boetersiella and chondrobasis arconada & ramos, 2001; girardi & boeters (2012) for corbellaria girardi & boeters, 2012; delicado et al. (2019) for deganta arconada & ramos in delicado et al., 2019; arconada et al. (2007) for iberhoratia arconada & ramos, 2007; arconada & ramos (2006) for josefus arconada & ramos, https://raxml-ng.vital-it.ch/#/ delicado d. et al., systematics of the hydrobiid genus arganiella 143 table 1 (continued on next page). taxon name, locality name, genbank numbers and original references for sequences analysed in this study. taxon locality genbank # 18s genbank # coi original reference agrafia wiktori agrafa mountains, sikiá, evrytania, greece (39.36861, 21.63139) jf906758 jf906765 szarowska & falniowski (2011) alzoniella finalina spring at the porra river, molino, liguria, savona, italy (44.219, 8.255) af367686 af367650 wilke et al. (2001) arganiella pescei santa susanna springs, rivodutri, latium, italy (42.49513, 12.84604) mw561453 mw553909 present study arganiella tabanensis taban spring, montenegro (42.52795, 19.21921) mw561454 mw553910 present study arganiella wolfi virgen de los ángeles hermitage, sierra de aracena, huelva, spain (37.8785, -6.66681) mh348095 mh350192 delicado et al. (2019) avenionia brevis spring of the fountain of st.-victor-la coste, gard, france (44.057, 4.636) af367670 af367638 wilke et al. (2001) belgrandia thermalis thermal channel near s. giuliano, tuscany, pisa, s. giuliano terme, italy (43.751, 10.440) af367684 af367648 wilke et al. (2001) belgrandiella kusceri rakovski potok [crab stream], near rakovski skocjan, rakek, slovenia (45.77750, 14.18556) jx970574 kt218520 wilke et al. (2013); falniowski & beran (2015) boetersiella sturmi la mata spring, mata bejid, jaén, spain (37.69503, -3.50703) mh348097 mh350200 delicado et al. (2019) bullaregia tunisiensis a spring in djebba, province béja, tunisia (36.47125, 9.09972) mn575709 kx821683 khalloufi et al. (2017, 2020) chondrobasis levantina caroche spring, teresa de cofrentes, valencia, spain (39.09608, -0.91972) mh348098 mh350203 delicado et al. (2019) corbellaria celtiberica manubles river, soria, spain (41.60997, -1.95586) mh348099 mh350207 delicado et al. (2019) daphniola graeca daphne spring, about 30 km north of larissa, greece (39.89111, 22.60722) ef070624 eu047764 szarowska (2006); falniowski et al. (2007) deganta azarum la fontona spring, borondes, asturias, spain (43.33189, -6.01494) mh348100 mh350208 delicado et al. (2019) fissuria boui spring near la prouveresse, alpes maritimes, france (43.64279, 6.88735) af367690 af367654 wilke et al. (2001) graecoarganiella parnassiana small spring south of eptalofos, parnassus mountains, greece (38.59278, 22.5039) jn202341 jn202348 falniowski & szarowska (2011a) graziana alpestris spring at the porra river, liguria, savona, molino, italy (44.219, 8.255) af367673 af367641 wilke et al. (2001) grossuana delphica kastalia spring at delphi, greece (38.48306, 22.50528) ef061917 ef061922 szarowska et al. (2007) hauffenia tellinii isonzo river near sagrado spring, friuli-venetia julia, gorizia, italy (45.8743, 13.4856) af367672 af367640 wilke et al. (2001) horatia klecakiana studenci spring, n of kučiće, in the valley of the cetina river, croatia (43.44481, 16.80708) kj159127 kj159128 szarowska & falniowski (2014) iberhoratia morenoi prado del rey spring, cádiz, spain (36.66042, -5.45028) mh348101 mh350210 delicado et al. (2019) european journal of taxonomy 750: 140–155 (2021) 144 2006 and milesiana arconada & ramos, 2006; quiñonero-salgado & rolán (2017) for shells of navalis quiñonero-salgado & rolán, 2017; boeters et al. (2019) for salaeniella boeters, quiñonero-salgado & ruiz-cobo, 2019; arconada & ramos (2002) for spathogyna arconada & ramos, 2002; ramos et al. (2000) for tarraconia ramos & arconada, 2000; falniowski & szarowska (2011a) for graecoarganiella falniowski & szarowska, 2011; radea et al. (2013) for isimerope radea & parmakelis, 2013; radea et al. (2016) for myrtoessa radea, 2016 and bodon et al. (2001) for the remaining genera. character and character states were coded following the terminology of hershler & ponder (1998). results molecular analyses the data matrix constructed of coi (658 bp) and 18s (492 bp) sequences yielded an alignment with a total length of 1150 bp. our ml and bi analyses generated similar tree topologies and branch supports (fig. 1). in both inferences, arganiella did not form a monophyletic group; instead, both a. wolfi and a. tabanensis were distantly related to the type species a. pescei. also in both analyses, a. wolfi resolved as the sister taxon to the genus iberhoratia from the iberian peninsula (bs = 100%, bpp = 1.00), and a. tabanensis, to the genus kerkia radoman, 1978 from the balkan peninsula and adjacent islands (bs = 91%, bpp = 0.95). the phylogenetic position of a. pescei was not well resolved in either of the phylogenetic analyses. sequence divergence among the three species of arganiella ranged from 14.5 to 16.7% for coi and 2.0 to 3.8% for 18s. divergence among genera ranged from 9.1 to 22.5% for coi and 0 to 4.5% for 18s. overall, our results show that dna sequence divergence values among the three species of arganiella fall within the range of inter-generic variation and that both a. wolfi and a. tabanensis not only have a sister group relationship with a genus from their respective mediterranean peninsulas (fig. 1) but also table 1 (continued). taxon locality genbank # 18s genbank # coi original reference islamia piristoma spring on right bank of magra river near romito, liguria, la spezia, arcola, italy (44.1042, 9.9337) af367671 af367639 wilke et al. (2001) josefus aitanica flores spring, requena, valencia, spain (39.48694, -1.12205) mh348107 mh350231 delicado et al. (2019) kerkia kusceri cave krska jama, krka, ivancna gorica, slovenia (45.89, 14.77111) ky087833 ky087867 rysiewska et al. (2017) mercuria similis la puebla, mallorca, spain (39.79111, 3.10472) af212913 af213346 wilke et al. (2000) milesiana schuelei spring in gaucin, málaga, spain (36.51878, -4.67703) – mh350247 delicado et al. (2019) spring in berchul, félix, almería, spain (36.87719, -2.66561) mh348108 – delicado et al. (2019) pauluccinella minima lago di piediluco, s. egidio, italy jx970578 jx970612 wilke et al. (2013) sadleriana fluminensis jadro river at solin near split, croatia (43.5453, 16.48780) af367683 – wilke et al. (2001) sava river, slovenia (46.17344, 14.41505) – kf193083 szarowska & falniowski (2013) spathogyna fezi roble spring, yémeda, cuenca, spain (39.74281, -0.28383) mh348109 mh350251 delicado et al. (2019) tarraconia gasulli la esperanza spring, navajas, castellón, spain (39.86990, -0.50719) mh348110 mh350254 delicado et al. (2019) delicado d. et al., systematics of the hydrobiid genus arganiella 145 clear morphological differences with these sister genera, indicating the two species are representatives of distinct genera (table 2). this evidence, altogether, demonstrates the need to revise the genus assignment of the non-type species a. wolfi and a. tabanensis. taxonomy phylum mollusca cuvier, 1795 class gastropoda cuvier, 1795 superorder caenogastropoda cox, 1960 superfamily truncatelloidea gray, 1840 family hydrobiidae stimpson, 1865 genus aretiana delicado & ramos gen. nov. urn:lsid:zoobank.org:act:7b207b6d-47e8-4211-8139-0cf98da353d7 type species boetersiella wolfi boeters & glöer, 2007. fig. 1. maximum likelihood tree based on the combined (coi, 18s) dataset. statistical support of the nodes is indicated when bayesian posterior probabilities ≥ 0.95 and bootstrap supports ≥ 75% (black dots). on the right, the morphology of shell, penis and distal female genitalia is presented for: a) arganiella wolfi boeters & glöer, b) a. pescei giusti & pezzoli, 1980 and c) a. tabanensis boeters, glöer & pešić, 2014. reproductive organs were re-drawn from arconada & ramos (2007a), bodon et al. (2001) and boeters et al. (2014) for the three species, respectively. scale bar below topology: substitutions per site. http://zoobank.org/urn:lsid:zoobank.org:act:7b207b6d-47e8-4211-8139-0cf98da353d7 european journal of taxonomy 750: 140–155 (2021) 146 revised diagnosis shell trochiform; whorls 3.5–4.0; aperture complete, rounded; outer lip narrow, straight in lateral profile; umbilicus wide. operculum corneous, yellowish, thin, pliable, oval to rounded, paucispiral with a central nucleus, without peg. two pairs of basal cusps on each central radular tooth. ctenidium well developed, with approximately 14 gill filaments. osphradium positioned opposite to approximately the middle of the ctenidium. stomach without gastric caecum; rectum forms a gentle u-shape in the mantle cavity. bursa copulatrix medium-sized, pyriform, pedunculated and protruding beyond the posterior edge of the albumen gland; bursal duct shorter than bursal length; unpigmented renal oviduct that makes a complete loop over the pallial gland; one elongated or pyriform seminal receptacle arising from the renal oviduct, just above the insertion point with the bursal duct. prostate gland bean-shaped, about twice as long as wide. penis small and simple, gradually tapering. nervous system unpigmented. etymology the genus is named after villa aretiana, the roman name of the town aracena, which gives its name to the mountain range where the genus was found (i.e., sierra de aracena); gender feminine. remarks aretiana delicado & ramos gen. nov. can be distinguished from arganiella as the former has a taller shell, a more oval operculum, pigmentation on the body and eyespots, fewer gill filaments, a narrower penis, a larger and pyriform bursa copulatrix located beyond the posterior edge of the albumen gland and fewer cusps on the lateral radular teeth (for comparison, see giusti & pezzoli 1980; boeters & glöer 2007; arconada & ramos 2007a). the new genus differs from the closely related genus iberhoratia by its taller shell with a narrower umbilicus, the absence of lobes on the inner edge of the penis and of a proximal seminal receptacle (sr2) and presence of two pairs of basal cusps on each central radular tooth (see arconada et al. 2007). mean coi divergence for aretiana delicado & ramos gen. nov. was 15.9% with arganiella and 11.2% with iberhoratia. genus docleiana delicado & pešić gen. nov. urn:lsid:zoobank.org:act:04639db3-91d2-46b7-bfab-ffc1ea3c27bd type species arganiella tabanensis boeters, glöer & pešić, 2014. revised diagnosis shell valvatiform; whorls ca 3; aperture complete, from rounded to ellipsoidal; outer lip narrow, straight in lateral profile; umbilicus wide. operculum corneous, yellowish, thin, pliable, rounded, paucispiral with a central nucleus, without peg. two pairs of basal cusps on each central radular tooth. ctenidium well developed, with 10–11 gill filaments. animal unpigmented. osphradium positioned opposite approximately to the middle of the ctenidium. stomach without gastric caecum; rectum forms a gentle v-shape in the mantle cavity. bursa copulatrix small, globular, pedunculated and positioned beyond the posterior edge of the albumen gland; bursal duct longer than bursal length; unpigmented renal oviduct; one pyriform seminal receptacle arising at the insertion point with the bursal duct loop. penis small and simple, gradually tapering. etymology the new genus is named after doclea, the name of the roman city located on the site of modern podgorica on whose municipal territory a new genus was found; gender feminine. http://zoobank.org/urn:lsid:zoobank.org:act:04639db3-91d2-46b7-bfab-ffc1ea3c27bd delicado d. et al., systematics of the hydrobiid genus arganiella 147 remarks docleiana delicado & pešić gen. nov. can be distinguished from arganiella as the former has a more ellipsoidal shell aperture, a narrower base of the penis, a smaller bursa copulatrix that is positioned beyond the posterior edge of the albumen gland, a larger seminal receptacle and more cusps on the lateral radular teeth (fig. 2; for comparison, see giusti & pezzoli 1980; boeters et al. 2014). the new genus differs from the closely related genus kerkia by its smaller shell, absence of lobes on the inner edge of the penis, a smaller bursa copulatrix, a more globular seminal receptacle and v-shaped rectum (see radoman 1978; bodon et al. 2001). mean coi sequence divergence for docleiana delicado & pešić gen. nov. was 14.5% with arganiella and 12.3% with kerkia. discussion previous taxonomic studies of valvatiform hydrobiids have relied on morphological similarities to classify species, a practice that can be difficult for this group because of their small size and simplified morphologies. considering this, more recent systematic revisions have incorporated molecular fig. 2. shells, operculum and radulae of docleiana tabanensis (boeters, glöer & pešić, 2014) (ugsb 18847). a–c. shells. d–e. operculum (d = inner side; e = outer side). f. protoconch. g–i. radulae. g. portion of radula ribbon. h. central radular teeth. i. inner and outer marginal teeth. european journal of taxonomy 750: 140–155 (2021) 148 methods to re-evaluate the taxonomy of valvatiform hydrobiid species and have indeed revealed genus misidentifications. some examples are the species hauffenia plana (bole, 1961), later transferred to the genus bracenica radoman, 1973 by hofman et al. (2020); horatia hadei gittenberger, 1982 transferred to daphniola radoman, 1973 by falniowski & szarowska (2011b); and neohoratia azarum boeters & rolán, 1988, later recognized as islamia azarum (boeters & rolán, 1988) by arconada & ramos (2006) and currently combined as deganta azarum by arconada & ramos in delicado et al. (2019). in the case of arganiella, a. wolfi and a. tabanensis, two endemic species from the iberian and balkan peninsulas, respectively, were classified within this genus on the basis of shell and genital similarities with the apennine species a. pescei (arconada & ramos 2007a, 2007b; boeters et al. 2014). our survey of dna sequence divergence within arganiella, although based on a limited number of samples and gene fragments, showed the substantial divergence of these two species from the type species a. pescei (> 14.5% for coi). this degree of dna sequence differentiation is comparable to those inferred between other recognized valvatiform genera (uncorrected p-distances 9.1–22.5% for coi). moreover, arganiella was not recovered as a monophyletic group by our phylogenetic inferences (fig. 1). however, incorrect systematic conclusions can be drawn from molecular phylogenies when the specimens have previously been misidentified (e.g., radomaniola/horatia in szarowska & falniowski, 2014). detailed morphological examinations of the studied material are therefore needed for a more reliable systematic interpretation. our comparative morphological study (table 2) indicated close similarities among the, until now, considered species of arganiella, especially in those characters related to the penis, ctenidium and radula and, to a lesser degree, the shell, pigmentation and distal genitalia of females. some character states found in species of arganiella, such as a simple penis without lobes, the presence of a single distal seminal receptacle and two cusps at both sides of the basis of the central radular tooth, are rarely present in other valvatiform genera (bodon et al. 2001; radea et al. 2016) and can, thus, lead to genus misidentification. however, differences in other characters can be observed among the three species. the most dissimilar species is a. wolfi: it has a larger and more trochiform shell, body pigmentation and a larger bursa copulatrix (arconada & ramos 2007a; boeters & glöer 2007). arganiella tabanensis is more similar to a. pescei than a. wolfi, especially in shell shape and body pigmentation (boeters et al. 2014). however, it differs from a. pescei in shell size, the shape of the bursa copulatrix and the bend type of the rectum. on the basis of this dna and morphologic evidence, we assign the species a. wolfi and a. tabanensis to two distinct genera. we base the establishment of the two new genera, aretiana delicado & ramos gen. nov. for arganiella wolfi and docleiana delicado & pešić gen. nov. for arganiella tabanensis, on the high level of dna sequence divergence and phylogenetic position of the two species (fig. 1), as well as on their unique combination of morphological characters (table 2). given the restricted geographic distribution of most valvatiform hydrobiids (bodon et al. 2001), we focus our taxonomic decision on comparisons with geographically close groups. most genera of iberian valvatiform hydrobiids were included in our phylogenetic study, except navalis, which is only known by empty shells, and salaeniella, for which the females are still unknown and no well-preserved specimens are available for molecular studies. regardless, these genera are morphologically very distinct from aretiana delicado & ramos gen. nov. and have distant distribution ranges within the iberian peninsula (quiñonero-salgado & rolán 2017; boeters et al. 2019). according to our comparative morphological study, the most similar iberian genus to aretiana delicado & ramos gen. nov. is boetersiella. these two genera are highly divergent genetically and display clear differences in shell and female genital features, validating them as different genera. dna sequence data of valvatiform genera living in the balkan springs are still scarce. although our phylogenetic analyses recovered docleiana delicado & pešić gen. nov. as an independent lineage sister to the balkan genus kerkia, this relationship may change as more valvatiform species are sequenced and analysed. until then, morphological characters support the placement of a. tabanensis into a new genus. the balkan genera hadziella kuščer, 1932 and dabriana radoman, 1974, which have not yet delicado d. et al., systematics of the hydrobiid genus arganiella 149 ta bl e 2 (c on tin ue d on n ex t p ag e) . m or ph ol og ic al c ha ra ct er s re co rd ed fr om th e va lv at if or m h yd ro bi id g en er a di st ri bu te d ov er th e ib er ia n, a pe nn in e an d b al ka n pe ni ns ul as . t hi s da ta s et is a n ex te ns io n of ta bl e 2 in cl ud ed in r ad ea e t a l. (2 01 6) . ta xa sl sw sk u w a s o p b ac b c sr (s ) sr p p p l p p p s c t e ye s r ec tu m ib er ia n pe ni ns ul a a re tia na g en . n ov . 1. 7 1. 7 0 2 0 0 2 3 1 0 0 0 0 0 1 1 s b oe te rs ie lla 1. 6 1. 5 0 1 1 0 1 2 1 0 0 0 0 0 0 1 u c ho nd ro ba si s 1. 3 1. 2 0 1 1 0 1 3 1 0 1 1 0 0 0 1 u c or be lla ri a 0. 5 1. 3 0 2 0 0 1 3 2 2 1 2 0 0 0 0 s d eg an ta 1. 7 1. 6 0 1 0 0 2 3 1 0 2 1, 2 0 0 1 1 u ib er ho ra tia 1. 5 1. 8 0 2 0 0 1 1 2 2 1 2 0 0 1 1 u o r s jo se fu s 1. 5 1. 6 0 1 1 0 2 0 2 1 1 4 0 0 0 1 u m ile si an a 1. 1 1. 5 0 2 0 0 2 0 2 1 1 2 0 0 1 1 u n av al is 0. 8 1. 0 1 2 0 – – – – – – – – – – – – sa la en ie lla 1. 2 1. 6 0 2 1 0 – – – – 1 2 0 0 1 0 s sp at ho gy na 1. 0 1. 4 0 2 1 0 1 1 2 2 1 2 0 0 1 1 v ta rr ac on ia 1. 8 1. 8 0 2 0 0 1 3 0 0 1 2 0 0 1 1 u a pe nn in e pe ni ns ul a a rg an ie lla 1. 0 2. 0 0 2 0 0 2 1 1 0 0 0 0 0 1 0 s f is su ri a* 1. 6 1. 8 0 2 0 0 1 1 2 2 3 1, 3 1 0 1 0 s h au ffe ni a* 1. 2 2. 3 0 2 0 1 1 3 1 0 0, 1 0, 4 0 1 0, 1 0 z p ez zo lia 1. 0 1. 4 0 2 1 0 0 0, 3 2 2 0 0 0 0 0 0 s b al ka n pe ni ns ul a d oc le ia na g en . n ov . 1. 1 1. 5 0 2 0 0 2 2 1 0 0 0 0 0 1 0 z b ra ce ni ca 1. 0 2. 0 0 2 0 1 1 3 2 2 1 2 0 0 – 0 – d ab ri an a 2. 2 2. 5 0 2 0 – 0 3 1 0 0 0 0 0 0 0 – d al m at el la 1. 6 2. 2 1 2 0 – – – – – – – – – – – – d ap hn io la 1. 3 1. 2 0 1 1 0 1 1 2 2 1 2 0 0 1 1 – g oc ea 0. 9 1. 6 0 2 1 1 1 3 1 0 0 0 0 0 0 0 z g ra ec oa rg an ie lla 1. 2 1. 5 0 2 1 0 1 3 2 2 1 1 1 0 0 1 – european journal of taxonomy 750: 140–155 (2021) 150 ta bl e 2 (c on tin ue d) . ta xa sl sw sk u w a s o p b ac b c sr (s ) sr p p p l p p p s c t e ye s r ec tu m h ad zi el la 0. 6 1. 8 0 2 1 1 1 1 1 0 0 0 0 0 0 0 z h or at ia 2. 1 2. 1 0 1 1 0 1 1 2 2 1 2 0 0 1 1 0 is im er op e 1. 6 2. 0 0 2 1 0 1 3 0 0 1 3 1 0 0 1 u is la m ia * 2. 0 1. 8 0 0 0 0 1 0 2 2 1 4 0 0 1 1 u k ar ev ia 1. 1 1. 7 1 2 2 0 1 3 2 2 1 3 0 0 – 1 – k er ki a 1. 4 2. 3 0 2 0 1 2 1 1 0 1 3 0 0 1 0 s ly nh id ia 1. 2 1. 1 0 0 2 0 1 1 1 0 1 4 0 0 – 1 – m yr to es sa 1. 1 1. 4 0 2 0 0 1 1 2 2 0 0 2 0 1 1 v o hr id oh au ffe ni a 1. 0 1. 5 1 2 1 0 1 1 2 2 1 3 0 0 – 1 – o hr ig oc ea 0. 8 1. 2 0 2 0 0 1 3 2 2 1 3 0 0 – 1 – p re sp ol ito re a 1. 0 1. 3 0 2 0 0 1 2 2 2 1 3 0 0 – 1 – p se ud oh or at ia 1. 3 1. 4 0 1 0 1 1 2 2 2 1 3 0 0 1 1 0 p se ud oi sl am ia 1. 1 1. 3 0 2 1 0 1 1 2 2 1 4 0 0 – 1 – st ru gi a 1. 6 1. 7 0 2 0 0 1 1 1 0 1 3 0 0 – 1 – za um ia 1. 5 1. 2 0 0 0 0 1 1 1 0 1 4 0 0 – 0 – * g en er a di st ri bu te d ou ts id e of th e ca te go ri ze d re gi on s. c on tin uo us (i n m m ) a nd d is cr et e (o r c at eg or ic al ) c ha ra ct er s: a s = ap er tu re s ha pe : r ou nd ed (0 ), ov al (1 ), sq ua re (2 ); b ac = p ai rs o f b as al c us ps o n th e ce nt ra l r ad ul ar to ot h; b c = b ur sa c op ul at ri x: a bs en t ( 0) , o vo id ( 1) , g lo bu la r (2 ), py ri fo rm ( 3) ; c t = c te ni di um : a bs en t ( 0) , p re se nt ( 1) ; e ye s: a bs en t ( 0) , p re se nt ( 1) ; o p = op er cu lu m pe g: a bs en t ( 0) , p re se nt ( 1) ; p = p en is : s im pl e w ith ou t l ob es ( 0) , w ith o ne lo be ( 1) , w ith tw o lo be s (2 ), w ith m or e th an tw o lo be s (3 ); p l = pe ni al lo be (s ): a bs en t ( 0) , ba sa l l ob e (1 ), m ed ia l l ob e (2 ), lo be a t ⅔ o f pe ni s le ng th ( 3) , a pi ca l l ob e (4 ); p p = pe ni al p ap ill a: a bs en t ( 0) , p re se nt ( 1) ; p s = pe ni al s ty le t: ab se nt ( 0) , p re se nt ( 1) ; r ec tu m : w ith ou t b en d (0 ), slik e (s ), u -l ik e (u ), v -l ik e (v ), z -l ik e (z ); s k = s he ll ke el s or o th er o rn am en ts : a bs en t ( 0) , p re se nt ( 1) ; s l = sh el l l en gt h (m ax im um re co rd ed v al ue fo r t he ty pe s pe ci es ); s r = s em in al re ce pt ac le : a bs en t ( 0) , o ne s em in al re ce pt ac le (1 ), tw o se m in al re ce pt ac le s (2 ); s r p = se m in al re ce pt ac le p os iti on : ab se nt (0 ), cl os e to e ac h ot he r ( 1) , d is ta nt ly p os iti on ed fr om e ac h ot he r ( 2) ; s w = s he ll w id th (m ax im um re co rd ed v al ue fo r t he ty pe s pe ci es ); u w = u m bi lic us w id th : ve ry n ar ro w (0 ), na rr ow (1 ), w id e (2 ); – = n o da ta . c on tin uo us v ar ia bl es p ro vi de d in m m . delicado d. et al., systematics of the hydrobiid genus arganiella 151 been sequenced, resemble docleiana delicado & pešić gen. nov. in penis morphology but differ in shell size and radular and female genital features; they also lack a ctenidium (table 2). the systematic findings suggesting erroneous assignment of the three geographically disjunct species to the same genus due to morphological similarities conflict with the use of traditional taxonomy to classify valvatiform hydrobiid taxa and highlight the need to integrate morphological and molecular data for more robust taxonomic assessments. acknowledgements we thank s. cianfanelli for collecting and depositing a sample of a. pescei in the ugsb collection at justus liebig university, giessen, and a. möbus and s. agel (imaging unit, biomedical research centre seltersberg, justus liebig university, giessen) for their assistance with the esem photomicrographs. we are also grateful to m. haase, another anonymous reviewer and the section editor t. backeljau for their helpful comments on earlier versions of the manuscript. the english was reviewed by m. modrell. this work was funded by a german science foundation (dfg) grant (de 2605/1-1) to d. delicado and the fauna iberica xii project (pgc2018-095851-b-c61) to m.a. ramos. references akaike h. 1974. a new look at the statistical model identification. ieee transactions on automatic control 19 (6): 716–723. https://doi.org/10.1109/tac.1974.1100705 arconada b. & ramos m.a. 2001. new data on hydrobiidae systematics: two new genera from the iberian peninsula. journal of natural history 35: 949–984. https://doi.org/10.1080/002229301300323884 arconada b. & ramos m.a. 2002. spathogyna, a new genus for valvata (? tropidina) fezi altimira, 1960 from eastern 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https://doi.org/10.1635/0097-3157%282001%29151%5b0001:msohmg%5d2.0.co%3b2 https://doi.org/10.1016/j.ympev.2012.10.025 https://doi.org/10.1093/molbev/msy073 https://doi.org/10.1016/s1055-7903%2802%2900326-3 delicado d. et al., systematics of the hydrobiid genus arganiella 155 published on: 21 may 2021 topic editor: rudy jocqué section editor: thierry backeljau desk editor: pepe fernández printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. a new species of gyrodactylus (monogenea, gyrodactylidae), an ectoparasite from the endemic iranocichla hormuzensis (teleostei, cichlidae), the only iranian cichlid maarten p.m. vanhove*,1,7, walter a. boeger2, fidel muterezi bukinga3, filip a.m. volckaert4, tine huyse5 & antoine pariselle6 1,4,5 laboratory of biodiversity and evolutionary genomics, department of biology, ku leuven, charles deberiotstraat 32, b-3000 leuven, belgium. emails: maarten.vanhove@bio.kuleuven.be, filip.volckaert@bio.kuleuven.be, tine.huyse@bio.kuleuven.be 2 laboratório de ecologia molecular e parasitologia evolutiva, departamento de zoologia, setor de ciências biológicas, universidade federal do paraná, caixa postal 19073, cep 81531-980, curitiba, pr, brazil. email: wboeger@gmail.com 3 section de parasitologie, département de biologie, centre de recherche en hydrobiologie, b.p. 73, uvira, democratic republic congo, via b.p. 254, bujumbura, burundi. email: fidelmuterezi@yahoo.fr 6 ise-m, umr5554 cnrs, ur226 ird, université montpellier ii – cc 063, f-34095 montpellier cedex 5, france. email: antoine.pariselle@ird.fr 7 ichthyology unit, african zoology department, royal museum for central africa, leuvensesteenweg 13, b-3080 tervuren, belgium. *corresponding author, email: maarten.vanhove@bio.kuleuven.be 1 urn:lsid:zoobank.org:author:f40a361d-f45c-4c91-86af-115962126120 2 urn:lsid:zoobank.org:author:318e38be-6ab8-4d77-bfa9-36f5e8a546e0 3 urn:lsid:zoobank.org:author:b521076b-7ed5-4532-8eea-cb33bf4019f7 4 urn:lsid:zoobank.org:author:ac726421-5168-4787-a939-0ccbcbee5114 5 urn:lsid:zoobank.org:author:72a8d565-7dcb-4103-a4b9-84248e88ff4b 6 urn:lsid:zoobank.org:author:de920c99-1505-4b1f-a523-e375b1dc6201 abstract. iranocichla hormuzensis occupies a biogeographically peculiar position. this endemic of southern iran is the only iranian cichlid. while it is phylogenetically related to african oreochromine members of the cichlid family, it remains unclear how it has dispersed into its current range. it is one of the many lasting enigmas of cichlid biogeography. monogenean fish parasites may provide useful additional information in such cases. therefore, i. hormuzensis was examined for these flatworms. a gyrodactylid parasite is reported and compared to congeners from the palearctic and from cichlids. in this way, we verify whether it shows affinities to parasites from fishes that are either biogeographically or phylogenetically close to iranocichla hormuzensis. the species is new to science and is described as gyrodactylus jalalii sp. nov. this is the first description of a parasite infecting i. hormuzensis. because of the fixation method or age of the material, dna could not be isolated. due to the lack of genetic data, no conclusions can be drawn on its phylogenetic positioning. indeed, gyrodactylus phylogeny cannot be european journal of taxonomy 30: 1-10 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2012.30 www.europeanjournaloftaxonomy.eu 2012 · vanhove m.p.m. et al.. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:489a6bc2-795c-4945-a642-440d4f92a126 1 mailto:maarten.vanhove%40bio.kuleuven.be?subject=maarten.vanhove%40bio.kuleuven.be mailto:filip.volckaert%40bio.kuleuven.be?subject=filip.volckaert%40bio.kuleuven.be mailto:tine.huyse%40bio.kuleuven.be?subject=tine.huyse%40bio.kuleuven.be mailto:wboeger%40gmail.com?subject=wboeger%40gmail.com mailto:fidelmuterezi%40yahoo.fr?subject=fidelmuterezi%40yahoo.fr mailto:antoine.pariselle%40ird.fr?subject=antoine.pariselle%40ird.fr mailto:maarten.vanhove%40bio.kuleuven.be?subject=maarten.vanhove%40bio.kuleuven.be http://zoobank.org/urn:lsid:zoobank.org:author:f40a361d-f45c-4c91-86af-115962126120 http://zoobank.org/urn:lsid:zoobank.org:author:318e38be-6ab8-4d77-bfa9-36f5e8a546e0 http://zoobank.org/urn:lsid:zoobank.org:author:b521076b-7ed5-4532-8eea-cb33bf4019f7 http://zoobank.org/urn:lsid:zoobank.org:author:ac726421-5168-4787-a939-0ccbcbee5114 http://zoobank.org/urn:lsid:zoobank.org:author:72a8d565-7dcb-4103-a4b9-84248e88ff4b http://zoobank.org/urn:lsid:zoobank.org:author:de920c99-1505-4b1f-a523-e375b1dc6201 http://dx.doi.org/10.5852/ejt.2012.30 http://www.europeanjournaloftaxonomy.eu/index.php/ http://creativecommons.org/licenses/by/3.0/ http://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:489a6bc2-795c-4945-a642-440d4f92a126 inferred from morphological characteristics alone. moreover, the congeners phenotypically reminiscent of the new species belong to a gyrodactylus clade which is highly diverse in geographic range and host choice. hence, there is no evidence linking the new species to an exclusively african or cichlid-bound gyrodactylus lineage. keywords. gyrodactylus jalalii sp. nov., oreochromini, platyhelminthes, perciformes, species description. vanhove m.p.m., boeger w.a., muterezi bukinga f., volckaert f.a.m., huyse t. & pariselle a. 2012. a new species of gyrodactylus (monogenea, gyrodactylidae), an ectoparasite from the endemic iranocichla hormuzensis (teleostei, cichlidae), the only iranian cichlid. european journal of taxonomy 29: 1-10. http://dx.doi.org/10.5852/ ejt.2012.30 introduction the fascination and scientific interest that cichlids raise in many scientists and hobbyists is well reviewed by barlow (2000). this author aptly shows the importance of cichlid fishes in a wide range of fields in evolutionary biology, from ecology to behavior and from developmental biology to speciation. numbering about 1350 described species, with hundreds remaining to be discovered (nelson 2006), these “model” fishes are the most species-rich vertebrate family worldwide (kocher 2004). counting only described species, cichlidae is the largest non-ostaryophysan freshwater fish family (kullander 2003). its species richness reaches its peak in the more than 900 african representatives. some relatives to african cichlids occur in the middle east (belonging to astatotilapia pellegrin, 1904; oreochromis günther, 1889; sarotherodon rüppell, 1852; tilapia smith, 1840 and tristramella trewavas, 1942) (werner & mokady 2004) and one in iran. the latter is iranocichla hormuzensis coad, 1982, the only representative of the monotypic iranocichla coad, 1982. the species was first reported by behnke (1975) and saadati (1977) before being formally described (coad 1982). this southern iranian endemic cichlid is found in mostly saline riverine systems draining into the strait of hormuz (persian gulf). the question rises how it arrived in this biogeographically isolated range. most cichlid species in the middle east have congeners among the african “tilapiine” (oreochromis, tilapia and sarotherodon: schwarzer et al. 2009) or haplochromine (astatotilapia: werner & mokady 2004) cichlids. tristramella and iranocichla, however, are endemic genera. phylogenetically, they are also placed in oreochromini, one of the tribes into which “tilapiine” cichlids were split by schwarzer et al. (2009). thus, they are closely affiliated to african cichlids. middle-eastern cichlids at present only occur in the levant and these iranian systems. this fact might indicate a relict-type consequence of a recent (pleistocene to recent) aridification event and decrease in temperature (overview in coad 1982). furthermore, the persian gulf was a river valley between 90,000 and 10,000 years ago. see also arndt et al. (2003) for a discussion on how the low level of the mediterranean during pleistocene episodes is suggested to have allowed freshwater dispersal from the nile to the levant. although those conditions would have allowed dispersal of cichlids, it is also possible that the marine environment would not represent a barrier to the spread of these animals, anyway (see references below). indeed, occupying mostly saline streams shows that i. hormuzensis is salt-tolerant (coad 1982). moreover, several “tilapiines” are known to disperse between rivers through brackish coastal waters (nelson 2006). conversely, murray (2001) favors marine dispersal through the tethys sea/indian ocean to coastal dispersal in view of the absence of cichlids along the rest of the region’s coastlines. either way, both freshwater and marine dispersal pathways could explain the presence of this cichlid in waters draining into the strait of hormuz. be it on a local or a global scale, the history behind current cichlid distribution across continents has not been unequivocally unraveled. the roles of intercontinental european journal of taxonomy 30: 1-10 (2012) 2 http://dx.doi.org/10.5852/ejt.2012.30 http://dx.doi.org/10.5852/ejt.2012.30 dispersal, on the one hand, and of vicariance (following the break-up of gondwana) and intra-continental freshwater dispersal, on the other hand, are still highly debated. a potential source of information that remains largely unexplored is the (monogenean) parasite fauna of cichlid species (reviewed in pariselle et al. 2011). many parasites have a close association with their host and a shorter generation time than their host, hence providing an alternative perspective on host evolution. ectoparasitic monogenean flatworms often do not tolerate substantial changes in salinity, as indicated by their distribution and phylogenetic patterns worldwide (although there are exceptions, e.g., this is less the case in gyrodactylidae, certainly when infecting euryhaline hosts, see below). one might therefore hypothesize that certain freshwater monogeneans rarely survive dispersal through barriers represented by marine environments. hence, the phylogenetic relationships within a lineage of monogenean ectoparasites reflect both continental patterns and host phylogeny. freshwater fish hosts most likely retain their monogenean assemblage when dispersing through continental surface waters. conversely, it was suggested that most freshwater fishes might lose their autochtonous parasites during marine dispersal events. this would allow subsequent recolonisation by “local” parasite species once arriving in freshwater at the other end of a marine barrier (pariselle et al. 2011). here, a gyrodactylus species parasitizing i. hormuzensis is described. we analyze its affinity to congeners infecting other cichlids and to species known from fishes of the same region. because of the ability to switch hosts, gyrodactylids are expected to mainly mirror continental affinities rather than the relationships between its host species (boeger et al. 2003), depending of course on the time-scale and the dispersal routes used. the result might hence shed light on the pathway cichlids used to reach southern iran. material and methods eight host fishes (fixed and stored in ethanol or in formaldehyde) were retrieved from collections of the royal museum for central africa (tervuren, belgium) (mrac b2-28-p-1-8). branchial arches, body, fins and recipient were inspected for parasites under a wild m8 stereomicroscope. monogeneans were removed with a dissection needle. they were treated with 10% sodium dodecyl sulphate for a couple of minutes, before being fixed using ammonium picrate-glycerine (malmberg 1957) and mounted on a slide under a cover-slip. measurements were adapted from shinn et al. (2004) using a zeiss axio imager z1 microscope at a magnification of 100 x (oil immersion, 10 x ocular) under differential interference contrast, with an axiocammr3 camera and axiovision v.4.2.8 software. for drawings, some specimens were stained with gomori’s trichrome and mounted in canada balsam for study of their soft anatomy; other specimens were cleared and mounted in hoyer’s or gray and wess’ media for study of their sclerotized structures (all solutions prepared as in humason, 1979). illustrations were prepared with the aid of a digital camera (5 megapixels) and a projector attached to an olympus bx51 microscope equipped with phase contrast. taxon and author names in this study follow eschmeyer (2012) for hosts and monodb (http://www.monodb.org) for monogenea. results a single (incomplete and therefore unfortunately unidentified) ancyrocephaline monogenean (dactylogyridae, ancyrocephalinae) was recovered. all other monogeneans found on the i. hormuzensis specimens belong to gyrodactylus. vanhove m.p..m. et al., gyrodactylus from the cichlid iranocichla hormuzensis 3 http://www.monodb.org class monogenea van beneden, 1858 subclass polyonchoinea bychowsky, 1937 order gyrodactylidea bychowsky, 1937 family gyrodactylidae van beneden & hesse, 1863 subfamily gyrodactylinae van beneden & hesse, 1863 genus gyrodactylus von nordmann, 1832 gyrodactylus jalalii sp. nov. urn:lsid:zoobank.org:act:a89d45dc-7ca2-4807-af2e-e34f9a9bb060 figs 1-2 etymology the species epithet, jalalii, honours prof. dr. behiar jalali jafari (1953-2010) (obituary in shamsi 2010). he was a researcher in aquatic animal health and fish parasitology at the veterinary department of the islamic azad university (iran) and a keen student of monogeneans. the authors express the hope that this patronym might serve as an indication for the respect and appreciation this kind man enjoyed from his colleagues. type material examined thirty-one specimens, twenty of which (ethanol-preserved) were used for measurements. the holotype (mnhn hel301) and paratypes (mnhn hel302–hel305) are deposited in the muséum national d’histoire naturelle (paris, france). paratypes are deposited in the natural history museum (london, united kingdom) (nhmuk 2012.9.10.1–2012.9.10.2), the royal museum for central africa (tervuren, belgium) (mrac mt: 37711–37713), the harold w. manter laboratory of parasitology (lincoln, nebraska) (hwml-49758) and the united states national parasite collection (beltsville, maryland) (usnpc 106050). type host iranocichla hormuzensis coad, 1982 (teleostei, perciformes, cichlidae). type locality mehran river, persian gulf basin (2009). infection site gill filaments, fins, eye. description (measurements in micrometres (µm) and angles in degrees (°); average ± standard deviation, followed by range and number of measurements in parentheses). body (fig. 1a) fusiform, 361.7 ± 64.4 (294.7-528.9; n = 13) long; greatest width 128.1 ± 21.3 (98.8164.0; n = 13). two head organs provided with single spicule each. cephalic glands lateral to pharynx. pharynx composed of two tandem, muscular bulbs. oesophagus short. digestive glands lateral to oesophagus. caeca two, non-confluent, reaching level of germarium. male copulatory organ (mco) (fig. 1b) armed with a broad-based, robust, recurved, apical spine 5.7 ± 1.1 (4.5-7.6; n = 8) long, 5-7 smaller flanking spines in a single row, becoming more slender from the terminal over the subterminal towards the median ones (terminology of garcía-vásquez et al. 2007). testis dorsal to germarium. european journal of taxonomy 30: 1-10 (2012) 4 http://zoobank.org/urn:lsid:zoobank.org:act:a89d45dc-7ca2-4807-af2e-e34f9a9bb060 germarium immediately posterior to uterus. uterus with up to 2 embryos. unicellular glands lateral to terminations of caeca, posterior to germarium. haptor elongate (fig. 1a, d). anchor (hamulus) (fig. 1e) 79.9 ± 4.4 (70.4-86.3; n = 20) long; point 33.9 ± 2.3 (28.3-38.4; n = 19) long; base (superficial root) 28.2 ± 3.9 (19.7-33.1; n = 20) long; deep root knob-like; groove proximally at the base of the anchor, serving as articulation to superficial (ventral) bar; shaft 49.5 ± 2.2 (45.6-55.0; n = 19) long, proximally 10.2 ± 0.8 (8.7-11.9; n = 20) wide, distally 5.7 ± 0.8 (4.5-7.4; n = 19) wide; point sharply curved, with aperture 30.5 ± 2.5 (26.2-35.4; n = 20), aperture angle 42.7 ± 3.2 (36.9-47.3; n = 19), inner aperture angle 46.9 ± 5.3 (30.1-53.3; n = 19); inner curve length 1.9 ± 0.8 (0.7-3.3; n = 18) with point curve angle 9.5 ± 4.5 (4.2-22.9; n = 18). ventral bar (fig. 1d) 33.5 ± 2.5 (29.8-37.7; n = 20) wide, 44.1 ± 3.9 (36.4-49.8; n = 20) long; anterior bilateral processes slender, pronounced and 6.0 ± 1.1 (4.2-7.8; n = 20) long with process to mid-length 10.4 ± 1.7 (7.5-14.8; n = 20). median portion of ventral bar fig. 1. gyrodactylus jalalii sp. nov. a. whole mount. b. male copulatory organ. c. marginal hook. d. anchor-bar complex. e. anchor. scale bars represent 50 µm (whole mount), 10 µm (marginal hook, mco) or 30 µm (anchor, anchor-bar complex). vanhove m.p..m. et al., gyrodactylus from the cichlid iranocichla hormuzensis 5 8.3 ± 1.2 (5.7-9.8; n = 20) long; shield (ventral bar membrane) 25.4 ± 3.3 (19.8-30.1; n = 20) long, subrectangular, clearly striated in much the same way as bar proper. deep (dorsal) bar 23.0 ± 2.8 (18.929.9; n = 20) wide, medially constricted and at connection to deep root of anchors. marginal hook (figs 1c, 2a, b) 31.8 ± 4.1 (25.8-43.5; n = 20) long, shank with small distal bulb, 26.9 ± 4.1 (22.1-40.0; n = 20) long; sickle (hooklet) 5.4 ± 0.3 (4.5-6.1; n = 20) long, 4.3 ± 0.4 (3.6-5.1; n = 20) wide proximally, 4.5 ± 0.5 (3.8-5.3; n = 20) distally; toe depressed, 2.0 ± 0.4 (1.4-3.0; n = 20) long; convex platform; concave base; round keel; point of sickle proper as long as shaft, forming an angle of about 90° from each other; aperture 5.2 ± 0.5 (4.5-6.4; n = 20); instep/arch height 0.6 ± 0.1 (0.4-0.8; n = 20). remarks in comparison with congeners parasitizing cichlids, the striated ventral bar proper and shield, as well as the conspicuous ventral bar processes, seem most reminiscent to g. zimbae vanhove, snoeks, huyse & volckaert, 2011. however, the anterolateral processes of the ventral bar of g. zimbae are more earshaped. in g. zimbae, the ventral bar shield is slender and rounded and the hooklet lacks an arched base (versus subrectangular shield and concave hooklet base in g. jalalii sp. nov.). other cichlid gyrodactylus with relatively large ventral bar processes include g. shariffi cone, arthur & bondad-reantaso, 1995 and g. yacatli garcía-vásquez, hansen, christison, bron & shinn, 2011. just like in g. jalalii sp. nov., point and shaft of their hooklet sickle are at a right angle. these species are easily distinguished from g. jalalii sp. nov. by the smaller size of their haptoral sclerites and the ventral bar in particular (e.g., anchor 47.5 and 48.4 long, ventral bar shield 14.4 and 8.5 long, in g. shariffi and g. yacatli respectively) (garcía-vásquez et al. 2011). it should be noted, however, that these two species were described from cultured oreochromis niloticus (linnaeus, 1758) in the philippines, resp. mexico. the authors describing g. yacatli consider accidental infection or host switch a more likely scenario than an african origin (garcía-vásquez et al. 2011). hence, g. zimbae seems to be the most comparable cichlid parasite whose natural distribution is certainly african. comparison to palearctic congeners followed pugachev et al. (2009). the rather large ventral bar processes, in combination with the length of the marginal hooks, and mco armed with one large apical spine and one row of smaller spines of similar size, resemble the morphology of g. ophiocephali gussev, 1955 from channa argus (cantor, 1842) (perciformes: channidae) and cyprinus carpio linnaeus, 1758 (cypriniformes, cyprinidae), and to g. tokobaevi ergens & karabekova, 1980 from gymnodiptychus fig. 2. a. micrograph detailing the marginal hooks of gyrodactylus jalalii sp. nov. (photographed in utero, with an olympus bx50 microscope, using phase contrast, and olympus dp-soft v.3.2 software). b. idem, detail. scale bars represent 8 µm. european journal of taxonomy 30: 1-10 (2012) 6 dybowskii (kessler, 1874) (cypriniformes, cyprinidae). however, in g. ophiocephali and g. tokobaevi, the processes are longer than the ventral bar proper (median length, i.e., without the shield), which is not the case in g. jalalii sp. nov. elongate antero-lateral processes, albeit not longer than the ventral bar proper, are also found in g. hrabei ergens, 1957 and g. mariannae winger, hansen, bachmann & bakke, 2008, parasites of cottus linnaeus, 1758 spp. (scorpaeniformes, cottidae). these species, however, have a comparatively shorter anchor root than g. jalalii sp. nov. the longitudinal striae on the ventral bar shield as well as an overlap in size of anchor and marginal hook are reminiscent of g. lotae gussev, 1953 from lota lota (linnaeus, 1758) (gadiformes, lotidae). this species can be distinguished from g. jalalii sp. nov. because the new species has blunter and larger ventral bar processes, and a marginal hook sickle toe which joins smoothly into the platform, whereas this transition leaves a sharp “bump” in the platform in g. lotae. discussion just as barlow (2000) praises cichlids, early students of gyrodactylus expressed their high expectations for the scientific interest of these flatworms, as they recognized the species richness of this genus and the amount of work and skill it would take to adequately study it (von nordmann 1832; van beneden & hesse 1863). given this scientifically challenging position of both cichlids and gyrodactylids, g. jalalii sp. nov. was described, a monogenean ectoparasite of the only iranian cichlid i. hormuzensis. to the best of our knowledge, it is the first parasite to be formally described from this fish species. it brings the total number of gyrodactylus species described from cichlid hosts to 15 (paperna 1979; boeger & popazoglo 1995; christison et al. 2005; přikrylová et al. 2009, 2012; garcía-vásquez et al. 2011; vanhove et al. 2011). jalali et al. (2001) observed that the highly distinct zoogeographical regions of iran and the high degree of endemicity of its freshwater fishes gave rise to a diverse monogenean fauna. these authors did not mention gyrodactylus in their overview. indeed, jalali et al. (2005) noted that representatives of this genus from the endemic freshwater fishes of the country are basically unknown and undescribed. they did not report gyrodactylids from iranocichla. this cichlid seems distributed mostly in the mesopotamian faunal region of iran, from which jalali et al. (2001) reported a rather unexplored, endemic and specific monogenean fauna, containing both palearctic and african elements. for this reason, g. jalalii sp. nov. was compared to palearctic and african congeners. morphological similarities are unsurprisingly apparent (see above). haptoral morphology alone does not allow reliable phylogenetic inferences for species of gyrodactylus (ziętara & lumme 2004). unfortunately, we were unsuccessful in amplifying dna from g. jalalii sp. nov. hence, genetic data are at present unavailable for the new species, as is the case for several congeners used in this comparison. however, it is noteworthy that some of the aforementioned species show affinities in molecular phylogenetic reconstructions. indeed, g. mariannae is close to g. hrabei, which is genetically similar to g. flesi malmberg, 1957 (winger et al. 2008). this species clusters with g. lotae and g. zimbae, among other species, in a gyrodactylus clade containing representatives from various subgenera as defined by malmberg (1970) and from many continents and host taxa (ziętara & lumme 2004; vanhove et al. 2011). despite the need for genetic data, assuming, that g. jalalii sp. nov. is indeed related to this lineage, this fact alone does not make it a very suitable candidate to provide information on biogeographical dispersal pathways of cichlid fishes. indeed, several species also belonging to this diverse clade have been observed to show a broad salinity tolerance (g. branchialis huyse, malmberg & volckaert, 2004 and g. ostendicus huyse & malmberg, 2004: huyse et al. 2006) or the ability to switch infection sites (g. arcuatus bychowsky, 1933: raeymaekers et al. 2008) or hosts (g. arcuatus: huyse et al. 2003, 2006). however, while there is hence no proof that g. jalalii sp. nov. belongs to an exclusively african or cichlid-infecting lineage, this broad ecological spectrum enhances the potential of these parasites as biogeographical markers, conditional to the availability of molecular data. vanhove m.p..m. et al., gyrodactylus from the cichlid iranocichla hormuzensis 7 acknowledgements m.p.m.v. is a phd fellow, and t.h. a post-doctoral fellow, of the research foundation – flanders (fwovlaanderen). w.a.b. is a research fellow of cnpq – brazil. f.m.b. received financial support from ird (best) for a training session in belgium, during which part of the morphological work for this study was performed. prof. l. schoofs, prof. l. arckens and dr. e. meelkop (ku leuven) are thanked for the use of the zeiss microscope, and dr. sh. shamsi (charles sturt university, wagga wagga) for help in obtaining literature. s. asadollah (isfahan university of technology) and the late prof. b. jalali jafari (islamic azad university, teheran) are acknowledged for kindly providing us with iranocichla specimens and with information on this fish. two anonymous reviewers provided useful suggestions. references arndt a., van neer w., hellemans b., robben j., volckaert f. & waelkens m. 2003. roman trade relationships at sagalassos (turkey) elucidated by ancient dna of fish remains. journal of archaeological science 30: 1095-1105. http://dx.doi.org/10.1016/s0305-4403(02)00204-2 barlow g.w. 2000. the cichlid fishes. nature’s grand experiment in evolution. perseus publishing, cambridge, massachusetts. behnke r.j. 1975. fishes from the qanats of iran. in: 55th annual meeting: 75. american society of ichthyologists and herpetologists, williamsburg, virginia. boeger w.a. & popazoglo f. 1995. neotropical monogenoidea. 23. two new species of gyrodactylus (gyrodactylidae) from a cichlid and an erythrinid fish of southeastern brazil. memórias do instituto oswaldo cruz 90 (6): 689-694. http://dx.doi.org/10.1590/s0074-02761995000600006 boeger w.a., kritsky d.c. & pie m.r. 2003. context of diversification of the viviparous gyrodactylidae (platyhelminthes, monogenoidea). zoologica scripta 32 (5): 437-448. http://dx.doi.org/10.1046/j.14636409.2003.00130.x christison k.w., shinn a.p. & van as j.g. 2005. gyrodactylus thlapi n. sp. 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(crassiclitellata, lumbricidae) daniel f. marchán 1,3,*, thibaud decaëns 2,*, darío j. díaz cosín 3, mickaël hedde 4, emmanuel lapied 5 & jorge domínguez 6 1,6 grupo de ecoloxía animal (gea), universidade de vigo, e-36310 vigo, spain. 2 cefe, univ montpellier, cnrs, ephe, ird, univ paul valéry montpellier 3, montpellier, france. 3 department of biodiversity, ecology and evolution, faculty of biology, universidad complutense de madrid, madrid, spain. 4 umr eco&sols, inrae–ird–cirad–supagro montpellier, 2 place viala, 34060 montpellier cedex 2, france. 5 taxonomia biodiversity fund, 7 rue beccaria, 72012, paris, france. * corresponding authors: daniel.fernandez.marchan@uvigo.es; thibaud.decaens@cefe.cnrs.fr 3 email: dadico@ucm.es 4 email: mickael.hedde@inra.fr 5 email: elapied@yahoo.com 6 email: jdguez@uvigo.es 1 urn:lsid:zoobank.org:author:3b3731b6-b5fb-409a-a7a3-99fd0f96d688 2 urn:lsid:zoobank.org:author:b61f61b2-3012-4526-8ff9-dc94d372af77 3 urn:lsid:zoobank.org:author:38538b17-f127-4438-9de2-f9d6c597d044 4 urn:lsid:zoobank.org:author:f4a219f7-7e75-4333-8293-3004b3cd62c5 5 urn:lsid:zoobank.org:author:b1fb8658-dfc3-481c-a0be-b8488a018611 6 urn:lsid:zoobank.org:author:167575d5-d2cc-4b37-8b1d-0233e6b154e5 abstract. the area comprising the pyrenees, northeast spain, southern france and corsica-sardinia supports a large part of the diversity of lumbricidae earthworms, including most species of the endemic genera prosellodrilus, cataladrilus and scherotheca. in this region, the probability of encountering new species for science is significant, especially in scarcely sampled localities. in this study, we describe two unidentified species recently collected in the hyères archipelago (france), which we assigned to the genera cataladrilus and scherotheca based on morphological characters and molecular phylogenetic analyses. other species of scherotheca from montpellier (including the type species of the genus, sc. gigas gigas) were included in the analysis to clarify their conflicting systematics. a reduced molecular marker set (coi, 16s, 28s and nd1) proved as successful as larger marker sets for identifying phylogenetic relationships within the lumbricidae. remarkable disjunctions between both cataladrilus porquerollensis marchán & decäens sp. nov., scherotheca portcrosana marchán & decäens sp. nov. and their most closely related relatives, suggesting a strong influence of paleogeographic events on the earthworm fauna of the area and a possible role of near-shore islands as refugia for relict taxa. genetic distances and branch lengths supported the elevation of some subspecies of scherotheca to https://doi.org/10.5852/ejt.2020.701 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:d9291955-f619-46ea-90e1-da756d1b7c55 mailto:daniel.fernandez.marchan%40uvigo.es?subject= mailto:thibaud.decaens%40cefe.cnrs.fr?subject= mailto:dadico%40ucm.es?subject= mailto:mickael.hedde%40inra.fr?subject= mailto:elapied%40yahoo.com?subject= mailto:jdguez%40uvigo.es?subject= http://zoobank.org/urn:lsid:zoobank.org:author:3b3731b6-b5fb-409a-a7a3-99fd0f96d688 http://zoobank.org/urn:lsid:zoobank.org:author:b61f61b2-3012-4526-8ff9-dc94d372af77 http://zoobank.org/urn:lsid:zoobank.org:author:38538b17-f127-4438-9de2-f9d6c597d044 http://zoobank.org/urn:lsid:zoobank.org:author:f4a219f7-7e75-4333-8293-3004b3cd62c5 http://zoobank.org/urn:lsid:zoobank.org:author:b1fb8658-dfc3-481c-a0be-b8488a018611 http://zoobank.org/urn:lsid:zoobank.org:author:167575d5-d2cc-4b37-8b1d-0233e6b154e5 european journal of taxonomy 701: 1–22 (2020) 2 specific status as well as the retention of other subspecies, highlighting the importance of testing for such delimitation with molecular methods. keywords. earthworms, molecular phylogenetics, phylogeography, systematics. marchán d.f., decaëns t., díaz cosín d.j., hedde m., lapied e. & domínguez j. 2020. french mediterranean islands as a refuge of relic earthworm species: cataladrilus porquerollensis sp. nov. and scherotheca portcrosana sp. nov. (crassiclitellata, lumbricidae). european journal of taxonomy 701: 1–22. https://doi.org/10.5852/ejt.2020.701 introduction in the palearctic, lumbricidae rafinesque-schmaltz, 1815 is the most diverse and widely distributed native earthworm family, including 44 genera and around 670 species (blakemore 2008) (but around 45 species have been described since then). due to the large number of endemic species and genera, this family is believed to have originated and diversified in southwestern europe (omodeo & rota 2008). several genera, including prosellodrilus bouché, 1972 and scherotheca bouché, 1972, occur in the pyrenees, northeast spain (catalonia, aragón, navarra), southeastern france (occitanie, provencealpes-côte d’azur), corsica-sardinia and northeastern italy (qiu & bouché 1998a, 1998c), hinting at a paleogeographic link between these terranes before the oligocene–miocene rifting and breakup (bache et al. 2010). the genus prosellodrilus comprises around 32 species and subspecies (blakemore 2012). this genus is subdivided into a conflictive subgenus system, proposed by qiu & bouché (1998a) and baha & berra (2001): prosellodrilus, pyrenodrilus qiu & bouché, 1998, kenleenus qiu & bouché, 1998 and maghrebiella baha & berra, 2001. those subgenera were later elevated to genus-status by blakemore (2012), with maghrebiella amended to cadanera blakemore, 2012. a closely related genus with a more restricted range is cataladrilus qiu & bouché, 1998. cataladrilus comprises eight species distributed in the eastern pyrenees (catalonia, andorra and pyrénées-orientales) and two species of the subgenus latisinella qiu & bouché, 1998 found in soria (spain) more than 200 km away from the main range of the genus (qiu & bouché 1998b). a molecular phylogenetic analysis of the family lumbricidae, including representatives of both genera, supported this close relationship, not identifying them as monophyletic but rather highlighting them as intermixed within a single clade (domínguez et al. 2015). in spite of this, a morphological distinction between them is rather clear, with cataladrilus possessing anterior spermathecae (intersegments 9/10, 10/11) and prosellodrilus possessing posterior spermathecae (between intersegments 12/13 and 14/15). the genus scherotheca includes 41 species and subspecies (qiu & bouché 1998c), half of which occur in occitanie, provence-alpes-côte d’azur and corsica. scherotheca gigas (dugès, 1828), the type species of the genus, includes several subspecies; most of them were established as species by qiu & bouché (1998d). domínguez et al. (2015) included 5 representatives of scherotheca in their phylogeny of lumbricidae, two of which were unidentified species. in addition, the specimens identified as sc. gigas were collected in navarra (spain), which is far removed from the type locality of the species (montpellier) raising doubts about their identification. thus, the systematics of this highly diverse genus would benefit from the corroboration provided by a comprehensive molecular phylogenetic analysis, which has not yet been performed. despite the intensive research on earthworm diversity in southern france, some endemic species in relatively remote, comparatively weakly explored areas may remain undescribed. one example of such an area is the hyères archipelago, which faces the coast of provence. these continental islands remained https://doi.org/10.5852/ejt.2020.701 marchán d.f. et al., two new species (crassiclitellata, lumbricidae) from french islands 3 connected to corsica and sardinia until late chattian–aquitanian – ca 24 ma – when the first marine ingression preceded the rotation and drifting of the corso-sardinian microplate (oudet et al. 2010). due to their proximity to the coast, the islands were intermittently connected to the french mainland during the pleistocene until 11 000–12 000 bc, when they became permanently isolated (médail et al. 2013). the port-cros national park was created in 1963 to protect the hyères archipelago, and two of its three main islands (porquerolles and port-cros) constitute its core areas. two undescribed earthworm species were collected during a sampling survey of the islands of porquerolles and port-cros (decaëns et al. 2020). further detailed morphological diagnoses and molecular phylogenetic analyses confirmed the novelty of both taxa, but also revealed that they should be assigned to the genera scherotheca and cataladrilus respectively. the inclusion of closely related congeneric species in the molecular phylogenetic reconstruction provided further insight into the systematics of these genera. in this work we relate the results of both of these morphological and molecular analyses, and propose the formal description of the two new species under the names scherotheca portcrosana marchán & decäens sp. nov. and cataladrilus porquerollensis marchán & decäens sp. nov. material and methods specimens, sampling and morphological description specimens described in this work were collected in a sampling survey carried out in the port-cros national park (france) in march 2018 (decaëns et al. 2020). the rest of the specimens were collected in montpellier (france) and its vicinities at different times between 2015 and 2019. the list of species and the localities where they were collected is shown in table 1. earthworms were obtained by soil digging and hand-sorting, rinsed with water and fixed in 100% ethanol to enable further molecular analyses. species classification and morphological diagnoses were carried using the set of external and internal morphological characters used by qiu & bouché (1998a, 1998b, 1998c, 1998d), and following the format established by domínguez et al. (2018). main external morphological characters were: average length, average number of segments, average weight, pigmentation, type of prostomium, setal arrangement, position of papillae, position of first dorsal pore, nephridial pore arrangement, position and development of male pores, position and development of female pores, position of spermathecal pores, position of clitellum, position of tubercula pubertatis. main internal anatomical characters were: position of oesophageal hearts, position and morphology of calciferous glands, position of crop, position of gizzard, type of typhlosole, shape of nephridial bladders, number and position of seminal vesicles, number and position of spermathecae. institutional acronyms earthworm holotypes and paratypes were deposited in the following institutions: cefe = center of functional and evolutionary ecology, montpellier, france ucmlt = earthworm collection of universidad complutense de madrid, spain dna isolation, sequencing and molecular analyses total genomic dna was extracted using the dneasy blood & tissue kit (qiagen) from ventral integument samples of approximately 5 × 5 mm. molecular barcoding (cytochrome c oxidase subunit i – coi) was performed for all the specimens of ca. porquerollensis marchán & decäens sp. nov. (30) and sc. portcrosana marchán & decäens sp. nov. (9) at the biodiversity institute of ontario using the primers from folmer et al. (1994) and following the standard protocol established for the international barcode of life project (http://ibol.org/) (see details in decaëns et al. 2016). regions of the nuclear 28s rrna and mitochondrial 16s rrna, nadh dehydrogenase (nd1) and coi (3348 bp) were amplified for two specimens of ca. porquerollensis marchán & decäens sp. nov. and sc. portcrosana marchán & decäens http://ibol.org/ european journal of taxonomy 701: 1–22 (2020) 4 sp. nov. and one specimen of sc. gigas gigas, sc. gigas mifuga bouché, 1972, sc. gigas heraultensis qiu & bouché, 1998, sc. rhodana bouché, 1972 and sc. monspessulensis idica bouché,1972 using the polymerase chain reaction (pcr), with primers and conditions described in pérez-losada et al. (2009, 2015). pcr products were purified and sequenced by the c.a.c.t.i genomics service (university of vigo). dna sequences obtained in this study (including the two new species and the above mentioned scherotheca spp), as well as their associated meta-data and genbank accession numbers are all available in the public dataset “ds-ewsppcnp” on the bold bioinformatics platform (https://doi.org/10.5883/ds-ewsppcnp). sequences reported by domínguez et al. (2015, 2018), pérez-losada et al. (2009, 2011, 2015), paoletti et al. (2016) and de sosa et al. (2019), including representatives from most of the lumbricidae genera and two members of the closest families (hormogastridae vejdovsky, 1884 and criodrilidae michaelsen, 1900) were downloaded from genbank and used as a reference dataset. included species are shown in appendix 1. sequences were aligned with mafft ver. 7 (katoh & standley 2013) with default settings and concatenated with bioedit (hall 1999), resulting in a matrix of 3348 bp. the best fitting evolutionary table 1. list of species studied in this work and their sampling localities. species location coordinates habitat cataladrilus porquerollensis sp. nov. porquerolles island, france; locality 1 (pqr1) 42.9988º n 6.20025º e meadow porquerolles island, france; locality 2 (pqr2) 42.9955º n 6.20609º e vineyard porquerolles island, france; locality 3 (pqr3) 42.9961º n 6.20393º e olive grove scherotheca portcrosana sp. nov. port-cros island, france; locality 1 (pcr3) 43.0042º n 6.39014º e evergreen oak forest port-cros island, france; locality 2 (pcr6) 43.0119º n 6.39384º e mixed pine/ evergreen oak forest port-cros island, france; locality 3 (pcr8) 43.0089º n 6.41176º e meadow scherotheca gigas gigas (dugès, 1828) bois de montmaur, montpellier, france 43.6443º n 3.865º e pine forest scherotheca gigas mifuga bouché, 1972 montpellier ecotron, prades le lez, france 43.6817º n 3.8762º e mediterranean garrigue scherotheca gigas heraultensis qiu & bouché, 1998 mont de la gardiole, vic-la-gardiole, france 43.49484º n 3.77569º e mediterranean garrigue scherotheca rhodana bouché, 1972 iut de montpellier, montpellier, france 43.635411º n 3.853335º e urban mixed forest scherotheca monspessulensis idica bouché, 1972 iut de montpellier, montpellier, france 43.635411º n 3.853335º e urban mixed forest http://dx.doi.org/10.5883/ds-ewsppcnp marchán d.f. et al., two new species (crassiclitellata, lumbricidae) from french islands 5 model for each partition was selected with jmodeltest ver. 2.1.3 (darriba et al. 2012) by applying the akaike information criterion (aic; akaike 1973), and bayesian information criterion (bic; schwarz 1978). gtr+i+g was selected as best-fitting evolutionary model for coi, 28s and nd1, and hky+i+g was selected for 16s. maximum likelihood analysis was performed with raxml-hpc ver. 8 (stamatakis 2014) as implemented in the cipres science gateway ver. 3.3 (miller et al. 2010), using gtr+i+g for each data partition with 10 alternative runs and estimating the support for the resulting topologies by 1000 rapid bootstrap replicates. bayesian inference of the phylogeny was estimated with mrbayes ver. 3.1.2 (ronquist & huelsenbeck 2003) as implemented in the cipres science gateway ver. 3.3. parameters were set to 50 million generations sampling every 5000th generation (10 000 trees). two independent runs with four chains each were performed and 20% of the trees were discarded as burn-in. the remaining trees were combined and summarized on a 50% majority-rule consensus tree. clade support (bootstrap and posterior probability) values over 70% and 90% respectively were considered as high (see marchán et al. 2018; de sosa et al. 2019). uncorrected average pairwise distances between the newly described species and their closest relatives for the molecular markers coi and 16s were calculated in mega x (kumar et al. 2018) in order to support their status as separate species. results molecular phylogenetic analyses the phylogenetic relationships recovered by the bayesian inference (fig. 1, appendix 2) and maximum likelihood analyses were generally consistent. a clade containing cataladrilus and prosellodrilus was well supported, but the relationship between them was not fully resolved. cataladrilus porquerollensis marchán & decäens sp. nov. was recovered as a sister species to ca. monticola qiu & bouché, 1998. scherotheca portcrosana marchán & decäens sp. nov. was recovered within a well-supported, fig. 1. detail of the clades including the species cataladrilus porquerollensis marchán & decäens sp. nov. and scherotheca portcrosana marchán & decäens sp. nov. obtained by bayesian phylogenetic analysis of the concatenated sequence of molecular markers coi–16s–nd1–28s. the complete phylogenetic tree is shown in appendix 2. posterior probability support values are shown besides corresponding nodes. european journal of taxonomy 701: 1–22 (2020) 6 monophyletic scherotheca, as a sister to two undescribed species from corsica. in addition, sc. rhodana, sc. monspessulensis idica, sc. gigas gigas and sc. savignyi guerne & horst, 1893 appeared separated from each other by comparatively long branches, while sc. gigas heraultensis and sc. gigas mifuga formed a well-supported, comparatively shallow clade with sc. gigas gigas. the specimen from navarra identified as sc. gigas by domínguez et al. (2015) was recovered as a sister taxon to sc. savignyi. uncorrected average pairwise genetic (uapg) distances for both the coi and 16s molecular markers are given in table 2. the uapg distances for coi between ca. porquerollensis marchán & decäens sp. nov. and its closest relatives (representatives of cataladrilus and prosellodrilus) ranged from 13.4 to 18.3%. the uapg distance for 16s was noticeably lower with congeneric ca. monticola qiu & bouché, 1998 (12.3%) than with species belonging to prosellodrilus (18.5%) and scherotheca (18.6–22%). the uapg coi distances between sc. portcrosana marchán & decäens sp. nov. and other representatives of the genus ranged from 11.8% (scherotheca sp. 1 corsica) to 14.6% (sc. rhodana). the uapg 16s distances with congeneric species ranged between 11.8% (scherotheca sp. 1 corsica) to 15.8% (scherotheca sp. 2 corsica), the values of which were much lower than the distances with other genera (prosellodrilus and cataladrilus, 18.3–21.3%). the uapg distances between sc. gigas gigas and the subspecies mifuga and heraultensis for the molecular markers coi (1.4–3.7%) and 16s (2.9–3.2%) were much lower than with the other species of scherotheca (coi: 11.1–12.9%, 16s: 9.3–13.5%). systematics phylum annelida lamarck, 1802 class oligochaeta grube, 1850 / clitellata michaelsen, 1919 order megadrili benham, 1890 / haplotaxida michaelsen, 1900 family lumbricidae rafinesque-schmaltz, 1815 genus cataladrilus qiu & bouché, 1998 type species cataladrilus monticola qiu & bouché, 1998 diagnosis lumbricinae rafinesque-schmaltz, 1815 of small to medium size. longitudinal furrows in peristomium and pygidium. closely paired or separate chaetae. aligned nephridial pores. spermathecal pores simple in 9/10, 10/11 (exception cataladrilus multhitecus qiu & bouché, 1998: 7/8–10/11). male pores in ½ 15 with developed porophores. calciferous gland in 11–15, usually with diverticles in 11. gizzard in 17–19 (exception cataladrilus annulatus qiu & bouché, 1998: (18)19–21). typhlosole simple, bifid or multifid. nephridial bladders u-shaped, reclinate (exception cataladrilus multhitecus qiu & bouché, 1998: v-shaped – “fourchué”). two pairs of seminal vesicles in 11, 12 (exception cataladrilus mrsici qiu & bouché, 1998: three pairs in 9, 11, 12). cataladrilus porquerollensis marchán & decäens sp. nov. urn:lsid:zoobank.org:act:de99e6a6-65ef-496a-aeaa-4f6fbafa19cb fig. 2, table 3 http://zoobank.org/urn:lsid:zoobank.org:act:de99e6a6-65ef-496a-aeaa-4f6fbafa19cb marchán d.f. et al., two new species (crassiclitellata, lumbricidae) from french islands 7 ta bl e 2. u nc or re ct ed a ve ra ge p ai rw is e ge ne tic (u a pg ) d is ta nc es b et w ee n th e sp ec ie s de sc ri be d in th is w or k (c at al ad ri lu s po rq ue ro lle ns is m ar ch án & d ec äe ns s p. n ov . a nd s ch er ot he ca p or tc ro sa na m ar ch án & d ec äe ns s p. n ov .) an d th ei r c lo se st re la tiv es fo r t he m ol ec ul ar m ar ke rs c o i ( un de r d ia go na l) an d 16 s (o ve r d ia go na l) . ca.porquerollensis sp. nov. ca. edwarsi ca. monticola pr. pyrenaicus pr. biserialis sc. sp 2 corsica sc. sp 1 corsica sc. corsicana sc. portcrosana sp. nov. sc. gigas gigas sc. gigas mifuga sc. gigas heraultensis sc. monspessulensis idica sc. rhodana sc. savignyi sc. cf. gigas navarra c a. p or qu er ol le ns is s p. n ov . 0. 6 – 12 .3 18 .5 – 21 .0 21 .9 19 .7 20 .8 22 .0 22 .6 21 .4 18 .6 18 .8 19 .4 19 .4 c a. e dw ar si 13 .4 – – – – – – – – – – – – – – – c a. m on tic ol a 14 .1 13 .6 – 13 .7 – 15 .5 15 .9 16 .2 18 .3 19 .7 19 .5 18 .4 15 .7 14 .1 17 .2 15 .3 p r. py re na ic us 15 .9 15 .6 14 .7 – – 20 .4 20 .5 20 .1 21 .3 24 .7 24 .3 24 .3 19 .4 19 .6 22 .5 20 .0 p r. bi se ri al is 18 .3 14 .2 14 .3 15 .2 – – – – – – – – – – – – sc . s p 2 c or si ca – – – – – – 10 .8 12 .8 15 .8 12 .9 13 .6 13 .4 12 .6 11 .5 13 .6 10 .5 sc . s p 1 c or si ca 14 .4 15 .3 17 .1 16 .4 18 .8 – – 10 .4 11 .8 11 .7 10 .3 10 .7 11 .1 10 .9 12 .9 8. 9 sc . c or si ca na 15 .3 13 .6 15 .2 17 .4 15 .7 – 13 .0 – 15 .1 12 .8 13 .0 12 .2 13 .9 12 .9 14 .6 11 .6 sc . p or tc ro sa na s p. n ov . 15 .5 12 .3 13 .7 15 .2 17 .1 – 11 .8 13 .1 0. 2 13 .5 13 .2 13 .2 13 .7 13 .3 13 .4 13 .8 sc . g ig as g ig as 15 .8 14 .9 16 .9 17 .4 18 .1 – 12 .1 11 .7 12 .9 – 2. 9 3. 2 11 .7 10 .8 11 .4 9. 3 sc . g ig as m ifu ga 16 .4 14 .9 17 .8 17 .6 17 .8 – 12 .4 11 .9 13 .0 1. 4 – 3. 9 11 .9 11 .4 12 .2 9. 9 sc . g ig as h er au lte ns is 17 .6 14 .8 17 .1 16 .8 18 .0 – 11 .3 12 .0 12 .3 3. 7 3. 4 – 11 .0 10 .6 11 .8 9. 5 sc . m on sp es su le ns is id ic a 16 .0 15 .8 16 .1 17 .2 19 .0 – 12 .5 11 .9 12 .4 11 .1 11 .0 11 .6 – 8. 9 12 .0 9. 9 sc . r ho da na 15 .0 16 .5 15 .9 17 .2 17 .3 – 11 .8 13 .0 14 .6 12 .0 12 .3 12 .4 12 .2 – 11 .5 9. 4 sc . s av ig ny i 16 .3 15 .1 16 .7 18 .8 19 .8 – 12 .6 12 .7 13 .8 12 .2 11 .9 13 .1 12 .4 13 .4 – 9. 0 sc . c f. gi ga s n av ar ra 15 .7 16 .0 16 .0 15 .4 16 .5 – 12 .7 11 .9 12 .8 12 .6 12 .1 12 .1 11 .3 12 .2 10 .5 – european journal of taxonomy 701: 1–22 (2020) 8 diagnosis specimens of cataladrilus porquerollensis marchán & decäens sp. nov. can be distinguished from the other known species of cataladrilus by the position of the clitellum in segments (19)20–32,33(34) and tubercula pubertatis in segments (28)29–31 (table 3). they resemble cataladrilus (latisinella) mrsici in the position of tubercula pubertatis and small body size, but differ in the position of the clitellum (22–32 in ca. mrsici), two pairs of seminal vesicles (three in ca. mrsici) and paired chaetae (separate in ca. mrsici). etymology the species name is derived from porquerolles, the island where this species was found. material examined holotype france • adult; provence-alpes-côte d’azur, var, porquerolles island; 42.9961º n, 6.20393º e [locality 3 (pqr3) (table 1)]; 14 mar. 2018; t. decaëns, e. lapied, m. hedde and m. zwicke leg.; olive grove; bold sample id: ew-pnpc-0098; ucmlt. paratypes france – provence-alpes-côte d’azur • 10 specs; same collection data as for holotype; bold sampleid: ew-pnpc-0036, ew-pnpc-0037, ew-pnpc-0038, ew-pnpc-0039, ew-pnpc-0040, ew-pnpc-0096, ew-pnpc-0097, ew-pnpc-0099, ew-pnpc-0100, ew-pnpc-0268; ucmlt • 10 specs; same collection data as for holotype; bold sampleid: ew-pnpc-0041, ew-pnpc-0042, ewpnpc-0043, ew-pnpc-0044, ew-pnpc-0045, ew-pnpc-0101, ew-pnpc-0102, ew-pnpc-0103, ew-pnpc-0104, ew-pnpc-0105; cefe • 1 spec.; var, porquerolles island; 42.9988º n, 6.20025º e [locality 1 (pqr1) (table 1)]; 14 mar. 2018; t. decaëns, e. lapied, m. hedde and m. zwicke leg.; meadow; bold sampleid: ew-pnpc-0173; cefe • 6 specs; var, porquerolles island; 42.9955º n, 6.20609º e [locality 2 (pqr2) (table 1)]; 14 mar. 2018; t. decaëns, e. lapied, m. hedde and m. zwicke leg.; vineyard; bold sampleid: ew-pnpc-0127, ew-pnpc-0128, ew-pnpc-0129, ew-pnpc-0130, ew-pnpc-0131, ew-pnpc-0132; ucmlt • 2 specs; same collection data as for preceding; bold sampleid: ew-pnpc-0133, ew-pnpc-0134; cefe. morphological description external morphology body pigmentation absent in live specimens. white-beige homogeneous color in fixed specimens (fig. 2). average length 48 mm (41–51 mm, n = 5 adults); diameter 2 mm (1.8–2.2 mm, n = 5 adults); body cylindrical in cross-section; average number of segments 196 segments (166–239, n = 5 adults; 239 segments in the holotype). average weight (alcohol fixed specimens): 0.14 g (0.12–0.17 g, n = 5 adults). prostomium epilobous. longitudinal furrows in first 2–3 segments. first dorsal pore at the intersegmental furrow 10/11–11/12. nephridial pores aligned in b. spermathecal pores at the intersegmental furrows 9/10, 10/11 in c. male pores in segment 15, surrounded by a well-developed porophore. female pores inconspicuous. clitellum saddle-shaped in segments (19)20–32,33(34). tubercula pubertatis in segments (28)29–31. chaetae very small and closely paired, with interchaetal ratio aa: 9, ab: 1, bc: 3, cd: 1, dd: 7 at segment 40. chaetophores/genital papillae in segments 11, 13. internal anatomy septa 5/6–9/10 thickened and muscular. hearts in segments 6–11, oesophageal. calciferous glands in segments 11–13, without clear diverticles or enlargements. crop in segments 15–16, gizzard in segments marchán d.f. et al., two new species (crassiclitellata, lumbricidae) from french islands 9 table 3. morphological characters of cataladrilus porquerollensis marchán & decäens sp. nov. and morphologically similar relatives within the genera cataladrilus qiu & bouché, 1998 and prosellodrilus bouché, 1972. length expressed in mm, weight expressed in mg. characters cataladrilus porquerollensis sp. nov. cataladrilus (latisinella) mrsici cataladrilus albus prosellodrilus dactylothecus length 45–51 35 120–145 92 n. segments 196 168 229–297 207 weight 140 102 1526 409 pigmentation absent absent absent absent prostomium epilobous epilobous epilobous epilobous setae closely paired separate closely paired closely paired setal arrangement 9:1:3:1:7 5:5:5:4:8 15:1:5:1:15 20:1:10:1:30 papillae 11, 13 absent 28–34 13, 14, 15 first dorsal pore 10/11–11/12 6/7 6/7 5/6 nephridial pores aligned, b aligned, b aligned, b aligned, b male pores ½ 15, b ½ 15, b ½ 15, b ⅔ 15 female pores ½ 14, inconspicuous ½ 14, b ½ 14, b ½ 14, b spermathecal pores 9/10, 10/11 simple in c 9/10, 10/11 simple in c 9/10, 10/11 simple in d 13/14, 14/15 in c clitelum (19)20–32, 33(34) 22–32 22–34 19–32 tubercula pubertatis (28)29–31 29–31 (½ 29)30–34 22–25 oesophageal hearts 6–11 6–11 6–11 6–11 calciferous glands 11–13 11–14, dilated in 12, 13 11–14, diverticles in 11 11–14, diverticles in 11 crop 15–16 15–16 15–16 15–16 gizzard 17–18 17–18 17–18 17–18 typhlosole bifid simple bifid bifid nephridial bladders u-shaped, reclinate u-shaped, reclinate u-shaped, reclinate u-shaped, reclinate, proximal ampulla seminal vesicles 11, 12 9, 11, 12 11, 12 11, 12 spermathecae simple, globular in 10, 11 simple, pyriform in 10, 11 simple, small globular in 10, 11 simple, pedunculate in 14, 15 european journal of taxonomy 701: 1–22 (2020) 10 17–18. typhlosole bifid with a small intermediate ridge of varying development. male sexual system holandric, testes and funnels (not enclosed in testes sacs, but with sperm present) located ventrally in segments 10 and 11. two pairs of small seminal vesicles in segments 11 and 12, with free seminal masses filling most of these segments. ovaries and female funnels in segment 13, ovarian receptacles (ovisacs) in segment 14. two pairs of globular, intracelomic spermathecae in segments 9 and 10 (intersegments 9/10, 10/11). nephridial bladders u-shaped, reclinate in segments 14, 20. distribution and ecology cataladrilus porquerollensis marchán & decäens sp. nov. is known from the island of porquerolles in the hyères archipelago, france. this species has been found in meadows, vineyards and olive groves, thus appears to inhabit moderately to highly anthropized habitats. genus scherotheca bouché, 1972 type species scherotheca gigas dùges, 1828. fig. 2. external morphology of cataladrilus porquerollensis marchán & decäens sp. nov. a. fixed specimen. b. schematic drawing. abbreviations: cl = clitellum; fp = female pore; mp = male pore; tp = tubercula pubertatis. marchán d.f. et al., two new species (crassiclitellata, lumbricidae) from french islands 11 diagnosis lumbricinae of large to very large size, post-clitellar trapezoidal section. pigmentation brown to dark brown. prostomium epilobous, closed. longitudinal furrows in the peristomium. nephridial pores “en solfège” (irregularly distributed). spermathecal pores at least in two intersegments, between 9/10 and 13/14, or 13/14 and 15/16, sometimes multiple. anterior septa strongly thickened. male pores in ½ 15, usually with porophores. gizzard in 17–20 (21, 22). typhlosole pinnate. two or four pairs of seminal vesicles in (9, 10) 11, 12. scherotheca portcrosana marchán & decäens sp. nov. urn:lsid:zoobank.org:act:663f18d2-a2ea-4b4e-8ed3-55a3fe5b7d57 fig. 3, table 4 diagnosis specimens of scherotheca portcrosana marchán & decäens sp. nov. can be distinguished from other known species of scherotheca by the position of the clitellum in segments 26–35 (½ 36) and tubercula pubertatis in segments 30–33, position of spermathecae in 12, 13 in addition to a smaller body size and faint pigmentation (table 4). etymology the species name is derived from port-cros, the island inhabited by this species. material examined holotype france • adult; provence-alpes-côte d’azur, var, port-cros island; 43.0089º n, 6.41176º e [locality 3 (pcr8) (table 1)]; 13 mar. 2018; t. decaens, e. lapied, m. hedde and m. zwicke leg.; meadow; bold sample id: ew-pnpc-0174; ucmlt. paratypes france – provence-alpes-côte d’azur • 1 spec.; var, port-cros island; 43.0042º n, 6.39014º e [locality 1 (pcr3) (table 1)]; 13 mar. 2018; t. decaens, e. lapied, m. hedde and m. zwicke leg.; evergreen oak forest; bold sampleid: ew-pnpc-0111; ucmlt • 5 specs; same collection data as for preceding; bold sampleid: ew-pnpc-0117, ew-pnpc-0119, ew-pnpc-0120, ew-pnpc-0121, ew-pnpc-0122; cefe • 1 spec.; var, port-cros island; 43.0119º n, 6.39384º e [locality 2 (pcr6) (table 1)]; 13 mar. 2018; t. decaens, e. lapied, m. hedde and m. zwicke leg.; mixed pine/evergreen oak forest; bold sampleid: ew-pnpc-0237; ucmlt • 1 spec.; same collection data as for preceding; bold sampleid: ew-pnpc-0242; cefe. morphological description external morphology body pigmentation very faint brown-grey. white-beige with dorsal brownish mid-segment brown bands in fixed specimens (fig. 3). average length 80 mm (75–85 mm, n = 2 adults); average diameter 7 mm (6.9–7.1 mm, n = 3 adults); body cylindrical in cross-section; average number of segments 163 (160–166, n = 2 adults; 166 segments in the holotype). average weight (fixed specimens): 2.13 g (1.89–2.36 g, n = 2 adults). prostomium epilobous, closed. longitudinal furrows in segments 1 and 2. first dorsal pore at intersegmental furrow 5/6. nephridial pores “en solfège” (irregularly distributed). spermathecal pores at intersegmental furrows 12/13 and 13/14 in c. male pores in segment 15, surrounded by a http://zoobank.org/urn:lsid:zoobank.org:act:663f18d2-a2ea-4b4e-8ed3-55a3fe5b7d57 european journal of taxonomy 701: 1–22 (2020) 12 well-developed porophore. female pores on segment 14. clitellum saddle-shaped in segments 26–35 (½ 36). tubercula pubertatis in segments 30–33. chaetae small and closely paired, with interchaetal ratio aa: 8, ab: 1, bc: 3.5, cd: 1, dd: 18 at segment 40. chaetophores/genital papillae in segments 11, 12, 14, 27, 29 and 34–38. fig. 3. external morphology of scherotheca portcrosana marchán & decäens sp. nov. a. live specimen (© t. decaëns). b. fixed specimen. c. schematic drawing. abbreviations: cl = clitellum; fp = female pore; mp = male pore; tp = tubercula pubertatis. marchán d.f. et al., two new species (crassiclitellata, lumbricidae) from french islands 13 table 4. morphological characters of scherotheca portcrosana marchán & decäens sp. nov. and morphologically similar relatives of genera scherotheca bouché, 1972 and eumenescolex qiu & bouché, 1998. length expressed in mm, weight expressed in mg. characters scherotheca portcrosana sp. nov. scherotheca corsicana corsicana scherotheca portonana eumenescolex simplex length 75–85 90–140 78–165 105–112 n. segments 160–166 177–215 193–208 99–154 weight 1890–2360 2000–3500 1588–2821 ? pigmentation brown (faint) dark brown brown ? prostomium epilobous epilobous epilobous pro-epilobous setae closely paired closely paired closely paired ? setal arrangement 8:1:3.5:1:18 8:1.5:4.5:1:19 11.6:1.6:8:1:33 ? papillae 11–14, 27, 29, 34–38 9–11, 27–37 11–14, 34, 35 ? first dorsal pore 5/6 9/10 10/11 11/12 nephridial pores en solfège en solfège en solfège ? male pores ½ 15, b ½ 15, b ½ 15, b ? female pores ½ 14, b ½ 14, b ½ 14, b ? spermathecal pores 12/13, 13/14 simple in c 12/13, 13/14 simple in c 12/13, 13/14 simple in c 13/14 simple in c clitelum 26–35(½ 36) (½ 26)1/27–½ 36(36) (25)26–35 26–½ 36 tubercula pubertatis 30–33 (½ 30) 31–½ 34(34) (28)29–32(33) 29–½ 33 oesophageal hearts 6/11 6/11 6/11 6–11 calciferous glands ½ 10–14, diverticles in 10 ½ 10–14, diverticles in 10 ½ 10–14, diverticles in 10 ? crop 15–16 15–16 15–16 ? gizzard 17–19 17–19 17–19 17–19 typhlosole pinnate pinnate pinnate ? nephridial bladders u-shaped, reclinate j-shaped, reclinate j-shaped, reclinate ? seminal vesicles 9, 10, 11, 12 9, 10, 11, 12 9, 10, 11, 12 9, 10, 11, 12 spermathecae simple, oval in 12, 13 simple, oval in 13, 14 simple, oval in 13, 14 ? european journal of taxonomy 701: 1–22 (2020) 14 internal anatomy septa 5/6–10/11 thickened and muscular. hearts in segments 6–11, oesophageal. calciferous glands in segments 10–14, with diverticula in segment 10. crop in segments 15–16, gizzard in segments 17–19. typhlosole pinnate. male sexual system holandric, testes and funnels (not enclosed in testes sacs, but with sperm present) located ventrally in segments 10 and 11. four pairs of reniform seminal vesicles in segments 9, 10, 11 and 12, with the latter two pairs being larger. ovaries and female funnels in segment 13, ovarian receptacles (ovisacs) in segment 14. two pairs of small globular spermathecae in segments 12 and 13 (intersegments 12/13, 13/14). nephridial bladders u-shaped, reclinate in segment 30. distribution and ecology scherotheca portcrosana marchán & decäens sp. nov. is known from the island of port-cros in the hyères archipelago, france. this species has been found in meadows, pine and evergreen oak forests, thus appears to have a preference for natural habitats. discussion systematic implications molecular phylogenetic analyses and genetic distances clearly support the morphology-based inclusion of ca. porquerollensis marchán & decäens sp. nov. within the genus cataladrilus. it is worth noting that the relationship between cataladrilus and prosellodrilus is currently not fully resolved based on molecular information, however the morphological characters clearly separate both genera (table 3) according to the number and position of spermathecae: two pairs in intersegments 9/10, 10/11 for cataladrilus (except for ca. multithecus qiu & bouché, 1998) instead of two to three pairs (sometimes multiple) in intersegments (12/13), 13/14, 14/15 for prosellodrilus (qiu & boché 1998a, 1998b). several species from cataladrilus and prosellodrilus are yet to be included in a detailed molecular phylogenetic analysis, with emphasis on including representatives from the different subgenera (possibly genera for prosellodrilus sensu lato according to blakemore (2012). this will be necessary to resolve the reciprocal monophyly of both genera and the status of the subgenera/genera included within them. cataladrilus porquerollensis marchán & decäens sp. nov. resembles cataladrilus (latisinella) mrsici, the most in the position of their tubercula pubertatis (segments 29–31) and their clitellum (segments (19)20–32, 33(34) and 22–32 respectively), but they both differ in the total extension of the clitellum, body size, disposition of their chaetae (closely paired vs separate) and number of seminal vesicles (two pairs in segments 11, 12 vs 3 pairs in segments 9, 11, 12). cataladrilus porquerollensis marchán & decäens sp. nov shows a remarkable external morphological similarity with allolobophora festae rosa, 1892 (sometimes attributed to genus prosellodrilus), however, they can be easily distinguished by the position of their spermathecae (9/10, 10/11 vs 12/13, 13/14). the inclusion of sc. portcrosana marchán & decäens sp. nov. within scherotheca is well supported by the phylogenetic trees. while french mainland species of scherotheca appear superficially very different from sc. portcrosana marchán & decäens sp. nov. (large body size, intense brown-grey pigmentation, very long clitellum), species of scherotheca from corsica share with sc. portcrosana marchán & decäens sp. nov. their relatively small body size, pale brown pigmentation and comparatively short clitellum. indeed, molecular analyses revealed a closer affinity of sc. portcrosana marchán & decäens sp. nov. with the corsican scherotheca over the mainland species. scherotheca portcrosana marchán & decäens sp. nov. appears very close to sc. corsicana corsicana pop, 1947 and sc. portonana qiu & bouché, 1998 according to their similar position of clitellum and tubercula pubertatis (table 4). however, it differs from both species by the exact position of clitellum marchán d.f. et al., two new species (crassiclitellata, lumbricidae) from french islands 15 and tubercula pubertatis, as well as being slightly smaller and less pigmented. scherotheca portcrosana marchán & decäens sp. nov. also shows a remarkable resemblance to eumenescolex simplex zicsi, 1981 (initially described as sc. corsicana simplex then transferred due to its single pair of spermathecae), but they clearly differ in the number and position of their spermathecae (in intersegments 12/13, 13/14 vs 13/14). this similarity highlights the importance of including representatives of eumenescolex qiu & bouché, 1998 in a molecular phylogenetic framework to test the close relationship among both genera that was previously suggested by qiu & bouché (1998d). qiu & bouché (1998d) elevated 9 subspecies of scherotheca to species while maintaining or creating others. genetic distances within the clade comprising sc. gigas gigas from montpellier and sc. gigas heraultensis and sc. gigas mifuga are way below the usual threshold established for divergence between conspecific taxa (less than 10% for coi, chang et al. 2009), supporting the subspecific status of these taxa; while sc. rhodana (described as sc. gigas rhodana) appears to be as divergent from sc. gigas gigas as from sc. monspessulensis idica or other species of the genus (with genetic distances above the aforementioned threshold). thus, the morphological criteria followed by qiu & bouché (1998d) appear supported by molecular data. however, larger sample sizes would be necessary to confirm this statement. these findings suggest that molecular markers are suitable for delimiting species and subspecies within scherotheca. if this approach performs consistently across different animal taxa, it may be advisable to implement it widely in order to avoid the dismissal of unrecognized species-level taxa in ecological studies and biodiversity assessment (for different examples see rutgers et al. 2016). scherotheca cf. gigas (from domínguez et al. 2015) collected from navarra appear to be unrelated to sc. gigas gigas from montpellier, but closely related to sc. savignyi. these earthworms may belong to one of the former subspecies of sc. gigas such as scherotheca aquitania bouché, 1972, which is relatively common in northeastern spain (qiu & bouché 1998d). preliminary re-inspection of these specimens showed external morphological characters compatible with sc. aquitania. thus, the inclusion of french specimens of sc. aquitania (ideally from the type locality) would enable confirmation of this hypothesis. this is just one of the many points highlighting the need for a comprehensive molecular phylogeny of the genus scherotheca. robust systematics of the many species of this genus in northeastern spainsouthern france-corsica-italy would contribute to the following: a) a better understanding of the impact on ecosystems of these large, deep burrowing anecic earthworms, which have been found to be important in both agricultural and unmanaged habitats (gavinelli et al. 2018), b) assessment of the conservation status of the genus, with several taxa being narrowly restricted and potentially vulnerable to human impacts (rida & bouché 1995) and c) insight into the historical biogeography of native earthworm families in this geologically complex area. phylogeographic implications for cataladrilus and scherotheca the discovery of a species of cataladrilus in the hyères archipelago is puzzling from a phylogeographic point of view. the main range of cataladrilus is restricted to catalonia (spain) and andorra, with two species in neighboring locations in france separated from the hyères archipelago by 340 km. this disjunct distribution could be explained by a failure to find closely related species in the geographic gap, which would suggest that southeastern france could still harbor other undescribed species. this, however, seems quite unlikely if considering the intense sampling already done by marcel bouché at the end of the past century (bouché 1972). alternatively, ca. porquerollensis marchán & decäens sp. nov. may be a relict from a formerly wider distribution. this would be consistent with paleogeographic events in the area: the eastern pyrenees and provence were connected through an exposed gulf of lion up to late oligocene–early miocene (around 24 ma) when a wide marine ingression separated them up to the present (except for the duration of the messinian salinity crisis) (sissingh 2006). the possibility of cataladrilus and prosellodrilus being synonyms (as the lack of reciprocal monophyly may suggest) european journal of taxonomy 701: 1–22 (2020) 16 would not change the observed disjunction, as the closest species of prosellodrilus (pr. tikalus qiu & bouché, 1998) is found 225 km to the west of porquerolles. the closer relationship between sc. portcrosana marchán & decäens sp. nov. and the corsican scherotheca than with french mainland scherotheca is also unexpected. however, several species of plants, arthropods and vertebrates are common to the hyères archipelago and corsica (médail et al. 2013). scherotheca portcrosana marchán & decäens sp. nov. may have diverged from its relatives before or during the rifting and drifting of the corso-sardinian microplate around 24 ma (oudet et al. 2010). corsica remained connected to the french mainland through the rotation to its current position (sissingh 2006), which would have allowed later migration to the hyères archipelago of a common ancestor of sc. portcrosana marchán & decäens sp. nov. and corsican scherotheca. however, under this second scenario, one would expect to find other relatives in eastern provence (where the land bridge was supposed to join both areas). the area of provence has been strongly disturbed by human activities, especially in the coastal area, while the small islands like porquerolles and port-cros remained relatively well preserved. it is possible that these islands acted as refugia while continental relatives got extinct or relegated to relict habitats. this phenomenon has already been suggested to explain the distribution of the gecko euleptes europaea gené, 1839, which is frequent in corsica and sardinia and rare in the coastal islands of provence and liguria, probably after a range retraction in the mainland (delauguerre et al. 2011). further survey focusing on relic natural ecosystems in the littoral area (cap lardier) or further into the mainland (i.e., sainte baume, plaine des maures, massif de l’esterel) could be performed to test this hypothesis. whatever explicative hypothesis is retained, these disjunct distributions, as previously observed in other earthworm genera (pérez losada et al. 2011; domínguez et al. 2018; marchán et al. 2018), highlight the strong connection between paleogeographic events and earthworm evolution and divergence. a more robust approach to time-calibrated phylogenies in earthworms (hindered by the lack of body fossils) and the integration of different paleogeographic reconstructions could illuminate both the origin of palearctic earthworms (hormogastridae, lumbricidae) and the geological history of the western mediterranean terranes from the late cretaceous to the neogene. acknowledgments this work 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meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. https://doi.org/10.1016/0929-1393(94)00032-3 https://doi.org/10.1093/bioinformatics/btg180 https://doi.org/10.1016/j.apsoil.2015.08.015 https://doi.org/10.1017/s0016774600077933 https://doi.org/10.1093/bioinformatics/btu033 european journal of taxonomy 701: 1–22 (2020) 20 species publication allolobophora chaetophora bouché, 1972 domínguez et al. 2015; pérez-losada et al. 2015 allolobophora chlorotica savigny, 1826 domínguez et al. 2015; pérez-losada et al. 2015 allolobophora dacica pop, 1938 domínguez et al. 2015; pérez-losada et al. 2015 allolobophora dubiosa orley, 1881 domínguez et al. 2015; pérez-losada et al. 2015 allolobophora mehadiensis rosa, 1895 domínguez et al. 2015; pérez-losada et al. 2015 allolobophora moebii michaelsen, 1895 domínguez et al. 2015; pérez-losada et al. 2015 allolobophora molleri rosa, 1889 pérez-losada et al. 2009, 2015; domínguez et al. 2015 allolobophora robusta rosa, 1895 domínguez et al. 2015; pérez-losada et al. 2015 aporrectodea caliginosa savigny, 1826 pérez-losada et al. 2009, 2015; domínguez et al. 2015 aporrectodea georgii michaelsen, 1890 domínguez et al. 2015; pérez-losada et al. 2015 aporrectodea jassyensis michaelsen, 1891 domínguez et al. 2015; pérez-losada et al. 2015 aporrectodea limícola michaelsen, 1890 pérez-losada et al. 2009, 2015; domínguez et al. 2015 aporrectodea longa ude, 1885 pérez-losada et al. 2009, 2015; domínguez et al. 2015 aporrectodea nocturna evans, 1946 pérez-losada et al. 2009, 2015; domínguez et al. 2015 aporrectodea rosea savigny, 1826 domínguez et al. 2015; pérez-losada et al. 2015 aporrectodea trapezoides duges, 1828 pérez-losada et al. 2009, 2015; domínguez et al. 2015 aporrectodea tuberculata eisen, 1874 pérez-losada et al. 2009, 2015; domínguez et al. 2015 carpetania elisae álvarez, 1977 domínguez et al. 2015; pérez-losada et al. 2015 cataladrilus edwarsi qiu & bouché, 1998 domínguez et al. 2015; pérez-losada et al. 2015 cataladrilus montícola qiu & bouché, 1998 domínguez et al. 2015; pérez-losada et al. 2015 cernosvitovia dudichi zicsi & sapkarev, 1982 domínguez et al. 2015; pérez-losada et al. 2015 cernosvitovia rebeli rosa, 1897 domínguez et al. 2015; pérez-losada et al. 2015 compostelandrilus bercianus domínguez, aira, porto, díaz cosín & pérez-losada, 2017 domínguez et al. 2018 compostelandrilus menciae domínguez, aira, porto, díaz cosín & pérez-losada, 2017 domínguez et al. 2018 criodrilus lacuum hoffmeister, 1845 domínguez et al. 2015; pérez-losada et al. 2015 dendrobaena attemsi michaelsen, 1902 domínguez et al. 2015; pérez-losada et al. 2015 dendrobaena cf. biblica rosa, 1893 domínguez et al. 2015; pérez-losada et al. 2015 dendrobaena illyrica cognetti de martiis, 1906 domínguez et al. 2015; pérez-losada et al. 2015 dendrobaena jastrebensis mrsic & sapkarev, 1987 domínguez et al. 2015; pérez-losada et al. 2015 dendrobaena octaedra savigny, 1826 domínguez et al. 2015; pérez-losada et al. 2015 dendrobaena pentheri rosa, 1905 domínguez et al. 2015; pérez-losada et al. 2015 dendrobaena pygmaea friend, 1923 domínguez et al. 2015; pérez-losada et al. 2015 dendrobaena veneta rosa, 1886 domínguez et al. 2015; pérez-losada et al. 2015 diporodrilus pilosus bouché, 1972 domínguez et al. 2015; pérez-losada et al. 2015 eisenia andrei bouché, 1972 domínguez et al. 2015; pérez-losada et al. 2015 eisenia balatonica pop, 1943 domínguez et al. 2015; pérez-losada et al. 2015 eisenia fétida savigny, 1826 domínguez et al. 2015; pérez-losada et al. 2015 eisenia lucens vaga, 1857 domínguez et al. 2015; pérez-losada et al. 2015 eiseniella tetraedra savigny, 1826 domínguez et al. 2015; pérez-losada et al. 2015 appendix 1 (continued on next page). species included in the reference dataset. marchán d.f. et al., two new species (crassiclitellata, lumbricidae) from french islands 21 appendix 1 (continued). species included in the reference dataset. species publication eiseniona albolineata díaz cosín, trigo & mato, 1989 domínguez et al. 2015; pérez-losada et al. 2015 eiseniona oliveirae rosa, 1894 domínguez et al. 2015; pérez-losada et al. 2015 eisenoides carolinensis michaelsen, 1910 domínguez et al. 2015; pérez-losada et al. 2015 eisenoides lonnbergi michaelsen, 1894 domínguez et al. 2015; pérez-losada et al. 2015 eophila crodabepis paoletti et al., 2016 paoletti et al. 2016, de sosa et al. 2019 eophila gestroi cognetti de martiis, 1905 paoletti et al. 2016, de sosa et al. 2019 eophila tellinii rosa, 1888 paoletti et al. 2016, de sosa et al. 2019 galiciandrilus bertae díaz cosín, mato & mascato, 1985 pérez-losada et al. 2011, 2015; domínguez et al. 2015 galiciandrilus morenoe díaz cosín, calvin & mato, 1985 pérez-losada et al. 2011, 2015; domínguez et al. 2015 helodrilus cernosvitovianus zicsi, 1967 domínguez et al. 2015; pérez-losada et al. 2015 helodrilus cortezi qiu & bouché, 1998 domínguez et al. 2015; pérez-losada et al. 2015 helodrilus patriarcalis rosa, 1893 domínguez et al. 2015; pérez-losada et al. 2015 lumbricus castaneus savigny, 1826 domínguez et al. 2015; pérez-losada et al. 2015 lumbricus rubellus hoffmeister, 1843 domínguez et al. 2015; pérez-losada et al. 2015 octodriloides boninoi omodeo, 1962 domínguez et al. 2015; pérez-losada et al. 2015 octodrilus complanatus duges, 1928 domínguez et al. 2015; pérez-losada et al. 2015 octodrilus exacystis rosa, 1896 domínguez et al. 2015; pérez-losada et al. 2015 octodrilus gradinescui pop, 1938 domínguez et al. 2015; pérez-losada et al. 2015 octodrilus pseudocomplanatus omodeo, 1962 domínguez et al. 2015; pérez-losada et al. 2015 octodrilus transpadanus rosa, 1884 domínguez et al. 2015; pérez-losada et al. 2015 octolasion cyaneum savigny, 1826 domínguez et al. 2015; pérez-losada et al. 2015 octolasion lacteum orley, 1885 domínguez et al. 2015; pérez-losada et al. 2015 octolasion montanum wessely, 1905 domínguez et al. 2015; pérez-losada et al. 2015 panoniona leoni michaelsen, 1891 domínguez et al. 2015; pérez-losada et al. 2015 postandrilus lavellei qiu & bouché, 1998 pérez-losada et al. 2011, 2015; domínguez et al. 2015 postandrilus majorcanus qiu & bouché, 1998 pérez-losada et al. 2011, 2015; domínguez et al. 2015 postandrilus medoakus qiu & bouché, 1998 pérez-losada et al. 2011, 2015; domínguez et al. 2015 postandrilus palmensis qiu & bouché, 1998 pérez-losada et al. 2011, 2015; domínguez et al. 2015 postandrilus sapkarevi qiu & bouché, 1998 pérez-losada et al. 2011, 2015; domínguez et al. 2015 proctodrilus antipai michaelsen, 1891 domínguez et al. 2015; pérez-losada et al. 2015 prosellodrilus biauriculatus bouché, 1972 domínguez et al. 2015; pérez-losada et al. 2015 prosellodrilus biserialis bouché, 1972 domínguez et al. 2015; pérez-losada et al. 2015 prosellodrilus pyrenaicus cognetti, 1904 domínguez et al. 2015; pérez-losada et al. 2015 scherotheca cf. gigas bouché, 1972 domínguez et al. 2015; pérez-losada et al. 2015 scherotheca corsicana pop, 1947 domínguez et al. 2015; pérez-losada et al. 2015 scherotheca savignyi bouché, 1972 domínguez et al. 2015; pérez-losada et al. 2015 scherotheca sp 1 pérez-losada et al. 2009, 2015; domínguez et al. 2015 scherotheca sp 2 pérez-losada et al. 2009, 2015; domínguez et al. 2015 zophoscolex cyaneus briones & díaz cosín, 1993 domínguez et al. 2018 european journal of taxonomy 701: 1–22 (2020) 22 appendix 2. phylogenetic tree obtained from the concatenated sequence of coi–16s–nd1–28s molecular markers. 1 european journal of taxonomy 789: 1–10 issn 2118-9773 https://doi.org/10.5852/ejt.2022.789.1623 www.europeanjournaloftaxonomy.eu 2022 · cavalcante b.m.s. & johanson k.a. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:6cde7b15-c377-40c9-9a76-345c0bd7085a three new species of oxyethira eaton, 1873 (trichoptera: hydroptilidae) from new caledonia bruna maria silva cavalcante 1,* & kjell arne johanson 2 1 laboratório de entomologia, departamento de zoologia, instituto de biologia, universidade federal do rio de janeiro, caixa postal 68044, 21941-971, rio de janeiro, rj, brazil. 1 programa de pós-graduação em zoologia, museu nacional, universidade federal do rio de janeiro, rj, brazil. 2 zoology department, swedish museum of natural history, box 50007, se-104 05 stockholm, sweden. * corresponding author: bmsilva.bio@gmail.com 2 email: kjell.arne.johanson@nrm.se 1 urn:lsid:zoobank.org:author:7c9d157e-b681-4372-88db-f79ee3a551eb 2 urn:lsid:zoobank.org:author:f2a38cf6-59eb-4f88-bfeb-761dbea7b01a abstract. oxyethira eaton, 1873 is one of the most diverse genera of hydroptilidae, comprising over 240 species distributed in all biogeographical regions. here three new species of oxyethira (trichoglene) neboiss, 1977 are described and illustrated from male specimens collected in new caledonia: o. (trichoglene) hamus sp. nov., recognized by the hook-shaped apex of the long inferior appendages in lateral view and by the posterior margin of segment ix with a trilobed appearance in ventral view; o. (trichoglene) rectangulata sp. nov., recognized by the rectangular shape of the inferior appendages, which are totally fused and with two pairs of small setae on the inner face; and o. (trichoglene) spiralis sp. nov., recognized by the strongly curvilinear shape of the subgenital process in dorsal and lateral views and by the long process spiralling around the ejaculatory duct at the phallus apex. key words. hydroptilidae, new caledonia, microcaddisflies, oxyethira, taxonomy. cavalcante b.m.s. & johanson k.a. 2022. three new species of oxyethira eaton, 1873 (trichoptera: hydroptilidae) from new caledonia. european journal of taxonomy 789: 1–10. https://doi.org/10.5852/ejt.2022.789.1623 introduction the microcaddisfly genus oxyethira eaton, 1873 (trichoptera: hydroptilidae: hydroptilinae) includes 247 extant and one fossil species (morse et al. 2021). it has been recorded from all zoogeographic regions, and is especially diverse in the new world, with more than 100 species registered (holzenthal & calor 2017). oxyethira species are currently grouped into 11 subgenera, but 29 species remain as incertae sedis (kelley 1984, 1989; oezdikmen 2007; morse et al. 2021). the subgenus trichoglene neboiss, 1977 (26 species) has an australasian distribution and occurs in australia, new caledonia and new zealand (wells & johanson 2015; morse et al. 2021). https://doi.org/10.5852/ejt.2022.789.1623 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:6cde7b15-c377-40c9-9a76-345c0bd7085a mailto:bmsilva.bio%40gmail.com?subject= mailto:kjell.arne.johanson%40nrm.se?subject= http://zoobank.org/urn:lsid:zoobank.org:author:7c9d157e-b681-4372-88db-f79ee3a551eb http://zoobank.org/urn:lsid:zoobank.org:author:f2a38cf6-59eb-4f88-bfeb-761dbea7b01a https://doi.org/10.5852/ejt.2022.789.1623 european journal of taxonomy 789: 1–10 (2022) 2 a total of 26 species of oxyethira are registered from new caledonia belonging to 3 subgenera: dampfitrichia mosely, 1937 (one species), pacificotrichia kelley, 1989 (13 species) and trichoglene neboiss, 1977 (11 species). another species, o. macropennis wells & johanson, 2015, remains unplaced to subgenus (wells & johanson 2015; johanson & wells 2019). the diagnostic characteristics for the males in the subgenus trichoglene are as follows: segment viii without excision (considered as a plesiomorphic feature); short titillator (even absent in some species); phallus with a curved spine-shaped process in the sub-distal region; and subgenital processes widely separated and partially fused with the pleural region of segment ix (the two latter are considered apomorphic features) (kelley 1984). three species groups are recognized among the new caledonian members of the subgenus trichoglene: the spinifera-group, with abdominal segment ix subquadrate; the caledoniensis-group, with venter of abdominal segment ix in ventral view produced anteriorly, proximally either rounded or tapered and somewhat triangular; and the insularis-group, with y-shaped inferior appendages (wells & johanson 2015). in this study, three new oxyethira (trichoglene) species are described, two belonging to the caledoniensis-group (o. spiralis sp. nov. and o. hamus sp. nov.) and one belonging to the spiniferagroup (o. rectangulata sp. nov.). material and methods the material was collected using malaise traps (gressit & gressit 1952) and preserved in 80% ethanol. for examination of the male genital structures, the abdomen was removed and cleared in 10% koh and mounted temporarily in glycerin or glycerin jelly on a slide for viewing and drawing. pencil sketches were made using a drawing tube mounted on a leitz laborlux s light microscope, and then scanned and used as templates to produce vector graphics in adobe illustrator ver. cs6. the removed abdomens were then stored permanently with the rest of their respective bodies in microvials with 80% ethanol. the terminology used in the descriptions follows that of kelley (1984). institutional abbreviations mnhn = muséum national d’histoire naturelle, paris, france nhrs = swedish museum of natural history, stockholm, sweden abbreviations of genital structures bp = bilobed process ej = ejaculatory duct ia = inferior appendages sg = subgenital processes viii = segment viii ix = segment ix cavalcante b.m.s. & johanson k.a., three new species of oxyethira eaton, 1873 3 results class insecta linnaeus, 1758 order trichoptera kirby, 1813 family hydroptilidae stephens, 1836 genus oxyethira eaton, 1873 oxyethira (trichoglene) hamus sp. nov. urn:lsid:zoobank.org:act:200e6d45-51b4-4012-a3b1-4fe519ed65f6 fig. 1a–d diagnosis this new species belongs to the group of oxyethira caledoniensis. it is similar to o. houailou wells & johanson, 2015 by the presence of strong sclerotized margins of the inferior appendages, each one bearing one seta in the mesal region as seen in ventral view; and by the general shape of the phallus apex, with a subdistal spine curved to the left in dorsal view. oxyethira (trichoglene) hamus sp. nov. is recognized by the hook-shaped apex of the long inferior appendages in lateral view (short and with pointed apex in o. houailou) and by the posterior margin of segment ix having a trilobed appearance in ventral view (forming a subtriangular projection in o. houailou). etymology ʻhamusʼ, ʻhookʼ in latin, refers to the shape of the inferior appendages in lateral view. material examined holotype new caledonia • ♂; province sud haute yaté fauna reserve, 1760 m s of bridge pont perignon, 50 m upstream from bridge over stream; 22.14954° s, 166.701211° e; alt. 180 m; 14 dec. 2003–13 jan. 2004; k.a. johanson leg.; loc#081; malaise trap; mnhn. paratypes 9 ♂♂; same collection data as for holotype; nhrs. description body. male antennae with 18 flagellomeres. forewing length 1.4–1.8 mm (n = 10, mean 1.5 mm); tibial spurs 0, 3, 4; forewings with venation typical for subgenus. abdominal sternite vii without mesoventral process. male genitalia (fig. 1a–d). abdominal segment viii almost cylindrical, with shallow incision at posterior margin in ventral view (fig. 1c). abdominal segment ix inserted within segment viii; anterior margin forming three lobes in ventral view (fig. 1c), surpassing anterior margin of segment viii (fig. 1a–b). segment x not visible. subgenital processes slightly curvilinear in dorsal view (fig. 1b), each with pointed apex curved downward in lateral view and one bilobed process at base (fig. 1a). inferior appendages fused, with strongly sclerotized margins, bearing one pair of small mesal setae at posterior margin visible in dorsal (fig. 1b) and ventral views (fig. 1c), and one pair of small setae at inner margin visible in dorsal view (fig. 1b); posterior margin slightly concave; in lateral view hook-shaped with apex curved backward and touching base of subgenital process (fig. 1a). setal lobes absent. aedeagus elongate; in subdistal region, slender spine pointing left in dorsal view. titillator absent (fig. 1d). http://zoobank.org/urn:lsid:zoobank.org:act:200e6d45-51b4-4012-a3b1-4fe519ed65f6 european journal of taxonomy 789: 1–10 (2022) 4 fig. 1. oxyethira (trichoglene) hamus sp. nov., male genitalia. a. lateral. b. dorsal. c. ventral. d. aedeagus, dorsal. abbreviations: see material and methods. scale bars = 0.1 mm. cavalcante b.m.s. & johanson k.a., three new species of oxyethira eaton, 1873 5 oxyethira (trichoglene) rectangulata sp. nov. urn:lsid:zoobank.org:act:c930eb33-5e37-49c2-8a94-882e97eaf988 fig. 2a–d diagnosis oxyethira (trichoglene) rectangulata sp. nov. belongs to the group of oxyethira spinifera. it is similar to o. tiwaka wells & johanson, 2015 by the quadrangular shape of segment ix in ventral view, and in lateral view by the inferior appendages being divided into a dorsal part forming a hump and a ventral part forming a pointed projection. this species can be recognized by the rectangular shape of the inferior appendages in dorsal and ventral views, which are totally fused, bearing two pairs of small setae on the inner face (in o. tiwaka these appendages are much smaller and the ventral part forms a quadrate-shaped sclerotized projection in ventral view); by the shape of the subgenital process in lateral view, which is thinner and almost straight in o. rectangulata sp. nov., and thicker, c-shaped in o. tiwaka. etymology ʻrectangulataʼ, from latin, ʻrectangularʼ, refers to the shape of the inferior appendages in dorsal and ventral view. material examined holotype new caledonia • ♂; province sud, haute yaté fauna reserve, 1760 m s of bridge pont perignon, 50 m upstream from bridge over stream; 22.14954° s, 166.701211° e; alt. 180 m; 14 dec. 2003–13 jan. 2004; k.a. johanson leg.; loc#081; malaise trap; mnhn. paratypes 2 ♂♂; same collection data as for holotype; nhrs. description body. male antennae with 20 flagellomeres. forewing length 1.7–1.8 mm (n = 3); tibial spurs 0, 3, 4; forewings with venation typical for subgenus. abdominal sternite vii with a small, pointed mesoventral process. male genitalia (fig. 2a–d). abdominal segment viii cylindrical with wide, v-shaped incision in ventral view (fig. 2c). abdominal segment ix inserted within segment viii; anterior margin slightly concave, not surpassing anterior margin of segment viii (fig. 2b–c). segment x not visible. subgenital processes almost straight in dorsal (fig. 2b) and ventral views (fig. 2c), each bearing one bilobed process at base. inferior appendages fused, rectangular in dorsal (fig. 2b) and ventral views (fig. 2c), bearing two pairs of small mesal setae at inner margin visible in dorsal view (fig. 2b); divided into two parts in lateral view (fig. 2a): dorsal part forming a truncate hump, ventral part forming pointed projection. setal lobes absent. aedeagus elongate, with slender apical spine curved left in dorsal view. ejaculatory duct extending freely at apex. titillator absent (fig. 2d). http://zoobank.org/urn:lsid:zoobank.org:act:c930eb33-5e37-49c2-8a94-882e97eaf988 european journal of taxonomy 789: 1–10 (2022) 6 fig. 2. oxyethira (trichoglene) rectangulata sp. nov., male genitalia. a. lateral. b. dorsal. c. ventral. d. aedeagus, dorsal. scale bars = 0.1 mm. cavalcante b.m.s. & johanson k.a., three new species of oxyethira eaton, 1873 7 oxyethira (trichoglene) spiralis sp. nov. urn:lsid:zoobank.org:act:5027693e-fc66-4256-b619-34c77ca3e6ff fig. 3a–d diagnosis based on general similarities, oxyethira (trichoglene) spiralis sp. nov. is tentatively placed in the group of oxyethira caledoniensis. it is similar to o. caledoniensis kelley, 1989 by the shape of segment ix in ventral and lateral views, with the anterior margin only slightly produced anteriorly and by the shape of the apparently fused inferior appendages. in o. caledoniensis the inferior appendages are widely separated and form bilobed sclerotized structures that are absent in o. spiralis sp. nov. the new species can be recognized in lateral view by the subquadrate shape of the inferior appendages, by the strongly curvilinear shape of the subgenital process in dorsal and lateral views, and by the long process spiralling around the ejaculatory duct at the phallus apex. etymology ʻspiralisʼ, from latin, ʻspiralʼ, referring to the shape of the long process present around the ejaculatory duct in this species. material examined holotype new caledonia • ♂; province nord, western slope of kugüö mt, xwé wida di stream, 2 km e of koh; 21°35.474′ s, 165°50.776′ e; alt. 154 m; 20 nov.–12 dec. 2003; k.a. johanson leg.; loc#036; malaise trap; mnhn. description body. male antennae with 22 flagellomeres. forewing length 1.7 mm (n = 1); tibial spurs 0, 2, 4; forewings with venation typical for subgenus. abdominal sternite vii with a small, pointed mesoventral process. male genitalia (fig. 3a–d). abdominal segment viii cylindrical without excisions. abdominal segment ix inserted in segment viii; anterior margin slightly produced anteriorly (fig. 3a–b). segment x not visible. subgenital processes with strongly sigmoid shape in dorsal (fig. 3b) and ventral views (fig. 3c), each bearing one bilobed process at base. inferior appendages fused and almost quadrate in lateral view (fig. 3a). setal lobes absent. aedeagus elongate, long process spiralling around ejaculatory duct; ejaculatory duct extending freely in middle of two weakly sclerotized projections at phallus apex. titillator absent (fig. 3d). discussion despite the great morphological variation present in the genitalia of oxyethira, all species described here present characteristics unique for the subgenus trichoglene (kelley 1984): segment viii without modification; and widely separated subgenital processes that are partially fused with the pleural region of segment ix. there are different distribution patterns for oxyethira species in new caledonia ranging from wide to localised, or even disjunct (north-south extremes) (wells & johanson 2015). oxyethira (trichoglene) hamus sp. nov. and o. (trichoglene) rectangulata sp. nov have their distribution restricted to the type locality in the southern region of new caledonia (province sud). other species previously described have a similar distribution pattern on the island: oxyethira (trichoglene) amieu wells & johanson, 2015, o. (trichoglene) insularis kelley, 1989, o. (trichoglene) parinsularis wells & johanson, 2015 http://zoobank.org/urn:lsid:zoobank.org:act:5027693e-fc66-4256-b619-34c77ca3e6ff european journal of taxonomy 789: 1–10 (2022) 8 and o. (trichoglene) perignonica wells & johanson, 2015. these species can be well localized or widely distributed in the sud province but are never found in the northern region (province nord) (wells & johanson 2015). oxyethira (trichoglene) spiralis sp. nov is only known from its type locality in the province nord. fig. 3. oxyethira (trichoglene) spiralis sp. nov., male genitalia. a. lateral. b. dorsal. c. ventral. d. aedeagus, dorsal. scale bars = 0.1 mm. cavalcante b.m.s. & johanson k.a., three new species of oxyethira eaton, 1873 9 wells & johanson (2015) reviewed the new caledanian oxyethira species and described 17 new species, bringing the total number of species for the island to 26. of these, 11 species belong to trichoglene. together with the herein described species the diversity of oxyethira on new caledonia is increased by about 10%, and of those in trichoglene by almost 30%. the three new species were found among several thousand new caledonian hydroptilidae stored in ethanol and collected at the end of 2003 and beginning of 2004. this means that new species can still be found in material from well-studied areas that have given rise to descriptions of dozens of new species in the family hydroptilidae (wells & johanson 2012, 2014, 2015; wells, johanson & mary-sasal 2013). this shows the potential for finding new hydroptilidae in new caledonia and that undescribed species are still available in collections or likely not yet collected. acknowledgments this work was possible thanks to the sandwich doctoral scholarship provided by capes (coordenação de aperfeiçoamento de pessoal de nível superior (process n. 88887.508066/2020-00)) and the scholarship provided by cnpq (conselho nacional de desenvolvimento científico e tecnológico (process n. 141121/2018-1)), to which we are highly grateful. references gressit j.l. & gressit m.k. 1952. an improved malaise trap. pacific insects 4: 87–90. holzenthal r.w. & calor a.r. 2017. catalog of the neotropical trichoptera (caddisflies). zookeys 654: 1–566. https://doi.org/10.3897/zookeys.654.9516 johanson k.a. 2011. the ecnomid caddisflies: generic composition and a male-based generic key, with description of caledomina noumea n. g. et n. sp. from new caledonia (trichoptera: ecnomidae). annales de la société entomologique de france (nouvelle série) 47: 344–349. https://doi.org/10.1080/00379271.2011.10697726 johanson k.a. & wells a. 2019. new caledonia’s trichoptera – present status of knowledge. zoosymposia 14: 87–102. https://doi.org/10.11646/zoosymposia.14.1.12 kelley r.w. 1984. phylogeny, morphology and classification of the micro-caddisfly genus oxyethira eaton (trichoptera: hydroptilidae). transactions of the american entomological society 110: 435–463. kelley r.w. 1989. new species of micro-caddisflies (trichoptera: hydroptilidae) from new caledonia, vanuatu and fiji. proceedings of the entomological society of washington 91 (2): 190–202. morse j.c. 2021. trichoptera world checklist. available from http://entweb.sites.clemson.edu/database/trichopt/index.htm [accessed 15 oct. 2021]. özdikmen h. 2007. a nomenclatural act: replacement names for two homonymous caddisfly generic names (trichoptera). munis entomology & zoology 2 (2): 443–444. wells a. & johanson k.a. 2012. review of the new caledonian species of paroxyethira mosely, 1924 (trichoptera: hydroptilidae). zootaxa 3478 (1): 330–344. https://doi.org/10.11646/zootaxa.3478.1.31 wells a. & johanson k.a. 2014. review of the new caledonian species of acritoptila wells, 1982 (trichoptera, insecta), with descriptions of 3 new species. zookeys 397: 1–23. https://doi.org/10.3897/zookeys.397.7059 wells a. & johanson k.a. 2015. review of new caledonian species of oxyethira eaton, with description of 17 new species, and new records for hydroptila dalman and hellyethira neboiss (trichoptera, hydroptilidae). zookeys 530: 37–90. https://doi.org/10.3897/zookeys.530.6047 https://doi.org/10.3897/zookeys.654.9516 http://dx.doi.org/10.1080/00379271.2011.10697726 https://doi.org/10.11646/zoosymposia.14.1.12 http://entweb.sites.clemson.edu/database/trichopt/index.htm https://doi.org/10.11646/zootaxa.3478.1.31 https://doi.org/10.3897/zookeys.397.7059 https://doi.org/10.3897/zookeys.530.6047 european journal of taxonomy 789: 1–10 (2022) 10 wells a., johanson k.a. & mary-sasal n. 2013. the new caledonian genus caledonotrichia sykora (trichoptera, insecta) reviewed, with descriptions of 6 new species. zookeys 287: 59–89. https://doi.org/10.3897/zookeys.287.4615 manuscript received: 17 september 2021 manuscript accepted: 17 november 2021 published on: 12 january 2022 topic editor: nesrine akkari section editor: helen m. barber-james desk editor: eva-maria levermann printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. https://doi.org/10.3897/zookeys.287.4615 a taxonomic review of the genus asterostegus (echinodermata: ophiuroidea), with the description of a new species – corrigendum masanori okanishi1,* & toshihiko fujita2 1 seto marine biological laboratory, field science education and research center, kyoto university, 459 shirahama, nishimuro, wakayama, 649-2211 japan 2 national museum of nature and science, 4-1-1 amakubo, tsukuba, ibaraki, 305-0005 japan * corresponding author. e-mail: mokanishi@tezuru-mozuru.com okanishi m. & fujita t. 2014. a taxonomic review of the genus asterostegus (echinodermata: ophiuroidea), with the description of a new species – corrigendum. european journal of taxonomy 84: 1–18. http://dx.doi. org/10.5852/ejt.2014.84 one sentence of the abstract (p. 1) should be corrected as follows. from: “a new species, asterostegus sabineae sp. nov. is described from off reunion island and two other species, a. tuberculatus mortensen, 1933 and a. maini, are redescribed.” to: “ a new species, asterostegus sabineae sp. nov. is described from off madagascar island and two other species, a. tuberculatus mortensen, 1933 and a. maini, are redescribed.” one sentence in materials and methods (p. 3) should be corrected as follows. from: “the holotype (mnhn mo-01) and four paratypes of the new species were fixed in 99% ethanol (smnh-type-8333; smnh-type-8536).” to: “ the holotype (mnhn ie-2013-8012) and four paratypes of the new species were fixed in 99% ethanol (smnh-type-8333; smnh-type-8536).” a catalogue number of one paratype specimen of asterostegus sabineae (smnh-type-85436) was misspelled. thus the type material account of asterostegus sabineae sp. nov. (p. 4) should be corrected as follows. from: “...holotype (mnhn ie-2013-8012), three paratypes (smnh-type-8333) and one paratype (smnh-type-85436), all ethanol preserved specimens...” to: “...holotype (mnhn ie-2013-8012), three paratypes (smnh-type-8333) and one paratype (smnh-type-8546), all ethanol preserved specimens...” material examined section of asterostegus tuberculatus (p. 12) should be corrected as follows. from: “smnh-123461, one ethanol preserved specimen, west coast of reunion island, 500 m, 2009.” to: “ smnh-123461, one ethanol preserved specimen, west coast of reunion island, 500 m, 2009, collected by t. molochau (aquarium reunion)” european journal of taxonomy 84: 1-2 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2014.84 www.europeanjournaloftaxonomy.eu 2014 · okanishi m. & fujita t. this work is licensed under a creative commons attribution 3.0 license. c o r r i g e n d u m the following corrections have been made to paper no. 76 (http://dx.doi.org/10.5852/ejt.2014.76) 1 mailto:mokanishi%40tezuru-mozuru.com?subject=mokanishi%40tezuru-mozuru.com http://dx.doi.org/10.5852/ejt.2014.84 http://dx.doi.org/10.5852/ejt.2014.84 http://dx.doi.org/10.5852/ejt.2014.84 http://www.europeanjournaloftaxonomy.eu/index.php/ejt https://creativecommons.org/licenses/by/3.0/ http://dx.doi.org/10.5852/ejt.2014.76 acknowledgements (p. 17) should be replaced by the following. “we are most grateful to sadie mills (niwa) for her critical reading of the manuscript and constructive comments. we wish to express our sincere gratitude to owen anderson and kareen schnabel (niwa) for their assistance with the examination of the holotype of asterostegus maini and to sabine stöhr (smnh) and marc eléaume (mnhn) for arranging the loans of examined specimens. we are grateful to thierry molochau from aquarium reunion for donating the specimen of a. tuberculatus to smnh. this work was supported by grants from the research institute of marine invertebrates (tokyo), the showa seitoku memorial foundation, and the japan society for the promotion of society (scientific research [c] no. 22570104, research fellowships for young scientists no. 22506). the atimo vatae expedition to south madagascar (principal investigator, philippe bouchet) was part of a cluster of mozambique-madagascar expeditions funded by the total foundation, prince albert ii of monaco foundation, and stavros niarchos foundation under “our planet reviewed”, a joint initiative of mnhn and pro natura international (pni) in partnership with institut d’halieutique et des sciences marines, university of toliara (ih.sm) and the madagascar bureau of wildlife conservation society (wcs). institut de recherche pour le developpement (ird) deployed its research catamaran antea.” published on: 23 may 2014 printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; national botanic garden of belgium, meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark. european journal of taxonomy 84: 1–2 (2014) 2 european journal of taxonomy 183: 1–26 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2016.183 www.europeanjournaloftaxonomy.eu 2016 · turner i.m. & utteridge t.m.a. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e 1 whither polyalthia (annonaceae) in peninsular malaysia? synopses of huberantha, maasia, monoon and polyalthia s.s. ian m. turner * & timothy m.a. utteridge royal botanic gardens kew, richmond, surrey, tw9 3ae, united kingdom * corresponding author: turner187@btinternet.com abstract. an updated classification of polyalthia in peninsular malaysia is presented. a synopsis (listing of species with synonymy and typification, and keys to species) is presented for the genera huberantha, maasia, monoon and polyalthia sensu stricto. one new species (polyalthia pakdin i.m.turner & utteridge sp. nov.) is described and a conservation assessment presented for it. monoon xanthopetalum merr. represents a new record for peninsular malaysia. six new lectotypes are designated. keywords. enicosanthum, huberantha, maasia, monoon, polyalthia. turner i.m. & utteridge t.m.a. 2016. whither polyalthia (annonaceae) in peninsular malaysia? synopses of huberantha, maasia, monoon and polyalthia s.s. european journal of taxonomy 183: 1–26. http://dx.doi. org/10.5852/ejt.2016.183 introduction when sinclair (1955) revised the annonaceae of the malay peninsula he recognised 32 species of polyalthia blume (including polyalthia evecta finet & gagnep. from peninsular thailand still unrecorded from peninsular malaysia, and the cultivated polyalthia longifolia (sonn.) thwaites). the 30 native species made polyalthia the largest genus in the family as represented in the malayan flora. the genus was characterised by sinclair largely in terms of floral morphology including subequal corolla whorls of spreading, relatively flat, petals, numerous flat-topped stamens and many carpels with 1–5 ovules each. sinclair considered the genus to be divisible into two sections that he called ‘polyalthia section eu-polyalthia’ (correctly polyalthia section polyalthia) and polyalthia section monoon (miq.) benth. & hook.f. sinclair appeared confident in the classification, noting the polyalthia was a large genus ‘but very uniform’. however, in recent years the homogeneity of polyalthia has come under close scrutiny with a growing conviction that it represented an unnatural grouping. molecular analyses have confirmed the polyphyletic nature of polyalthia sensu lato leading to a process of dismemberment as species have been transferred to various different genera. of relevance to the species included in polyalthia by sinclair, section monoon has been raised again to generic rank (xue et al. 2012) (14 spp. of sinclair’s native polyalthia species) and includes enicosanthum becc. (7 spp. recognised by sinclair). three of sinclair’s section monoon species are now recognised in the new genus maasia mols et al. (mols et al. 2008) and two in huberantha chaowasku in chaowasku et al. [chaowasku et al. 2015, originally named hubera chaowasku in chaowasku et al. (chaowasku et al. 2012) but this was considered to be an illegitimate later homonym of huberia dc. (melastomataceae)]. the 17 species sinclair recognised in ‘section eu-polyalthia’ are all now deemed to belong in polyalthia sensu stricto. http://dx.doi.org/10.5852/ejt.2016.183 http://www.europeanjournaloftaxonomy.eu https://creativecommons.org/licenses/by/3.0/ mailto:turner187@btinternet.com http://dx.doi.org/10.5852/ejt.2016.183 http://dx.doi.org/10.5852/ejt.2016.183 european journal of taxonomy 183: 1–26 (2016) 2 some asian polyalthia species have also been transferred to marsypopetalum scheff. (xue et al. 2011), but this has not affected any of the species native to peninsular malaysia. in this paper, we review these changes in the taxonomy of polyalthia and associated genera in peninsular malaysia. this is done by providing synopses of the four genera huberantha, maasia, monoon miq. and polyalthia including keys to the species. one new species of polyalthia is described and a conservation assessment for it provided. materials and methods material in the herbarium of the royal botanic gardens kew (k) and a loan of annonaceae specimens from the herbarium of the forest research insitute of malaysia (kep) was examined. for the new species, specimens were geo-referenced using hamidah et al. (2011), and a conservation assessment was made following iucn criteria (iucn 2012) using geocat to estimate the extent of occurrence (eoo) and area of occupancy (aoo) (http://geocat.kew.org; bachman et al. 2011). results order magnoliales juss. ex bercht. & j.presl family annonaceae juss. subfamily malmeoideae chatrou & al. tribe miliuseae hook.f. & thomson key to genera 1. lower surface of leaves densely covered with minute papillae giving a white or glaucous appearance .............................................................................................................................................. maasia – leaves not white or glaucous below .................................................................................................. 2 2. leaf venation herringbone (eucamptodromous) with insertion of lateral veins to the midrib decurrent; intercostal venation scalariform to subscalariform ............................................................. monoon – leaf venation looping (brochidodromous) with insertion of the lateral veins to the midrib not decurrent; intercostal venation reticulate ........................................................................................... 3 3. leaf base generally more or less symmetrical, not (sub)cordate. ovules 1 per carpel. seeds 1 per monocarp ...................................................................................................................... huberantha – leaf base generally more or less asymmetrical, often (sub)cordate. ovules usually 2–6 per carpel. seeds usually 2 or more per monocarp ............................................................................ polyalthia maasia mols et al. systematic botany 33: 493 (mols et al. 2008). – type: maasia hypoleuca (hook.f. & thomson) mols et al. key to species of maasia 1. leaves generally not exceeding 8 cm long, 3 cm wide. flowering pedicel to 5 mm long, petals to 6 mm long. monocarps ellipsoidal, few (5 or less), not exceeding 2 cm long .............. m. hypoleuca – leaves generally more than 8 cm long, 3 cm wide. flowering pedicel more then 5 mm long, petals more than 6 mm long. monocarps globose or if ellipsoidal then many (generally 10 or more) and more then 2 cm long at maturity ........................................................................................................ 2 http://geocat.kew.org turner i.m. & utteridge t.m.a., whither polyalthia (annonaceae) in peninsular malaysia? 3 2. flowering pedicel more than 17 mm long, c. 1 mm thick, petals more than 1 cm long. monocarps ellipsoidal ...................................................................................................................... m. sumatrana – flowering pedicel to 17 mm long, c. 0.5 mm thick, petals to 8 mm long. monocarps globose ............. ............................................................................................................................................. m. glauca maasia glauca (hassk.) mols et al. systematic botany 33: 493 (mols et al. 2008). – uvaria glauca hassk., flora (beiblätter) 25, 2 (2): 31 (hasskarl 1842). – guatteria glauca (hassk.) miq., flora van nederlandsch indië 1 (2): 49 (miquel 1858), nom. illegit., non g. glauca ruiz & pav. (ruiz & pavon 1798). – monoon glaucum (hassk.) miq., annales musei botanici lugdunu-batavi 2: 19 (miquel 1865). – polyalthia glauca (hassk.) f.muell., descriptive notes on papuan plants. appendix: 95 (von mueller 1877). – lectotype (designated by turner 2011: 46): indonesia, java, j.k. hasskarl s.n. (l (l 0188666)). – epitype (designated by turner 2011: 46): indonesia, java: udjung kulon reserve, mt pajung, 17 nov. 1960, a.j.g.h. kostermans et al. unesco 165 (holo-: l (l 0188665); iso-: a, bo, g, k). guatteria hypoleuca miq., flora van nederlandsch indië. supplementum primum. prodromus florae sumatranae: 381 (miquel 1861). – lectotype (designated by rogstad 1989: 209): indonesia, sumatra: priaman, s. dat., h. diepenhorst s.n. [herb. bogor. no. 2095] (u (u 0000368)). unona merrittii merr., philippine journal of science 1, supplement iii: 190 (merrill 1906). — polyalthia merrittii (merr.) merr., philippine journal of science. section c, botany 10: 250 (merrill 1915). – lectotype (designated by turner 2011: 46): philippines, mindoro, bongabong river, feb. 1906, h.n. whitford 1447 (hololecto-: k (k000691629); isolecto-: us). polyalthia parkinsonii hutch., bulletin of miscellaneous information, kew 1917: 25 (hutchinson 1917). – lectotype (designated by turner 2011: 46): india, andaman islands, bom-ling-la, 11 feb. 1916, c.e. parkinson 943 (hololecto-: k (k000691470); isolecto-: k (k000691469)). description and distribution sinclair (1955: 321), kochummen (1972: 88), rogstad (1989: 209–212), turner (2014: 74–75). maasia hypoleuca (hook.f. & thomson) mols et al. systematic botany 33: 493 (mols et al. 2008). – polyalthia hypoleuca hook.f. & thomson, the flora of british india 1: 63 (hooker & thomson 1872). – lectotype (designated by rogstad 1989: 227): singapore, sep. 1867, a.c. maingay 1516a [kew distrib. no. 50] (k (k000691444)). description and distribution sinclair (1955: 320–321), kochummen (1972: 88), rogstad (1989: 227–229), turner (2014: 75–76). maasia sumatrana (miq.) mols et al. systematic botany 33: 493 (mols et al. 2008). – guatteria sumatrana miq., flora van nederlandsch indië. supplementum primum. prodromus florae sumatranae: 380 (miquel 1861). – monoon sumatranum (miq.) miq., annales musei botanici lugdunu-batavi 2: 19 (miquel 1865). – polyalthia sumatrana (miq.) kurz, journal of the asiatic society of bengal. part ii (natural history &c.) 43: 53 (kurz 1874). – lectotype (designated by rogstad 1989: 220): indonesia, sumatra, priaman province, h. diepenhorst s.n. [herb. bogor. 2342] (u (u 0000372)). description and distribution sinclair (1955: 319–320), kochummen (1972: 91), rogstad (1989: 229–221), turner (2014: 76–78). european journal of taxonomy 183: 1–26 (2016) 4 huberantha chaowasku in chaowasku et al. kew bulletin 70 (2)-23: 1 (chaowasku et al. 2015). – hubera chaowasku in chaowasku et al., phytotaxa 69: 46 (chaowasku et al. 2012), non huberia dc. (de candolle 1828). – type: hubera cerasoides (roxb.) chaowasku. key to species of huberantha 1. sepals suborbicular, to 3 mm long; petals drying uniformly pale brown ........................................... ............................................................................................................…......................… h. jenkinsii – sepals triangular, c. 6 mm long or more; petals often drying with powdery yellow patches ……..... .......................................................................................................................................… h. rumphii huberantha jenkinsii (hook.f. & thomson) chaowasku in chaowasku et al. kew bulletin 70 (2)-23: 2 (chaowasku et al. 2015). – guatteria jenkinsii hook.f. & thomson, flora indica: 141 (hooker & thomson 1855). — polyalthia jenkinsii (hook.f. & thomson) hook.f. & thomson, the flora of british india 1: 64. (hooker & thomson 1872). – hubera jenkinsii (hook.f. & thomson) chaowasku in chaowasku et al., phytotaxa 69: 48 (chaowasku et al. 2012). – lectotype (designated by turner 2011: 73): india, assam, s.dat., f. jenkins s.n. (k (barcode no. k000691481)). polyalthia andamanica kurz ex king, journal of the asiatic society of bengal. part ii (natural history &c.) 61: 53 (king 1892). – lectotype (designated by turner 2011: 74): india, andaman islands, south andaman, namanagha, 13 dec. 1890, king’s collector [h.h. kunstler] s.n. (hololecto-: k (k000739193); isolecto-: k). polyalthia havilandii boerl., icones bogorienses 1: 107, t. 66 (boerlage 1899a). – type: borneo, sarawak, near kuching, 15 feb. 1894, g.d. haviland 2 (holo-: bo (sheet no. bo-1352593); iso-: l (l 0188829)). polyalthia cumingiana merr., publications of the bureau of science government laboratories 35: 71 (merrill 1905). – lectotype (designated by turner 2011: 74): philippines, luzon, prov. tayabas, h. cuming 827 (hololecto-: k (k000691639); isolecto-: k [×2], ny, wu). unona agusanensis elmer, leaflets of philippine botany 5: 1743 (elmer 1913). – polyalthia agusanensis (elmer) merr., philippine journal of science. section c, botany 10: 250 (merrill 1915). – lectotype (designated by turner 2011: 74): philippines, mindanao, province of agusan, cabadbaran (mt urdaneta), sep. 1912, a.d.e. elmer 13654 (hololecto:ny (00026313); isolecto-: a, k, l). description and distribution sinclair (1955: 305–306), kochummen (1972: 88), turner (2014: 71–72). huberantha rumphii (blume ex hensch.) chaowasku in chaowasku et al. kew bulletin 70 (2)-23: 2 (chaowasku et al. 2015). — guatteria rumphii blume ex hensch., vita g. e. rumphii: 153 (henschel 1833). – polyalthia rumphii (blume ex hensch.) merr., an enumeration of philippine flowering plants 2: 162 (merrill 1923). – hubera rumphii (blume ex hensch.) chaowasku in chaowasku et al., phytotaxa 69: 50 (chaowasku et al. 2012). – type: entirely based on arbor nigra parvifolia of rumphius, herbarium amboinense 3: 10, t 4, f 2, t 5 (rumphius 1743). guatteria canangioides rchb.f. & zoll. in zollinger, linnaea 29: 322–323 (zollinger 1858). – monoon canangioides (rchb.f. & zoll.) miq., annales musei botanici lugdunu-batavi 2: 18 (miquel 1865). – polyalthia canangioides (rchb.f. & zoll.) boerl., catalogus plantarum phanerogamarum quae in horto botanico bogoriensi coluntur herbaceis exceptis (1): 19 (boerlage 1899b). – lectotype (designated by turner 2011: 78): indonesia, sumatra, province of lampong, h. zollinger 3047 (hololecto-: p (p01983362); isolecto-: ?bm, ?p). turner i.m. & utteridge t.m.a., whither polyalthia (annonaceae) in peninsular malaysia? 5 guatteria parveana miq., flora van nederlandsch indië 1 (2): 48 (miquel 1858). – polyalthia canangioides var. parveana (miq.) boerl., catalogus plantarum phanerogamarum quae in horto botanico bogoriensi coluntur herbaceis exceptis (1): 20 (boerlage 1899b). type: based on manuscript description by h. zollinger of uvaria parveana. unona borneensis miq., annales musei botanici lugdunu-batavi 2: 11 (miquel 1865). – desmos borneensis (miq.) merr. journal of the straits branch of the royal asiatic society, special number: 255 (merrill 1921). – lectotype (designated by turner 2011: 78): borneo, poeloe lampei, p.w. korthals s.n. (l (l 0189189)). polyalthia scortechinii king, journal of the asiatic society of bengal. part ii (natural history &c.) 61: 56 (king 1892). – lectotype (designated by turner 2011: 78): peninsular malaysia, perak, larut, may 1884, king’s collector [h.h. kunstler] 6125 (hololecto-: k (k000739187); isolecto-: cal, dd, k (k000739186)). polyalthia kunstleri king, journal of the asiatic society of bengal. part ii (natural history &c.) 61: 55 (king 1892). – lectotype (designated by turner 2011: 78): peninsular malaysia, perak, larut, jan. 1883, king’s collector [h.h. kunstler] 3767 (hololecto-: k (k000691450); isolecto-: cal, dd, k (k000691451), mel). polyalthia glandulosa merr., philippine journal of science. section c, botany 10: 247 (merrill 1915). – lectotype (designated by turner 2011: 78): philippines, mindanao, district of zamboanga, santa maria, near mount pulangbato, 4 oct. 1912, j. reillo bur. sci. 16460 (hololecto-: us (us00098663); isolecto-: k (k000691642), l, p (p00372685)). description and distribution sinclair (1955: 306–307), kochummen (1972: 89) turner (2014: 72). monoon miq. annales musei botanici lugdunu-batavi 2: 15 (miquel 1865). – lectotype (designated by saunders & xue 2011: 236): monoon lateriflorum (blume) miq. enicosanthum becc., nuovo giornale botanico italiano 3: 183 (beccari 1871). – type: enicosanthum paradoxum becc. marcuccia becc., nuovo giornale botanico italiano 3: 181, t. 3 (beccari 1871). – type: marcuccia grandiflora becc. griffithia maingay ex king, annals of the royal botanic garden (calcutta) 4: 8 (king 1893), non griffithia wight & arn. (wight & arnott 1834). – griffithianthus merr., philippine journal of science. section c, botany 10: 231 (merrill 1915). – lectotype (designated by merrill 1915: 231): griffithia magnoliipetala maingay ex king. woodiella merr., journal of the straits branch of the royal asiatic society 85: 187 (merrill 1922), non woodiella sacc. & p.syd. (sydow 1899). – woodiellantha rauschert, taxon 31: 555 (rauschert 1982). – type: woodiella sympetala merr. cleistopetalum h.okada, acta phytotaxonomica et geobotanica 47: 4 (okada 1996). – type: cleistopetalum borneense h.okada. key to species of monoon 1. inflorescences on trunk or restricted to base of trunk ........................................................................ 2 – inflorescences in leaf axils, or on twigs behind leaves (sometimes on branches) .............................. 6 2. inflorescences borne on slender, flexible, branched basal flagellae ........................... m. hypogaeum – not so ................................................................................................................................................ 3 3. inflorescences on stout, sparsely branching axes to at least 7 cm long ....................... m. praestigiosum european journal of taxonomy 183: 1–26 (2016) 6 – inflorescences on woody tubercles or much-branched axes less than 7 cm long ............................... 4 4. inflorescences on woody tubercles .............................................................................. m. anomalum – inflorescences on much-branched axes ............................................................................................. 5 5. leaves with 8–10 pairs of lateral nerves. outer petals to 2.5 cm long, 0.5 cm wide. monocarps with blunt beak ...................................................................................................................... m. borneense – leaves with c. 20 pairs of lateral nerves. outer petal to 7 cm long, 2.5 cm wide. monocarps with hard, sharp beak ................................................................................................................ m. congregatum 6. leaves membranous. inflorescences axillary, few-flowered (generally more than one). sepals more than 8 mm long. monocarps globose to ellipsoidal with rounded to slightly blunt apex ..................... ............................................................................................................................ m. membranifolium – leaves chartaceous to coriaceous. inflorescences single to multi-flowered but if few-flowered and axillary then with sepals to 6 mm long. monocarps ellipsoidal with blunt to beaked apex .............. 7 7. young flowers with conical ‘bud’ stage; sepals more than 6 mm wide ............................................. 8 – young flowers not with a conical ‘bud’ stage; sepals less than 6 mm wide ...................................... 11 8. leaves with generally more than 20 pairs of lateral nerves. flower pedicels more than 2.5 cm long .... ................................................................................................................................... m. macranthum – leaves with up to 20 pairs of lateral nerves. flower pedicels to 1 cm long ...................................... 9 9. twigs densely tomentose. leaves with dense hairs on nerves below. flower pedicel to 12 mm long ... ............................................................................................................................................ m. fuscum – twigs glabrescent. leaves glabrous to glabrescent beneath. flower pedicel generally more than 12 mm long ...................................................................................................................................... 10 10. leaves to 17 cm long, 7 cm wide. sepals much broader than long. monocarps woolly hairy, apex rounded ............................................................................................................................ m. cupulare – leaves generally more than 17 cm long, 7 cm wide. sepals about as long as broad. monocarps not woolly hairy, apex beaked .................................................................................... m. magnoliiflorum 11. petals distinctly longitudinally nerved. monocarps subsessile, pale tomentose ....... m. pachyphyllum – petals not longitudinally nerved. monocarps stipitate, glabrous to tomentose ............................... 12 12 (leaves more than 24 cm long, 7 cm wide). flowers borne distichously on sparsely branched hairy axes to 2 cm long, 4 mm thick. petals thick and fleshy. monocarps with stipe to 14 mm long, shorter than seed-bearing portion ........................................................................................... m. malayanum – (leaves generally smaller). flowers borne in fascicles or on very short and relatively slender axes. petals relatively thin. monocarps with stipe generally exceeding length of seed-bearing portion ... 13 13. flowering pedicel c. 0.5 mm thick; sepals to 2 mm long; petals to 3 mm wide .............. m. asteriellum – flowering pedicel at least 1 mm thick; sepals more than 2 mm long; petals 4 mm wide or more ... 14 14. petals 4–5 mm wide. monocarps densely short brown tomentose, stipe attachment eccentric. .............. ............................................................................................................................... m. xanthopetalum – petals 8 mm or more wide. monocarps more or less glabrous, stipe attachment not eccentric ........ 15 15. outer petals obovate (widest above midway), claw and blade distinct ........................................... 16 turner i.m. & utteridge t.m.a., whither polyalthia (annonaceae) in peninsular malaysia? 7 – outer petals lanceolate to oblong-linear (widest at or below midway), little distinction between claw and blade ......................................................................................................................................... 17 16. leaves obovate, reticulations obscure from above ..................................................... m. hookerianum – leaves elliptic, reticulations distinct from above ............................................................ m. glabrum 17. leaves generally less than 20 cm long and 5 cm wide; lateral nerves 9–13 pairs. sepals 3–4 mm long .. ................................................................................................................................. m. sclerophyllum – leaves generally more than 20 cm long and 5 cm wide; lateral nerves 14–15 pairs. sepals to 3 mm long............................................................................................................................. m. lateriflorum monoon anomalum (becc.) b.xue & r.m.k.saunders taxon 61: 1030 (xue et al. 2012). – polyalthia anomala becc., nuovo giornale botanico italiano 3: 188 (beccari 1871). – type: borneo, sarawak, mt mattan, may 1866, o. beccari p.b. 1605 (holo-: fi-b (erb. becc. no. 481)). polyalthia clavigera king, journal of the asiatic society of bengal. part ii (natural history &c.) 61: 60 (king 1892). – lectotype (designated by turner 2010a): peninsular malaysia, perak, waterfall hill, jun. 1888, l. wray 2075 (hololecto-: k (k000691553); isolecto-: cal, sing). polyalthia glomerata king, journal of the asiatic society of bengal. part ii (natural history &c.) 61: 61 (king 1892). – lectotype (designated by sinclair 1955: 318): peninsular malaysia, perak, jan. 1883, king’s collector [h.h. kunstler] 3817 (hololecto-: k (k000691552); isolecto-: bm, l, p). polyalthia sumatrana var. macrocarpa kochummen & whitmore in kochummen et al., federation museums journal 13: 134 (kochummen et al. 1970). – lectotype (designated here): peninsular malaysia, johore, labis, compartment 81, 15 apr 1967, t. suppiah kep 104976 (hololecto-: kep (fruiting material, explicitly excluding the foliage material); isolecto-: k). cleistopetalum sumatranum h.okada, acta phytotaxonomica et geobotanica 47: 5, fig. 4 (okada 1996). – type: indonesia, sumatra, west sumatra, gunung gadut, bukit pinang pinang, c. 20 km east of padang, 14 dec. 1987, h. okada 4607 (holo-: bo; iso-: kyo). description and distribution sinclair (1955: 318–319), turner (2014: 110). remarks note that polyalthia sumatrana var. macrocarpa was described from a mixed collection. fruiting material of monoon anomalum was confused with foliage of maasia sumatrana. as the diagnosis focused on the characters of the fruits, we here designate the fruiting material as lectotype and exclude the foliage material. monoon asteriellum (ridl.) b.xue & r.m.k.saunders taxon 61: 1030 (xue et al. 2012). – polyalthia asteriella ridl., journal of the straits branch of the royal asiatic society 82: 169 (ridley 1920). lectotype (designated here): peninsular malaysia, perak, larut hills, 1892, h.n. ridley 2986 (hololecto-: k (k000691443); isolecto-: cal, sing (sing 0048679)). description and distribution sinclair (1955: 309–310). european journal of taxonomy 183: 1–26 (2016) 8 monoon borneense (h. okada) b.xue & r.m.k.saunders taxon 61: 1030 (xue et al. 2012). – cleistopetalum borneense h.okada, acta phytotaxonomica et geobotanica 47: 4, fig. 3 (okada 1996). – polyalthia sinclairiana i.m.turner, gardens’ bulletin, singapore 58: 275 (turner 2007). – type: borneo, east kalimantan, sungai menubar, 12 dec. 1980, kato & wiriadinata 7119 (holo-: kyo; iso-: bo, l). polyalthia macropoda king, journal of the asiatic society of bengal. part ii (natural history &c.) 61: 60 (king 1892), non p. macropoda (miq.) f.muell. (von mueller 1877). – lectotype (designated by turner 2007): peninsular malaysia, perak, jun. 1883, king’s collector [h.h. kunstler] 4279 (hololecto-: k (k000691557); isolecto-: bm, l, p). description and distribution sinclair (1955: 316–317). monoon congregatum (king) b.xue & r.m.k.saunders taxon 61: 1031 (xue et al. 2012). – polyalthia congregata king, journal of the asiatic society of bengal. part ii (natural history &c.) 61: 61 (king 1892). – enicosanthum congregatum (king) airy shaw, bulletin of miscellaneous information, kew 1939: 277 (airy-shaw 1939). – lectotype (designated by xue et al. 2012: 1031): peninsular malaysia, perak, gopeng, aug. 1883, king’s collector [h.h. kunstler] 4831 (hololecto-: k (k000691419); isolecto-: cal, dd, sing (sing0059443)). description and distribution sinclair (1955: 194). monoon cupulare (king) b.xue & r.m.k.saunders taxon 61: 1031 (xue et al. 2012). – griffithia cupularis king, annals of the royal botanic garden (calcutta) 4: 9, t. 219 (king 1893). – griffithianthus cupularis (maingay ex hook.f. & thomson) merr., philippine journal of science. section c, botany 10: 231 (merrill 1915). – enicosanthum cupulare (king) airy shaw, bulletin of miscellaneous information, kew 1939: 277 (airy-shaw 1939). – lectotype (designated by xue et al. 2012: 1031): peninsular malaysia, perak, larut, may 1885, king’s collector [h.h. kunstler] 7630 (hololecto-: k (k000691422); isolecto-: bm (bm001050387), cal). description and distribution sinclair (1955: 188–189). monoon fuscum (king) b.xue & r.m.k.saunders taxon 61: 1031 (xue et al. 2012). – griffithia fusca king, annals of the royal botanic garden (calcutta) 4: 10, t. 220 (king 1893). – griffithianthus fuscus (maingay ex hook.f. & thomson) merr., philippine journal of science. section c, botany 10: 231 (merrill 1915). enicosanthum fuscum (king) airy shaw, bulletin of miscellaneous information, kew 1939: 277 (airy-shaw 1939). – lectotype (designated by xue et al. 2012: 1031): peninsular malaysia, perak, ulu bubong, jul. 1886, king’s collector [h.h. kunstler] 10404 (hololecto-: k (k000691418); isolecto-: bm (bm001100029), cal). description and distribution sinclair (1955: 189–190). turner i.m. & utteridge t.m.a., whither polyalthia (annonaceae) in peninsular malaysia? 9 monoon glabrum (hook.f. & thomson) b.xue & r.m.k.saunders taxon 61: 1031 (xue et al. 2012). – ellipeia glabra hook.f. & thomson, the flora of british india 1: 52 (hooker & thomson 1872). – polyalthia glabra (hook.f. & thomson) j.sinclair, gardens’ bulletin, singapore 14: 315 (sinclair 1955). – type: peninsular malaysia, malacca, 14 dec. 1867, a.c. maingay 3007 [kew distribution no. 66] (holo-: k [×2] (k000691571, k000691570); iso-: cal). polyalthia curtisii ridl., journal of the straits branch of the royal asiatic society 54: 11 (ridley 1910). – lectotype (designated here): peninsular malaysia, penang, telok bahang, jun. 1901, c. curtis 3644 (hololecto-: k (k000691572); isolecto-: sing). description and distribution sinclair (1955: 315). monoon hookerianum (king) b.xue & r.m.k.saunders taxon 61: 1032 (xue et al. 2012). – polyalthia hookeriana king, journal of the asiatic society of bengal. part ii (natural history &c.) 61: 57 (king 1892). – lectotype (designated by turner 2011: 72): peninsular malaysia, perak, feb. 1884, king’s collector [h.h. kunstler] 5550 (hololecto-: k (k000691563); isolecto-: cal, p (p00601064)). description and distribution sinclair (1955: 312–315), turner (2014: 113–115). monoon hypogaeum (king) b.xue & r.m.k.saunders taxon 61: 1032 (xue et al. 2012). – polyalthia hypogaea king, journal of the asiatic society of bengal. part ii (natural history &c.) 61: 62 (king 1892). – lectotype (designated by turner 2011: 72): peninsular malaysia, perak, larut, 1881, king’s collector [h.h. kunstler] 2437 (hololecto-: k (k000691555); isolecto-: cal). description and distribution sinclair (1955: 315–316), turner (2014: 115–116). monoon lateriflorum (blume) miq. annales musei botanici lugdunu-batavi 2: 19 (miquel 1865). – guatteria lateriflora blume, bijdragen tot de flora van nederlandsch indië: 20 (blume 1825). – lectotype (designated by turner 2011: 74): indonesia, java: s.dat., anon. s.n. (l (l 0188880)). description and distribution sinclair (1955: 310–311), kochummen (1972: 88–89), turner (2014: 116). monoon macranthum (king) b.xue & r.m.k.saunders taxon 61: 1032 (xue et al. 2012). – polyalthia macrantha king, journal of the asiatic society of bengal. part ii (natural history &c.) 61: 54 (king 1892). – enicosanthum macranthum (king) j.sinclair, gardens’ bulletin, singapore 14: 190 (sinclair 1955). – lectotype (designated by xue et al. 2012: 1032): peninsular malaysia, perak, king’s collector [h.h. kunstler] 6654 (hololecto-: l (l0037926); isolecto-: cal, dd). european journal of taxonomy 183: 1–26 (2016) 10 description and distribution sinclair (1955: 190–191). monoon magnoliiflorum (maingay ex hook.f. & thomson) b.xue & r.m.k.saunders taxon 61: 1032 (xue et al. 2012). – polyalthia magnoliiflora maingay ex hook.f. & thomson, the flora of british india 1: 64 (hooker & thomson 1872). – griffithia magnoliipetala king, annals of the royal botanic garden (calcutta) 4: 9, t. 218 (king 1893), as ‘magnoliaepetala’, nom. superfl. – griffithianthus magnoliiflorus (maingay ex hook.f. & thomson) merr., philippine journal of science. section c, botany 10: 231 (merrill 1915). – enicosanthum magnoliiflorum (maingay ex hook.f. & thomson) airy shaw, bulletin of miscellaneous information, kew 1939: 277 (airy-shaw 1939). – lectotype (designated by xue et al. 2012: 1032): malacca, 6 feb. 1868, a.c. maingay 3259 [kew distrib. no. 93] (k (k000691423)). description and distribution sinclair (1955: 187–188). monoon malayanum i.m.turner & utteridge webbia 70: 99 (turner & utteridge 2015). – type: peninsular malaysia, trengganu, ulu trengganu, ulu telemong forest reserve, batu kota, 16 sep. 1969. h.s. loh fri 13446 (holo-: kep; iso-: k [×2]). description and distribution turner & utteridge (2015: 99). monoon membranifolium (j.sinclair) b.xue & r.m.k.saunders taxon 61: 1032 (xue et al. 2012). – enicosanthum membranifolium j.sinclair, gardens’ bulletin, singapore 14: 191 (sinclair 1955). – lectotype (designated by xue et al. 2012: 1032): peninsular malaysia, kelantan, gua panjang at gua ninik, 21 oct. 1927, m.r. henderson sfn 19522 (hololecto-: sing (sing0048664); isolecto-: k (k000691413), sing (sing0048665)). description and distribution sinclair (1955: 191). monoon pachyphyllum (king) b.xue & r.m.k.saunders taxon 61: 1033 (xue et al. 2012). – polyalthia pachyphylla king, journal of the asiatic society of bengal. part ii (natural history &c.) 61: 66 (king 1892). – lectotype (designated by xue et al. 2012: 1033): peninsular malaysia, perak, king’s collector [h.h. kunstler] 6655 (hololecto-: k (barcode no. k000691559); isolecto-: bm (bm000754027), cal, dd, sing). description and distribution sinclair (1955: 308). monoon praestigiosum (j.sinclair) b.xue & r.m.k.saunders taxon 61: 1033 (xue et al. 2012). – enicosanthum praestigiosum j.sinclair, gardens’ bulletin, singapore 14: 192 (sinclair 1955). – lectotype (designated by xue et al. 2012: 1032): peninsular malaysia, turner i.m. & utteridge t.m.a., whither polyalthia (annonaceae) in peninsular malaysia? 11 johore, sungei kayu, 10 oct. 1936, kiah sfn 32017 (hololecto-: sing (sing00048666); isolecto-: a (a00039311), bo, bkf, k [×2] (k000691412, k000898010), kep (kep 8626), l, lae, pnh). description and distribution sinclair (1955: 192–194). monoon sclerophyllum (hook.f. & thomson) b.xue & r.m.k.saunders taxon 61: 1033 (xue et al. 2012). – polyalthia sclerophylla hook.f. & thomson, the flora of british india 1: 65 (hooker & thomson 1872). – lectotype (designated here): peninsular malaysia, malacca, 9 may 1867, a.c. maingay 1340 [kew distrib. no. 101] (hololecto-: k [×2] (k000691575, k000691576)). polyalthia purpurea ridl., journal of the straits branch of the royal asiatic society 82: 168 (ridley 1920). – lectotype (designated by turner 2012: 242): singapore, cultivated in singapore botanic gardens, jun. 1904, h.n. ridley 12014 (hololecto-: k (k000691573); isolecto-: sing). description and distribution sinclair (1955: 311–312). monoon xanthopetalum (merr.) b.xue & r.m.k.saunders taxon 61: 1034 (xue et al. 2012). – polyalthia xanthopetala merr., journal of the straits branch of the royal asiatic society 85: 185 (merrill 1922). – lectotype (designated by turner 2011: 80): borneo, sabah, batu lima near sandakan, sep.–dec. 1920, m. ramos 1705 (hololecto-: k (k0006916180); isolecto-: a (a00242651), bm (bm000754105), bo, l (l0189515), us (us00098685)). remarks in studying material of monoon from peninsular malaysia a group of specimens was found that did not match any of the species recorded from this area. the specimens were notable for the persistent goldenbrown tomentum on the twigs, the often dark brown shade of the dry leaves, petals linear-lanceolate c. 4 cm long and c. 5 mm wide, monocarps ellipsoidal, covered with a short dense pale brown tomentum with the stipe attached slightly eccentrically giving a similar appearance to those of some uvaria species previously included in the genus ellipeia. one of us (imt) having worked on the borneo species (turner 2014), the specimens were reminiscent of monoon xanthopetalum merr. comparison of the specimens from peninsular malaysia with collections from borneo (cf. turner 2014: fig. 26) produced a close match. we have no doubt that monoon xanthopetalum does occur in peninsular malaysia, representing a major range extension for the species. it is interesting to note that the peninsular malaysia collections of m. xanthopetalum are all from the east coast which fits with the presence of the species in borneo. specimens examined peninsular malaysia: johore, mersing, hutan simpan lenggor, 2 nov. 2010, l.e. teo & din kl 5758, (kep). pahang, rompin, gunong lesong forest rerve, 2 may 1956, forester lindong kep 83466 (k); lesong forest reserve, 27 jun. 1972, y.c. chan fri 19827 (k, kep). trengganu, 26 oct. 1994, c. wiart & l.e. teo kl 4434 (kep); kemaman, kajang, ulu bendong, 6 nov. 1935, e.j.h. corner sfn 30291 (k). polyalthia blume flora javae (anonaceae): 68 (blume 1830). – lectotype (designated by hutchinson 1923: 259): polyalthia subcordata (blume) blume. european journal of taxonomy 183: 1–26 (2016) 12 sphaerothalamus hook.f., transactions of the linnean society of london 23: 156 (hooker 1860). – type: sphaerothalamus insignis hook.f. key to the species of polyalthia 1. inflorescences cauliflorous, ramiflorous or axillary ........................................................................... 2 – inflorescences subopposite leaves ..................................................................................................... 8 2. flower pedicel 2–5(–7) mm long; petals narrowly linear (generally more than 10 times long as broad). monocarps subsessile (stipes to 5 mm long) ......................................................................... 3 – flower pedicel (6–)10–30 mm long; petals not narrowly linear (rarely to 10 times as long as broad). monocarps stipitate (stipes 10 mm long or more)............................................................................... 6 3. subshrub to 30 cm tall. petals not exceeding 3 cm long ....................................................... p. pumila – tree to 6 m or more tall. petals to 5 cm or more long ........................................................................ 4 4. petals 5 mm wide or more. monocarps densely brown woolly hairy ........................... p. cinnamomea – petals 2–5 mm wide. monocarps pubescent, becoming glabrous with maturity ............................... 5 5. inflorescences mostly on trunk and branches. twigs persistently brown hairy. sepals 15–17 mm long. monocarps 2 cm wide ........................................................................................... p. stenopetala – inflorescences mostly among leaves. twigs becoming glabous with age. sepals 7 mm long. monocarps less than 1 cm wide ................................................................................... p. angustissima 6. inflorescences mostly cauliflorous, or if among leaves then pedicel generally 20 mm long or more ..... .......................................................................................................................................... p. cauliflora – inforescences not cauliflorous, flower pedicel to 12 mm long .......................................................... 7 7. leaves drying grey. flower pedicel more than 1 mm wide, widening distally; petals 2–5 cm long. monocarps more or less cylindrical, 6–7 mm diameter; stipe c. 1 mm diameter ................. p. lateritia – leaves drying brown or grey-brown. flower pedicel c. 0.5 mm wide; petals less than 1 cm long. monocarps more or less globose, c. 15 mm diameter; stipe c. 2 mm diameter..................... p. obliqua 8. leaves with many lateral nerves (at least 18 pairs); lamina bullate .................................................. 9 – leaves with up to 17 pairs of lateral nerves; lamina not bullate ...................................................... 10 9. twigs villose. base of leaf auriculate or cordate, often extending below twig ..................... p. bullata – twigs glabrous or hairy, not villose. base of leaf not auriculate or cordate, never extending below twig ............................................................................... p. pakdin i.m.turner & utteridge sp. nov. 10. leaves narrowly elliptic or lanceolate to 2.5 cm wide, apex long acuminate ..................... p. dumosa – leaves generally more than 2.5 cm wide, apex not long acuminate ............................................... 11 11. leaf apex obtuse. branchlets with thick corky bark. flower pedicel slender (5–25 mm long, < 1 mm thick) ................................................................................................................................. p. suberosa – leaf apex generally acute to acuminate. branchlets not corky-barked. flower pedicels generally shorter and/or thicker ....................................................................................................................... 12 12. flowers subsessile (pedicel to 2 mm long) ..................................................................................... 13 – flowers distinctly pedicellate (pedicel 5 mm long or more) ........................................................... 14 13. leaves glabrous; elliptic-lanceolate; apex blunt. petals coriaceous, fleshy ...................... p. parviflora turner i.m. & utteridge t.m.a., whither polyalthia (annonaceae) in peninsular malaysia? 13 – leaves pubescent beneath; lanceolate; acuminate. petals thin .......................................... p. hirtifolia 14. twigs, leaf undersides, flowers and fruits furry .......................................................... p. chrysotricha – twigs, leaf undersides, flowers and fruits glabrous or hairy but not all furry .................................. 15 15. leaves generally with 10 or fewer pairs of lateral nerves. flower pedicel to 5 mm long. monocarps to 6, stipe shorter than seed-bearing portion ................................................................ p. brunneifolia – leaves with 12 or more pairs of lateral nerves. flower pedicel 5 mm or more long. monocarps 10 or more, stipe generally longer than seed-bearing portion .................................................................. 16 16. petals relatively thin; sparsely hairy outside. stipe of monocarps c. 0.5 mm thick ................. p. socia – petals relatively thick and leathery; densely hairy outside. stipe of monocarps c. 1 mm thick ...... 17 17. pedicel 3–6 cm long .......................................................................................................... p. oblonga – pedicel to 1 cm long ....................................................................................................... p. motleyana polyalthia angustissima ridl. journal of the straits branch of the royal asiatic society 54: 11 (ridley 1910). – lectotype (designated by bunchalee & chantaranothai 2006: 3): singapore: bukit timah, feb. 1896, h.n. ridley 8050 (hololecto-: sing (sing0048677); isolecto-: k). description and distribution sinclair (1955: 286). polyalthia brunneifolia j.sinclair gardens’ bulletin, singapore 14: 301 (sinclair 1955). – lectotype (designated here): peninsular malaysia, selangor, bukit enggang, kajang, 8 apr. 1930, c.f. symington 24179 (hololecto-: k (k000691554); isolecto-: sing (sing0048680). description and distribution sinclair (1955: 301–302). polyalthia bullata king journal of the asiatic society of bengal. part ii (natural history &c.) 61: 64 (king 1892). – lectotype (designated by turner 2009: 92): peninsular malaysia, perak, gopeng, kinta, aug. 1883, king’s collector [h.h. kunstler] 4804 (hololecto-: cal; isolecto-: k (k000691434)). description and distribution sinclair (1955: 300–301), turner (2014: 145). polyalthia cauliflora hook.f. & thomson flora indica: 138 (hooker & thomson 1855). – type: singapore, oct. 1822, n. wallich s.n. [eic 6476] (holo-: k-w (k001123992)). guatteria teysmannii miq., flora van nederlandsch indië. supplementum primum. prodromus florae sumatranae: 378 (miquel 1861). – monoon teysmannii (miq.) miq., annales musei botanici lugdunubatavi 2: 19 (miquel 1865). – polyalthia teysmannii (miq.) king, journal of the asiatic society of bengal. part ii (natural history &c.) 61: 67 (king 1892). – lectotype (designated by turner 2011: european journal of taxonomy 183: 1–26 (2016) 14 69): indonesia, sumatra, palembang, batu radja, s. dat., j.e. teijsmann s.n. [herb. bogor. 3901] (u (u 0000373)). unona desmantha hook.f. & thomson, the flora of british india 1: 61 (hooker & thomson 1872). – desmos desmanthus (hook.f. & thomson) saff., bulletin of the torrey botanical club 39: 508 (safford 1912). – polyalthia cauliflora var. desmantha (hook.f. & thomson) j.sinclair, gardens’ bulletin, singapore 14: 295 (sinclair 1955). – lectotype (designated by turner 2010b: 268): malaysia, peninsular malaysia, malacca, 1865-1866, a.c. maingay 1045 [kew distribution no. 48] (k (k000691431)). polyalthia beccarii king, journal of the asiatic society of bengal. part ii (natural history &c.) 61: 65 (king 1892). – polyalthia cauliflora hook.f. & thomson var. beccarii (king) j.sinclair, gardens’ bulletin, singapore 14: 294 (sinclair 1955). – lectotype (designated by turner 2009: 93): malaysia, peninsular malaysia, perak, gopeng, kinta, jul. 1883, king’s collector [h.h. kunstler] 4522 (hololecto-: k (k00069131); isolecto-: cal, dd, u). unona pycnantha hook.f. & thomson, the flora of british india 1: 60 (hooker & thomson 1872). – polyalthia pycnantha (hook.f. & thomson) king, journal of the asiatic society of bengal. part ii (natural history &c.) 61: 67 (king 1892). – lectotype (designated by turner 2010b: 268): malaysia, peninsular malaysia: malacca, 9 jan. 1865 or 1866, a.c. maingay 1491 [kew distribution no. 48] (k (k000691459)). unona wrayi hemsl., hooker’s icones plantarum 16: t. 1553 (hemsley 1887). – polyalthia wrayi (hemsl.) ridl., the flora of the malay peninsula 1: 49 (ridley 1922). – polyalthia cauliflora hook.f. & thomson var. wrayi (hemsl.) j.sinclair, gardens’ bulletin, singapore 14: 296 (sinclair 1955). – type: peninsular malaysia, perak, ulu kenring, jan. 1884, l. wray 560 (holo-: k (k000691466)). description and distribution sinclair (1955: 292–297), turner (2014: 145–146). polyalthia chrysotricha ridl. the flora of the malay peninsula 1: 57 (ridley 1922). – type: peninsular malaysia, selangor, ulu langat, menuang gasing, feb. 1912, c.b. kloss s.n. (holo-: k (k000691577)). description and distribution sinclair (1955: 302–303), turner (2014: 147). polyalthia cinnamomea hook.f. & thomson flora indica: 138 (hooker & thomson 1855). – lectotype (designated by airy-shaw 1939: 282): singapore, anon. [?n. wallich] s.n. [eic 6444] (hololecto-: k-w (k001123933)). unona cauliflora hook.f. & thomson, flora indica: 137 (hooker & thomson 1855), non u. cauliflora blanco (blanco 1845). – type: singapore, t. lobb s.n. (holo-: k (k000691457) [excl. material in packet from eic = k000691458]). polyalthia velutinosa ridl., journal of the straits branch of the royal asiatic society 59: 64 (ridley 1911). – lectotype (designated here): peninsular malaysia, lankawi, gunong raya at 1500 feet, feb. 1911, (aniff) haniff 15552 (hololecto-: sing (sing 0058808); isolecto-: bm, k (k000691439), bm). description and distribution sinclair (1955: 286–288), kochummen (1972: 87–88), turner (2014: 148). turner i.m. & utteridge t.m.a., whither polyalthia (annonaceae) in peninsular malaysia? 15 polyalthia dumosa king journal of the asiatic society of bengal. part ii (natural history &c.) 61: 52 (king 1892). – lectotype (designated by bunchalee & chantaranothai 2006: 3): peninsular malaysia, perak, maxwell’s hill, l. wray 2628 (hololecto-: sing; isolecto-: cal). description and distribution sinclair (1955: 297). polyalthia hirtifolia j.sinclair gardens’ bulletin, singapore 14: 300 (sinclair 1955). – polyalthia hirta ridl., journal of the straits branch of the royal asiatic society 82: 168 (ridley 1920), non p. hirta (miq.) diels (diels 1912). – lectotype (designated by turner 2012: 241): peninsular malaysia, penang, pulau butong reserve, mar. 1892, c. curtis 2745 (hololecto-: k (k000691452); isolecto-: bm, k (barcode no. k000691453), sing). description and distribution sinclair (1955: 300). polyalthia lateritia j.sinclair gardens’ bulletin, singapore 14: 290 (sinclair 1955). – type: peninsular malaysia, perak, waterloo new road, may 1890, c. curtis 2704 (holo-: sing (sing0048683); iso-: k (k000691549), sing (sing0051296)). description and distribution sinclair (1955: 290–291), polyalthia motleyana (hook.f.) airy shaw bulletin of miscellaneous information, kew 1939: 280 (airy-shaw 1939). – oxymitra motleyana hook.f., transactions of the linnean society of london 23: 155 (hooker 1860), as ‘molleyana’. – lectotype (designated by turner 2011: 76): borneo, labuan, e.s. barber 228 (hololecto-: k (k000691612); isolecto-: k (k000691613)). polyalthia sarawakensis diels, notizblatt des botanischen gartens und museums zu berlin-dahlem 11: 80 (diels 1931). – type: borneo, sarawak, mt mattan, apr. 1866, o. beccari p.b. 1623 (holo-: fi-b (erbario collesioni beccari no. 415)). polyalthia motleyana var. glabrescens airy shaw, bulletin of miscellaneous information, kew 1939: 282 (airy-shaw 1939). – type: borneo, sarawak, o. beccari p.b. 1813 (holo-: k (k000691611); iso-: fi-b). description and distribution sinclair (1955: 303–305), turner (2014: 157–159). polyalthia obliqua hook.f. & thomson flora indica: 138 (hooker & thomson 1855). – lectotype (designated by turner 2011: 77): peninsular malaysia, malacca, s. dat., w. griffith s.n. (k (k000691435)). european journal of taxonomy 183: 1–26 (2016) 16 polyalthia similis merr., philippine journal of science. section c, botany 8: 371 (merrill 1913). – lectotype (designated by turner 2011: 77): philippines, leyte, dagami, aug. 1912, m. ramos bur. sci. 15185 (hololecto-: k (k000691620); isolecto-: bm, us (us00098679)). polyalthia lopadantha diels, notizblatt des botanischen gartens und museums zu berlin-dahlem 11: 81 (diels 1931). – type: borneo, sarawak, o. beccari p.b. 1819 (holo-: fi-b (erb. coll. becc. no. 419); iso-: a[×2], b[×2] (b 10 0186733, b 10 0186734 (fragment)), k (k000691615), m). description and distribution sinclair (1955: 290), turner (2014: 159–160). polyalthia oblonga king journal of the asiatic society of bengal. part ii (natural history &c.) 61: 65 (king 1892). – polyalthia motleyena var. oblonga (king) j.sinclair, gardens’ bulletin, singapore 14: 304 (sinclair 1955). – lectotype (designated by turner 2011: 77): peninsular malaysia, perak, larut, may 1885, king’s collector [h. h. kunstler] 7671 (hololecto-: k; isolecto-: cal, dd, sing). description and distribution sinclair (1955: 304), turner (2014: 160). polyalthia pakdin i.m.turner & utteridge sp. nov. figs 1–2 urn:lsid:ipni.org:names:77153637-1 diagnosis with its large, multi-nerved leaves the new species has generally been confused with polyaltha bullata king, but it lacks the villose tomentum on the twigs and the strongly auriculate to cordate leaf bases. in comparison with p. bullata, the flowers are borne on shorter pedicels (< 1 versus 2 cm) with shorter sepals (3–4 versus 7 mm) and petals (to 1 cm versus up to 4 cm); and with monocarps on shorter stipes (to 6 mm versus up to 10 mm). etymology we are pleased to commemorate the late prof. kamarudin mat salleh (1959–2009). a leading malaysian student of the annonaceae, he was widely known as ‘pak din’ (uncle din in malay, din being the common abbreviation of names with that ending). the epithet represents a compounding of this soubriquet. kamarudin was native to the state of kelantan, which is part of the range of the new species. type peninsular malaysia: trengganu, jambu bongkok forest reserve [4°50’n 103°22’e], 21 mar. 1972, l.e. teo & g. pachiappan kl 3033 [t & p 433] (holo-: k). additional specimens examined peninsular malaysia: kelantan, chabang tongkat forest reserve [5°49’n 102°17’e], compt 77, alt. 250’ 28 apr. 1967, k.m. kochummen fri 2317 (kep). trengganu, dunggun, jambu bongkok [4°50’n 103°21’e], 12 mar. 1961, j. carrick s.n. (k ×2); ibid., sekayu, bukit lanjut forest reserve [4°58’n 102°58’e], 18 sept. 1969, h.s. loh fri 13467 (kep); ibid., dungun, near 36th mile, jerangau road [4°56’36”n 103°10’42”e], 22 sept. 1955, j. sinclair & kiah bin salleh sfn 40941 (k, kep); ibid., sekayu forest reserve [4°58’n 102°57’e], compt 43, 22 sept. 1969, t. suppiah fri 11853 (kep); http://www.ipni.org/urn:lsid:ipni.org:names:77153637-1 turner i.m. & utteridge t.m.a., whither polyalthia (annonaceae) in peninsular malaysia? 17 fig. 1. polyalthia pakdin i.m.turner & utteridge sp. nov. a. leafy twig with flowers. b. inflorescence and base of leaf (abaxial view). c. flower. d. magnified view of abaxial surface of outer petal. e. calyx from below. f. stamen. g. carpel. h. infructescense with immature monocarp. i. mature monocarp. j. seed. scale bars: single bar = 1 mm, graduated single bar = 5 mm; double bar = 1 cm, graduated double bar = 5 cm. drawn from carrick s.n. (a, c–e), kl 3033 (b, f, g), sfn 40941 (h), fri 11853 (i), fri 13467 (j). european journal of taxonomy 183: 1–26 (2016) 18 ibid., bukit bauk forest reserve, compartment 8a [4°45’n 103°21’e], 17 june 1967, t.c. whitmore fri 3903 (kep). description shrub 1–4 m tall. young twigs drying red-brown, rather smooth and shiny, with some irregular longitudinal ridges with rounded tops, varying from glabrous to densely pale short hairy, twigs becoming black to brown with age. leaves chartaceous to subcoriaceous (tend to look more leathery than they are to the touch), often bullate, drying grey or grey-brown, more rarely dark brown, above, brown or greybrown below with brown venation, midrib and laterals sunken above in dry leaves, prominent beneath, more or less glabrous though sometimes with pale hairs along midrib below near base, lamina oblongoblanceolate to narrowly oblong-elliptic, 17–36 × 3.5–9 cm, apex acuminate, base obtuse, generally slightly asymmetric, lateral nerves 19–24 pairs, arching and looping within margin to form a scalloped intramarginal nerve, reticulations distinct. petiole drying dark brown to black, 6–10 mm long, 2–4 mm in diameter, sometimes with short erect pale hairs. inflorescences subopposite leaves, singleor fewflowered. flowering pedicel 2–8 mm long, 1 mm diameter widening distally, drying brown with minute pale hairs [some 4-merous flowers present on kl 3033], sepals ovate c. 3–4 × 2–3 mm, outside brown, minutely bumpy with short pale hairs, inside glabrous, petals thin-textured, ovate to ovate-lanceolate 6–10 × 2.5–3 mm, drying dark brown to blackish, outside minutely bumpy with adpressed pale hairs, inside glabrous, stamens many c. 1.5 mm long, carpels many. fruiting pedicel to 2 mm thick, sepals often persisting, monocarps to 9 or more, irregularly ellipsoidal, 8–10 × 7–9 mm, drying red-brown, surface minutely pimply with pale hairs, apiculate, stipe to 6 mm long, 1 mm diameter, pimply, pale hairy. seeds generally two, lenticular, c. 7 × 6 × 4 mm, drying golden brown with a longitudinal groove running parallel to the flat face. field notes leaves bullate (sfn 40941), dark glossy green above, paler green beneath (sfn 40941), shiny dark green above, slightly yellowish green below (kl 3033); flowers yellowish (kl 3033), petals reddish orange (sfn 40941), petals and sepals orange (fri 2317), stamens paler reddish orange (sfn 40941), no scent with flowers (sfn 40941); fruits green (fri 3903), fruits green turning red (fri 11853), fruits red (fri 13467, sfn 40941); seeds brown (sfn 40941). distribution and habitat recorded from the eastern side of peninsular malaysia from kelantan and, with most records, trengganu (fig. 2). it occurs in lowland tropical rain forest. conservation status polyalthia pakdin sp. nov. is restricted to the lowland forests of eastern peninsular malaysia and currently known from seven collections from five localities (two are from the sekayu area and two from jambu bongkok). the extent of occurrence (eoo) is 1,856 km2 falling within the endangered (en) threshold of less than 5,000 km2, and the area of occupancy (aoo; based on a user-defined cell width of 2 km) is 28 km2 falling within the endangered (en) threshold. whilst none of the collections is contemporary (the most recent is 1972), nearly all are from forests reserves, and, in addition, one collection is from ‘roadside forest’ (j. sinclair & kiah bin salleh sfn 40941). whilst the eoo is within the en threshold, the presence of the species within protected areas would not place the species within this category because recent satellite imagery shows the protected areas still with relatively large areas of intact forest (using the most recent google earth imagery). the sekayu collections, for example, are on the edge of the very poorly known and under-collected taman negara national park and more field collections are needed in such areas to better understand the distribution of this species. without any other information, we must assume that the species is still present in these protected areas and that the protected areas still turner i.m. & utteridge t.m.a., whither polyalthia (annonaceae) in peninsular malaysia? 19 harbour the appropriate habitat for p. pakdin sp. nov. it is difficult to demonstrate trends in the reduction of habitat etc. without modern collections, and for these reasons we assign a preliminary conservation assessment of near threatened (nt). affinities polyalthia bullata, the species with which p. pakdin sp. nov. was formerly confused, was included in the informal group of polyalthia species termed the polyalthia insignis group by johnson & murray (1999). they characterised the group as follows: shrubs or small trees; leaves oblique, subcordate or auriculate at base, often drying grey above, brown below; inflorescences internodal to leaf opposed or sometimes cauliflorous; petals orange or red; apex of ovary woody and persistant as a short beak on the monocarp; stigmas capitate and cohering to drop as a cap at the end of anthesis, several ovules per carpel, laterally attached in vertical row and seed not pitted but with encircling groove. while polyalthia pakdin sp. nov. does exhibit some of these characters, it does not have auriculate or cordate leaf bases and the stigmas do not seem to cohere. in fact, p. pakdin sp. nov. also comes close to the bornean polyalthia polyphlebia diels which was not included in the p. insignis group by johnson & murray. the leaf form of p. pakdin sp. nov. and p. polyphlebia are similar but p. polyphlebia generally has greyish corky twigs, longer pedicels (8–15 versus 2–8 mm) with the flowers larger (outer petals 13–14 mm versus 6–10 mm) and the petals fleshy rather than thin in texture; with the monocarps on longer stipes (to 25 mm versus 6 mm). polyalthia parviflora ridl. journal of the straits branch of the royal asiatic society 61: 49 (ridley 1912). – lectotype (designated by turner 2012: 241): peninsular malaysia, pulau langkawi, kuala malacca, sep. 1890, c. curtis 2533 (hololecto-: sing (sing 0059296); isolecto-: cal). polyalthia rufa craib, bulletin of miscellaneous information, kew 1924: 82 (craib 1924). – type: thailand, nakawn sawawn, mê wong, 25 may 1922, a.f.g. kerr 6022 (holo-: k[×2] (k00595497, k00595496); iso-: abd, bm (000553972(bm))). fig. 2. map showing the collecting localities of polyalthia pakdin i.m.turner & utteridge sp. nov. european journal of taxonomy 183: 1–26 (2016) 20 description and distribution sinclair (1955: 299). polyalthia pumila ridl. journal of the straits branch of the royal asiatic society 54: 12 (ridley 1910). – lectotype (designated by turner 2012: 241): peninsular malaysia, perak, dindings, sera woods, mar. 1896, h.n. ridley 7996 (hololecto-: sing (sing 0059298); isolecto-: k (k000691442)). description and distribution sinclair (1955: 289). polyalthia socia craib bulletin of miscellaneous information, kew 1925: 10 (craib 1925). – type: thailand, pattani, bukit, 400 m, evergreen forest, 7 jul. 1923, a.f.g. kerr 7101 (holo-: k (k000595494); iso-: abd[×2], bk (257670), bm (000553974(bm)), tcd (tcd0009808)). description and distribution sinclair (1955: 291–292). polyalthia stenopetala (hook.f. & thomson) finet & gagnep. mémoires de la société botanique de france 4: 96 (finet & gagnepain 1906). – unona stenopetala hook.f. & thomson, flora indica: 136 (hooker & thomson 1855). – desmos stenopetalus (hook.f. & thomson) saff., bulletin of the torrey botanical club 39: 507 (safford 1912). – lectotype (designated by sinclair 1955: 285): java, t. lobb 414 (hololecto-: k (k000691463); isolecto-: cge). unona crinita hook.f. & thomson, the flora of british india 1: 61 (hooker & thomson 1872). – desmos crinitus (hook.f. & thomson) saff., bulletin of the torrey botanical club 39: 507 (safford 1912). – polyalthia crinita (hook.f. & thomson) ridl., journal of the straits branch of the royal asiatic society 75: 6 (ridley 1917). – type: peninsular malaysia, malacca, 18 jan. 1865 or 1866, a. c. maingay 1513 [kew distribution no. 41] (holotype: k[×2] (k000691461, k000691462)). description and distribution sinclair (1955: 285), turner (2014: 162). polyalthia suberosa (roxb.) thwaites enumeratio plantarum zeylaniae 5: 398 (thwaites 1864). – uvaria suberosa roxb., plants of the coast of coromandel 1: 31, t. 34 (roxburgh 1795). – guatteria suberosa (roxb.) dunal, monographie de la famille des anonacées: 128 (dunal 1817). – lectotype (designated by huber 1985: 41): roxburgh’s plate t. 34. description and distribution sinclair (1955: 298). discussion the changes in the species recognised for peninsular malaysia between sinclair’s revision and the current compilation are summarised in table 1. three species have been added to the list in the turner i.m. & utteridge t.m.a., whither polyalthia (annonaceae) in peninsular malaysia? 21 sinclair 1955 current paper enicosanthum congregatum monoon congregatum enicosanthum cupulare monoon cupulare enicosanthum fuscum monoon fuscum enicosanthum macranthum monoon macranthum enicosanthum magnoliiflorum monoon magnoliiflorum enicosanthum membranifolium monoon membranifolium enicosanthum praestigiosum monoon praestigiosum polyalthia jenkinsii huberantha jenkinsii polyalthia rumphii huberantha rumphii polyalthia glauca maasia glauca polyalthia hypoleuca maasia hypoleuca polyalthia sumatrana maasia sumatrana polyalthia asteriella monoon asteriellum polyalthia clavigera monoon anomalum polyalthia glabra monoon glabrum polyalthia hookeriana monoon hookerianum polyalthia hypogaea monoon hypogaeum polyalthia lateriflora monoon lateriflorum polyalthia macropoda monoon borneense polyalthia pachyphylla monoon pachyphyllum polyalthia sclerophylla monoon sclerophyllum monoon malayanum monoon xanthopetalum polyalthia angustissima polyalthia angustissima polyalthia bullata polyalthia bullata polyalthia brunneifolia polyalthia brunneifolia polyalthia cauliflora polyalthia cauliflora polyalthia chrysotricha polyalthia chrysotricha polyalthia cinnamomea polyalthia cinnamomea polyalthia dumosa polyalthia dumosa polyalthia hirtifolia polyalthia hirtifolia polyalthia lateritia polyalthia lateritia polyalthia motleyana polyalthia motleyana polyalthia motleyana var. oblonga polyalthia oblonga polyalthia obliqua polyalthia obliqua polyalthia parviflora polyalthia parviflora polyalthia pumila polyalthia pumila polyalthia socia polyalthia socia polyalthia stenopetala polyalthia stenopetala polyalthia suberosa polyalthia suberosa polyalthia pakdin sp. nov. table 1. comparison of accepted names of species in polyalthia sensu lato native to peninsular malaysia between sinclair (1955) and the current paper. european journal of taxonomy 183: 1–26 (2016) 22 intervening 60 years which is perhaps fewer than might be expected and must reflect sinclair’s thorough study of the specimens available to him. there have been a large number of name changes, with fewer than half the species still having the same accepted name – all in polyalthia s.s. the reason for this is primarily to create a more phylogenetically accurate taxonomy. it is clear that polyalthia sensu lato was heterogeneous: uniting species with a roughly similar floral morphology. users of taxonomy, such as foresters and ecologists, generally dislike name changes so there may not be much enthusiasm from them after these changes. the splitting up of polyalthia will possibly have some practical advantages as large genera can be unwieldy and daunting to non-specialists. however, expanding the list of genera in a family that already has a large number to deal with may also discourage the users of taxonomy. fortunately in this case, one of the three ‘new’ genera – maasia – is easy to recognise because of the white or glaucous underside to the leaves giving a consistent gestallt to specimens that is soon learnt. monoon and polyalthia sensu stricto can be distinguished fairly readily, though some would doubtless have preferred the 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(cnidaria: myxosporea) parasitizing the gallbladder of elachistocleis cesarii from brazil, based on morphological and molecular analyses diego henrique mirandola dias vieira  1,*, letícia pereira úngari  2, edna paulino de alcantara  3, enzo emmerich 4, andré luiz quagliatto santos  5, lucia helena o´dwyer  6 & reinaldo josé da silva  7 1,2,3,4,6,7 são paulo state university (unesp), institute of biosciences, dbbvpz, section of parasitology, rua professor doutor antônio celso wagner zanin 250, botucatu, são paulo 18618-689, brazil. 5 laboratório de ensino e pesquisa em animais silvestres, faculdade de medicina veterinária, universidade federal de uberlândia, minas gerais, brazil. * corresponding author: diegovieira.50@gmail.com 2 email: letspungari@hotmail.com 3 email: ednnapaulinos@gmail.com 4 email: enzo_emmerich@hotmail.com 5 email: quagliatto.andre@gmail.com 6 email: lucia.odwyer@unesp.br 7 email: reinaldo.silva@unesp.br 1 urn:lsid:zoobank.org:author:89297795-fdd3-40e5-8658-d5a15eadd535 2 urn:lsid:zoobank.org:author:cb714126-46d6-4d5b-bca9-ae04ad47da04 3 urn:lsid:zoobank.org:author:688931cf-0846-4c21-93be-f9832b40fa6e 4 urn:lsid:zoobank.org:author:3c9f80e1-75a2-48a1-8576-5c287b4f1f26 5 urn:lsid:zoobank.org:author:659b987a-e16c-4947-8a18-408e99256add 6 urn:lsid:zoobank.org:author:501b3588-c93b-485d-acad-e0c4d9ed2ed2 7 urn:lsid:zoobank.org:author:419d6833-6175-48be-b371-65878984c032 abstract. numerous pseudoplasmodia containing myxospores belonging to the genus cystodiscus were found in the gallbladder of elachistocleis cesarii from mato grosso state, brazil. herein, we describe cystodiscus elachistocleis sp. nov., using morphological and small subunit ribosomal dna sequences. the mature myxospores were ellipsoid to ovoid, measuring in average 10.6 (9.8–11.2) μm in length and 6.2 (5.6–6.6) μm in width. polar capsules were pyriform and equal in size measuring in average 3.6 (2.8–4.3) μm in length and 2.6 (2.2–3.1) μm in width. polar filaments had 2–4 coils. the myxospores had 3–5 transverse ridges. the phylogenetic analysis showed cystodiscus elachistocleis sp. nov. as a sister species of cystodiscus cf. immersus 1, in a subclade formed by species that parasitize the amphibians gallbladder. this is the first species of cystodiscus described parasitizing a species of elachistocleis and the third species of cystodiscus described in brazil. keywords. elachistocleis, frog, phylogeny, microhylidae, myxidiidae. https://doi.org/10.5852/ejt.2021.775.1549 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index http://zoobank.org/urn:lsid:zoobank.org:pub:485c88f5-e2f2-4627-a865-0a534bb542d0 https://orcid.org/0000-0002-6137-1353 https://orcid.org/ 0000-0002-2982-9336 https://orcid.org/0000-0002-7814-531x https://orcid.org/0000-0002-2807-4592 https://orcid.org/0000-0002-9367-368x https://orcid.org/0000-0002-3426-6873 mailto:diegovieira.50@gmail.com mailto:letspungari%40hotmail.com?subject= mailto:ednnapaulino@gmail.com mailto:enzo_emmerich@hotmail.com mailto:quagliatto.andre%40gmail.com?subject= mailto:reinaldo.silva%40unesp.br?subject= mailto:reinaldo.silva@unesp.br http://zoobank.org/urn:lsid:zoobank.org:author:89297795-fdd3-40e5-8658-d5a15eadd535 http://zoobank.org/urn:lsid:zoobank.org:author:cb714126-46d6-4d5b-bca9-ae04ad47da04 http://zoobank.org/urn:lsid:zoobank.org:author:688931cf-0846-4c21-93be-f9832b40fa6e http://zoobank.org/urn:lsid:zoobank.org:author:3c9f80e1-75a2-48a1-8576-5c287b4f1f26 http://zoobank.org/urn:lsid:zoobank.org:author:659b987a-e16c-4947-8a18-408e99256add http://zoobank.org/urn:lsid:zoobank.org:author:501b3588-c93b-485d-acad-e0c4d9ed2ed2 http://zoobank.org/urn:lsid:zoobank.org:author:419d6833-6175-48be-b371-65878984c032 european journal of taxonomy 775: 107–118 (2021) 108 vieira d.h.m.d., pereira úngari l., de alcantara e.p., emmerich e., quagliatto santos a.l., o´dwyer l.h. & da silva r.j. 2021. description of cystodiscus elachistocleis sp. nov. (cnidaria: myxosporea) parasitizing the gallbladder of elachistocleis cesarii from brazil, based on morphological and molecular analyses. european journal of taxonomy 775: 107–118. https://doi.org/10.5852/ejt.2021.775.1549 introduction myxozoans are cnidarians parasites with a complex life cycle (lom & dyková 2006). lutz (1889) proposed the genus cystodiscus lutz, 1889 to accommodate cystodiscus immersus lutz, 1889 based on a large disc-like appearance found in the amphibious gallbladder. the genus was abandoned and classified as a synonym of myxidium bütschli, 1882; however, hartigan et al. (2012), performing phylogenetic molecular analyses, reerected the genus cystodiscus as a monophyletic group instead of the polyphyletic nature of myxidium. they have a global distribution and are endoparasites of a great diversity of amphibian hosts (hartigan et al. 2012a, 2012b, 2016). cystodiscus spp. could be pathogenic but some hosts do not show pathological signs and release myxospores over time without any impact on the health of their hosts (hartigan et al. 2012b). in pathogenic cases, the hosts may show inflammation in the nervous tissue leading to behavioral changes and even spontaneous death. also, in cases of liver diseases, they can cause inflammation and hyperplasia, affecting the metamorphosis of tadpoles, as well as metabolism and immune function (hartigan et al. 2012b). due to its pathogenic potential, the parasite can have an important ecological impact on the conservation of amphibians (hartigan et al. 2012c). microhylidae günther, 1858 is one of the most diverse families of extant amphibians, distributed across most of the tropics (frost 2020). three subfamilies of microhylids are recognized: adelastinae peloso, frost, richards, rodrigues, donnellan, matsui, raxworthy, biju, lemmon, lemmon & wheeler, 2016, gastrophryninae fitzinger, 1843, and otophryninae wassersug & pyburn, 1987 (de sá et al. 2012; peloso et al. 2014). elachistocleis parker, 1927 (subfamily gastrophryninae) is a genus of microhylid frogs widespread distributed with 19 species, most of which occur east of the andes in south america (frost 2020; sánchez-nivicela et al. 2020). in brazil, 12 species of elachistocleis were reported (segalla et al. 2019). elachistocleis cesarii (miranda-ribeiro, 1920), an endemic species in brazil, occurs from southeastern to central brazil in eastern são paulo, south-central minas gerais, goiás, and mato grosso states (toledo et al. 2010; frost 2020). this species has sexual dimorphism and females are larger than males. they have a fossorial habit, spending most of the year sheltered under the ground, and have a marked pattern of seasonal activity (toledo et al. 2010). this species is not on the international union for conservation of nature (iucn) red list of endangered animals. during a study on the biodiversity of anuran parasites in mato grosso state, brazil, pseudoplasmodia containing myxospores morphologically consistent with cystodiscus were observed in the gallbladder of e. cesarii. the present study aims to describe this new species of cystodiscus parasitizing e. cesarii from brazil, based on morphological and molecular analyses, thus increasing the knowledge of brazilian anurans parasites. material and methods anuran collection and morphological analysis in february 2020, one female anuran, e. cesarii measuring 39 mm snout-vent length and weight of 5.8 g, was captured for a major project of biodiversity of parasites in amphibians (fapesp 2018/096234; fapesp 2018/00754-9). a new sampling effort was carried out at different times within the major https://doi.org/10.5852/ejt.2021.775.1549 vieira d.h.m.d. et al., new species of cystodiscus from brazil 109 project, but it was not successful in collecting more specimens of e. cesarii. the specimen was collected on a dirt road in the municipality of araguaiana, mato grosso state, brazil (14°35′5.29″ s, 51°41′19.15″ w). the host was deposited in the herpetologica collection of the regional university of ceará (num. chufc a9761). the anuran was killed using 50 mg/kg of sodium thiopental (thiopentax®), a commercial anesthetic administered intracerebrally, following the guidelines of sebben (2007) and the animal use ethics committee (ibama license 60640-1; ceua-unesp 1061). the coelom was opened via a midventral longitudinal incision and the gall bladder was examined for cysts or signs of myxosporean infection. gallbladder contents were removed by puncturing the bladder wall and pipetting out the contents (hartigan et al. 2016). the fresh smears containing pseudoplasmodia were evaluated with a differential interference contrast microscope (leica dmlb 5000, leica microsystems, wetzlar, germany) at 1000 × magnification. the morphological measurements of myxospores freshly preserved followed the recommendations of lom & arthur (1989). measurements of 30 myxospores were obtained and presented as mean ± standard deviation. molecular analysis two pseudoplasmodia from the gallbladder were fixed in absolute ethanol and used for the molecular analyses. the access to the genetic data was authorized by the brazilian ministry of environment (sisgen aa4666fa). the two isolated pseudoplasmodia were collected from the bile material with finetipped sterile needles. dna isolation was carried out following the animal tissue protocol of the dneasy blood & tissue kit (qiagen, valencia, ca, usa). the partial ssu rdna gene was amplified with pcr using general eukaryotic and myxozoan primers (table 1). amplification was performed in a bio-rad mycycler (bio-rad laboratories pty ltd., gladesville, australia), with initial denaturation at 95°c for 3 min, followed by 35 cycles of 95°c for 1 min, 55°c for 45 s, 72°c for 2 min and a final extension at 72°c for 7 min. pcr reactions were performed in 25 μl of solution containing 3 μl of extracted dna and 1 μl of each pcr primer at 10 pmol, using pcr ready-to-go beads (pure taqtmready-to-gotm beads, ge healthcare, chicago, usa). the solution consisted of stabilizers, bsa, datp, dctp, dgtp, dttp, ± 2.5 units of puretaq dna polymerase and reaction buffer. each bead was reconstituted to a final volume of 25 μl. pcr products were analysed by electrophoresis on 1% agarose gel stained with gelred and visualised under uv light. the products of the pcr reaction for the ssu rdna gene were purified and precipitation reaction by ethanol/edta/ sodium acetate according to the protocol suggested by the manufacturer was performed, and then sequenced with a bigdye® terminator ver. 3.1 cycle sequencing kit (applied). automatic sequencing by capillary electrophoresis was performed on the abi3730xl dna analyzer (applied biosystems). the partial sequences obtained were assembled and edited using a sequenchertm ver. 5.2.4 (gene codes, ann arbor, mi) to obtain a consensus sequence. the newly generated partial sequences of ssu rdna were aligned using geneious ver. 7.1.3 (kearse et al. 2012) with the clustalw algorithm (larkin et al. 2007) and default settings with related genetic sequences that appeared on blastn search (table 2). the bayesian inference (bi) and maximum-likelihood (ml) analyses were performed using mrbayes ver. 3.1.2 (ronquist & huelsenbeck 2003) markov chain monte carlo (mcmc) chains were run for 10 million generations and the log-likelihood scores were plotted. the ‘burn-in’ was set to 30%. phyml 3.1 (guindon et al. 2010) software was used to perform ml analysis, using bootstrap confidence calculated with 1000 replications and gtr+i+g evolutionary model which was chosen by jmodeltest (posada 2008) as the best model for this analysis. phylogenetic trees were generated and edited in figtree ver. 1.4 (rambaut 2012). chloromyxum trilineatum sekyia, rosyadi, zhang & sato, 2019 (lc417364) was used as an out group. the aligned sequences of myxosporean parasites were compared using a pair-wise distance (p-distance) matrix (table 3). european journal of taxonomy 775: 107–118 (2021) 110 institutional abbreviations inpa = instituto nacional de pesquisas da amazônia results taxonomy phylum cnidaria hatschek, 1888 unranked subphylum myxozoa grassé, 1970 class myxosporea bütschli, 1881 order bivalvulida shulman, 1959 family myxidiidae thélohan, 1982 genus cystodiscus lutz, 1889 cystodiscus elachistocleis sp. nov. urn:lsid:zoobank.org:act:e2ae8907-e225-4804-a2f9-e3920ad6298d figs 1–3 type host elachistocleis cesarii (miranda-ribeiro, 1920). site of infection gallbladder. etymology the species epithet is derived from the genus elachistocleis of the host. material examined hapantotypes brazil • 10+ myxospores; mato grosso state, araguaiana municipality; 14°35′5.29″ s, 51°41′19.15″ w; glycerogelatin slide; genbank mz645740-mz645741; inpa79. description numerous pseudoplasmodia were found free in the bile. pseudoplasmodia (fig. 1c) of approximately 2 mm were rounded in shape, appeared to be formed by a ʻgelatinousʼ substance, and were floating in the host’s bile containing several myxospores in its interior. the morphology of myxospores found table 1. primers used for the amplification and sequencing of the ssu rdna of cystodiscus elachistocleis sp. nov. found parasitizing the gallbladder of elachistocleis cesarii (miranda-ribeiro, 1920) from araguaiana, mato grosso state, brazil. primer sequence 5’-3’ paired with reference erib1 acctggttgatcct act1r barta et al. (1997) act1r aatttcacctctcgctgcca erib1 hallett & diamant (2001) myxgen4f gtgccttgaataaatcagag erib10 diamant et al. (2004) erib10 cttccgcaggttcacctacgg myxgen4f barta et al. (1997) mx5 ctgcggacggctcagtaaatcagt mx3 andree et al. (1999) mx3 ccaggacatcttagggcatcacaga mx5 andree et al. (1999) http://zoobank.org/urn:lsid:zoobank.org:act:e2ae8907-e225-4804-a2f9-e3920ad6298d vieira d.h.m.d. et al., new species of cystodiscus from brazil 111 ta bl e 2. m or ph om et ri c co m pa ri so n of m at ur e m yx os po re s of c ys to di sc us e la ch is to cl ei s sp . no v. ( in pa 79 ) w ith c ys to di sc us s pp . a bb re vi at io ns : ss l = m yx os po re le ng th ; s w = m yx os po re w id th ; p l = po la r c ap su le le ng th ; p w = p ol ar c ap su le w id th ; t r = tr an sv er se ri dg es ; p t = p ol ar fi la m en t tu rn s. a ll di m en si on s ar e gi ve n in µ m . sp ec ie s sl sw pl pw t r pt si te o f in fe ct io n ty pe h os t c ou nt ry r ef er en ce c . e la ch is to cl ei s sp . n ov . 10 .6 ± 0 .3 (9 .8 –1 1. 2) 6. 2 ± 0. 5 (5 .6 –6 .6 ) 3. 6 ± 0. 4 (2 .8 –4 .3 ) 2. 6 ± 0. 3 (2 .2 –3 .1 ) 3– 5 2– 4 ga llb la dd er e la ch is to cl ei s ce sa ri i ( m ir an da -r ib ei ro , 1 92 0) b ra zi l pr es en t s tu dy c . i m m er su s l ut z, 1 88 9 11 .8 –1 3. 3 7. 5– 8. 6 3. 5– 4. 2 – 7– 9 – ga llb la dd er r hi ne lla m ar in a (l in na eu s, 17 58 ) b ra zi l k ud o & s pr ag ue (1 94 0) c . l yn do ye ns e c ar in i, 19 32 11 .0 –1 2. 0 7. 5– 8. 0 4. 0 (i n di am et er ) – – – ga llb la dd er r hi ne lla m ar in a (l in na eu s, 17 58 ) b ra zi l c ar in i ( 19 32 ) c . m el le ni (j ir ku , b ol ek , w hi pp s, j an ov y, k en t & m od ry , 2 00 6) 12 .3 (1 2. 0– 13 .5 ) 7. 6 (7 .0 – 9. 0) 5. 2 (4 .8 –5 .5 ) 4. 2 (3 .8 –4 .5 ) 2– 5 6– 7 ga llb la dd er p se ud ac ri s tr is er ia ta w ie dn eu w ie d, 1 83 8 u sa ji rk u et a l. (2 00 6) c . s er ot in us (k ud o & s pr ag ue , 1 94 0) 16 .0 –1 8. 0 9. 0 5. 0– 5. 5 (i n di am et er ) – 10 – 13 3– 5 ga llb la dd er r an a pi pi en s sc hr eb er , 1 78 2 u sa k ud o & s pr ag ue (1 94 0) c . t yp ho ni us (g ra y, 1 99 3) 10 .9 (9 .8 –1 2. 2) 7. 2 (5 .7 –8 .9 ) 3. 8 (2 .5 –5 .5 ) 3. 6 (3 .3 –5 .2 ) 9– 11 4– 5 ga llb la dd er b uf o m ar ga ri tif er (l au re nt i, 17 68 ) pe ru g ra y (1 99 3) c . h al da ri (s ar ka r, 19 82 ) 10 .8 (1 0. 0– 12 .0 ) 6. 7 (6 .5 –7 .0 ) 3. 6 (3 .0 –4 .0 ) – – – ga llb la dd er h yl a ar bo re a l in na eu s, 1 75 8 in di a sa rk ar (1 98 2) c . l es m in te ri (d el vi nq ui er , m ar ku s & p as sm or e, 1 99 2) 12 .5 (9 .5 –1 5. 0) 6. 5 (5 .7 –8 .0 ) – – 1– 2 – ga llb la dd er to m op te rn a kr ug er en si s pa ss m or e & c ar ru th er s, 1 97 5 so ut h a fr ic a d el vi nq ue r e t a l. (2 01 2) c . a xo ni s h ar tig an , f ia la , d yk ov á, r os e, ph al en & š la pe ta , 2 01 2 14 .1 (1 3. 0– 15 .5 ) 8. 5 (8 .0 –1 0. 5) 3. 8 (3 .0 –5 .0 ) 3. 7 (3 .0 –5 .0 ) 5– 12 4– 5 ga llb la dd er an d bi le du ct s li to ri a ra ni fo rm is k ef er st ei n, 18 67 a us tr al ia h ar tig an e t a l. (2 01 2c ) c . a us tr al is h ar tig an , f ia la , d yk ov á, r os e, ph al en & š la pe ta , 2 01 2 16 .0 (1 5. 0– 18 .0 ) 8. 7 (8 .0 –1 0. 0) 5. 3 (5 .0 –6 .0 ) 4. 8 (4 .0 –5 .5 ) 5– 11 5– 6 ga llb la dd er an d bi le du ct s li m no dy na st es p er on ii d um ér il & b ib ro n, 1 84 1 a us tr al ia h ar tig an e t a l. (2 01 2c ) european journal of taxonomy 775: 107–118 (2021) 112 in the gallbaldder of e. cesarii corresponded to the genus cystodiscus. the myxospores (figs 1a–b, 2) were ellipsoid to ovoid. the two myxospore valves were joined by a raised ridge that appeared straight or slightly curved (s-shaped in sutural view) along the medial axis of the myxospore. transverse depressions on the surface of the myxospores appeared connected to the sutural ridge depression. their measurements presented as mean ± sd (range) were: myxospore length 10.6 ± 0.3 (9.8–11.2) μm, myxospore width 6.2 ± 0.5 (5.6–6.6) μm. the polar capsules were pyriform and equal in size, situated each one in a myxospore extremity and measured 3.6 ± 0.4 (2.8–4.3) μm in length and 2.6 ± 0.3 (2.2– 3.1) μm in width. the polar filament present within the polar capsule had 2–4 coils. the myxospores showed 3–5 transverse ridges and a binucleated sporoplasm. filiform polar appendages were not observed. remarks cystodiscus elachistocleis sp. nov. was morphometrically compared to all cystodiscus spp. described worldwide (table 2). the species that most resembled c. elachistocleis sp. nov. was c. haldari (sarkar, 1982), which showed similarity in all measurements available for comparison. however, the geographical distance (brazil vs india) and the different type host (e. cesarii vs hyla arborea (linnaeus, 1758)) allow us to separate the two species. furthermore, c. haldari does not present any comparative values such as the number of turns of the polar filament and the number of transverse ridges, which could differentiate the two species. cystodiscus thyponius gray (1993) also presented measures of length and width of myxospores similar to those found in c. elachistocleis sp. nov. however, differences were observed in the polar capsule width (2.6 ± 0.3 (2.2–3.1) vs 3.6 (3.3–5.2)) and the number of transverse ridges (3–5 vs 9–11). regarding the cystodiscus spp. described from brazil, c. elachistocleis sp. nov. was morphometrically compared with two species parasitizing the gallbladder of rhinella marina (linnaeus, 1758). cystodiscus immersus, kudo & sprague (1940) presented a longer body length than that found in c. elachistocleis sp. nov. (11.8–13.3 vs 10.6 ± 0.3 (9.8–11.2)), in addition to a greater number of transverse ridges (7–9 vs 3–5). cystodiscus lyndoyense carini, 1932 showed the length of the body (11.0–12.0 vs 10.6 ± 0.3 (9.8– 11.2)) and the body width (7.5–8.0 vs 6.2 ± 0.5 (5.6–6.6)) longer than that found for c. elachistocleis sp. nov. the other cystodiscus spp. already described presented a body length longer than that observed in c. elachistocleis sp. nov. molecular analyses fragments of 1730-bp and 1916-bp of the ssu rdna gene were generated. the fragments showed 100% similarity when aligned. the blast search of the sequences did not reveal a direct match with myxozoan sequences available in genbank. the genetically closest species was c. immersus, which exhibited a similarity of 97.3%, and a difference of 24 out of 885 nucleotides (table 3). table 3. the similarity in ssu rdna sequences of cystodiscus spp. the upper triangle shows the percentage of nucleotide similarity, while the lower triangle shows the actual nucleotide difference. host isolate 1 2 3 4 5 6 1 cystodiscus elachistocleis sp. nov. mz645740 97.3 94.1 94.9 94.0 91.4 2 cystodiscus cf. immersus hq822162 24 95.1 96.2 94.9 93.0 3 cystodiscus cf. immersus 2 hq822159 52 43 84.6 93.8 91.4 4 cystodiscus melleni dq003031 89 34 48 93.1 92.3 5 cystodiscus australis hq822149 53 45 55 61 91.0 6 cystodiscus axonis hq822165 76 62 76 68 80 vieira d.h.m.d. et al., new species of cystodiscus from brazil 113 a well-supported phylogenetic tree divided into three main groups was obtained (fig. 3). a monophyletic group composed of sphaeromyxa thélohan, 1892 spp. that parasitizes fish, a polyphyletic group composed of zschokkella auerbach, 1909 and myxidium bütschli, 1882 spp. that infect fish, and fig. 1. myxospores and pseudoplasmodia of cystodiscus elachistocleis sp. nov. (inpa79) found parasitizing the gallbladder of elachistocleis cesarii (miranda-ribeiro, 1920) from araguaiana, mato grosso state, brazil. a. front view of c. elachistocleis sp. nov. b. front and side view of c. elachistocleis sp. nov. c. pseudoplasmodium (p) containing several myxospores of c. elachistocleis sp. nov. highlights myxospores (m) through the pseudoplasmodium-forming tissue. european journal of taxonomy 775: 107–118 (2021) 114 finally a monophyletic group composed of cystodiscus spp. that parasitize amphibians. still, there is a small monophyletic group composed of myxidium ssp. found in reptiles, fish, and birds. cystodiscus elachistocleis sp. nov. appears as a sister species of cystodiscus cf. immersus 1, in a subclade formed by species that parasitize the amphibians gallbladder. discussion considering the myxospore morphology, the morphometric data, and ssu rdna gene partial sequence obtained in the present study, we described c. elachistocleis sp. nov. this finding contributes to our knowledge of the biodiversity of cystodiscus in amphibians from brazil. to our knowledge, c. elachistocleis sp. nov. is the first record of a myxozoan species parasitizing e. cesarii and the third species of cystodiscus described from brazil. brazil is one of the most species-rich areas for amphibians globally (segalla et al. 2017). however, amphibians are also considered the most threatened vertebrate group with many species facing extinction (wake & vredenburg 2008; rebouças et al. 2021). amphibians also are hosts of a variety of endoparasites and ectoparasites, which in some cases influence their fitness, behavior, feeding, reproduction, and fertility (barta & desser 1984; lainson et al. 2003; muñoz-leal et al. 2017). for this study, only one specimen of e. cesarii was collected despite the extensive sampling effort performed. this could be due to the low presence of the host at the collection site or its ecology, which makes collecting it more difficult than other species. despite this, several studies describe new myxozoans from a single infected host (hartigan et al. 2012a; chen et al. 2020; mathews et al. 2021). regarding myxozoan parasites, there are only two species of cystodiscus described on brazilian anurans, c. immersus, and c. lyndoyense. however, the morphological and morphometric data differ from the species in the present study. the specificity and the host organ combined with myxospore morphology and molecular data are essential for a determination of a new myxozoan species. following this taxonomic strategy, we described a new species of cystodiscus, a gallbladder parasite of e. cesarii. the phylogenetic analysis performed here supports previous work that suggests that cystodiscus are amphibian gallbladder species (hartigan et al. 2012a). the monophyletic subclade composed of cystodiscus spp. was well supported. the low values of posterior probability found in some nodes within the subclade can be explained by the low number of described species that have their partial fig. 2. schematic drawing of cystodiscus elachistocleis sp. nov. (inpa79) found parasitizing the gallbladder of elachistocleis cesarii (miranda-ribeiro, 1920) in front and side view. vieira d.h.m.d. et al., new species of cystodiscus from brazil 115 sequences of the ssu rdna gene available. cystodiscus cf. immersus 1 and cystodiscus cf. immersus 2 appear with different sister species in the subclade, indicating that they are two distinct species. the genus cystodiscus is globally distributed and may have ecological implications outside of brazil (hartigan et al. 2012c). these parasites are associated with inflammation of nervous tissue and hepatic disease in several threatened or common frog species (hartigan et al. 2012b). although the situation of e. cesarii in the list of endangered species is ʻleast concern lcʼ, a parasite that causes serious pathologies in these amphibians could further decrease the population of amphibians in brazil, something that has been happening gradually (subirá et al. 2012; ceballos et al. 2020). despite this, in the present study, no pathologies were observed in the gallbladder or any other host organ associated with parasitism by cystodiscus elachistocleis sp. nov. acknowledgments we are grateful to fapesp (fundação de amparo à pesquisa do estado de são paulo) and fapemig (fundação de amparo à pesquisa do estado de minas gerais) for financial support. r.j. da silva is supported by fapesp #2016/50377-1, cnpq #309125/2017-0, cnpq-protax #440496/2015-2. d.h.m.d. vieira is supported by fapesp 2019/19060-0. l. pereira úngari is supported by fapesp #2018/00754-9 and 2018/09623-4. l. o´dwyer is supported by fapesp #2018/09623-4. all applicable international, national, and/or institutional guidelines for the care and use of animals were followed fig. 3. phylogenetic tree of bayesian analysis based on partial ssu rdna partial sequences showing the position of cystodiscus elachistocleis sp. nov. 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museum, prague, czech republic. european journal of taxonomy 751: 24–37 issn 2118-9773 https://doi.org/10.5852/ejt.2021.751.1377 www.europeanjournaloftaxonomy.eu 2021 · schneider c.w. et al. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e 24 the pseudodichotomous dasya sylviae sp. nov. (delesseriaceae, ceramiales) from 60–90 m mesophotic reefs off bermuda craig w. schneider 1,*, margaret m. cassidy 2 & gary w. saunders 3 1,2 department of biology, trinity college, hartford, ct 06106, usa. 3 centre for environmental & molecular algal research, department of biology, university of new brunswick, fredericton, new brunswick e3b 5a3, canada. * corresponding author: cschneid@trincoll.edu 2 email: margaret.cassidy@trincoll.edu 3 email: gws@unb.ca abstract. the red alga dasya sylviae c.w.schneid., m.m.cassidy & g.w.saunders sp. nov. is described from mesophotic depths of 60–90 m off bermuda. genetic sequences (coi-5p, rbcl) and morphological characteristics show that this species is distinct from other known pseudodichotomous species of dasya. of ten current species in the genus reported from bermuda, only three, d. collinsiana m.howe, d. cryptica c.w.schneid., quach & c.e.lane and d. punicea (zanardini) menegh., share the overall pattern of pseudodichotomous branching in their axes; however, key morphological features easily distinguish them from d. sylviae sp. nov. the species most similar in habit to d. sylviae sp. nov. is d. crouaniana j.agardh (type locality west indies), but it bears shorter pseudolateral branches, and broader and longer tetrasporangial stichidia than the new species. unique among the species of dasya, d. sylviae sp. nov. lacks post-sporangial cover cells in tetrasporangial stichidia. keywords. western atlantic, dasya, mesophotic zone, rhodophyta. schneider c.w., cassidy m.m. & saunders g.w. the pseudodichotomous dasya sylviae sp. nov. (delesseriaceae, ceramiales) from 60–90 m mesophotic reefs off bermuda. european journal of taxonomy 751: 24–37. https://doi.org/10.5852/ejt.2021.751.1377 introduction since joining the 2016 nekton xl catlin cruise of the r / v baseline explorer, the mesophotic collections of macroalgae taken off the coast of bermuda have yielded a growing list of species new to science (richards et al. 2018; schneider et al. 2018, 2019a, 2019b, 2020). in the present paper, we report another new species, this a member of the genus dasya c.agardh that was abundantly collected at several offshore reef sites from depths of 60–90 m (stefanoudis et al. 2018: fig. 53, as dasya sp.). the genus dasya presently hosts 90 species from throughout the world’s temperate and tropical seas (guiry & guiry 2020). its axial growth is sympodial with a main leading axis being continually displaced to one side by a new axis forming from a lateral bud near the apex. the resulting lateral axis takes over as the new main axis, a pattern that is repeated over and over in axial development (parsons 1975). https://doi.org/10.5852/ejt.2021.751.1377 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ https://orcid.org/0000-0003-0506-3791 https://orcid.org/0000-0003-4813-6831 mailto:cschneid%40trincoll.edu?subject= mailto:margaret.cassidy%40trincoll.edu?subject= mailto:gws%40unb.ca?subject= https://doi.org/10.5852/ejt.2021.751.1377 schneider c.w. et al., dasya sylviae sp. nov. from bermuda 25 based on appearance and not development, species of dasya mostly look to have alternate indeterminate branching patterns. however, some species of dasya have sympodial branching patterns that appear to be dichotomous or pseudodichotomous (subdichotomous), and they are described as having this type of branching (e.g., schneider et al. 2017; howe 1918; huisman 2018). in these species, the former main axis is not as suppressed by the newly produced lateral axis as in the case of most species of dasya and often grows nearly as long as the new leading axis, in some species to a greater degree than in others. of the ten species historically reported from bermuda (schneider 2003; schneider et al. 2017), only three, d. collinsiana m.howe, d. cryptica c.w.schneid., quach & c.e.lane and d. punicea (zanardini) menegh., have an at least partial pseudodichotomous branching pattern appearance of their indeterminate axes. a morphological comparison was conducted to compare all of the known large species of the genus demonstrating mostly or partially pseudodichotomous branching with collections discovered in deep water off bermuda demonstrating pseudodichotomous branching. using both morphological comparisons and molecular data from the offshore samples, we were able to determine that the mesophotic specimens represent a species of dasya new to science. material and methods on the 2016 nekton xl catlin cruise of the r / v baseline explorer (bex) off the coast of bermuda, collections from living low-profile reefs in the mesophotic zone (stefanoudis et al. 2018) were made by a team of technical rebreather divers (global underwater explorers (gue), high springs, florida, usa) equipped with closed-circuit jj-ccr ce edition rebreathers (jj-ccr aps, copenhagen, denmark) modified to gue configuration, and triton 1000-2 class submersibles (vero beach, florida, usa) with mechanical collecting arms. approximately 30 specimens of a common species of dasya were taken at five collecting sites from depths of 60–90 m. vouchers of type specimens are deposited in the herbaria noted in the material examined section below; herbarium abbreviations follow the online index herbariorum (thiers, continuously updated). collection site locations on the bex were recorded using a beier radio dp1 (dynamic positioning, beier integrated systems, mandeville, louisiana, usa) to receive shipboard gps. after living specimens were chosen for dna analysis, they were photographed using a canon powershot s90 digital camera (canon inc., tokyo, japan), and fragments of each were then dried in silica gel for later dna extraction. the remainder of the dna specimens were dried on herbarium paper as permanent vouchers. hand-cut sections were mounted in 30% corn syrup with acidified 1% aniline blue in a ratio of 20 :1. dried specimens were scanned on an epson et-2650 scanner (seiko epson corporation, suwa, nagano, japan), and photomicrographs were taken using a zeiss axioskop 40 microscope (oberkochen, germany) equipped with a spot idea 28.2 5mp digital camera (diagnostic instruments, sterling heights, michigan, usa). specimens generated for use in our molecular analysis are listed in table 1. when quick drying single or multiple isolates of associated field collection numbers, the silica gel samples were designated with unique ‘bda’ numbers. dna extractions of bda numbers followed saunders & mcdevit (2012) and pcr amplification and sequencing of coi-5p and rbcl were as detailed in saunders & moore (2013). these markers initially identified five specimens assignable to a new genetic group. to place this new species into a wider phylogenetic context, additional coi-5p and rbcl sequences were generated for a variety of species of dasya and the taxonomically related genera dasysiphonia i.k.lee & j.a.west, heterosiphonia mont. and rhodoptilum j.agardh from canada, australia and the us (table 1). to expand further our analyses, coi-5p and rbcl data were also downloaded from genbank for additional taxonomically related species, and their accession numbers are included in fig. 1. two single-gene alignments were generated: coi-5p with 28 sequences of 664 bp (base pairs) and rbcl with 36 sequences and 1272 bp. these alignments were analysed separately in geneious ver. 2021.0.1 (https://www.geneious.com; kearse et al. 2012) with maximum likelihood (gtr+i+g) using raxml (stamatakis 2014) with partitioning by codon and 500 bootstrap replicates. since no conflicts were detected, a concatenated coi-5p and http://www.geneious.com european journal of taxonomy 751: 24–37 (2021) 26 sp ec ie s v ou ch er n o. b o l d n o. c ol le ct or s c ol l. da te c ou nt ry p ro vi nc e /  s ta te l oc al it y °l at it ud e °l on gi tu de c o i5p rb cl d as ya b ai llo uv ia na (s .g .g m el .) m on t. g w s0 12 46 6 a b m m c 14 29 310 k . d ix on 21 a ug . 2 01 0 c an ad a n ov a sc ot ia so ut h h ar bo ur , c ap e b re to n is la nd 46 .8 67 23 7 -6 0. 44 71 24 h q 91 94 72 m w 69 87 13 d as ya ‘b ai llo uv ia na ’ c w s/ tr p 1 617 -4 (b d a 20 13 ) a b m m c 22 69 516 c . s ch ne id er , t. p op ol iz io 3 a ug . 2 01 6 b er m ud a sp itt al , o ff sh or e of m id oc ea n po in t 32 .3 20 00 00 -6 4. 65 80 56 — m w 69 87 01 d as ya c er am io id es h ar v. g w s0 14 83 8 a b m m c 71 12 -1 0 g . k ra ft , g . s au nd er s 14 j an . 2 01 0 a us tr al ia v ic to ri a q ue en sc liff j et ty , po rt p hi lli p h ea ds -3 8. 26 69 14 4. 66 78 h m 91 74 37 m w 69 87 11 d as ya c la vi ge ra (w om er sl ey ) m .j .p ar so ns g w s0 14 85 3 a b m m c 71 25 -1 0 g . k ra ft , g . s au nd er s 14 j an . 2 01 0 a us tr al ia v ic to ri a q ue en sc liff j et ty , po rt p hi lli p h ea ds -3 8. 26 69 14 4. 66 78 h m 91 74 45 m w 69 87 23 d as ya c or ym bi fe ra j. a ga rd h c w s/ c e l 10 -2 610 (b d a 04 22 ) b e r m r 25 210 c . s ch ne id er , c . l an e, d . m cd ev it, t. p op ol iz io 24 a ug . 2 01 0 b er m ud a sh ar k h ol e, h ar ri ng to n so un d, b er m ud a i. 32 .3 37 08 3 -6 4. 70 38 89 k x 91 33 29 m w 69 87 10 d as ya s es si lis y am ad a g w s0 11 95 5 a b m m c 70 65 -1 0 g . s au nd er s, t. a be , n . y ot su ku ra 2 d ec . 2 00 8 ja pa n k ik on i ( si te o f o ld m ar in e st at io n) 41 .7 00 75 14 0. 52 39 9 h m 91 74 02 m w 69 87 15 d as ya s p. 1 a u s g w s0 02 58 8 a b m m c 52 85 -0 9 g . s au nd er s 22 j an . 2 00 5 a us tr al ia ta sm an ia sn ug p ar k -4 3. 06 61 5 14 7. 26 45 h m 91 61 29 m w 69 87 00 d as ya s p. 2 g w s g w s0 14 96 9 a b m m c 72 19 -1 0 g . s au nd er s, k . d ix on 18 j an . 2 01 0 a us tr al ia ta sm an ia w in dm ill p oi nt , g eo rg e to w n -4 1. 10 97 14 6. 81 69 9 h m 91 75 14 m w 69 87 16 d as ya s p. 2 ta s g w s0 25 12 4 o z se a 50 810 g . s au nd er s, k . d ix on 11 n ov . 2 01 0 a us tr al ia w es te rn a us tr al ia c an al r oc ks -3 3. 66 93 5 11 4. 99 53 1 m w 69 97 59 m w 69 87 05 d as ya s p. 3 w a g w s0 24 88 8 o z se a 51 410 g . s au nd er s, k . d ix on 10 n ov . 2 01 0 a us tr al ia w es te rn a us tr al ia c oz y c or ne r (k no bb y pt .) -3 4. 25 59 5 11 5. 02 77 7 m w 69 97 59 m w 69 87 03 d as ya s p. 4 w a g w s0 24 89 3 o z se a 44 310 g . s au nd er s, k . d ix on 10 n ov . 2 01 0 a us tr al ia w es te rn a us tr al ia c oz y c or ne r (k no bb y pt .) -3 4. 25 59 5 11 5. 02 77 7 m w 69 97 63 m w 69 87 08 d as ya s pi nu lig er a c ol lin s & h er v. c w s/ c e l 10 -1 56 (b d a 02 10 ) b e r m r 13 110 c . s ch ne id er , c . l an e, d . m cd ev it, t. p op ol iz io 21 a ug . 2 01 0 b er m ud a g at es f or t, st . g eo rg e’ s h ar bo ur 32 .3 78 78 -6 4. 66 33 9 k x 91 33 27 m w 69 87 14 1 d as ya s yl vi ae c .w .s ch ne id ., m .m . c as si dy & g .w .s au nd er s sp . n ov . c w s/ tr p 1 611 -1 (b d a 19 69 ) a b m m c 22 68 816 c . s ch ne id er , t. p op ol iz io 28 j ul . 2 01 6 b er m ud a l ed ge n or th n or th ea st of s t. g eo rg e’ s i. 32 .4 79 93 33 -6 4. 59 48 05 m w 69 97 58 m w 69 87 04 c w s/ tr p 1 611 -1 0 (b d a 19 82 ) a b m m c 22 69 216 c . s ch ne id er , t. p op ol iz io 28 j ul . 2 01 6 b er m ud a l ed ge n or th n or th ea st of s t. g eo rg e’ s i. 32 .4 79 93 33 -6 4. 59 48 05 m w 69 97 65 — c w s/ tr p 1 612 -2 (b d a 19 92 ) a b m m c 22 69 316 c . s ch ne id er , t. p op ol iz io 29 j ul . 2 01 6 b er m ud a l ed ge n or th n or th ea st of s t. g eo rg e’ s i. 32 .4 82 70 -6 4. 58 75 31 m w 69 97 61 — c w s/ tr p 1 617 -2 (b d a 20 11 ) a b m m c 22 69 416 c . s ch ne id er , t. p op ol iz io 3 a ug . 2 01 6 b er m ud a sp itt al , o ff sh or e of m id oc ea n po in t 32 .3 20 00 0 -6 4. 65 80 56 m w 69 97 66 — ta bl e 1 (c on tin ue d on n ex t p ag e) . c ol le ct io n da ta fo r i so la te s w ith n ew ly g en er at ed g en b an k ac ce ss io n nu m be rs in cl ud ed in th e m ol ec ul ar a na ly se s of th is st ud y. 1 = ou r r bc l -3 p se qu en ce g en er at ed h er e w as li nk ed to rb cl -5 p da ta a va ila bl e in g en b an k (k x 91 33 53 ) f or d as ya sp in ul ig er a c ol lin s & h er v. schneider c.w. et al., dasya sylviae sp. nov. from bermuda 27 sp ec ie s v ou ch er n o. b o l d n o. c ol le ct or s c ol l. da te c ou nt ry p ro vi nc e /  s ta te l oc al it y °l at it ud e °l on gi tu de c o i5p rb cl d as ya s yl vi ae c .w .s ch ne id ., m .m . c as si dy & g .w .s au nd er s sp . n ov . c w s/ tr p 1 621 -1 4 (b d a 20 30 ) is ot yp e a b m m c 22 69 816 c . s ch ne id er , t. p op ol iz io 11 a ug . 2 01 6 b er m ud a sp itt al , o ff sh or e of m id oc ea n po in t 31 .3 20 55 55 -6 4. 65 88 89 m w 69 97 69 — c w s/ tr p 1 621 -1 4 (b d a 20 31 ) h ol ot yp e a b m m c 22 69 916 c . s ch ne id er , t. p op ol iz io 11 a ug . 2 01 6 b er m ud a sp itt al , o ff sh or e of m id oc ea n po in t 31 .3 20 55 55 -6 4. 65 88 89 — m w 69 87 21 d as ya te nu is m .j .p ar so ns & w om er sl ey g w s0 25 51 7 o z se a 48 510 g . s au nd er s, k . d ix on 14 n ov . 2 01 0 a us tr al ia w es te rn a us tr al ia b la ck w al l r ea ch , sw an r iv er -3 2. 02 10 1 11 5. 78 31 6 m w 69 97 64 m w 69 87 09 d as ys ip ho ni a ja po ni ca (y en do ) h .-s .k im g w s0 30 12 1 a b m m c 16 70 712 a . s av oi e 18 a pr . 2 01 2 u sa m as sa ch us et ts g ar ba ge b ea ch b re ak w at er , w oo ds h ol e 41 .5 25 18 -7 0. 67 25 6 k c 15 85 82 m w 69 87 22 d as ys ip ho ni a sp . 1 w a g w s0 24 41 6 o z se a 52 910 g . s au nd er s, k . d ix on 7 n ov . 2 01 0 a us tr al ia w es te rn a us tr al ia l itt le b ea ch -3 4. 97 16 3 11 8. 19 59 9 m w 69 97 62 m w 69 87 07 d as ys ip ho ni a sp . 2 w a g w s0 24 79 8 o z se a 53 610 g . s au nd er s, k . d ix on 9 n ov . 2 01 0 a us tr al ia w es te rn a us tr al ia c ap e l ee uw in l ig ht ho us e -3 4. 37 16 7 11 5. 13 63 m w 69 97 60 m w 69 87 06 h et er os ip ho ni a ca lli th am ni um (s on d. ) f al ke nb . g w s0 24 72 6 o z se a 53 310 g . s au nd er s, k . d ix on 9 n ov . 2 01 0 a us tr al ia w es te rn a us tr al ia c ap e l ee uw in l ig ht ho us e -3 4. 37 16 7 11 5. 13 63 m w 69 97 72 — g w s0 24 72 7 o z se a 53 410 g . s au nd er s, k . d ix on 9 n ov . 2 01 0 a us tr al ia w es te rn a us tr al ia c ap e l ee uw in l ig ht ho us e -3 4. 37 16 7 11 5. 13 63 m w 69 97 71 m w 69 87 20 h et er os ip ho ni a cr as si pe s (h ar v. ) fa lk en b. g w s0 23 91 1 o z se a 54 310 g . s au nd er s, r . w ith al l 26 n ov . 2 01 0 a us tr al ia n ew s ou th w al es y el lo w r oc k, l or d h ow e i. -3 1. 51 5 15 9. 03 44 m w 69 97 67 m w 69 87 17 h et er os ip ho ni a pl um os a (j .e lli s) b at te rs g w s0 40 69 0 a b m m c 21 92 816 g . s au nd er s, t. b ri ng lo e 9 ju n. 2 01 6 n or w ay k le pp es jo en 60 .1 84 74 5. 14 93 8 m n 18 42 31 m n 18 45 25 h et er os ip ho ni a de ns iu sc ul a k yl in g w s0 28 12 8 a b m m c 15 50 211 g . s au nd er s, k . d ix on 7 ju l. 20 11 c an ad a b ri tis h c ol um bi a m ur ch is on i. , e as t e nd , g w ai i h aa na s 52 .6 05 85 -1 31 .4 32 89 m w 69 97 70 m w 69 87 19 h et er os ip ho ni a sp . 2 a u s g w s0 16 47 6 a b m m c 82 55 -1 0 g . s au nd er s, l . k ra ft , k . d ix on 29 j an . 2 01 0 a us tr al ia ta sm an ia st an le y b re ak w at er -4 0. 76 73 1 14 5. 30 58 3 m w 69 97 56 m w 69 87 02 r ho do pt ilu m pl um os um (h ar v. & b ai le y) k yl in g w s0 36 81 5 a b m m c 21 03 215 g . s au nd er s, m . b ru ce , t. b ri ng lo e 11 j ul . 2 01 5 c an ad a b ri tis h c ol um bi a sm yt he p as sa ge e as t (m ac ro cy st is b ed ), h ai da g w ai i 54 .0 67 91 -1 32 .5 19 58 m w 69 97 68 m w 69 87 18 ta bl e 1 (c on tin ue d) . c ol le ct io n da ta fo r i so la te s w ith n ew ly g en er at ed g en b an k ac ce ss io n nu m be rs in cl ud ed in th e m ol ec ul ar a na ly se s of th is s tu dy . european journal of taxonomy 751: 24–37 (2021) 28 rbcl alignment was constructed (38 sequences, 1936 bp) with analyses as described for the single gene alignments, but with partitioning by gene and codon and with 1000 bootstrap replicates (fig. 1). in all cases, the trees were rooted on the lineage defined as the heterosiphonia group (choi et al. 2002). results phylogenetic analysis in the speciose genus dasya worldwide, there is a total of eight known corticated species taller than 3 cm at maturity that bear indeterminate axes appearing mostly or partially pseudodichotomously branched. a comparative review of these species is summarized in table 2. none of these bear a suite of characteristics that is comparable with the mesophotic specimens collected off bermuda. we successfully generated coi-5p (664 bp) for five individuals of this novel species and all had identical sequences except for one substitution in bda2011, or 0 –0.15% divergence within this species. the nearest neighbor identified through a blast search in genbank was dasya adela heggøy, rueness & sjøtun that demonstrated a 6% divergence. similarly, we generated an rbcl sequence for two specimens, which were identical over 1358 bp and 2.9% divergent from d. adela. phylogenetic analyses placed the new genetic group solidly in a clade with d. adela and an undescribed species from the euphotic zone off bermuda (fig. 1, dasya sp. 1bda). of further note, none of the genera included in the current tree were monophyletic except for the monospecific rhodoptilum, which nonetheless fell solidly in a group with the generitype of dasya, d. baillouviana (s.g.gmel.) mont. (fig. 1). clearly considerable taxonomic work remains to be done on the genera included in our phylogenetic analyses, but this does not detract from our clear discovery of a novel species best included in the genus dasya. therefore, based upon our molecular comparisons and phylogenetic analysis, we here describe the following unique mesophotic species of dasya for bermuda and the western atlantic. taxonomic treatment phylum rhodophyta wettst. subphylum eurhodophytina g.w.saunders & hommers. class florideophyceae cronquist subclass rhodymeniophycidae g.w.saunders & hommers. order ceramiales oltm. family delesseriaceae bory subfamily dasyoideae de toni genus dasya c.agardh nom. cons. dasya sylviae c.w.schneid., m.m.cassidy & g.w.saunders sp. nov. fig. 2 diagnosis differing from most species of dasya by its pronounced pseudodichotomous branching pattern (fig. 2a– c), and from its most similar congener in habit, d. crouaniana j.agardh, by its longer pseudolaterals, narrower and shorter tetrasporangial stichidia and axes fully covered with pseudolaterals to barely denuded proximal axes. the new taxon differs from all species of dasya by its lack of post-sporangial cover cells. etymology the species is named after dr sylvia a. earle, pioneering phycologist, scientist and open-water diver, 50 years after she led the first all-female team of aquanauts in tektite ii on the floor of the caribbean sea (earle 1972a, 1972b). schneider c.w. et al., dasya sylviae sp. nov. from bermuda 29 d . a na st om os an s d . c ar te ri d . c ol lin si an a d . c ro ua ni an a d . c ry pt ic a d . h ai tia na d . p un ic ea d . r os yl ni ae d . s yl vi ae s p. n ov . p la nt h ei gh t ( cm ) 3– 6( –2 0) to 1 1 1– 3( –5 +) to 7 4– 5 5– 9 5– 10 to 1 0 4– 17 b ra nc hi ng p at te rn ps eu do di ch ot om ou s ir re gu la rl y ps eu do di ch ot om ou s al te rn at e, ir re gu la r, to so m ew ha t p se ud odi ch ot om ou s w id el y ps eu do di ch ot om ou s to ir re gu la r ps eu do di ch ot om ou s ps eu do di ch ot om ou s ps eu do di ch ot om ou s to ir re gu la r al te rn at e to p se ud odi ch ot om ou s an d ir re gu la r ps eu do di ch ot om ou s a xi al c or ti ca ti on co m pl et e co m pl et e m os tly c om pl et e co m pl et e co m pl et e co m pl et e co m pl et e he av ily c or tic at ed to s ev er al s eg m en ts fr om a pe x co m pl et e a xi al d ia m et er (m m ) 1– 3 to 3 0. 5– 0. 75 0. 5– 0. 7 1– 2 0. 5– 0. 6 0. 4– 1. 2 1. 8 0. 3– 0. 8 a pi ce s of m ai n ax es ta pe ri ng s lig ht ly to tip s, b ra nc h en ds re cu rv ed ta pe ri ng d is ta lly ta pe ri ng d is ta lly ta pe ri ng s lig ht ly to ti ps , b ra nc h en ds re cu rv ed ta pe ri ng d is ta lly ta pe ri ng d is ta lly ta pe ri ng s lig ht ly to ti ps p se ud ol at er al s m on os ip ho no us m on os ip ho no us m on os ip ho no us m on os ip ho no us m on os ip ho no us m on os ip ho no us an d po ly si ph on ou s m on os ip ho no us m on os ip ho no us fe w w ith p ol ys ip ho no us ba se s m on os ip ho no us d is po si ti on ra nd om – sp ir al ed ra nd om ra nd om sp ir al ed su bve rt ic ill at e sp ir al ed ra nd om a xi al c ov er ag e de ns e co ve ra ge , de nu de d in pr ox im al a xe s of ol de r p la nt s de ns e co ve ra ge , de nu de d in pr ox im al a xe s of ol de r p la nt s de ns e co ve ra ge u p to th e la st 2 –3 di ch ot om ie s, lig ht er a bo ve de ns e co ve ra ge di st al ly , m os tly w ith lo w er h al f co m pl et el y de nu de d de ns e co ve ra ge , de nu de d in pr ox im al a xe s of ol de r p la nt s de ns e co ve ra ge , de nu de d in pr ox im al a xe s of ol de r p la nt s de ns el y co ve re d di st al ly , n ak ed pr ox im al ly de nu de d of m on os ip ho no us la te ra ls fo r m uc h of th ei r l en gt h de nu de d on ly in m os t p ro xi m al a re a of o ld er p la nt s b ra nc hi ng 1– 3 tim es fr om ba sa l p or tio ns 1– 4( –5 ) t im es ps eu do di ch ot om ou sl y br an ch ed n ea r ba se 5– 8 tim es di va ri ca te ly ps eu do di ch ot om ou s 2– 3 tim es di ch ot om ou sl y 2– 4 tim es fr om ba sa l p or tio ns – 2– 3 tim es di ch ot om ou sl y ne ar b as e 5 tim es d ic ho to m ou s to 7 ti m es ir re gu la rl y di ch ot om ou s o ve ra ll le ng th (m m ) 2– 5 to 3 to 0 .9 1– 2 2– 4 – 2. 0– 4. 4 0. 8 1. 7– 5. 5 n o. c el ls b as e to a pe x 31 –4 4 30 –7 0 – – – 25 –1 02 b as al c el l di am et er (μ m ) 15 –4 4 12 –1 4 10 0– 13 0 18 –2 0 12 –3 3 – to 4 4 21 –3 7 b as al c el l le ng th (μ m ) 14 –4 5 – w id er th an lo ng 20 –4 0 24 –4 8 – – 12 –3 0 su pr ab as al c el l di am et er (μ m ) 7– 14 ta pe ri ng – – 6– 16 – 20 –4 0 – 19 –3 2 su pr ab as al c el l le ng th (μ m ) 18 –3 3 – – – 1– 38 – 50 –8 0 – 8– 32 m ed ia n ce ll di am et er (μ m ) 13 –1 5 8– 12 to 7 5 – 14 –1 8 – – – 9– 22 ta bl e 2 (c on tin ue d on n ex t p ag e) . a m or ph ol og ic al c om pa ri so n of c or tic at ed p se ud od ic ho to m ou s d as ya s pp . t al le r t ha n 3 cm . european journal of taxonomy 751: 24–37 (2021) 30 d . a na st om os an s d . c ar te ri d . c ol lin si an a d . c ro ua ni an a d . c ry pt ic a d . h ai tia na d . p un ic ea d . r os yl ni ae d . s yl vi ae s p. n ov . m ed ia n ce ll le ng th (μ m ) 50 –6 0 2. 5 to 2 d ia m et er s 4 tim es a s lo ng a s br oa d 75 –9 2 – – – 27 –1 02 a pi ca l c el l di am et er (μ m ) 4– 12 8– 12 45 –5 5 5– 12 5– 18 – 12 – 8– 19 a pi ca l c el l le ng th (μ m ) 10 –2 4 90 –1 10 to 3 0 11 –4 3 – – – 32 –1 27 te tr as po ra ng iu m di am et er (μ m ) 32 –4 5 35 –4 0 30 –4 0 18 –3 0 30 –5 0 60 –7 0( –8 0) 30 –4 0 to 4 0 21 –2 9 te tr as po ra ng ia l st ic hi di um lin ea r l an ce ol at e la nc eo la te ov at ela nc eo la te , of te n cu rv ed la nc eo la te to el on ga te d cy lin dr ic al lin ea r l an ce ol at e la nc eo la te a nd sl ig ht ly c ur ve d ov at e to la nc eo la te ov at e to la nc eo la te la nc eo la te to na rr ow ly e lli pt ic al d ia m et er (μ m ) 50 –1 00 – 90 –1 30 80 –1 20 60 –8 0 70 –2 00 70 –9 0 12 0 73 –8 0 l en gt h (μ m ) 21 0– 40 0 – 30 0– 50 0 to 1 00 0 34 0– 10 40 30 0– 70 0 30 0– 40 0 to 6 40 26 9– 30 5 n o. fe rt ile s eg m en ts  / st ic hi di um 10 –1 3 15 – – 10 –2 7 – – to 1 2 10 –1 2 n o. s po ra ng ia  / fe rt ile s eg m en t 3– 5 4 – – 4( –5 ) 5 5 6 3– 5 p os tsp or an gi al co ve r ce lls 1( –2 ) 2( –3 ) 3 – 1 (2 –) 3 – 3 ab se nt sp er m at an gi al st ic hi di um – – – la nc eo la te to el on ga te d cy lin dr ic al – – – lin ea r l an ce ol at e la nc eo la te to na rr ow ly e lli pt ic al d ia m et er (μ m ) – – 24 –3 6 60 –7 0 – – – 50 –6 0 46 –6 9 l en gt h (μ m ) – – 60 –1 00 15 0– 22 5 – – – to 4 25 19 2– 25 8 ty pe lo ca lit y in do ne si a w es te rn a us tr al ia b er m ud a w es t i nd ie s b er m ud a h ai ti it al y n ew s ou th w al es b er m ud a r ef er en ce s sc hn ei de r e t a l. 20 17 h ui sm an 2 01 8 h ow e 19 18 ; l itt le r & l itt le r 20 00 ; pr es en t s tu dy ta yl or 1 92 8; l itt le r & l itt le r 20 00 ; d aw es & m at hi es on 2 00 8 sc hn ei de r e t a l. 20 17 fr ed er ic q & n or ri s 19 86 ta yl or 1 96 0; l itt le r & l itt le r 2 00 0; l óp ez -p iñ er o & b al la nt in e 20 01 m ill ar 1 99 6 pr es en t s tu dy ta bl e 2 (c on tin ue d) . a m or ph ol og ic al c om pa ri so n of c or tic at ed p se ud od ic ho to m ou s d as ya s pp . t al le r t ha n 3 cm . schneider c.w. et al., dasya sylviae sp. nov. from bermuda 31 fig. 1. concatenated coi-5p and rbcl raxml tree, partitioned by gene and codon, gtr+i+g with 1000 bootstrap replications (only bootstrap values > 75% shown). asterisks (*) denote 100% bootstrap support. sequences taken from genbank are indicated by including their accession numbers in parentheses (rbcl / coi-5p). the novel genetic group, dasya sylviae sp. nov., is presented in bold type. european journal of taxonomy 751: 24–37 (2021) 32 material examined type bermuda • spittal, south of cooper’s i. off castle harbour; 31°19.23333′ n, 64°39.53333′ w; depth 63.8 m; on rhodoliths; 11 aug. 2016; schneider & popolizio 16-21-14; genbank nos: mw698721 (holotype), mw699769 (isotype); holotype (fig. 2a): mich [bda2031]; isotypes (fig. 2b–c): bermuda natural history museum, mich, ny, unb, us, herb. cws [bda2030]. additional material bermuda • ledge north northeast of st. george’s i.; 32°28.79600′ n, 64°35.68833′ w; depth 90 m; 28 jul. 2016; schneider & popolizio 16-11-1, ⊕ [bda1969] • loc. cit.; depth 60 m; 28 jul. 2016; schneider & popolizio 16-11-10 [bda1982] • loc. cit.; 32°28.96200′ n, 64°35.25183′ w; depth 90 m; 29 jul. 2016, schneider & popolizio 16-12-2 [bda1992] • spittal, offshore of midocean point; 32°19.20000′ n, 64°39.48333′ w; depth 62.7 m; 3 aug. 2016; schneider & popolizio 16-17-2 [bda2011]. description plants epilithic, erect to 17 cm tall, carmine red, arising from small discoidal holdfasts; indeterminate axes sympodially branched, appearing pseudodichotomously branched throughout, only slightly tapering from base to apex (fig. 2 a–c), 0.3–0.8 mm diam. in median to lower portions and completely corticated by rhizoidal downgrowth (fig. 2d); indeterminate axes densely covered throughout with determinate, lightly pigmented, monosiphonous dichotomously branched axes (pseudolaterals; fig. 2e), except in some older plants in the most basal portions; pseudolaterals 1.7 –5.5 mm in length, 25–91 cells from base to apex, irregularly dichotomously branched from the first to the seventh cell of the pseudolateral, upper portions unbranched and slightly tapering (fig. 2e); basal cells of pseudolaterals initially globose to ellipsoid (fig. 2d), 21–37 µm diam. and 12–30 µm long, then slightly elongating; suprabasal cells 19–32 µm diam. and 8–32 µm long, elongating more centrally, 9–22 um diam. and 27–102 µm long, and reaching greatest lengths distally, 8–19 µm diam. and 32–127 µm long; tetrasporangial stichidia single, borne terminally on 3–7-celled unbranched pseudolaterals (fig. 2f) or terminating basal dichotomies of pseudolaterals (fig. 2g), lanceolate to narrowly elliptical in outline (fig. 2f–g), 73–80 µm diam. and 269–305 µm in length at maturity, composed of 10–12 fertile segments, acropetally producing then releasing sporangia (fig. 2f–g); sporangia globose, 21–29 µm diam., tetrahedrally divided, 3–5 per fertile segment, sporangia borne on 2-celled whorled branches, post-sporangial cover cells lacking (fig. 2f); one to two spermatangial stichidia terminating a basal dichotomy on 4–10-celled pedicels (pseudolaterals), narrowly elliptical to lanceolate in outline (fig. 2h), 46–69 µm diam. and 192–258 µm in length at maturity; carpogonial branches and cystocarps unknown. distribution and habitat at present, endemic to mesophotic reefs off bermuda, western atlantic ocean. discussion the genus dasya is characterized by the development of 2–4 preand post-sporangial cover cells that partially cover tetrasporangia in their stichidium (parsons 1975). dasya sylviae sp. nov. appears to be unique among its congeners as it completely lacks these cover cells, thus tetrasporangia sit naked on their whorl branches (table 2, fig. 2g). among the presently accepted 90 species of dasya (guiry & guiry 2020), only eight corticated species of dasya that reach at least 3 cm tall at maturity, including three from bermuda (d. collinsiana, d. cryptica and d. punicea), appear to share an overall axial pattern of pseudodichotomous branching with d. sylviae sp. nov. (table 2, fig. 2a–c). however, key morphological characteristics easily distinguish them from the new species presented here. among the few species with longer pseudolaterals approaching the length of those in the new species (to 5.5 mm schneider c.w. et al., dasya sylviae sp. nov. from bermuda 33 fig. 2. dasya sylviae c.w.schneid., m.m.cassidy & g.w.saunders sp. nov. a. holotype specimen, cws / trp 16-21-14 (bda2031); mich. b–c. isotype specimens. d. whole mount near apex with rhizoidal cortication and pseudolateral branches, cws / trp 16-17-2. e. whole mount of axis with emerging pseudolaterals, cws / trp 16-21-14. f. tetrasporangial stichidia on long pedicels, cws / trp 16-11-1 (bda1969). g. acropetal development in a tetrasporangial stichidium at base of pseudolateral branch with whorled two-celled branches bearing sporangia that lack post-sporangial cover cells, cws / trp 16-11-1 (bda1969). h. spermatangial stichidia at axis apex, cws / trp 16-21-14. scale bars: a–c = 2 cm; d, f = 50 µm; e = 500 µm; g–h = 100 µm. european journal of taxonomy 751: 24–37 (2021) 34 long), d. anastomosans (weber bosse) m.j.wynne and d. cryptica demonstrate a short bushy habit with more dense pseudolateral growth, larger tetrasporangia and longer stichidia (table 2). the species most similar to dasya sylviae sp. nov. in overall habit is d. crouaniana (type locality west indies), but the latter species is characterized by its striking loss of deciduous pseudolaterals in the lower half of its main axes contrasted by densely enveloped distal portions as illustrated in taylor (1928: pl. 35 fig. 5; 1960: pl. 71 fig 1). these pseudolaterals are shorter in length (1–2 µm) but more densely packed than the longer ones of d. sylviae sp. nov. (1.7–5.5 µm). furthermore, the tetrasporangial stichidia of d. crouaniana are both longer and broader (to 1 mm × 80–120 µm) than in the new species (269–305 µm × 73–80 µm). unfortunately, we do not have genetic information for this species, but its morphological differences distinguish it from d. sylviae sp. nov. while dasya punicea (type locality venice) appears to be somewhat similar to d. sylviae sp. nov. in habit, the mediterranean species can be differentiated by its subverticillate pseudolaterals, slightly longer tetrasporangial stichidia (300–400 µm vs 269–305 µm) and slightly larger tetrasporangia (30–40 µm vs 21–29 µm). its long pseudolaterals (to 4.4 mm) are reminiscent of the new species. dasya punicea was reported from bermuda by collins & hervey (1917) who stated that their specimens had a tendency to issue ramelli [pseudolaterals] “in more or less distinct whorls,” a condition dissimilar to that in specimens from the eastern atlantic where the pseudolaterals were spirally arranged (maggs & hommersand 1993). ballantine & aponte (2004) argued that the entity first reported in the western atlantic from bermuda by collins & hervey (1917) as d. punicea was different from eastern atlantic and mediterranean isolates. three archival specimens left by a. hervey as d. punicea (collins & hervey 1917; ny 2178604) are an admixture of two species on a single sheet, one representing a young dasya spinuligera collins & herv., the remaining representative of wrangelia c.agardh (wrangeliaceae j.agardh), not dasya. the mesophotic specimens described here as a new taxon were collected along with two other species of dasya at these depths, d. cf. baillouviana (s.g.gmel.) mont. (58–77 m) and d. spinuligera (60 m). unlike d. sylviae sp. nov., both of these species with different and distinctive morphologies are also known on shallow subtidal reefs in bermuda. including d. collinsiana, d. cryptica and d. spinuligera, d. sylviae sp. nov. represents the fourth species of the genus with its type locality in bermuda (collins & hervey 1917; howe 1918; schneider et al. 2017). genetically, dasya sylviae sp. nov. falls in a clade with the recently described d. adela, a species discovered in a landlocked fjord in norway (sjøtun et al. 2016), and the alternately to irregularly branched dasya sp. 1bda from the shallow subtidal of bermuda (schneider et al. 2017) (fig. 1). dasya adela is significantly smaller (to 3 cm) than d. sylviae sp. nov., and develops “radially to irregularly set side [indeterminate] branches” (sjøtun et al. 2016) and cover cells for tetrasporangia in stichidia. these two species are morphologically easy to differentiate even if their habitats weren’t also disparate. acknowledgements the xl catlin deep ocean survey, nekton’s mission to the northwest atlantic and bermuda aboard the bex, allowed for the 2016 collections. nekton gratefully acknowledges the support of xl catlin and the garfield western foundation. work on the r / v baseline explorer (bex) would have been impossible without the assistance of capt. larry bennett and his crew, brownies global logistics, triton submersibles and pilots, and the several volunteer technical divers of global underwater explorers led by dr todd kincaid. we thank dr thea popolizio for helping collect aboard the bex and multiple members of the saunders lab over the years, notably cody brooks and tanya moore, for sequencing the specimens of dasya reported on here. christopher flook of the bermuda institute of ocean sciences (bios) and roger simmons of the bermuda aquarium, natural history museum and zoo (bamz) provided logistical support while in bermuda. the genetic work at unb was supported by discovery and schneider c.w. et al., dasya sylviae sp. nov. from bermuda 35 accelerator grants to gws from the natural sciences and engineering research council of canada, as well as funding from the canada foundation for innovation, the new brunswick innovation foundation, and the nekton foundation. this is contribution no. 286 to the bermuda biodiversity project (bbp) of bamz, department of environment & natural resources, and nekton contribution no. 19. references ballantine d.l. & aponte n.e. 2004. dasya abbottiana sp. nov. 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meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. http://sweetgum.nybg.org/science/ih/ 1 european journal of taxonomy 775: 1–14 issn 2118-9773 https://doi.org/10.5852/ejt.2021.775.1539 www.europeanjournaloftaxonomy.eu 2021 · tshernyshev s.e. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:c57bc1ca-1ecf-41dc-b7b3-2d4de8137e3f pectotibialis paghmanensis tshernyshev gen. nov. – a new genus and species of soft-winged flower beetle (coleoptera, cleroidea, malachiidae) from afghanistan sergei e. tshernyshev 1,2 1 institute of systematics and ecology of animals, siberian branch of the russian academy of sciences, frunze street 11, 630091 novosibirsk, russia. 2 tomsk state university, lenina prospekt 36, tomsk 634050, russia. email: sch-sch@mail.ru urn:lsid:zoobank.org:author:c9cc4a86-bd1f-499a-b831-f727a111bbaa abstract. a new genus with a new species of soft-winged flower beetle, pectotibialis paghmanensis tshernyshev gen. et sp. nov. are described from afghanistan. the new genus can be distinguished from the congeners of the tribe apalochrini by the dark pectination in the apices of tibiae in both sexes, and the anterior tibiae which are hollowed at distal half, flattened and rounded femora, bituberculate basal parts of head and pronotum, two lamellate metathoracic appendages, tarsal comb above second tarsomere of anterior legs, and serrate antennae in the male. based on the metathoracic appendages and comb in anterior legs would attribute this new species to the new genus dromanthomorphus pic, 1921, but all the other above-mentioned characters define its independent status and the designation of a new genus; pectotibialis tshernyshev gen. nov. the external appearance, special male characters and genitalia of the type species of the new genus are illustrated, and a distribution map is provided. a key to the apalochrus-section of the tribe apalochrini is provided. keywords. afghanistan, soft-winged flower beetle, apalochrini, pectotibialis, new genus, new species, systematics. tshernyshev s.e. 2021. pectotibialis paghmanensis – a new species and a new genus of soft winged flower beetle of the tribe apalochrini (coleoptera, cleroidea, malachiidae) from afghanistan. european journal of taxonomy 775: 1–14. https://doi.org/10.5852/ejt.2021.775.1539 introduction amongst the soft-winged flower beetles obtained by my colleague oleg pak in afghanistan were several specimens of narrow, parallel-sided malachiids resembling representatives of the genus apalochrus erichson, 1840 and belonging to the tribe apalochrini. the male has with a comb above the second tarsomere in the anterior legs, and this character allows this species to be placed near protapalochrus evers, 1987. both genera, apalochrus erichson and protapalochrus evers, are distributed in central asia, and two species, a. flavicollis schaufuss, 1870 and p. (p.) fedtschenkoi (solsky, 1882), are known from afghanistan (mayor 2003, 2007; tshernyshev 2015c, 2016b); however, the newly found taxon https://doi.org/10.5852/ejt.2021.775.1539 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:c57bc1ca-1ecf-41dc-b7b3-2d4de8137e3f http://orcid.org/0000-0001-5931-9241 mailto:sch-sch@mail.ru http://zoobank.org/urn:lsid:zoobank.org:author:c9cc4a86-bd1f-499a-b831-f727a111bbaa https://doi.org/10.5852/ejt.2021.775.1539 2 european journal of taxonomy 775: 1–14 (2021) differs from them by the monochromous dark colouration of its surface which lacks yellow-orange spots. further study of the male has revealed characters previously unknown in any members of the tribe apalochrini mulsant & rey, 1867. the metathorax is swollen and impressed in the middle, with two lamellate appendages located below, near distal edge, and above, closer to mesothorax. the metathoracic appendages allow placing this species near dromanthomorphus pic, 1921, but species of this genus have only one appendage at the base of metathorax which is not lamellate and a different shape, and they also have swollen intermediate tibiae. these characters show that the species found in afghanistan is different from dromanthomorphus pic. another genus of the tribe with a thorn-like metathoracic appendage, mimapalochrus tshernyshev, 2015, occurs in se asia and is distinguishable by the elytral apices impressed near the suture, swollen anterior and an intermediate femora, simple not emarginate anterior tibiae slightly widened posteriorly, the presence of pheromone glands on intermediate tibiae, and extremely large eyes. hence, the newly found species does not belong to any known genera of the tribe apalochrini and represents a new genus and new species. the tribe apalochrini unites soft-winged flower beetles with typical antennae possessing a second antennomere which is extremely small, round or oval, usually almost completely hidden by the scape, so that the antennae appear to be 10-segmented. the highest species diversity of the tribe occurs in se asia, africa and australia where species have a wide spectrum of male characters of importance to differentiate taxa. the taxonomic structure of the tribe has been recently discussed with special reference to southern asian genera, and, a number of recent articles devoted to, or pertaining to, the malachiidae fleming, 1821 fauna of the himalayas and an area of south-east asia and australia have been published (yoshitomi 2008, 2010, 2014; asano & kojima 2009, 2013, 2014; tshernyshev 2009, 2012b, 2015a–e, 2016a–d, 2018, 2020a–c; asano & kawashima 2010; yoshitomi & lee 2010; asano & yoshitomi 2011; asano et al. 2011a, 2011b, 2018; geiser 2011; asano 2013a, 2013b, 2015, 2017, 2018; asano & okajima 2013; plonski 2013, 2014a, 2014b, 2015, 2016, 2017; plonski & geiser 2014; plonski & michael 2014; plonski & puchner 2014; constantin 2015; liu et al. 2015, 2016, 2017; yoshitomi et al. 2015; tshernyshev & kopetz 2018; tshernyshev & shcherbakov 2020); however, further revision is still necessary to clarify our knowledge of the taxonomy. amongst those genera currently known in the tribe aplochrini, four generic groups are considered, namely apalochrus-group (apalochrus erichson, 1840, paratinoides l. medvedev, 1964, protapalochrus evers, 1987, pectapalochrus tshernyshev, 2016, oculapalochrus tshernyshev, 2015), collops-group (collops erichson, 1840, protocollops evers, 1991, troglocollops wittmer, 1965, simoderus abeille de perrin, 1891), laius-group (laius guérin-méneville, 1831, intybia pascoe, 1866, troglointybia tshernyshev, 2015, dicranolaius champion, 1921, eulaius wittmer, 1996), and dromanthomorphusgroup (dromanthomorphus pic, 1921, hadrocnemus kraatz, 1895, mimapalochrus tshernyshev, 2015). representatives of the four groups can be divided into two sections based on the modification of the basal antennomeres in the male. a key to the genera of the tribe apalochrini with simple unmodified 1st, 3rd and/or 4th antennomeres, the apalochrus-section, is given below. the species from afghanistan discussed above belongs to this section; the key to the other section is given in another article, currently awaiting publication. however, the complex of male special characters for the new afghanistan species necessitates a new genus, pectotibialis tshernyshev gen. nov., as described below, with pectotibialis paghmanensis tshernyshev sp. nov. selected as the type species. material and methods for descriptions, special male structures and genitalia were studied; ‛special male structures’ refer to the sculptured head and pronotum, swollen, impressed and appendiculate metathorax, and excavate anterior tibiae. illustrations for the species were prepared using specimens from the type locality: 3 tshernyshev s.e., pectotibialis paghmanensis tshernyshev gen. et sp. nov. from afghanistan pectotibialis paghmanensis tshernyshev sp. nov. holotype, male and paratype, female — afghanistan, kabul province, near paghman village. the specimens are currently deposited in the following institution, which is subsequently referred to by the acronym: sch_isea = the author’s collection, which is housed in the institute of animal systematics and ecology, siberian branch of the russian academy of sciences, novosibirsk, russia. the beetles were studied using an amscope trinocular stereomicroscope (ultimate trinocular zoom microscope 6.7x-90x model zm-2ty), and digital photographs were taken using a carl zeiss stemi 2000 trinocular microscope and the axiovision programme. male genitalia, embedded in dmhf (dimethyl hydantoin formaldehyde), were mounted onto a transparent card and pinned under the specimen. results taxonomy class insecta linnaeus, 1758 order coleoptera linnaeus, 1758 suborder polyphaga emery,1886 superfamily cleroidea latreille, 1802 family malachiidae fleming, 1821 subfamily malachiinae fleming, 1821 tribe apalochrini mulsant & rey, 1867 pectotibialis tshernyshev gen. nov. urn:lsid:zoobank.org:act:d03dbce5-498e-4993-97ef-c0f6b741d9e4 type species pectotibialis paghmanensis tshernyshev sp. nov. fixed by monotypy in the present paper. diagnosis moderately small sized (ca 3.8–4.1 mm) soft-winged flower beetles with the body elongate, narrow, parallel-sided, slightly expanded posteriorly, with two or three apical ventrites of the abdomen not covered by elytra. metallic green upperside with yellow or yellow-brown parts, i.e., labrum, labium, mandibles, palpi, mouthparts and antennae, tibiae and tarsi in all legs, coxae and trochanters of anterior legs and areas of joining between them in intermediate and posterior legs. antennae weakly flabellate, not long, expanding over the base of elytra, with the 3rd antennomere wide triangular. head small, narrower than pronotum, interocular area slightly impressed and sculptured, frons tuberculate, eyes simple, not large, slightly protruding. pronotum almost completely equilateral, narrowly transverse, impressed near basal margin, disk convex with wide subtriangular tubercle in the middle and looking bituberculate, basal side and basal half of lateral sides slightly flattened and distinctly marginate, and anterior side indistinctly marginate. elytra parallel or slightly widened just below the middle, not covering two or three apical segments of the abdomen, coarsely punctured, with distinct and weakly protruding shoulders; apices evenly rounded, simple. hind wings normally developed (in both sexes). anterior tarsi with distinct transverse small comb above the 2nd tarsomere and wide triangular 1st tarsomere; claws very short and hidden by round transparent plates at base. anterior tibiae strongly excavate dorsally in apical half with elongate tip looking like initial tarsomere, not swollen; intermediate and posterior tibiae simple, not http://zoobank.org/urn:lsid:zoobank.org:act:d03dbce5-498e-4993-97ef-c0f6b741d9e4 4 european journal of taxonomy 775: 1–14 (2021) swollen or emarginate, with the setae similar to that in tarsal comb arranged in apical edge near tarsus. femora wide, roundly swollen, anterior with a dent in middle. anterior trochanters oval and impressed to the basal half of the femur. metathorax swollen and impressed in the middle with two appendages, one lamellate transverse above, and the other longitudinal with a vertical round semi-transparent plate at apex. pygidium elongate, equilateral, evenly rounded distally, ultimate abdominal ventrite bilaciniate, narrow, transverse, evenly narrowed dorsally and emarginate in the middle, aedeagus simple, weakly curved dorsally, narrow, with elongate narrow apical lamella, endophallus with a few denticles at apex of inner sac, tegmen long, narrow, with thin, short pointed parameres. comparison special male characters differentiating pectotibialis tshernyshev gen. nov. from all other members in the tribe apalochrini are given in the key below. the new genus is similar to the genera dromanthomorphus pic and oculapalochrus tshernyshev in the presence of appendages on the metathorax and tarsal comb above the second tarsomere in the anterior legs of the male, but typical characters that differentiate it are as follows: distinguished from dromanthomorphus pic by simple, not widened intermediate tibiae and from oculapalochrus tshernyshev by simple eyes; the strongly excavate anterior tibiae in combination with the abbreviated elytra of the new genus are different from both genera. notes this species and genus are known only from afghanistan, central asia. list of species in the genus pectotibialis: 1. p. paghmanensis tshernyshev gen. et sp. nov. (afghanistan: paghman). pectotibialis paghmanensis tshernyshev sp. nov. urn:lsid:zoobank.org:act:93ec6ffd-b1d1-409f-b12a-081e1d5b96f5 figs 1–2 material examined holotype afghanistan • ♂; kabul province, near paghman village, 34°35′ n, 68°57′ e, 2600 m a.s.l.; 12 jul. 2009; o. pak leg.; sch_isea_000123. paratypes afghanistan • 6 ♀; same collection data as for holotype; sch_isea_000124–129. description holotype, male (fig 1a–b) body. elongate, subparallel, slightly expanded posteriorly, elytra not covering two ultimate segments of the abdomen. labrum, labium, mandibles, palpi, mouthparts and antennae. yellow-brown, except for black spots on upper side of the 1st antennomere (fig. 2d), tibiae and tarsi in all legs, coxae and trochanters in anterior legs and commissure parts between them in intermediate and posterior legs and membranes of abdominal ventrites yellow, other body parts black–brown; upper surface of head, pronotum and elytra with a weak green metallic tint. surface evenly covered with yellow-white dense fine long erect and curved hairs. vesicles white-yellow, and thoracic mesepimera black. http://zoobank.org/urn:lsid:zoobank.org:act:93ec6ffd-b1d1-409f-b12a-081e1d5b96f5 5 tshernyshev s.e., pectotibialis paghmanensis tshernyshev gen. et sp. nov. from afghanistan fig. 1. pectotibialis paghmanensis tshernyshev gen. et sp. nov. a, b, d–i. holotype, ♂ (sch_ isea_000123). c. allotype, ♀. a, c. external appearance, dorsal view. b. external appearance, lateral view. d. right antenna. e. head, pronotum and scutellum, subdorsal view. f. metathorax, subdorsal view. g. metathorax, lateral view. h. left anterior tibia and tarsus, ventral view. i. left anterior tibia and tarsus, dorsal view. scale bars: 0.5 mm. 6 european journal of taxonomy 775: 1–14 (2021) head. not wider than pronotum, interocular area slightly impressed and edged with thin carina, frons with two tubercles, eyes small, oval, weakly protruding; genae short and straight; clypeus narrow, transverse, straight; labrum narrow, transverse; palpae simple with apical segment slightly widened and securiform; surface of head dull, sparsely indistinctly punctured lacking microsculpture, evenly covered with light fine long and curved pubescence. antennae. weakly flabellate from 6th to 11th antennomeres (fig. 1d), 1.7 mm long, reaching the base of elytra; 1st antennomere enlarged, round-oval, 2nd antennomere small, round, almost completely invisible, hidden by the 1st antennomere, 3rd and 4th antennomeres triangular, the 3rd is 1.2 times as long and wide as the 4th antennomere, 5th antennomere triangular with elongate outer edge and is as long as the 3rd antennomere, but considerably narrower than it, 6th to 10th antennomeres with extended outer edges, apical segment elongate, thin, subcylindrical, slightly curved in basal third; surface evenly covered with short, yellowwhite adpressed pubescence with single erect longer light hairs on outer sides of the antennomeres. pronotum. almost completely equilateral, narrowly transverse impressed near basal margin, disk convex with wide subtriangular tubercle in a middle and look bituberculate at base; anterior margin slightly convex; posterior margin straight; lateral sides almost completely straight with rounded angles; basal side and basal half of lateral sides slightly flattened and distinctly marginate, anterior side indistinctly marginate; surface glabrous, with sparse smoothed punctures lacking microsculpture, evenly covered with fine long curved light hairs. scutellum. small, triangular with rounded tip, almost completely covered by pronotum, sparsely punctured and covered with light semi-erect hairs. elytra. oblong, parallel or slightly widened just behind the middle and evenly rounded distally, not covering three apical segments of the abdomen, at base not wider than pronotum; humeri small, slightly protruding; apices evenly rounded, simple; disc indistinctly marginate on external sides and along suture, lateral edge in middle near epipleura slightly impressed; surface dull, coarsely punctured and wrinkled, with smoothed microsculpture, evenly covered with yellow-white long erect hairs. hind wings. normally developed. legs. not long; posterior femora not reaching elytral apices; anterior tibiae not swollen but widened distally and with a strong rectangular excavation inwardly in apical half so that the tip of the tibiae looks like a 1st tarsomere (fig. 1h–i), intermediate and posterior tibiae simple, not swollen or emarginate, with the setae similar to that in tarsal comb arranged in apical edge near tarsus, femora wide, roundly swollen, anterior with tooth in middle (fig. 2a–b), anterior trochanter oval and impressed to the basal half of the femur, all tarsi 5-segmented, slightly compressed and elongate, 2nd tarsomere in anterior tarsi with small transverse comb above, the 1st tarsomere enlarged, triangular, 1.3 times as long as the 2nd tarsomere, 2 times as long as the 3rd tarsomere and 3 times as long as the 4th tarsomere, claw-segment flattened, as long as the 1st tarsomere, claws very short and thin, almost completely hidden by round semi-transparent membrane at base. surface of legs covered with light short semi-erect pubescence and sparse white long erect hairs. ventral body surface. sparsely punctured, shining, evenly and sparsely covered with long, fine, semierect white hairs; metathorax swollen and complicatedly sculptured: impressed in a middle, and with two protuberances, one lamellate transverse above, and the other longitudinal and possessing vertical round semi-transparent plate at apex (fig. 1f–g). pygidium slightly elongate, almost equilateral, evenly rounded distally, with long dark erect hair on outer side (fig. 2c), ultimate abdominal ventrite is bilaciniate, narrow, transverse, evenly narrowed dorsally and emarginate in the middle (fig. 2d), aedeagus simple, weakly curved dorsally, narrow, with elongate narrow and ventrally curved apical lamella, endophallus with three 7 tshernyshev s.e., pectotibialis paghmanensis tshernyshev gen. et sp. nov. from afghanistan denticles at apex near lamella (fig. 2f), tegmen long, narrow, with thin and short pointed parameres those are 2.3 times as short as the base of tegmen (fig. 2g). length. 3.8 mm, width (at elytral base) 0.8 mm. female differs in having simple head and pronotum lacking sculpture, anterior tarsi simple, lacking a comb above the 2nd tarsomere, antennae dentate with the 3rd and 4th antennomeres narrow-triangular, and of the equal shape and length, anterior tibiae lacking excavation, but with stretched outer edge bearing the 1st tarsomere, anterior femora lacking tooth, metathorax evenly convex lacking appendages, inner side of the apex of anterior and intermediate tibiae with a row of setae similar to that in tarsal comb of male, abdomen long, with four segments extending beyond apex of the elytra. length. 4.3 mm, width (at elytral base) 1.0 mm. fig. 2. pectotibialis paghmanensis tshernyshev gen. et sp. nov., holotype, ♂. a. femur of left anterior leg. b. left anterior leg, ventral view. c. left intermediate leg, ventral view. d. pygidium. e. ultimate abdominal ventrite. f. aedeagus, dorsal view. g. tegmen. h. distribution map. scale bars: 0.5 mm. 8 european journal of taxonomy 775: 1–14 (2021) distribution known from type locality only, afghanistan: paghman environs (fig. 2h). key to the genera of the tribe apalochrini with simple not modified 1st, 3rd and 4th antennomeres 1. male special structures are indistinct, antennae filiform or flabellate .....................................................2 – male special structures distinct and located on head, antennae, pronotum, metathorax, sternites or legs ..........................................................................................................................................................10 2. second segment of anterior tarsi lacking a comb above, elytra with green or blue metallic lustre .............................................................................................................. apalochrus erichson, 1840 – second segment of anterior tarsi with a comb above, elytra entirely black, sometimes with metallic lustre ...........................................................................................................................................................3 3. body of staphylinoid shape, narrow, black, lacking metallic lustre, elytra strongly shortened and not covering abdomen, wings reduced, antennae filiform ............................. paratinoides medvedev, 1964 – body typical of malachiidae, elytra completely covering the abdomen .................................................4 4. anterior tibiae simple, not widened or impressed, antennae flabellate of filiform .................................5 – anterior tibiae widened or impressed distally, antennae flabellate .........................................................9 5. antennae filiform ......................................................................................................................................6 – antennae flabellate ...................................................................................................................................7 6. antennomeres simple, not widened and slightly flattened, lacking impressions or grooves, anterior femora simple, lacking emarginations; surface of the body covered with long erect and short adpressed black pubescence ......................................................................................... protapalochrus evers, 1987 – intermediate 3rd and 6th antennomeres widened, dorsal side of 5th–10th antennomeres sulcate in middle, anterior femora with oval emargination in apical half covered with white pubescence; surface of the body covered with short semi-erect and adpressed black-brown pubescence ...... acroapalochrus tshernyshev, 2020 7. pronotum simple, lacking longitudinal grooves .............................. pectapalochrus tshernyshev, 2016 – pronotum bisulcate in middle ........................................................................... airomalachius pic, 1950 9. anterior tibiae compressed and widened in apical half, intermediate tibiae impressed and flattened in outer side with weakly widened and curved apices, pronotum convex and narrowly depressed at base, elytra and pronotum uniformly dark with green or blue metallic lustre and covered with long erect grey hairs ................................................................................................... mesapalochrus tshernyshev, 2020 – anterior tibiae slightly widened anteriorly and weakly impressed dorsally, intermediate tibiae simple, not widened and not flattened, pronotum convex, not depressed at base, orange with longitudinal greenbronze stripe in a middle, elytra orange-yellow, each with two bronze metallic oval-triangular spots, surface covered with short erect fine white pubescence ................................nudopectinus evers, 1987 10. anterior tibiae simple, not widened or emarginate, male special structure is located in apical half of the pronotum as transverse fissure with elevate side and protuberance behind, anterior tarsi with a comb above the 2nd tarsomere, posterior tibiae slightly but distinctly swollen in basal fourth .................................................................................................. simoderus abeille de perrin, 1891 – anterior tibiae slightly but distinctly widened, simple or emarginate, pronotum simple or with protruding process in anterior side, male special structures located in femora or tibiae, head, metathorax or abdominal ventrites; anterior tarsi with a comb above 2nd tarsomere or lacking it ..........................11 11. anterior tibiae simple, not emarginate, anterior tarsi with a comb above 2nd tarsomere ......................15 – anterior tibiae emarginate at apical part, anterior tarsi with a comb above 2nd tarsomere or lacking it ...............................................................................................................................................................12 12. head with emarginations and protuberances, antennae serrate or flabellate, abdomen simple ............13 – head simple, lacking emarginations and protuberances, antennae filiform ..........................................14 13. pronotum with narrow horn-like process protruding over the head, antennae serrate .............................................................................................................. troglolaius wittmer, 1952 – pronotum simple, antennae flabellate ......................................................... trogloapalochrus pic, 1919 9 tshernyshev s.e., pectotibialis paghmanensis tshernyshev gen. et sp. nov. from afghanistan 14. 4th and 5th abdominal ventrites modified and with thorn-like process in the middle ............................15 – 4th and 5th abdominal ventrites simple, lacking process .........................................................................16 15. intermediate tibiae slightly widened and swollen, with the inner side emarginate at apical quarter, trochanters of anterior legs simple .......................................................... opisthapalochrus evers, 1987 – intermediate tibiae simple, not widened or emarginate, trochanters of anterior legs dentate ..............................................................................................................spinapalochrus pic, 1919 16. 2nd tarsomere in anterior legs with a comb above ..................................................................................17 – 2nd tarsomere in anterior legs lacking a comb above, antennae serrate, anterior tibiae evenly emarginate, intermediate tibiae widened and narrowly emarginate, with pheromone glands .................................................................. mesopezus jacobson, 1911 (= epitinus evers, 1987) 17. eyes extremely large, antennae flabellate ..............................................................................................18 – eyes simple, not large .............................................................................................................................19 18. metathorax swollen with appendage in the middle, elytral apices impressed near suture, anterior and intermediate femora swollen, intermediate tibia slightly widened posteriorly and possessing pheromone glands ............................................................................................... mimapalochrus tshernyshev, 2015 – metathorax weakly swollen, lacking appendage in a middle, elytral apices not impressed, simple, all legs simple, not widened or curved, without glands ....................... oculapalochrus tshernyshev, 2015 19. metathorax strongly swollen and possessing gladiate or rudimentary appendage with hairs in the middle ......................................................................................................................................................20 – metathorax slightly swollen, lacking appendage or hairs .....................................................................18 20. anterior tibiae slightly emarginate, intermediate tibiae swollen with apical margin simple and lacking setae, pronotum and head simple, lacking relief sculpture, metathorax possesses gladiate appendage in the middle directed to intermediate coxae, elytra completely covering the abdomen .........................................................dromanthomorphus pic, 1921 (= pectinus evers, 1987) – anterior tibiae deeply emarginate in apical quarter, apical margin of anterior and intermediate legs surrounded with pectinate setae similar to that in tarsal comb; intermediate tibiae slightly widened, not stout, with longitudinal groove within, pronotum with protuberance in basal half, forehead bituberculate, metathorax slightly swollen and impressed in the middle with two appendages, one lamellate transverse above, and the other appendage longitudinal with vertical round semi-transparent lamellate apex below, elytra not covering two apical segments of the abdomen ................pectotibialis tshernyshev gen. nov. 21. anterior tibiae strongly modified, deeply emarginated with lamellate processes, inner spurs of the anterior legs modified, antenna flabellate or filiform .............................................................................22 – anterior tibiae slightly widened or weakly emarginate, simple, spurs short and simple, antenna filiform .....................................................................................................................................................23 22. 1st tarsomere in anterior legs enlarged and compressed, leaf-shaped, dorsally with a row of golden hairs forming a tuft, 2nd tarsomere very short, almost completely covered by the vane-shape longitudinal comb, anterior tibiae strongly emarginate, with a stretched apex bearing enlarged hook-like spur, intermediate tibiae strongly widened and modified, emarginate within and possessing a row of long hairs inside the emargination, antennae flabellate .................................hapalochrops bourgeois, 1908 – 1st tarsomere in anterior legs slightly enlarged and swollen, round-quadrate, 2nd tarsomere with a simple short comb above, outer spur in anterior tibiae long and strong, thorn-like, intermediate tibiae with deep emargination in apical quarter, and a long narrow appendage bearing hair tuft in apex of inner side of the emargination, antennae filiform ........................................................tibipectinus tshernyshev, 2020 23. anterior tibiae weakly widened distally, not emarginate, intermediate tibiae widened in apical half and sulcate in the middle of inner side, lacking emarginations .................protopectinus tshernyshev, 2020 – anterior tibiae weakly emarginate in dorsal or ventral side, intermediate tibiae stout and emarginate or thin and not emarginate ..........................................................................................................................24 24. anterior tibiae emarginate in dorsal side, intermediate tibiae strongly dilate and convex, with triangular perpendicular process in a middle of inner side and strongly emarginate beneath it .....................................................................................................................hadrocnemus kraatz, 1895 10 european journal of taxonomy 775: 1–14 (2021) – anterior tibiae emarginate in ventral side, intermediate tibiae not widened and with a row of long erect thin hairs on inner side ................................................................................... setapalochrus evers, 1988 discussion 38 genera are currently listed in the tribe apalochrini worldwide, of which 23 genera, acroapalochrus tshernyshev, 2020, airomalachius pic, 1950, apalochrus erichson, 1840, dromanthomorphus pic, 1921 (= pectinus evers, 1987), hadrocnemus kraatz, 1895, hapalochrops bourgeois, 1908, mesapalochrus tshernyshev, 2020, mesopezus jacobson, 1911 (= epitinus evers, 1987) (tshernyshev, 2003), mimapalochrus tshernyshev, 2015, nudopectinus evers, 1987, oculapalochrus tshernyshev, 2015, opisthapalochrus evers, 1987, paratinoides medvedev, 1964, pectapalochrus tshernyshev, 2016, pectotibialis tshernyshev gen. nov., protapalochrus evers, 1987, protopectinus tshernyshev, 2020, setapalochrus evers, 1988, simoderus abeille de perrin, 1891, spinapalochrus pic, 1919, trogloapalochrus pic, 1919, troglolaius wittmer, 1952 and tibipectinus tshernyshev, 2020, are belonging to the group “with simple not modified 1st, 3rd and 4th antennomeres in male” and 15 genera, collops erichson, 1840, dicranolaius champion, 1921, eulaius wittmer, 1996, flabellolaius wittmer, 1952, heterolaius champion, 1920, intybia pascoe, 1866, laius guérin-méneville, 1831 (= nossibeus evers, 1994), myrmecospectra motschulsky, 1858 (= myrmecophasma bourgeois, 1885), notointybia liu, ślipiński & pang, 2020, protocollops evers, 1991, stenolaius wittmer, 1995, syndesmolaius evers, 1986, zelotypus abeille de perrin, 1900, troglocollops wittmer, 1965 and troglointybia tshernyshev, 2015, belong to the group “with specific male structures located on 1st, 3rd and/or 4th antennomeres”. representatives of the first group are typical residents of eurasia, indo-china, south east asia, australia and africa with high species diversity in southern regions (tshernyshev 2011, 2012a). species of the second group are also occurring in these regions, but most diverse in australia and also represented in american continent. due to the absence of their fragments in fossil remains, the tribe apalochrini can be considered as the youngest in the family malachiidae, and, probably originated from the tribe illopini, that is also similar to malachiini due to the small size of the second antennomere. the wide species distribution, from africa to arctic region of eurasia, allows a high evolutionary potential of the tribe in adaptation to different type of habitats to be assumed. the tribe is at present being actively studied, and the description of several new species from southern regions is to be expected in the nearest future. the present generic revision can help in the understanding of the taxonomic diversity of the family malachiidae and the remarkable tribe apalochrini. acknowledgments the author is grateful to dr roman yakovlev (barnaul state university, russia) for his help in obtaining the malachiidae, to mr oleg pak (donetsk, ukraine) for collecting soft-winged flower beetles in afghanistan, to isidor plonski (vienna, austria) for help regarding literature sources and discussion of malachiidae taxonomy, and to prof. mark seaward (bradford university, uk.) for advice and linguistic revision of the text. this study was supported by the russian foundation for basic research (grant no. 19-04-00465-a), as well as the program of basic scientific research (fni) of the state academies of sciences, project no. 0247-2021-0004. references asano m. 2013a. early instar larvae of intybia niponicus (lewis) (coleoptera, malachiidae) and comparison with a clerid lst instar: the foetomorphic larva in malachiidae, ii. japanese journal of systematic entomology 19 (1): 21–27. asano m. 2013b. new combinations and redescriptions of six species of the tribe ebaeini portevin (coleoptera, malachiidae) in japan. japanese journal of systematic entomology 19 (2): 275–284. 11 tshernyshev s.e., pectotibialis paghmanensis tshernyshev gen. et sp. nov. from afghanistan asano m. 2015. taxonomic notes on some malachiid beetles of taiwan, with description of a new species of the genus intybia pascoe (coleoptera: malachiidae). japanese journal of systematic entomology 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(2022) performed a five-loci phylogenetic analysis (nrits, nrlsu, rpb1, rpb2 and mtssu) of the lecideaceae chevall. in order to test the monophyly of immersaria. the analysis showed two clades within immersaria: one with species with lecideine apothecia, and another with species with lecanorine apothecia. they excluded the lecanorine species from immersaria and proposed a new genus, lecaimmeria c.m.xie, lu l.zhang & li s.wang to accommodate them (xie et al. 2022). lecaimmeria could be distinguished from related genera by its waxy glossy, orange or red-brown thallus with an amyloid medulla, its immersed apothecia with a crypto-thalline margin, its orange epihymenium with an epinecral layer, and its porpidia-type asci with eight halonate, non-amyloid ascospores. species of the genus frequently grow on granite or sandstone, with the exception of one species, l. tuberculosa c.m.xie & xin y.wang, which grows on jade. the genus is represented by 10 species, distributed in alpine areas, high-latitude steppe or high-altitude dessert-steppe areas (xie et al. 2022). neither of the two genera had been recorded from pakistan previously. https://doi.org/10.5852/ejt.2022.834.1901 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ mailto:kamranhabiib%40gmail.com?subject= mailto:rizwanamughal6%40gmail.com?subject= mailto:drankhalid%40gmail.com?subject= https://doi.org/10.5852/ejt.2022.834.1901 habib k. et al., lecaimmeria pakistanica sp. nov. from azad jammu and kashmir, pakistan 95 during our study on the lichen biota of the state of azad jammu and kashmir, pakistan, a novel species of lecaimmeria was discovered. we present a brief diagnosis, an extensive description, illustrations, and a phylogenetic analysis based on its-sequence data. material and methods collection and preservation specimens were collected during surveys in the state of azad jammu and kashmir, pakistan in 2019. the specimens have been deposited in the lah herbarium of the institute of botany, university of the punjab, lahore. acronyms of herbaria follow index herbariorum (thiers continuously updated). morphological and chemical characterization macroand micromorphology of the specimens was examined under a stereo microscope (meiji techno, emz-5tr, japan) and a compound microscope (swift m4000-d). thalline chemistry was analyzed by spot tests, using 10% potassium hydroxide (k) and calcium hypochlorite (c), and thin layer chromatography (tlc) (solvent c) according to the method proposed by orange et al. (2001). anatomical characterization and measurement of anatomical features were done by preparing and observing hand-cut apothecial sections mounted in water, k and lugol’s solution (iki) under the compound microscope. molecular characterization and phylogenetic analysis genomic dna was extracted directly from a portion of the thallus with apothecia from each specimen using a modified 2% ctab method (gardes & bruns 1993). the its nrdna region was amplified using the primer pair its1f, as forward primer (5’cttggtcatttagaggaagtaa3’) (gardes & bruns 1993) and its4, as reverse primer (5’tcctccgcttattgatatgc3’) (white et al. 1990), following the amplification protocol of khan et al. (2018). the amplified dna fragments (pcr products) were visualized with the help of 1% agarose gel using ethidium bromide and a gel documentation system (sambrook & russel 2001). the amplified products were then sequenced from tsingke biotech company beijing, china, and the sequences deposited in genbank. phylogenetic analysis bidirectional sequences (its1 and its4) were assembled by using bioedit ver. 7.2.5 (hall 2005). comparative its sequences for the analysis were identified and retrieved from genbank using the basic local alignment search tool (blast) (altschul et al. 1990), evaluating maximum percent identification and query coverage. the two newly generated sequences were aligned with 18 sequences retrieved from genbank (table 1) using mafft ver. 7 (katoh et al. 2019). all sequences were trimmed terminally using bioedit ver. 7.2.5. the phylogenetic tree was constructed by mega x (kumar et al. 2018) using the maximum likelihood (ml) method. the optimal model for nucleotide sequences was estimated by mega x (kumar et al. 2018). the kimura 2-parameter (k2p) was found to be the best model for a phylogenetic tree construction. bellemerea alpina (sommerf.) clauzade & cl.roux and koerberiella wimmeriana (körb.) stein were chosen as an outgroup. results phylogenetic analysis the data matrix had 521 unambiguously aligned nucleotide positions of which 347 were conserved, 172 variables, 117 parsimony-informative and 54 were singletons. the new its nrdna sequences nested within the phylogenetic branch of newly proposed genus lecaimmeria (fig. 1). the sequences of our new species formed a well-supported (bs 92) separate clade outside a group comprised of l. orbicularis european journal of taxonomy 834: 94–101 (2022) 96 c.m.xie & lu l.zhang, l. lygaea c.m.xie & lu l.zhang and l. tibetica c.m.xie & xin y.wang, demonstrating its status as an independent species. taxonomic treatment kingdom fungi (l.) r.t.moore subkingdom dikarya hibbett, t.y.james & vilgalys division ascomycota (berk.) caval.sm. subdivision pezizomycotina o.e.erikss. & winka class lecanoromycetes o.e.erikss. & winka subclass lecanoromycetidae p.m.kirk, p.f.cannon, j.c.david & stalpers order lecideales vain. family lecideaceae chevall. genus lecaimmeria c.m.xie, lu l.zhang & li s.wang lecaimmeria pakistanica k.habib, r.zulfiqar & khalid sp. nov. mb844738 fig. 2 diagnosis distinguished from all the known species of the genus by having large ascospores (20–32 × 10–16 μm), and relatively taller hymenium. all the other species of the genus have ascospore dimensions in the range of 14–22 × 5–14 μm. also separated from other species of the genus by its nrdna sequence data. species genbank accession no. voucher no. country lecaimmeria botryoides mz227405 kun 20-66713 china lecaimmeria botryoides mz227406 kun 20-66721a china immersaria sp. mf149862 malicek 7717 macedonia lecaimmeria iranica kr061347 sdnu 20117663 china lecaimmeria iranica kr061348 sdnu 20117623 china lecaimmeria lygaea mz227458 kun 20-69054 china lecaimmeria mongolica mz227397 sdnu 20117613 china lecaimmeria mongolica mz227398 sdnu 20117399 china lecaimmeria orbicularis mz227415 kun 20-66803 china lecaimmeria orbicularis mz227414 kun 20-66801 china lecaimmeria pakistanica sp. nov. mw508503 lah-36674 pakistan lecaimmeria pakistanica sp. nov. mw508504 lah-36675 pakistan lecaimmeria qinghaiensis mz227454 kun 20-68696 china lecaimmeria qinghaiensis mz227455 kun 20-68698 china lecaimmeria tibetica mz227474 kun xy19-1288i china lecaimmeria tibetica mz227475 kun xy19-1288a china lecaimmeria tuberculosa mz227476 kun 18-58856 china lecaimmeria tuberculosa mz227477 kun 18-58857 china bellemerea alpina af332117 1999, hafellner 46531 (gzu) austria koerberiella wimmeriana mk812168 o-l-163472 norway table 1. species used in the phylogenetic analysis. pakistani collections are marked in bold. https://www.mycobank.org/mb/mb844738 habib k. et al., lecaimmeria pakistanica sp. nov. from azad jammu and kashmir, pakistan 97 etymology the specific epithet ʻpakistanicaʼ refers to country in which the new species was discovered. material examined holotype pakistan • azad jammu and kashmir, muzaffarabad, peer chinasi; 34°23′ n, 73°32′ e; alt. 2924 m; on rocks; 9 aug. 2018; t. saifullah and k. habib leg.; pc-21; lah[lah-36674]; genbank no.: mw508503. paratype pakistan • azad jammu and kashmir, muzaffarabad, peer chinasi; 34°23′ n, 73°32′ e; alt. 2700 m, on rocks; 22 jul. 2019; t. saifullah and k. habib leg.; pc-22; lah[lah-36675]; genbank no.: mw508504. description thallus crustose, areolate, up to 6 cm wide, in section 200–280 μm thick, upper surface yellow-brown to brown, no change when wet. areoles separate, flat to weakly convex, irregular to angular, slightly fig. 1. molecular phylogenetic analysis of lecaimmeria pakistanica k.habib, r.zulfiqar & khalid sp. nov. by the maximum likelihood method based on nrdna sequences, including its1, 5.8s and its2. numbers below branch node represent ml bootstrap (> 50%) based on 1000 replicates. sequences generated from pakistani collections are marked with black circle. european journal of taxonomy 834: 94–101 (2022) 98 fig. 2. lecaimmeria pakistanica k.habib, r.zulfiqar & khalid sp. nov. a. areolate thallus. b. wet thallus. c. apothecia and areoles. d. section of areole. e. section of apothecium. f. asci. g. ascospores. habib k. et al., lecaimmeria pakistanica sp. nov. from azad jammu and kashmir, pakistan 99 pruinose near margin, glossy, adnate, without fissures, marginal areoles slightly larger, up to 1.5 mm across, up to 0.6 mm thick, rarely with whitish margins. prothallus visible between areoles, blackish. cortex two layered, ca 40–60 μm thick, paraplectenchymatous, cells 8–12 μm in diam., upper layer paler brown, 10–16 μm thick, lower layer hyaline, 30–40 μm thick, epinecral layer distinct, up to 20 μm high. algal layer 80–120 μm thick, chlorococcoid, cells globose to subglobose, 10–20 μm in diam. medulla: hyphae white, 15–30 μm thick, corresponding with areole, iki+ blue. apothecia crypto-lecanorine, frequent at center of thallus, 1–4 per areole, immersed, sometimes not completely surrounded by the areole. disc contiguous to separate, flat to concave, reddish brown, rounded at first becoming irregular, sometimes surrounded by a white rim, up to 0.8 mm diam., thinly to rarely pruinose, margin pruinose. proper exciple thin, poorly differentiated, reduced, hyaline, 10–25 μm thick. hymenium hyaline, 130–160 μm tall including the epihymenium, which is pale brown to brown, 10– 15 μm thick, epinecral layer 3–7 μm thick; paraphyses apically branched, anastomosing, 1–2 μm wide, apically slight swollen, apices 2.5–3.5 μm wide. hypothecium 60–100 μm tall, light grayish brown, containing algal cell in the lower part. asci porpidia-type, clavate, 80–130 × 25–45 μm, amyloid wall 4–6 μm thick, 8-spored; ascospores hyaline, ellipsoid to broadly ellipsoid, mature with germ tube, 20–32 × 10–16 μm. spot tests cortex and medulla k-, c-, kc-; medulla iki+ blue; tlc none detected. ecology growing on sun-exposed rocks in a dense forest at an altitude of 2900 m. topography is mountainous in himalayan region. dominant tree species are pinus roxburghii sarg., pinus wallichiana a.b.jacks., cedrus deodara (roxb.) g.don, picea smithiana (wall.) boiss, abies pindrow royle. maximum and minimum temperature of 32°c and -8°c, respectively. annual rainfall varying between 1000–1500 mm. discussion during recent explorations of lichens from azad jammu and kashmir, pakistan, we observed specimens that could not be readily assigned to any known species. a phylogenetic analysis of the its nrdna region confirms their position within the genus lecaimmeria, and morphological data showed their distinctness from other known species of the genus. we therefore describe these specimens as a new species, lecaimmeria pakistanica sp. nov. lecaimmeria pakistanica sp. nov. is superficially similar to l. tibetica c.m.xie & xin y.wang, which was recently described from china (xie et al. 2022). the species have a similar thallus and apothecia coloration with no substance being detected by tlc, but l. pakistanica differs morphologically in having areoles up to 1.5 mm across (vs 0.3–0.5 mm), and apothecia up to 0.8 mm diam. (vs 0.25–0.5 mm). the anatomical differences between these two species include the size of ascospores and the type of paraphyses. ascospores are large and wider (20–32 × 10–16 μm) and paraphyses branched and anastomosing in l. pakistanica, whereas in l. tibetica, ascospores are small (12.5–15.0 × 5.0–6.0 μm) and paraphyses are unbranched and not anastomosing. another superficially similar taxon is l. mongolica c.m.xie & lu l.zhang, which also has the same thallus and apothecia coloration but has small areoles (0.4–0.8 mm), apothecia 0.25–0.75 mm diam., parapahyses unbranched and not anastomosing, small ascospores 10–17.5 × 6.0–7.5 μm and contains gyrophoric acid. the phylogenetically close taxon l. botryoides c.m.xie & li s.wang differs from the new taxon in having a red brown thallus, apothecia densely crowded while immature (3–6 / areolae), paraphyses only branched at the top and not anastomosing, comparatively very small ascospores 7.5–8.0 × 4.0–6.0 μm, and the presence of gyrophoric acid. european journal of taxonomy 834: 94–101 (2022) 100 the diagnostic features distinguishing l. pakistanica sp. nov. from the related species of the genus are presented in table 2. azad jammu and kashmir (ajk) is a state of pakistan that exhibits a large altitudinal variation, with climatic conditions and a diverse vegetation that supports a diverse and conspicuous lichen biota. the nature reserves have abundant biological resources, it is expected that more new species of lichen may be discovered in the azad jammu and kashmir in the future. acknowledgements we are grateful to dr gerhard rambold (university of bayreuth, germany) for his comments that greatly improved the manuscript and to dr xin yu wang (kunming institute of botany) for confirmation of the species. references altschul s.f., gish w., miller w., myers e.w. & lipman d.j. 1990. basic local alignment search tool. journal of molecular biology 215 (3): 403–410. https://doi.org/10.1016/s0022-2836(05)80360-2 calatayud v. & rambold g. 1998. two new species of the lichen genus immersaria (porpidiaceae). the lichenologist 30 (3): 231–244. https://doi.org/10.1006/lich.1997.0133 gardes m. & bruns t.d. 1993. its primers with enhanced specificity for basidiomycetes‐application to the identification of mycorrhizae and rusts. molecular ecology 2 (2): 113–118. https://doi.org/10.1111/j.1365-294x.1993.tb00005.x characters / species l. pakistanica sp. nov. l. tibetica l. mongolica l. botryoids l. lygaea l. cupreoatra thallus (colour) yellow-brown to brown orange-brown orange red-brown dark red-brown to dark brown brown areoles size (mm) up to 1.5 0.3–0.5 0.4–0.8 0.25–1.0 0.5–1.0 0.3–0.8 size (mm) and shape of apothecial disc up to 0.8 flat to concave 0.25–0.5 flat to slightly convex 0.25–0.75 flat to slightly convex 0.25–1.25 flat to concave 0.25–0.75 flat to concave 0.2–0.4 flat hymenium (µm) 130–160 105.0–137.5 62.5–82.5 67.5–100.0 (–155.0) 75.0–92.5 100–110 paraphyses branched and anastomosing unbranched and not anastomosing unbranched and not anastomosing only branched at the top, not anastomosing unbranched, not anastomosing branched and anastomosing size of ascospores (µm) 20–32 × 10–16 12.5–15.0 × 5.0–6.0 10.0–17.5 × 6.0–7.5 7.5–8.0 × 4.0–6.0 12.5–20.0 × 5.0–7.5 5–10 × 5–9 chemistry no substance detected no substance detected gyrophoric acid gyrophoric acid unknown fatty acid gyrophoric acid references this paper xie et al. 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(2011), https:// italic.units.it/ index.php table 2. comparison of l. pakistanica sp. nov. and related species of lecaimmeria c.m.xie, lu l.zhang & li s.wang. https://doi.org/10.1016/s0022-2836(05)80360-2 https://doi.org/10.1006/lich.1997.0133 https://doi.org/10.1111/j.1365-294x.1993.tb00005.x https://italic.units.it/index.php https://italic.units.it/index.php https://italic.units.it/index.php habib k. et al., lecaimmeria pakistanica sp. nov. from azad jammu and kashmir, pakistan 101 hall t.a. 2005. bioedit version 7.0.4. department of microbiology. north carolina state university. available from https://bioedit.software.informer.com [accessed 26 jul. 2022]. katoh k., rozewicki j. & yamada k.d. 2019. mafft online service: multiple sequence alignment, interactive sequence choice and visualization. briefings in bioinformatics 20 (4): 1160–1166. https://doi.org/10.1093/bib/bbx108 khan m., khalid a.n. & lumbsch h.t. 2018. a new species of lecidea (lecanorales, ascomycota) from pakistan. mycokeys 38: 25–34. https://doi.org/10.3897/mycokeys.38.26960 kumar s., stecher g., li m., knyaz c.& tamura k. 2018. mega x: molecular evolutionary genetics analysis across computing platforms. molecular biology and evolution 35 (6): 1547–1549. https://doi.org/10.1093/molbev/msy096 orange a., james p. & white f.j. 2001. microchemical methods for the identification of lichens. first ed. british lichen society, london. sambrook j. & russell d.w. 2001. detection of dna in agarose gels. in: molecular cloning. a laboratory manual (3rd ed.): 5–14. cold spring harbor laboratory press, new york. thiers b. continuously updated. index herbariorum: a global directory of public herbaria and associated staff. new york botanical garden’s virtual herbarium. available from http://sweetgum.nybg.org/science/ih/ [accessed 27 jul. 2022]. valadbeigi t., sipman h.j. & rambold g. 2011. the genus immersaria (lecideaceae) in iran, including i. iranica sp. nov. the lichenologist 43 (3): 203–208. https://doi.org/10.1017/s0024282911000077 white t.j., bruns t., lee s. & taylor j. 1990. amplification and direst sequencing of fungal ribosomal rna genes for phylogenetics. in: innis m.a., gelfand d.h., sninsky j.j. & white. j.t. (eds) pcr protocols. a guide to the methods and applications: 315–322. academic press. https://doi.org/10.1016/b978-0-12-372180-8.50042-1 xie c.m., wang l.s., zhao z.t., zhang y.y., wang x.y. & zhang l.l. 2022. revision of immersaria and a new lecanorine genus in lecideaceae (lichenised ascomycota, lecanoromycetes). mycokeys 87: 99. https://doi.org/10.21203/rs.3.rs-645064/v1 manuscript received: 24 july 2021 manuscript accepted: 11 july 2022 published on: 16 august 2022 topic editor: frederik leliaert desk editor: radka rosenbaumová printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; leibniz institute for the analysis of biodiversity change, bonn – hamburg, germany; national museum, prague, czech republic. https://bioedit.software.informer.com https://doi.org/10.1093/bib/bbx108 https://doi.org/10.3897/mycokeys.38.26960 https://doi.org/10.1093/molbev/msy096 http://sweetgum.nybg.org/science/ih/ https://doi.org/10.1017/s0024282911000077 https://doi.org/10.1016/b978-0-12-372180-8.50042-1 https://doi.org/10.21203/rs.3.rs-645064/v1 17 european journal of taxonomy 787: 17–31 issn 2118-9773 https://doi.org/10.5852/ejt.2021.787.1611 www.europeanjournaloftaxonomy.eu 2021 · lo russo v. & pastor de ward c.t. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:f386403a-8dda-4bc3-9197-a57e7aa05732 three new species of mesacanthion filipjev, 1927 (nematoda: thoracostomopsidae) from argentine coasts virginia lo russo 1,* & catalina t. pastor de ward 2 1,2 laboratorio de meiofauna marina (lameima) – instituto de diversidad y evolución austral (ideaus-conicet), blvd. alte. brown 2915, u9120acf puerto madryn, chubut, argentina. * corresponding author: lorusso@cenpat-conicet.gob.ar 2 email: pastor@cenpat-conicet.gob.ar 1 urn:lsid:zoobank.org:author:a67891f9-e1bd-407d-aa67-e2ed5c722a21 2 urn:lsid:zoobank.org:author:421da5cb-6fdd-4219-bc2c-79ae72a2e8f1 abstract. three new species of mesacanthion filipjev, 1927 were found along patagonian coasts (argentina). mesacanthion bifidum sp. nov. is characterized by short labial and cephalic setae, onchia of equal size, spicule arcuate, and gubernaculum with caudal apophysis, ending in two teeth. the species is related to m. virile (ditlevsen, 1930) de coninck & schuurmans stekhoven, 1933. however, the spicules and gubernaculum of both species are different in shape. mesacanthion longigubernaculum sp. nov. is characterized by its long and slender body, striated cuticle, relatively long cephalic and cervical setae, onchia of different sizes, amphidial fovea lentil-shaped, spicule arcuate, gubernaculum surrounding the spicule, and tail conical-cylindrical with terminal setae. mesacanthion sanantoniensis sp. nov. is characterized by its long and stout body, striated cuticle, long cephalic setae, onchia of different sizes, amphidial fovea pouch-shaped, spicule arcuate, gubernaculum with dorsal apophysis, and tail conical without terminal setae. following the key of jeong et al. (2019), the last two species are related to m. pali wieser, 1959 and m. longissimesetosum wieser, 1953, so we provide a key to differentiate the four species. keywords. free-living marine nematodes, diversity, taxonomy, patagonia, coastal area. lo russo v. & pastor de ward c.t. 2021. three new species of mesacanthion filipjev, 1927 (nematoda: thoracostomopsidae) from argentine coasts. european journal of taxonomy 787: 17–31. https://doi.org/10.5852/ejt.2021.787.1611 introduction since the middle of the 1970s, the free-living marine nematodes have been studied uninterruptedly in the argentine coasts, the southernmost continental region of south america. more than 400 species have been distinguished along the coastal area that extends over 4725 km. the purpose of the studies was not only for ecological investigations but taxonomical. on the other hand, as most of the species found were new to science, intense work has been carried out over the years to try to describe them. to https://doi.org/10.5852/ejt.2021.787.1611 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:f386403a-8dda-4bc3-9197-a57e7aa05732 https://orcid.org/0000-0001-8141-2450 mailto:lorusso%40cenpat-conicet.gob.ar?subject= mailto:pastor%40cenpat-conicet.gob.ar?subject= http://zoobank.org/urn:lsid:zoobank.org:author:a67891f9-e1bd-407d-aa67-e2ed5c722a21 http://zoobank.org/urn:lsid:zoobank.org:author:421da5cb-6fdd-4219-bc2c-79ae72a2e8f1 https://doi.org/10.5852/ejt.2021.787.1611 european journal of taxonomy 787: 17–31 (2021) 18 date, descriptions of 74 species have been published. among the species, some belonging to the genus mesacanthion filipjev, 1927 were found. the family thoracostomopsidae filipjev, 1927 consists of three subfamilies: thoracostomopsinae filipjev, 1927, trileptiinae gerlach & riemann, 1974 and enoplolaiminae de coninck, 1965. enoplolaiminae can be distinguished from the other subfamilies by their buccal cavity with three mandibles and three teeth. it has 17 genera (excluding hyptiolaimus cobb, 1930). six species of enoplolaiminae have been described in argentina as new to science. they belong to the genera enoplolaimus de man, 1893 (1), epacanthion wieser, 1953 (2), mesacanthoides wieser, 1953 (1) and parasaveljevia wieser, 1953 (2). mesacanthion belongs to enoplolaiminae. the most recent revision of mesacanthion was from jeong et al. (2019). an emended diagnosis is given as well as a list of valid species and species inquirenda and nomina nuda, a table with a comparison of diagnostic morphological characters of all species and a tabular and pictorial key to species with spicules shorter than two anal body diameters. after jeong et al. (2019) the genus mesacanthion has 39 valid species, 38 from marine environments and one from a freshwater environment. three species were originally described in south america: mesacanthion longissimesetosum wieser, 1953 (chile); m. proximum gerlach, 1957 (brazil); and m. rigens gerlach, 1957 (brazil). another five species have been recorded for the region: m. hirsutum gerlach, 1953; m. infantile (ditlevsen, 1930) de coninck & schuurmans stekhoven, 1933; m. longispiculum gerlach, 1954; m. majus (filipjev, 1927) gerlach & riemann, 1974; and m. virile (ditlevsen, 1930) de coninck & schuurmans stekhoven, 1933. in the present work, we describe the first three new species of mesacanthion for the argentine coast. material and methods description of the study sites samples were collected from three different areas along the patagonian atlantic coast, argentina (fig. 1). the northernmost site, the city of san antonio oeste (40°43′ s, 64°58′ w), is in san antonio bay, located in the northwest part of san matías gulf. about 220 kilometres to the south are two other sites, puerto madryn city (42°45′ s, 65°02′ w) and bahía kaiser (42°46′ s, 64°59′ w), both in nuevo gulf. the southernmost site, rada tilly (45°55ʹ07″ s, 67°32ʹ79″ w), is located in the middle part of san jorge gulf. sample collection and treatment samples were collected using a cylindrical plexiglass corer, fixed in 5% formaldehyde prepared in filtered seawater with rose bengal, and then the fixed nematodes were sieved through both 500 μm and 50 μm mesh sieves. the nematodes retained on the 50 μm mesh were separated from the sediment by ludox® tm and transferred to pure glycerin through a solution of ethanol : water : glycerin in 2:2:1 proportions and left at least one week in a desiccator. after that, they were mounted on glass slides sealed with canadax resin. specimen analysis morphometric data were obtained from camera lucida drawings using a zeiss standard wl microscope (d-7082 oberkochen) with differential interference contrast (dic). the measurements are given in micrometers. images were taken using a leica dm2500 microscope with incorporated camera. the literature was obtained from nemys (bezerra et al. 2021). the de man’s ratios ‘a’, ‘b’ and ‘c’ used in this paper were calculated as usual. institutional abbreviations cnp-nem = collection of nematodes of the patagonia national center (centro nacional patagónico) lo russo v. & pastor de ward c.t., new species of mesacanthion from argentina 19 macn-in = invertebrate collection of the argentine museum of natural science ‘bernardino rivadavia’ (museo argentino de ciencias naturales ‘bernardino rivadavia’) abbreviations for morphological terms a = body length / maximum body diameter abd = anal body diameter b = body length / pharynx length c = body length / tail length cʹ = tail length / body width at level of cloacal opening or anus cbd = corresponding body diameter l = total body length results phylum nematoda potts, 1932 class enoplea inglis, 1983 order enoplida filipjev, 1929 suborder enoplina chitwood & chitwood, 1937 superfamily enoploidea dujardin, 1845 family thoracostomopsidae filipjev, 1927 subfamily enoplolaiminae de coninck, 1965 genus mesacanthion filipjev, 1927 type species mesacanthion lucifer (filipjev, 1927) gerlach & riemann, 1974 diagnosis (after jeong et al. 2019) outer labial and cephalic setae situated at middle or anterior end of cephalic capsule. mandibles welldeveloped, provided with claws, arch-shaped, each consisting of two rod-like columns anteriorly united by a curved bar. teeth shorter than mandibles. spicule mostly short, unipartite and symmetrical, sometimes long, bipartite (divided by a seam: m. ditlevseni (filipjev, 1927) gerlach & riemann, 1974) and asymmetrical (anisomorphic and anisometric: m. diplechma (southern, 1914) filipjev, 1927). if long, usually gubernaculum present with caudal apophysis. marine and freshwater. mesacanthion bifidum sp. nov. urn:lsid:zoobank.org:act:62e01288-0803-428a-bb5e-d51b592d1591 figs 2, 5d etymology in reference to the proximal part of the gubernaculum that ends in two small pointed teeth, from the latin word ‘bifidum’. material examined holotype argentina • ♂; chubut, nuevo gulf, puerto madryn beach; 42°45′ s, 65°02′ w; 1 mar. 2003; c. pastor and v. lo russo leg.; high tide; fine sand sediments; cnp-nem 936. http://zoobank.org/urn:lsid:zoobank.org:act:62e01288-0803-428a-bb5e-d51b592d1591 european journal of taxonomy 787: 17–31 (2021) 20 m. bifidum sp. nov. m. longigubernaculum sp. nov. m. sanantoniensis sp. nov. character holotype ♂ ♀♀ holotype ♂ ♂♂ holotype ♂ ♂♂ number of specimens 1 3 1 2 1 2 l 2990 3363.3 (3150–3515) 1450 1445.7 (1380–1507) 2540 2466.7 (2120–2740) a 19.9 25 (22.8–26.2) 55.8 52.9 (47.8–55.8) 20.8 24.7 (20.8–29.4) b 4.9 4.9 (4.6–5.2) 4.1 4.5 (4.1–5.4) 4.2 4.1 (3.7–4.3) c 13.8 14.7 (13.8–15.7) 13.9 14.6 (13.9–15.1) 12.3 13.1 (12.3–13.9) inner labial setae length 7 8 (7–9) 8.5 8 (7.5–8.5) 7.5 7.3 (7–7.5) outer labial setae length 23 20.5 (20–21) 18 19 (16–23) 36 33 (28–36) cephalic setae length 18 13 (12–14) 13 12 (9–14) 21 20.2 (17.5–22) buccal cavity width 13 22.3 (18–27) 8 10.3 (8–13) 16 15 (14–16) buccal cavity length 18 19.3 (17–21) 12 10.7 (10–12) 16 16.7 (16–17) cephalic diameter at cephalic setae level 29 39.3 (38–41) 18 18.8 (18–20) 34 31.3 (29–34) amphid aperture width 2 1.9 (1.8–2) 3 3.5 (3–4) 2 1.8 (1.5–2) body diameter at amphid level 36 50.7 (48– 52) 21 21 (21–21) 38 36.3 (32–39) amphid width/cbd (%) 5.6 3.8 (3.5–4.2) 14.3 16.7 (14.3–19) 5.3 5.1 (3.8–6.3) cephalic capsule length 29 34 (32–36) 17 16 (14–17) 29 27.8 (26.5–29) cervical setae 10 15 (14–16) 9.5/20 13.3 (6.5–20) 9/13 10.2 (7.5– 14) nerve ring from anterior end 205 241.7 (210–265) 125 110 (92–125) 210 200 (170–220) body diameter at nerve ring level 70 89.7 (87–94) 25 26.7 (24–31) 76 67 (48–77) pharynx length 615 686.7 (680–700) 350 325.3 (256–370) 600 603.3 (570–640) body diameter at pharynx base level 96 121.7 (115–132) 24 25.7 (23–30) 102 91.3 (60–112) maximum body diameter 150 134.7 (132–138) 26 27.5 (25–31.5) 122 103 (72– 122) anal/cloacal distance from anterior end 2774 3133.7 (2922–3291) 1345.5 1346.5 (1287–1407) 2334 2277 (1967–2530) anal/cloacal body diameter 63 59.7 (57–63) 22.5 22.5 (20–25) 55 46.7 (36–55) spicule length as arc 93 – 22 22.7 (22–23) 71.5 71.2 (69–73) spicule length as cloacal diameter 1.5 – 1 1 (0.9–1.1) 1.3 1.6 (1.3–1.9) gubernaculum length 24 – 14 13.7 (10–17) 33 30.3 (27–33) gubernaculum apophysis length 21 – – – – – table 1 (continued on next page). measurements (μm) of mesacanthion bifidum sp. nov., m. longigubernaculum sp. nov. and m. sanantoniensis sp. nov., including mean values (range). lo russo v. & pastor de ward c.t., new species of mesacanthion from argentina 21 paratypes argentina • 1 ♀; same collection data as for holotype; cnp-nem 937 • 2 ♀♀; chubut, nuevo gulf, bahía kaiser; 42°46′ s, 64°59′ w; 8 m b.s.l.; oct. 1997; c. pastor leg.; cnp-nem 938 to 939. description measurements see table 1. male (holotype) large and stout body. cuticle smooth. few short (about 9–12 µm long) somatic setae scattered along body. cephalic region set-off with presence of cephalic capsule (17 µm in height). cephalic capsule with same thickness throughout its rough surface. anterior edge located at level of cephalic setae. posterior end with no real incisions forming lobes but with slightly scalloped edge. three rounded lips. each lip carries two inner slender labial setae (9 µm long). six outer labial setae (18 µm long) and four barely shorter cephalic setae (17 µm long) located at anterior of cephalic capsule, arranged in single crown. immediately posterior to cephalic capsule four short subdorsal and subventral subcephalic setae (about 4 µm) and further posteriorly (about 30 µm after cephalic capsule), arranged in four subdorsal and subventral groups of two or three, larger cervical setae present (15 µm long). amphideal aperture circular and amphideal fovea pouch-shaped, small (8% of cbd), laterally located just posterior to capsule end. metanemes not seen. funnel-shaped buccal cavity with wide opening widens at level of mandibles. its armature consists of three mandibles, each composed of two vertical rods (16 µm) united by arcuate bar (7 µm), at top ending as left and right claws. each mandible has tooth (13 µm). the three teeth equal in size. no ocellus nor pigment spots. pharynx cylindrical with irregular contours, cardia not visible. nerve ring lying at about 33% of pharynx length from anterior end. excretory-secretory system not visible. reproductive system diorchic, with opposed and outstretched testes in right position relative to intestine. one precloacal supplement, bar-shaped, located about 3 abd above cloaca. no precloacal setae present. spicules paired, arcuate, with slight manubrium (1.4 abd). gubernaculum small (26% of spicule, 0.4 abd), embracing spicule, two dorso-caudal apophyses present. the gubernaculum divided into two parts. proximal part with two small pointed teeth, surrounds tips of spicules. arched dense zone, shaped like ring, separates it from distal triangular part. tail 3.4 abd long, conical with end part cylindrical (1/5 approximately). few short caudal setae can be seen. caudal gland bodies in pre-anal region. terminal setae not present. cuticle around spinneret hardly englobed. m. bifidum sp. nov. m. longigubernaculum sp. nov. m. sanantoniensis sp. nov. character holotype ♂ ♀♀ holotype ♂ ♂♂ holotype ♂ ♂♂ gubernaculum as spicule length (%) 25.8 – 63.6 60.3 (43.5–73.9) 60 59.1 (42.5–75) precloacal organ length 34 – 5 4.7 (4–5) 21 19.3 (17–21) precloacal organ from anterior end 2520 – 1290 1317 (1233– 1428) 2310 2163.3 (1790–2390) vulva from anterior end/body length (%) – 51.5 (51.1–52) – – – – tail length 216 229.7 (224–237) 104.5 99.2 (93–104.5) 206 189.7 (153–210) c’ 3.4 3.9 (3.6–4.2) 4.6 4.4 (4–4.7) 3.7 4.1 (3.7–4.3) table 1 (continued). european journal of taxonomy 787: 17–31 (2021) 22 female similar to males in general body shape, anterior sensilla, amphideal fovea and cuticle. females longer and larger than males. lips not rounded but pointed. short somatic setae present all along body (9–12 µm) in greater quantity than in males. reproductive system didelphic amphidelphic, with two antidromously reflexed ovaries, positioned left of intestine. vulva and vagina conspicuous, with associated musculature. vulva not sclerotized. tail conical, ending in 1/5 cylindrical part, but this part less differentiated than in males. caudal gland bodies in pre-anal region. many short caudal setae. terminal setae not present. cuticle around spinneret englobed. fig. 1. map showing the study sites; map of sampling areas. a. ʻsan antonio oeste river estuaryʼ, río negro province. b. ʻpuerto madryn cityʼ and ʻbahía kaiserʼ, chubut province. c. ʻrada tillyʼ, chubut province. lo russo v. & pastor de ward c.t., new species of mesacanthion from argentina 23 fig. 2. mesacanthion bifidum sp. nov. a. cephalic sense organs on anterior end of female paratype (cnpnem 983). b. cephalic sense organs on male holotype (cnp-nem 936). c. female paratype (cnpnem 983) showing vulva and gonadal apparatus. d. precloacal supplement organ of male holotype (cnp-nem 936). e. posterior end of male holotype (cnp-nem 936). f. copulatory apparatus, spicules and gubernaculum of male holotype (cnp-nem 936). g. gonads of male holotype (cnp-nem 936). h. posterior end of female paratype (cnp-nem 983). scale bars: 1 = 20 μm; 2 = 100 μm; 3 = 200 μm. european journal of taxonomy 787: 17–31 (2021) 24 differential diagnosis mesacanthion bifidum sp. nov. is characterized by its smooth cuticle, relatively short labial and cephalic setae, onchia of equal size, amphidial fovea pouch-shaped, spicule arcuate, gubernaculum with pointed tips at the proximal end and dorso-caudal apophysis, and tail conical-cylindrical without terminal setae. following the key of jeong et al. (2019), our new species is closely related to m. virile. both species share some characteristics, such as spicules length less than 2 abd and gubernaculum with dorso-caudal apophysis and triangular shape with two parts. the male in m. bifidum sp. nov. has the outer labial setae less than 1 cephalic diameter in length, whereas in m. virile they are about 1.25 cephalic diameters. the position of the precloacal supplementary organ in m. virile is closer to the cloaca (1.1 spicule length distant from the anus) than in m. bifidum sp. nov. (2.7 spicule lengths distant from the anus) and the spicules of both species are quite different. m. virile has an l-shaped spicule with a mid-projection for muscle insertion, whereas m. bifidum sp. nov. has an arcuate spicule without projections. on the other hand, the gubernaculum also has differences in shape. although in both species the distal part is rather similar, the proximal part is rod-like in m. virile, whereas in m. bifidum sp. nov. two-pointed structures surround the spicule tip in the posterior position. mesacanthion longigubernaculum sp. nov. urn:lsid:zoobank.org:act:cbd8b1b5-e4c4-44a4-8b45-3180296c8ac3 figs 3, 5a, c etymology in reference to the length of the gubernaculum. material examined holotype argentina • ♂; chubut, san jorge gulf, rada tilly; 45°55′07″ s, 67°32′79″ w; 1 apr. 2006; c. pastor leg.; high littoral; fine sand sediments; cnp-nem 27442. paratypes argentina • 2 ♂♂; same collection data as for holotype; cnp-nem 27259 and 27443. description measurements see table 1. male (holotype) long and slender body. cuticle slightly striated. cephalic region set off with presence of delicate cephalic capsule (7 µm in height). cephalic capsule with same thickness throughout its smooth surface. anterior edge located above level of cephalic setae. posterior end of cephalic capsule almost straight. no ocellus nor pigment spots. three rounded lips, higher than mandibles. each lip carries two inner labial setae (8.5 µm long). six outer labial setae (18 µm long), longer than four cephalic setae (13 µm long), located at half the total height of cephalic capsule. posterior to cephalic capsule one crown of subcephalic setae formed by six pairs of setae with one short (7 µm) and one long seta (20 µm). then, crown of cervical setae (20 µm behind first) formed by four setae, two in subdorsal position and two in subventral position, and eight setae (7–20 µm) in same position as first crown. after nerve ring, another crown composed of groups of 4–5 short and scattered setae (about 4 µm) arranged in rows beside one longer seta (20 µm). rest of body presents few long somatic setae (about 14 µm) until cloaca. amphideal aperture lentil-shaped, small (14% of cbd) located just posterior to capsule end. metanemes not seen. http://zoobank.org/urn:lsid:zoobank.org:act:cbd8b1b5-e4c4-44a4-8b45-3180296c8ac3 lo russo v. & pastor de ward c.t., new species of mesacanthion from argentina 25 fig. 3. mesacanthion longigubernaculum sp. nov., ♂, holotype (cnp-nem 27442). a. cephalic sense organs on anterior end. b. posterior end, showing copulatory apparatus and precloacal supplement organ. c. gonads. d. copulatory apparatus, spicules and gubernaculum. scale bars: 1 = 20 μm; 2 = 100 μm; 3 = 10 μm. european journal of taxonomy 787: 17–31 (2021) 26 funnel-shaped buccal cavity with wide opening diminishing its width towards level of mandibles to their base. mandibles composed of two vertical rods (10 µm) united by delicate arcuate bar (4 µm) at top and ending as claws. dorsal tooth slightly smaller than two ventrosublateral teeth. all teeth pointed. pharynx cylindrical with irregular contours. cardia triangular and going into intestine. nerve ring lying at about 36% of pharynx length from anterior end. secretory-excretory system not visible. reproductive system diorchic, with opposed and outstretched testes in right position relative to intestine. one small precloacal bar-shaped supplement located about 3 abd above copulatory organ, almost parallel to body wall. subventral precloacal setae present. spicules paired, with distal portion arcuate (⅔) and proximal portion straight and dorsally directed (⅓). gubernaculum without apophysis, barely evident surrounding spicule, 63% of spicule. tail 4.6 abd long, conical in first part (⅓) with end part cylindrical (⅔ approximately). caudal gland bodies in pre-anal region. few caudal setae of same length as somatic setae and two small (3 µm) terminal setae present. female unknown. differential diagnosis mesacanthion longigubernaculum sp. nov. is characterized by its long and slender body, striated cuticle, relatively long cephalic and cervical setae, onchia of different sizes, amphidial fovea lentil-shaped, spicule arcuate, gubernaculum surrounding the spicule, and tail conical-cylindrical with terminal setae. following the key of jeong et al. (2019), m. longigubernaculum sp. nov. is similar to m. longissimesetosum and m. pali. however, m. longigubernaculum sp. nov. differs from m. longissimesetosum in having shorter inner and outer labial setae (8 and 19 vs 12 and 65–70 µm, respectively) and cephalic setae (12 vs 65 µm), a higher a index (52.9 vs 30), and a different shape of the gubernaculum. mesacanthion pali has longer inner and outer labial setae (24 and 84 vs 8 and 19 µm, respectively), lower b and c′ indices (b = 3 vs 4.5; c′ = 2.8 vs 4.4), a longer cephalic capsule (26 vs 16 µm), a gubernaculum with well-developed apophysis and the precloacal organ located at more than 2 spicule lengths from the anus. mesacanthion sanantoniensis sp. nov. urn:lsid:zoobank.org:act:79d5ac77-6294-4397-a91c-8dc83e8f02db figs 4, 5b, e etymology named after the locality where this species was found. material examined holotype argentina • ♂; río negro, san matías gulf, san antonio oeste; 40°43ʹ s, 64°58′ w; 10 oct. 2006; c. pastor leg.; lower littoral; fine sand sediments; cnp-nem 849. paratypes argentina • 1 ♂; same collection data as for holotype; cnp-nem 848 • 1 ♂; río negro, san matías gulf, san antonio oeste; 40°43ʹ s, 64°56′ w; 17 jun. 2006; c. pastor leg.; lower littoral; fine sand sediments; macn-in 43853. description measurements see table 1. http://zoobank.org/urn:lsid:zoobank.org:act:79d5ac77-6294-4397-a91c-8dc83e8f02db lo russo v. & pastor de ward c.t., new species of mesacanthion from argentina 27 fig. 4. mesacanthion sanantoniensis sp. nov., ♂, holotype (cnp-nem 849). a. cephalic sense organs on anterior end. b. posterior end, showing copulatory apparatus and precloacal supplement organ. c. copulatory apparatus, spicules and gubernaculum. d. gonads. scale bars: 1 = 20 μm; 2 = 100 μm. european journal of taxonomy 787: 17–31 (2021) 28 fig. 5. a. pharyngeal region of mesacanthion longigubernaculum sp. nov., ♂, holotype (cnp-nem 27442). b. pharyngeal region of m. sanantoniensis sp. nov., ♂, paratype (cnp-nem 848). c. detail of m. longigubernaculum sp. nov., ♂, holotype (cnp-nem 27442), showing cuticular striation. d. gubernaculum of m. bifidum sp. nov., ♂, holotype (cnp-nem 936). e. detail of m. sanantoniensis sp. nov., ♂, paratype (cnp-nem 848), showing cuticular striation. lo russo v. & pastor de ward c.t., new species of mesacanthion from argentina 29 male (holotype) large and stout body. cuticle slightly striated. cephalic region set off with presence of cephalic capsule (16 µm in height). cephalic capsule with same thickness throughout its rough surface. anterior edge located above level of cephalic setae. posterior end of cephalic capsule almost straight. no ocellus nor pigment spots. three high lips with points facing outward. each lip carries two inner labial setae (7 µm long). six outer labial setae (32 µm long) double length of four cephalic setae (16 µm long), located at middle of cephalic capsule arranged in single crown. near and posterior to cephalic capsule, short subcephalic setae (about 8–9 µm), two in subventral position, two lateral and two in subdorsal position. about 25 µm after cephalic capsule, toward nerve ring, pairs of cervical setae in subventral and subdorsal positions (same size as previous). same size setae present between nerve ring and end of oesophagus, but not in pairs and scattered. rest of body presents few shorter somatic setae (about 4 µm) until cloaca, except for longer one (8 µm) located just before precloacal organ. amphideal aperture pore-shaped, small (7% of cbd), located at middle of cephalic capsule. metanemes not seen. funnel-shaped buccal cavity with wide opening that diminishes its width towards level of mandibles to their base. its armature consists of three mandibles composed of two vertical rods (15 µm) united by arcuate bar (9 µm) at top and ending as claws. each mandible associated with tooth forming a unit. dorsal tooth slightly smaller (9 µm) than two ventrosublateral teeth (11 µm). tip of teeth not very pointed. pharynx cylindrical with irregular contours. cardia barely visible, triangular and going into intestine. nerve ring lying at about 33% of pharynx length from anterior end. secretory-excretory system not visible. reproductive system diorchic, with opposed and outstretched testes in right position relative to intestine. one ventral precloacal bar-shaped supplement located about 2.4 abd anterior to cloaca. subventral precloacal setae present. spicule paired, arcuate, with slight seam forming manubrium (1.5 abd). gubernaculum with dorsal apophysis (29% of spicule), perpendicular to body axis. tail 4.1 abd long, conical. it presents few setae in subventral and subdorsal positions (8 µm). caudal gland bodies in anal region. terminal setae not present. female unknown. differential diagnosis mesacanthion sanantoniensis sp. nov. is characterized by its long and stout body, striated cuticle, long cephalic setae, onchia of different sizes, amphidial fovea pouch-shaped, spicule arcuate, gubernaculum with dorsal apophysis, and tail conical without terminal setae. as m. longigubernaculum sp. nov., m. sanantoniensis sp. nov. is similar to m. longissimesetosum and m. pali, according to the key of jeong et al. (2019). they have males with a supplementary organ lying further away from the cloaca, presence of gubernaculum, and presence of subventral precloacal setae. mesacanthion sanantoniensis sp. nov. has shorter inner and outer labial setae than m. pali (7.3 and 33 vs 24 and 92 µm, respectively), shorter cephalic setae (20.2 vs 84 µm), lower a index (24.7 vs 50), and higher b (4.1 vs 3.3) and c′ (4.1 vs 2.8) indices, shorter mandibles (15 vs 25 µm) and a smaller precloacal organ (19.3 vs 31 µm). mesacanthion longissimesetosum and m. sanantoniensis sp. nov. have differences in mandible shape. mesacanthion sanantoniensis sp. nov. has mandibles ending in jaws, posteriorly straight and slightly smaller. mesacanthion longissimesetosum has mandibles posteriorly arcuate and larger. mesacanthion sanantoniensis sp. nov. has a distinct amphid and three crowns of cervical setae. mesacanthion longissimesetosum has no amphid, and the cervical setae are not arranged in crowns. mesacanthion longissimesetosum has two tiny precloacal setae just before the spicule vs none in m. sanantoniensis sp. nov. at the posterior part of the tail, the new species has two pairs of setae plus one single seta, and m. longissimesetosum has none. european journal of taxonomy 787: 17–31 (2021) 30 considering the similarities between m. longigubernaculum sp. nov., m. sanantoniensis sp. nov., m. longissimesetosum and m. pali, we provide a small key for an easy differentiation. key to species of mesacanthion with simple spicules shorter than 2 abd and subventral precloacal setae present this key can be taken as an addition of the last entrance (22) in the key to species of mesacanthion in jeong et al. (2019) to include two new species. 1. precloacal supplement organ less than 2 spicule lengths distant to cloaca ...... m. pali wieser, 1959 – precloacal supplement organ more than 2 spicule lengths distant to cloaca .................................... 2 2. gubernaculum without dorsal apophysis ........................................ m. longigubernaculum sp.nov. – gubernaculum with dorsal apophysis ............................................................................................... 3 3. de man´s b index greater than 5 .............................................. m. longissimesetosum wieser, 1953 – de man´s b index smaller than 5 .............................................................m. sanantoniensis sp. nov. discussion with the new species described in this study, the total number of mesacanthion species amounts to 42. for south america, the number of species present is 11 including new and previously known species. the present work contributes with three new species to the general knowledge of the taxonomy of freeliving marine nematodes and the genus of mesacanthion, in particular. species of mesacanthion are distributed all around the world, mostly in fine sand and mud sediments. each one of the new species in this study was found in a different gulf on the patagonian coasts. this results in an expansion in the distribution of the genus to new areas. they were found at different horizontal littoral and sublittoral zones, but all were inhabiting fine sand. this is also relevant for the biodiversity and ecology information of south america and argentina. acknowledgments this work was partly supported by gef-pnud project 02/018 a-b-55. the authors thank the two anonymous reviewers, whose recommendations improved the present work. references bezerra t.n., eisendle-flöckner u., hodda m., holovachov o., leduc d., mokievsky v., peña santiago r., sharma j., smol n., tchesunov a., venekey v., zhao z. & vanreusel a. 2021. nemys: world database of nematodes. available from http://nemys.ugent.be/ [accessed 30 jun. 2021]. cobb n.a. 1930. marine free-living nemas. scientific reports of the australasian antartic expedition (1911–1914). ser. c. zoology & botany 6 (7): 1–28. de coninck l.a. 1965. classe des nématodes: généralités. in: grassé p.-p. (ed.) traité de zoologie: anatomie, systématique, biologie, tome iv, fascicule ii: némathelminthes (nématodes): 1–217. de coninck l.a. & schuurmans stekhoven j.h. 1933. the freeliving marine nemas of the belgian coast ii. with general remarks on the structure and the system of nemas. mémoires du musée royal d’histoire naturelle de belgique 58: 3–163. de man j.g. 1893. cinquième note sur les nématodes libres de la mer du nord et de la manche. mémoires de la société zoologique de france 6: 81–125. http://nemys.ugent.be/ lo russo v. & pastor de ward c.t., new species of mesacanthion from argentina 31 ditlevsen h. 1930. marine free-living nematodes from new zealand. videnskabelige meddelelser fra dansk naturhistorisk forening 87: 201–242. filipjev i.n. 1927. les nématodes libres des mers septentrionales appartenant à la famille des enoplidae. archiv für naturgeschichte 91 (a): 1–216. gerlach s.a. 1954. nématodes marins libres des eaux souterraines littorales de tunisie et d‘algérie. vie et milieu 4: 221–237. gerlach s.a. 1957. die nematodenfauna des sandstrandes an der küste von mittelbrasilien (brasilianische meerse-nematoden iv). mitteilungen aus dem zoologischen museum in berlin 33: 411–459. https://doi.org/10.1002/mmnz.19570330206 gerlach s.a. & riemann f. 1974. the bremerhaven checklist of aquatic nematodes. a catalogue of nematoda adenophorea excluding the dorylaimida. part 2. veröffentlichungen des instituts für meeresforschung in bremerhaven, bremen. jeong r., tchesunov a.v. & lee w. 2019. bibliographic revision of mesacanthion filipjev, 1927 (nematoda: thoracostomopsidae) with description of a new species from jeju island, south korea. peerj 7: e8023. https://doi.org/10.7717/peerj.8023 wieser w. 1953. free-living marine nematodes i. enoploidea. acta universitatis lundensis, new series 49: 1–155. wieser w. 1959. free living nematodes and other small invertebrates of puget sound beaches. university of washington press, seattle. manuscript received: 7 july 2021 manuscript accepted: 8 november 2021 published on: 28 december 2021 topic editor: tony robillard desk editor: eva-maria levermann printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. https://doi.org/10.1002/mmnz.19570330206 https://doi.org/10.7717/peerj.8023 european journal of taxonomy 62: 1-18 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2013.62 www.europeanjournaloftaxonomy.eu 2013 · coleman c.o. & lörz a.-n. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:f38609c6-9244-424b-b692-d18df5febebd 1 iphimediidae of new zealand (crustacea, amphipoda) charles oliver coleman 1, 3,* & anne-nina lörz 2, 4 1 museum für naturkunde, leibniz institute for research on evolution and biodiversity, invalidenstraße 43, 10115 berlin, germany * corresponding author, e-mail: oliver.coleman@mfn-berlin.de 2 national centre for marine biodiversity & biosecurity, national institute of water and atmospheric research, private bag 14901, wellington 6021, new zealand e-mail: anne-nina.loerz@niwa.co.nz 3 urn:lsid:zoobank.org:author:1ec18609-2d14-462b-8e59-b1ce40166faf 4 urn:lsid:zoobank.org:author:9442484e-43a4-4383-a1a6-ae493087bca1 abstract. new zealand species of iphimediidae, amphipoda, are revised. based on new material from the chatham rise, east of new zealand, two new species are described in detail: labriphimedia meikae sp. nov. and labriphimedia martinae sp. nov. a key to the six species belonging to three genera of new zealand iphimediidae is provided. key words. crustacea, biodiversity, taxonomy, new species, chatham rise. coleman c.o. & lörz a.-n. 2013. iphimediidae of new zealand (crustacea, amphipoda). european journal of taxonomy 62: 1-18. http://dx.doi.org/10.5852/ejt.2013.62 introduction over 100 species of iphimediidae are known worldwide, yet only four were known from new zealand waters. many iphimediids have been recorded from australian waters (coleman & lowry 2006) to the northwest and in antarctic waters (coleman 2007) south of new zealand. so it is expected that discovering more species of iphimediids in new zealand is outstanding. improved sampling techniques, using small meshed gear in the understudied deep sea, gained two species of iphimediids new to science described here in detail. the new species belong to the genus labriphimedia k.h. barnard, 1931; the two other genera present in new zealand are iphimedia rathke, 1843 and anisoiphimedia g.s. karaman, 1980. anisoiphimedia is a dubious genus (coleman & lowry 2006), that will be revised when more material becomes available. anisoiphimedia haurakiensis (hurley, 1954) from subtropical northern new zealand was originally described in the genus iphimedia. in contract, the species from cold southern new zealand waters, iphimedia spinosa (hurley, 1954), was originally described as belonging to the genus panoploea thomson, 1880. http://dx.doi.org/10.5852/ejt.2013.62 http://www.europeanjournaloftaxonomy.eu/index.php/ejt http://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:f38609c6-9244-424b-b692-d18df5febebd mailto:oliver.coleman%40mfn-berlin.de?subject=oliver.coleman%40mfn-berlin.de mailto:anne-nina.loerz%40niwa.co.nz?subject=anne-nina.loerz%40niwa.co.nz http://zoobank.org/urn:lsid:zoobank.org:author:1ec18609-2d14-462b-8e59-b1ce40166faf http://zoobank.org/urn:lsid:zoobank.org:author:9442484e-43a4-4383-a1a6-ae493087bca1 http://dx.doi.org/10.5852/ejt.2013.62 european journal of taxonomy 62: 1-18 (2013) 2 four of the five species of labriphimedia are known from new zealand waters. the species labriphimedia hinemoa (hurley, 1954) from shallow waters of the hauraki gulf, northern new zealand, was originally described in the genus maoriphimedia. labriphimedia pulchridentata (stebbing, 1883) was described from heard island at 136 m but recently found at the macquarie ridge, south of new zealand in about 700 m depth (lörz 2012). labriphimedia vespucci barnard, 1931 is known from the falkland islands at 105-115 m depth, and the two species new to science described herein were sampled on chatham rise east of new zealand in 520-640 m depth. for sampling location of all iphimediids in new zealand waters see fig. 1. as the genus concept of the iphimediidae is still under construction, a key directly from family level iphimediidae to species level is provided. fig. 1. iphimediidae recorded from new zealand, from subtropical to subantarctic waters. coleman c.o. & lörz a.-n., iphimediidae of new zealand (crustacea, amphipoda) 3 material and methods the new iphimediid amphipoda were collected during the ocean survey 20/20 with r/v tangaroa on the chatham rise in 2007, east of new zealand. the material was sampled via an epibenthic sledge with a 500 µm mesh size. the iphimediids were fixed in 90% ethanol and later transferred into glycerol and mounted on slides for the preparation of the drawings. pencil drawings were made with a camera lucida on a leica m 205c dissecting microscope and a leica dmlb compound microscope. the line drawings were made using the technique described in coleman (2003, 2009). length measurements were made along the outline of the animals, beginning at the tip of the rostrum to the end of the telson. the descriptions were generated from a delta database (dallwitz 1993 onwards) of the iphimedia species and related genera like coboldus of the world, used in coleman & lowry (2006). the delta text-output was modified to improve readability and adding characters that are relevant for the new taxa presented herein. the type material is deposited in the invertebrate collection of the national institute of water and atmospheric research (niwa) in wellington, new zealand. results key to new zealand iphimediid species 1 coxa 5-7 posteriorly bicuspidate ..................................labriphimedia pulchridentata stebbing, 1883 coxa 5-7 posteriorly smooth or bearing single spine .........................................................................2 2 four pairs of spines dorsally ...............................................................................................................3 three pairs of spines dorsally, pleonite 3 not bearing paired spines ......................................................4 3 coxa 1 ventrally acute; urosomite 2 dorsally smooth .....................labriphimedia meikae sp. nov. coxa 1 ventrally rounded; urosomite 2 bearing dorsal carina ............................................................... ..................................................................................................labriphimedia hinemoa (hurley, 1954) 4 pleonites 1-2 mid-dorsally smooth prior to paired spines; coxa 7 rounded ............................................5 pleonites 1-2 with mid-dorsal pointy protrusion prior to paired teeth; coxa with 7 posteroventral margin pointed ..................................................................................labriphimedia martinae sp. nov. 5 coxa 1 anterodistally rounded; coxa 6 posteriorly pointed ...................................................................... ..........................................................................................anisoiphimedia haurakiensis (hurley, 1954) coxa 1 anterodistally truncate; coxa 6 posteriorly rounded .........iphimedia spinosa (hurley, 1954) systematics amphipoda latreille, 1816 iphimediidae boeck, 1871 labriphimedia k.h. barnard, 1931 labriphimedia k.h. barnard, 1931: 427 (type species: labriphimedia vespuccii k.h. barnard, 1931, original designation). maoriphimedia hurley, 1954: 771 (type species: maoriphimedia hinemoa hurley, 1954, original designation). — karaman & barnard, 1979: 111 (synonomy). type species labriphimedia vespucci k.h. barnard, 1931 diagnosis epistome short and broad. upper lip wider than long, emarginated or elongate and tapering. european journal of taxonomy 62: 1-18 (2013) 4 mandible tapering to a smooth spoon-shaped or spatulate apex, molar rudimentary, no spine row. lower lip without inner lobes. maxilla 1, palp 2 articles, stout or 1-articulate and minute. maxilliped outer plate broad, palp slender, 2nd article not medially produced. species composition labriphimedia hinemoa (hurley, 1954) labriphimedia martinae sp. nov. labriphimedia meikae sp. nov. labriphimedia pulchridentata (stebbing, 1883) labriphimedia vespucci k.h. barnard, 1931 labriphimedia meikae sp. nov. figs 2-6 urn:lsid:zoobank.org:act:bd138120-dfeb-4939-8848-7e6d79ee9b4c material examined holotype ♂, 6 mm, niwa 84598. additional material two paratypes of unknown sex, 5.2 mm and 5 mm, niwa 31826, same station data as holotype. locus typicus new zealand: tan0705/251, 24 april 2007, 42°59ˮ45’ s, 178°59ˮ44’ e – 42°59ˮ28’ s, 179°0ˮ19’ e, 520-530 m. etymology this species is named for ms meike snyder, the dear aunt of the senior author. description length. based on male holotype, 6 mm. head (fig. 2a). eyes reniform; rostrum only weakly curved; anterior margin with small rounded sculptured spine (which looks pointed in dorsal view), anteroventral corner with rounded lobe. antenna 1 (fig. 2b) peduncular article 1 with a long posteroventral spine reaching the distal margin of the peduncular article 2 and a shorter dorsal spine; peduncular article 2 with short spine; peduncular article 3 and flagellum unknown. antenna 2 (fig. 2d) peduncular article 2 with short rounded gland cone and longer bifid medial process; peduncular article 4 slightly drawn out laterally; peduncular article 5 and flagellum unknown. upper lip (fig. 2g) wider than long, emarginate. mandibles (fig. 2e) with smooth incisors and a spine-like lacinia mobilis on the left mandible (the right mandible apparently does not have a lacinia mobilis); palp 3-articulate, ratios of articles 1-3 1:1.4:1.2. lower lip (fig. 2c) inner lobes absent, distally rounded. maxilla 1 (fig. 3a) inner plate with 6 plumose setae on the inner margin; outer plate with 11 spine-like serrate setae; palp 2-articulate, well developed, longer than outer plate, with 9 apical setae. maxilla 2 (fig. 2f) inner plate slightly shorter and subequal in width compared to outer plate. maxilliped (fig. 3b-e) inner plate long, rectangular with slender plumose setae apically and 2 rows mediomarginally; outer plate with tapering apex, bordered with plumose setae; palp article 1 as long as articles 2 and 3 combined; article 2 weakly produced medially, not guarding along article 3; there may be an inconspicuous 4th article. http://zoobank.org/urn:lsid:zoobank.org:act:bd138120-dfeb-4939-8848-7e6d79ee9b4c coleman c.o. & lörz a.-n., iphimediidae of new zealand (crustacea, amphipoda) 5 fig. 2. labriphimedia meikae nov. sp., holotype ♂, 6 mm, niwa 84598. a. habitus. b. antenna 1, peduncular articles 1-2. c. lower lip. d. antenna 2, peduncular articles 1-4. e. left mandible. f. maxilla 2. g. upper lip. scale bars: a = 1 mm, b-g = 100 µm. a b c d e f g european journal of taxonomy 62: 1-18 (2013) 6 fig. 3. labriphimedia meikae nov. sp., holotype ♂, 6 mm, niwa 84598. a. maxilla 1. b. outlines of maxilliped. c. palp of maxilliped. d. outer plate of maxilliped. e. inner plate of maxilliped. scale bars: a, c-e = 100 µm, b = 200 µm. a b c d e coleman c.o. & lörz a.-n., iphimediidae of new zealand (crustacea, amphipoda) 7 pereon (fig 2a). pereonite 1 enlarged, about 3-4 x as long as pereonite 2. pereonites 5-7 with pointed and drawn out posteroventral angles. pereonite 7 without mid-dorsal carina, dorsodistal margin with 2 large narrow pointed spines, space between spines wide. gnathopod 1 (fig. 4a) coxa ventrally acute, ventral and anterior margin smooth apart from 2 minute notches with microtrichs; basis longest; ischium fig. 4. labriphimedia meikae nov. sp., holotype ♂, 6 mm, niwa 84598. a. gnathopod 1. b. chela of gnathopod 1. c. gnathopod 2. scale bars: a, c = 100 µm. a b c european journal of taxonomy 62: 1-18 (2013) 8 elongate, slightly longer than merus; carpus slightly longer than propodus; dactylus and propodus forming a chela, inner margin of dactylus serrate apically; setation, apart some setae on basis, only on chela. gnathopod 2 (fig. 4b) larger than gnathopod 1; coxa ventrally acute, ventral and anterior margin smooth apart from 2 minute notches with microtrichs; basis longest, ischium longer than merus; merus expanded distally; carpus and propodus subequal in length; propodus with groups of setae posteromarginally and apically; distal propodus extension wider than dactylus; chelate. pereopod 3 (fig. 5a) coxa longer than preceding coxae, posteroventrally acute, ventral margin apart from 2 minute fig. 5. labriphimedia meikae nov. sp., holotype ♂, 6 mm, niwa 84598. a. pereopod 3. b. pereopod 4. c. pereopod 5. scale bars: a-b = 100 µm. c = 200 µm. a b c coleman c.o. & lörz a.-n., iphimediidae of new zealand (crustacea, amphipoda) 9 a b c d e f g fig. 6. labriphimedia meikae nov. sp., holotype ♂, 6 mm, niwa 84598. a. pereopod 6. b. coxa of pereopod 7. c. basis to merus of pereopod 7. d. uropod 3. e. uropod 2. f. uropod 1. g. telson. scale bars: a-c = 200 µm, d-g = 100 µm. european journal of taxonomy 62: 1-18 (2013) 10 notches with microtrichs smooth; basis and ischium with rounded lobe on apical margin. pereopod 4 (fig. 5b) coxa pointed distally and posteromarginally, ventral margin smooth apart from some minute notches with microtrichs; basis and ischium with rounded lobe on apical margin. pereopod 5 (fig. 5c) coxa with long pointed posteroventral spine; basis posterior margin with 2 spines: posterodorsal spine pointed, posteroventral corner subquadrate; posterior margin weakly serrate; ischium anterodistally pointed; merus posteroventral lobe elongate; carpus to dactylus unknown. pereopod 6 (fig. 6a) of similar shape as pereopod 5, but basis wider with a pointed posteroventral corner and bearing an angular posterodorsal process. pereopod 7 (fig. 6b-c) coxa small with pointed posteromarginal spine; basis posteroventrally with spine, directed ventrally, posterior margin weakly serrate; merus posteroventral lobe elongate. pleon (fig. 2a). pleonite 1 without mid-dorsal carina, with 2 large, pointed spines, space between spines widely excavate, similar to the arrangement on pereonite 7. epimeron 1 posterior margin with 1 mid-lateral spine, posteroventral corner with short point. pleonite 2 without mid-dorsal carina; with 2 large, pointed spines, space between spines wide. epimeron 2 posterior margin with 1 mid-lateral spine, posteroventral corner pointed. pleonite 3 without mid-dorsal carina; dorsodistal margin with 2 straight spines, shorter than those on pereonite 7 and pleonites 1-2. epimeron 3 posterior margin with 1 well developed lateral, smooth spine, posteroventral corner produced into smooth spine. urosomite 1 longest, smooth. urosomite 2 shortest. telson (fig. 6g) emarginate. distribution chatham rise, new zealand (south-western pacific), 520-530 m. labriphimedia martinae sp. nov. figs 7-11 urn:lsid:zoobank.org:act:d8ed7489-8474-4e50-9856-27119a69e933 material examined holotype ♂, 6.5 mm, niwa 84743. locus typicus new zealand: tan0705/136, 14 april 2007, 43°17’25ˮ s, 175°33’08ˮ w – 43°17’36ˮ s, 175°33’47ˮ e, 638-644 m. etymology this species is named for the amiable ms martine willcox, the mother of the son-in-law of the senior author. description length. based on male holotype, 6.5 mm. head (fig. 7a). eyes ovate; rostrum strongly curved; anterior margin with angular spine, anteroventral corner with an acute long narrow spine. antenna 1 (fig. 7c-d) peduncular article 1 with a long distomedial spine and a small anterodistal spine; peduncular article 2 with longer dorsal and small spine, with an anterodistal spine; accessory flagellum present, accessory flagellum 1-articulate; article 3 with anterodistal process; article 4 subequal to 3. antenna 2 (fig. 7g) articles 2 and 3 serrate distomedially. mouthparts arranged in a cone. upper lip (fig. 7b) tapering distally, apex asymmetrical. mandibular body (fig. 7e, f, i, h) tapering distally; incisor not dentate; lacinia mobilis on right side rather stout; http://zoobank.org/urn:lsid:zoobank.org:act:d8ed7489-8474-4e50-9856-27119a69e933 coleman c.o. & lörz a.-n., iphimediidae of new zealand (crustacea, amphipoda) 11 fig. 7. labriphimedia martinae nov. sp., holotype ♂, 6.5 mm, niwa 84743. a. habitus. b. upper lip. c. antenna 1 peduncular articles 1-2. d. peduncular article 3 and flagellum of antenna 1. e. left mandible, setation of palp omitted. f. right mandibular palp. g. antenna 2. h. right mandibular body. i. apical mandibular palp article. scale bars: a = 1 mm, b-g = 100 µm. a b c d e f g h i european journal of taxonomy 62: 1-18 (2013) 12 a b c d e f g h i j fig. 8. labriphimedia martinae nov. sp., holotype ♂, 6.5 mm, niwa 84743. a. lower lip. b. outer plate of maxilla 2. c. inner plate of maxilla 2. d. left maxilla 1. e. palp of right maxilla 1. f. outlines of maxilla 2. g. palp of maxilliped. h. inner plate of maxilliped. i. outer plate of maxilliped. j. outline of maxillipeds. scale bars: a-j = 100 µm. coleman c.o. & lörz a.-n., iphimediidae of new zealand (crustacea, amphipoda) 13 a b c d e f fig. 9. labriphimedia martinae nov. sp., holotype ♂, 6.5 mm, niwa 84743. a. gnathopod 1. b. chela of gnathopod 1. c. gnathopod 2. d. chela of gnathopod 2. e. pereopod 3. f. presumed carpus to dactylus of pereopod 3. scale bars: a, c, e-f = 100 µm. european journal of taxonomy 62: 1-18 (2013) 14 fig. 10. labriphimedia martinae nov. sp., holotype ♂, 6.5 mm, niwa 84743. a. pereopod 4. b. presumed carpus to dactylus of pereopod 4. c. pereopod 7. d. pereopod 6. scale bars: a-d = 100 µm. a b c d coleman c.o. & lörz a.-n., iphimediidae of new zealand (crustacea, amphipoda) 15 molar not triturative, but protruding; palp 3 articulate, ratios of articles 1-3 1:2.75:1.5. lower lip (fig. 8a) inner lobes absent; lobes tapering distally into pointed apex. maxilla 1 (fig. 8d-e) palp 1-articulate, much shorter than outer plate, very small scale with an apical seta. maxilla 2 (fig. 8b-c, f) inner lobe considerably shorter than outer. maxilliped (fig. 8g-j) inner lobe elongate, surpassing the distal margin of 1st palp article, with setae on the apex and 2 rows of setae along the medial margin; outer plate fig. 11. labriphimedia martinae nov. sp., holotype ♂, 6.5 mm, niwa 84743. a. pereopod 5. b. pleopod 1. c. right uropod 2. d. uropod 1. e. uropod 3. f. telson. scale bars: a-f = 100 µm. b c d e f a european journal of taxonomy 62: 1-18 (2013) 16 tapering distally, bordered with plumose setae apically; palp consisting of 3 long articles and 1 minute distolateral article; 2nd palp article expanded distomedially. pereon (fig. 7a). pereonite 1 enlarged, 3-4 x as long as pereonite 2. pereonites 5-6 posteroventral angle acute; that of pereonite 7 pointed and drawn out. pereonite 7 without mid-dorsal carina, dorsodistal margin with 2 large pointed spines, space between spines widely excavate, with 1 posteroventral spine. gnathopod 1 (fig. 9a-b) coxa subrectangular, ventrally truncated and serrate; basis slightly longer than coxa; carpus somewhat longer than propodus, both carpus and propodus narrow; setation restricted to anterior margin of basis and region of chela. gnathopod 2 (fig. 9c-d) slightly shorter than gnathopod 1; coxa tapering, ventrally truncated, ventral margin with 2 notches with microtrichs; carpus and propodus subequal in length, strongly setose; pereopod 3 (fig. 9e-f) coxa ventrally rounded, ventral margin weakly serrate; basis and ischium with rounded lobe distomarginally. pereopod 4 (fig. 10a-b) coxa wide, apically rounded, with angular posteroventral corner, ventral margin serrate; posterior margin with rounded lobe; basis and ischium with rounded lobe distomarginally. pereopod 5 (fig. 11a) coxa wider than long, bilobate, posterior margin pointed; basis subrectangular, posterior margin straight, weakly serrate, posteroventral corner subquadrate, anterodistal angle pointed; ischium anterodistal angle pointed; merus drawn out posterodistally; carpus expanded distally, slightly shorter than propodus; dactylus not curved much, with small unguis. pereopod 6 (fig. 10d) coxa bilobate, anterior lobe shorter, with posteroventral spine; basis anterodistally pointed, posterodorsal corner subquadrate, posterior margin weakly serrate, posteroventral corner with small spine; ischium and merus as for pereopod 5. pereopod 7 (fig. 10c) coxa with posteroventral spine; basis anterodistally pointed, posterodorsal corner rounded, posterior margin weakly serrate, posteroventral corner with large spine; merus and carpus as for pereopod 5. pleon (fig. 7a). pleonite 1 with a low, truncated mid-dorsal carina with a small posterior point, with 2 large, pointed spines, space between spines wide. epimeron 1 posterior margin with 1 mid-lateral spine, posteroventral corner acutely with short point. pleonite 2 with a low truncated mid-dorsal carina with a small posterior point, dorsodistal margin with 2 large, pointed spines, space between spines wide. epimeron 2 posterior margin with 1 mid-lateral spine, posteroventral corner pointed. pleonite 3 with a broadly rounded, large mid-dorsal carina. epimeron 3 posterior margin without well developed lateral spines, posteroventral corner produced into 1 smooth spine, its dorsal part minutely serrate. urosomite 1 large with broadly rounded, large mid-dorsal carina with a shallow depression. urosomite 2 shortest, smooth. telson (fig. 11f) truncate, with several small notches along the distal margin. distribution chatham rise, new zealand (south-western pacific), 638-644 m. discussion the classification of the two new species turned out to be difficult. it was clear that they were new species after comparison with all known iphimediid species, but it was very difficult to decide in which genus they belong. the systematics of iphimediids is mostly based on the spination of the body, the coxae and the bases of the appendages, and on the mouthpart morphology, more specifically the shape and dentation of the incisor, and the development of the palps of maxilla 1 and the maxilliped. the mandible of l. meikae sp. nov. is similar to that of labriphimedia hinemoa (hurley, 1954). in both species the incisor is toothless and apically rounded and medially excavate. also some other characters on the mouthparts correspond in the shapes: of antenna 1 peduncular article 1, upper lip, lower lip, maxilla 1 and maxilliped. the heads of both species are very different: l. meikae sp. nov. has a weakly curved rostrum (vs. strongly curved), large eyes (vs. small), anterior head margin rounded in lateral view (vs. pointed) and the ventral head margin with a sculptured rounded lobe (vs. longer subacute lobe). coleman c.o. & lörz a.-n., iphimediidae of new zealand (crustacea, amphipoda) 17 the coxal plates 1-4 are not so pointed ventrally in l. hinemoa. the shapes of the pereopod 5-7 bases are similar, but the anterior margin in l. meikae sp. nov. is not so densely covered with setae as in l. hinemoa. both species share the 4 pairs of mid-dorsal spines, but the shape differs. in l. meikae sp. nov. the dorsal spines are slender seen from laterally and have a wide u-shaped excavation seen from dorsally. however, in l. hinemoa the spines are rather wide from lateral aspect and form a v-shaped narrow slit in the dorsal view. the telson of l. hinemoa appears to be deeper notched and has subacute lobes compared to l. meikae sp. nov., where the telson lobes are truncate. labriphimedia martinae sp. nov. is more difficult to classify. this species has some superficial resemblance to labriphimedia vespucci k.h. barnard, 1931 in the shape of coxae 1-4 and in the prominent carina on pleonite 3 and urosomite 1, which is, however, notched in lateral view in the latter species. also, both species have a serrate margin of epimeron 3 just dorsal of the posteroventral angle, but in l. vespucci there is an additional spine on the posterior margin. compared to l. vespucci, in l. martinae sp. nov. the paired dorsal spines on pleonite 3 are missing, the eyes are much larger, covering more than half of the head (vs. less than half) and the mouthparts are very different: the mouthparts are arranged into a pointed bundle, upper lip, mandibles and lower lip are pointed, also maxilla 1 is elongate, but its palp is reduced to a small scale. in l. vespucci, however, the mouthparts do not seem to be elongate, the upper lip is much broader than long and maxilla 1 looks similar to that of l. meikae sp. nov. (not elongate, with a long 2-articulate palp). a shortened palp of maxilla 1 is known in many iphimediid species, especially in the genus iphimedia, but there the palp is 2-articulate. a 1-articulate palp of maxilla 1 within iphimediidae is known in the genus coboldus krapp-schickel, 1974 and we were initially attracted by the idea that l. martinae sp. nov. could be classified in the genus coboldus. in favour of coboldus would count the shortened 1-articulate palp of maxilla 1, the pointed mouthpart bundle and the elongate 2nd article of the mandibular palp, which some species have (e.g., c. laetifucatus just, 1990). however, other characteristics of coboldus are not present in l. martinae: a strongly produced 2nd article of the maxillipedal palp guarding along article 3, the absence of carinae on pleonites 1-3 and urosomite 1 and the presence of a lateral spine on the posterior margin of epimeron 3. it may be that there is some phylogenetic connection between both genera, coboldus and labriphimedia, which we hope to resolve in a future project. the genus labriphimedia is so far only known from the southern hemisphere, and four of the now known five species of labriphimedia occur in new zealand waters. acknowledgements we like to thank ms makena löhr who did several pencil drawings of appendages of labriphimedia martinae sp. nov. erika mackay (niwa) created the map. the material was supplied by the niwa invertebrate collection and collected during the ocean survey 20/20 chatham/challenger biodiversity and seabed habitat project, jointly funded by the new zealand ministry of fisheries, land information new zealand, niwa, and the department of conservation. the second author was supported by niwa through coast and oceans research programme 2, biodiversity & biosecurity, taxonomy project cobr1402. references coleman c.o. 2003. “digital inking”: how to make perfect line drawings on computers. organism diversity and evolution 3, electronic supplement 14: 1-14. (http://senckenberg.de/odes/03-14.htm). http://dx.doi.org/10.1078/1439-6092-00081 http://senckenberg.de/odes/03-14.htm http://dx.doi.org/10.1078/1439-6092-00081 european journal of taxonomy 62: 1-18 (2013) 18 coleman c.o. 2007. synopsis of the amphipoda of the southern ocean; volume 2: acanthonotozomellidae, amathillopsidae, dikwidae, epimeriidae, iphimediidae, ochlesidae and vicmusiidae. bulletin de l’institut royal des sciences naturelles de belgique, biologie 77, supplement 2: 1-142. coleman c.o. 2009. drawing setae the digital way. zoosystematics and evolution 85 (2): 305-310. http://dx.doi.org/10.1002/zoos.200900008 coleman c.o. & lowry j.k. 2006. australian iphimediidae (crustacea: amphipoda). organisms diversity & evolution 6, electronic supplement 9: 1-44. http://www.senckenberg.de/odes/06-09.htm dallwitz m.j. 1993 onwards. applications and documentation of the delta system. http://deltaintkey.com lörz a.n. 2012. first records of epimeriidae and iphimediidae (crustacea, amphipoda) from macquarie ridge, with description of a new species and its juveniles. zootaxa 3200: 49-60. manuscript received: 1 august 2013 manuscript accepted: 27 september 2013 published on: 6 november 2013 topic editor: rudy jocqué desk editor: kristiaan hoedemakers printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; national botanic garden of belgium, meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark. http://dx.doi.org/10.1002/zoos.200900008 http://www.senckenberg.de/odes/06-09.htm http://delta-intkey.com http://delta-intkey.com european journal of taxonomy 314: 1–8 issn 2118-9773 https://doi.org/10.5852/ejt.2017.314 www.europeanjournaloftaxonomy.eu 2017 · van ginneken m. et al. this work is licensed under a creative commons attribution 3.0 license. c o r r i g e n d u m 1 the following corrections have been made to paper no. 310 (https://doi.org/10.5852/ejt.2017.310) morphometry and dna barcoding reveal cryptic diversity in the genus enteromius (cypriniformes: cyprinidae) from the congo basin, africa – corrigendum marjolein van ginneken 1,†, eva decru 2,*,†, erik verheyen 3 & jos snoeks 4 1 department of biology, systemic physiological and ecotoxicological research, university of antwerp, groenenborgerlaan 171, 2020 antwerpen, belgium. 2,4 royal museum for central africa, section vertebrates, ichthyology, leuvensesteenweg 13, 3080 tervuren, belgium. 2,4 department of biology, laboratory of biodiversity and evolutionary genomics, ku leuven, charles deberiotstraat 32, 3000 leuven, belgium. 3 royal belgian institute of natural sciences, od taxonomy and phylogeny, vautierstraat 29, 1000 brussels, belgium; department of biology, evolutionary ecology group, university of antwerpen, campus drie eiken, building d, room d.150 universiteitsplein 1, 2610 antwerpen, belgium. † equally contributing authors * corresponding author: eva.decru@africamuseum.be 1 email: marjolein.vanginneken@uantwerpen.be 3 email: everheyen@naturalsciences.be 4 email: jos.snoeks@africamuseum.be van ginneken m., decru e., verheyen e. & snoeks j. 2017. morphometry and dna barcoding reveal cryptic diversity in the genus enteromius (cypriniformes: cyprinidae) from the congo basin, africa – corrigendum. european journal of taxonomy 314: 1–8. https://doi.org/10.5852/ejt.2017.314 in the original publication, figs 3–9 were accidentally published in low resolution. the high resolution figures with their captions are published below. figs 8–9 showed a white rectangle for ‘ituri 8’, which should have been a black rectangle, herewith updated. the captions have not changed. https://doi.org/10.5852/ejt.2017.314 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/3.0/ https://doi.org/10.5852/ejt.2017.310 mailto:eva.decru%40africamuseum.be?subject= mailto:marjolein.vanginneken%40uantwerpen.be?subject= mailto:everheyen%40naturalsciences.be?subject= mailto:jos.snoeks%40africamuseum.be?subject= https://doi.org/10.5852/ejt.2017.314 european journal of taxonomy 314: 1–8 (2017) 2 fig. 3. scatterplot of pc2 against pc1 for a pca on 17 log-transformed measurements (n = 177) of enteromius cope, 1867: e. cf. miolepis (boulenger, 1902) (◊), e. cf. brazzai (pellegrin, 1901) (♦), e. cf. pellegrini (poll, 1939) (∆), and e. cf. atromaculatus (nichols & griscom, 1917) (▲). also shown are the type specimens examined of: e. miolepis (boulenger, 1902) (○), e. holotaenia (boulenger, 1904) (●), e. eutaenia (boulenger, 1904) (□), e. kerstenii (peters, 1868) (■), e. brazzai (pellegrin, 1901) ( ), e. tshopoensis (de vos, 1991) (▼), e. pellegrini (poll, 1939) (+), and e. atromaculatus (nichols & griscom, 1917) ( ). van ginneken m. et al., unexpected cryptic diversity in enteromius 3 fig. 4. scatterplot of pc2 against pc1 for a pca on 10 meristics (n = 177) of enteromius: e. cf. miolepis (boulenger, 1902) (◊), e. cf. brazzai (pellegrin, 1901) (♦), e. cf. pellegrini (poll, 1939) (∆), and e. cf. atromaculatus (nichols & griscom, 1917) (▲). also shown are the type specimens examined of: e. miolepis (boulenger, 1902) (○), e. holotaenia (boulenger, 1904) (●), e. eutaenia (boulenger, 1904) (□), e. kerstenii (peters, 1868) (■), e. brazzai (pellegrin, 1901) ( ), e. tshopoensis (de vos, 1991) (▼), e. pellegrini (poll, 1939) (+), and e. atromaculatus (nichols & griscom, 1917) ( ). european journal of taxonomy 314: 1–8 (2017) 4 fig. 5. scatterplot of pc2 against pc1 for a pca on 10 meristics (n = 36) of e. cf. miolepis specimens from the lower congo: inkisi (◊), luki 1 (♦) and luki 2 (∆). also shown are the type specimens examined of: e. miolepis (boulenger, 1902) (○), e. holotaenia (boulenger, 1904) (●), e. eutaenia (boulenger, 1904) (□) and e. kerstenii (peters, 1868) (■). van ginneken m. et al., unexpected cryptic diversity in enteromius 5 fig. 6. scatterplot of pc2 against pc1 for a pca on 8 meristics (n = 60) of e. cf. miolepis (boulenger, 1902) specimens from the congo basin (excluding types): ‘kisangani region’ 1 (◊), ituri 1 (♦), itimbiri (∆), léfini (▲), epulu 1 (○), inkisi (●), luapula 1 (□), luki 1 (■), luapula 2 ( ), luapula 3 (▼), ituri 2 (+), and luki 2 ( ). specimens from luapula 1 and luapula 2 can be separated from each other based on a pca on the log-transformed measurements; specimens of luki 2 fall separated when barbel lengths are included; specimens from ‘kisangani region’ 1 and itimbiri can be distinguished based on colour pattern. european journal of taxonomy 314: 1–8 (2017) 6 fig. 7. scatterplot of pc2 against pc1 for a pca on 10 meristics (n = 22) of e. cf. brazzai (pellegrin, 1901): ‘kisangani region’ 2 (◊), ituri 3 (♦) and ‘kisangani region’ 3 (∆). also shown are the type specimens examined of e. brazzai (pellegrin, 1901) (○) and e. tshopoensis (de vos, 1991) (●). van ginneken m. et al., unexpected cryptic diversity in enteromius 7 fig. 8. scatterplot of pc2 against pc1 for a pca on 10 meristics (n = 42) of e. cf. atromaculatus (nichols & griscom, 1917): ituri 5 (◊), ituri 6 (♦), ituri/‘kisangani region’ (∆), epulu 2 (▲), and ituri 8 ( ). also shown are the type specimens of e. atromaculatus (nichols & griscom, 1917) (○). european journal of taxonomy 314: 1–8 (2017) 8 published on: 28 april 2017 printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain. fig. 9. scatterplot of pc2 against pc1 for a pca on 10 meristics (n = 36) of e. cf. atromaculatus (nichols & griscom, 1917): epulu 2 (▲), and ituri 8 ( ). also shown are the type specimens of e. atromaculatus (nichols & griscom, 1917) (○). european journal of taxonomy 879: 188–189 issn 2118-9773 https://doi.org/10.5852/ejt.2023.879.2171 www.europeanjournaloftaxonomy.eu 2023 · pham n.t. et al. this work is licensed under a creative commons attribution license (cc by 4.0). c o r r i g e n d u m urn:lsid:zoobank.org:pub:c348e8a3-e8d4-48d6-ab5c-f6696bb58da5 188 a review of the genus enicospilus stephens (ichneumonidae: ophioninae) from vietnam, with descriptions of ten new species – corrigendum nhi thi pham  1,*, phu van pham  2, rikio matsumoto  3, so shimizu  4 & gavin r. broad  5 1,2 institute of ecology and biological resources, vietnam academy of science and technology, 18 hoang quoc viet, hanoi, vietnam. 3 osaka museum of natural history, nagai park 1-23, higashisumiyoshi-ku, osaka, japan. 4 institute for agro-environmental sciences, naro, kannondai 3-1-3, tsukuba 305-8604, japan. 4 laboratory of insect biodiversity and ecosystem science, graduate school of agricultural science, kobe university, kobe, hyogo, japan. 5 the natural history museum, cromwell road, london sw7 bd, uk. 1 urn:lsid:zoobank.org:author:67079e55-d341-48f2-aedb-71aa8d40ad2d 2 urn:lsid:zoobank.org:author:09b6f968-3b83-4052-89a1-1db2d91933cf 3 urn:lsid:zoobank.org:author:9d30e868-4c4b-4f40-a6c8-781f185fa0e4 4 urn:lsid:zoobank.org:author:d60bfe76-b686-4fb2-a720-a02de0a2c2b1 5 urn:lsid:zoobank.org:author:d06689de-526f-4cfa-8beb-9fb38850754a * corresponding author: ptnhi2@yahoo.com 2 email: phamphu93.k56@gmail.com 3 email: rikio@mus-nh.city.osaka.jp 4 email: parasitoidwasp.sou@gmail.com 5 email: g.broad@nhm.ac.uk pham n.t., pham p.v., matsumoto r., shimizu s. & broad g.r. 2023. a review of the genus enicospilus stephens (ichneumonidae: ophioninae) from vietnam, with descriptions of ten new species – corrigendum. european journal of taxonomy 879: 188–189. https://doi.org/10.5852/ejt.2023.879.2171 the present corrigendum corrects an error that occurred in pham et al. (2023). https://doi.org/10.5852/ejt.2023.873.2133 inadvertently, the classification of the publication stated on the title page as “research article” is incorrect. the correct classification should be “monograph”. we apologize for any confusion this error may have caused. the content of the paper remains unchanged. https://doi.org/10.5852/ejt.2023.879.2171 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ https://zoobank.org/urn:lsid:zoobank.org:pub:c348e8a3-e8d4-48d6-ab5c-f6696bb58da5 https://orcid.org/0000-0001-9304-9863 https://orcid.org/0000-0001-9369-6649 https://orcid.org/0000-0003-3979-9713 https://orcid.org/0000-0002-5202-4552 https://orcid.org/0000-0001-7223-5333 https://zoobank.org/urn:lsid:zoobank.org:author:67079e55-d341-48f2-aedb-71aa8d40ad2d https://zoobank.org/urn:lsid:zoobank.org:author:09b6f968-3b83-4052-89a1-1db2d91933cf https://zoobank.org/urn:lsid:zoobank.org:author:9d30e868-4c4b-4f40-a6c8-781f185fa0e4 https://zoobank.org/urn:lsid:zoobank.org:author:d60bfe76-b686-4fb2-a720-a02de0a2c2b1 https://zoobank.org/urn:lsid:zoobank.org:author:d06689de-526f-4cfa-8beb-9fb38850754a mailto:ptnhi2%40yahoo.com?subject= mailto:phamphu93.k56%40gmail.com?subject= mailto:rikio%40mus-nh.city.osaka.jp?subject= mailto:parasitoidwasp.sou%40gmail.com?subject= mailto:g.broad%40nhm.ac.uk?subject= https://doi.org/10.5852/ejt.2023.879.2171 https://doi.org/10.5852/ejt.2023.873.2133 pham n.t. et al., enicospilus from vietnam – corrigendum 189 reference pham n.t., pham p.v., matsumoto r., shimizu s. & broad g.r. 2023. a review of the genus enicospilus stephens (ichneumonidae: ophioninae) from vietnam, with descriptions of ten new species. european journal of taxonomy 873: 1–151. https://doi.org/10.5852/ejt.2023.873.2133 published on: 14 july 2023 printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; leibniz institute for the analysis of biodiversity change, bonn – hamburg, germany; national museum of the czech republic, prague, czech republic. https://doi.org/10.5852/ejt.2023.873.2133 1 european journal of taxonomy 676: 1–15 issn 2118-9773 https://doi.org/10.5852/ejt.2020.676 www.europeanjournaloftaxonomy.eu 2020 · olivar j.e.c. et al. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e cyrtandra argentii, a new species of cyrtandra (gesneriaceae) from the philippines, and a review of the c. villosissima group jay edneil c. olivar 1,*, hannah j. atkins 2, frank hauenschild 1,3 & alexandra n. muellner-riehl 1,4 1 department of molecular evolution and plant systematics & herbarium (lz), institute of biology, leipzig university, johannisallee 21–23, 04103 leipzig, germany. 2 royal botanic garden edinburgh, 20a inverleith row, eh3 5lr, united kingdom. 3 leipzig university, centre for teacher training and school research, prager str. 38–40, 04317 leipzig, germany. 4 german centre for integrative biodiversity research (idiv) halle-jena-leipzig, leipzig, germany. * corresponding author: jay_edneil.olivar@uni-leipzig.de 2 email: hatkins@rbge.org.uk 3 email: frank.hauenschild@uni-leipzig.de 4 email: muellner-riehl@uni-leipzig.de abstract. cyrtandra argentii olivar, h.j.atkins & muellner sp. nov., endemic to the philippines and named after george argent, is herein described and illustrated. collections associated with this new species are often confused with three other species, namely c. ferruginea merr., c. villosissima merr., and c. hirtigera h.j.atkins & cronk. distinguishing characters including keys, updated descriptions, distribution maps, and photos of live specimens are provided to aid identification of the four species. following the international union for the conservation of nature (iucn) criteria, c. argentii sp. nov. is considered to be near threatened (nt) due to its distribution in a zone susceptible to anthropogenic pressure and the lack of any formal protection. keywords. cyrtandra, c. argentii sp. nov., george argent, iucn, philippines. olivar j.e.c., atkins h.j., hauenschild f. & muellner-riehl a.n. 2020. cyrtandra argentii, a new species of cyrtandra (gesneriaceae) from the philippines, and a review of the c. villosissima group. european journal of taxonomy 676: 1–15. https://doi.org/10.5852/ejt.2020.676 introduction cyrtandra j.r.forst. & g.forst. (forster & forster 1775) is the largest genus of ca 800 spp. in the family gesneriaceae rich. & juss. (de candolle 1816). the genus is recognized by possessing two fertile stamens and fruits that are indehiscent, either hard capsules or fleshy berries (cronk et al. 2005; atkins et al. 2013). members of the genus exhibit diverse growth forms, ranging from herbs and shrubs, to climbers and small trees distributed throughout the malesian region and across the pacific (johnson et al. 2017; kartonegoro et al. 2018). species of cyrtandra are important rainforest elements, thriving in habitats with high humidity, low light intensity, and constant moisture supply (gillett 1967). despite a https://doi.org/10.5852/ejt.2020.676 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ mailto:jay_edneil.olivar%40uni-leipzig.de?subject= mailto:hatkins%40rbge.org.uk?subject= mailto:frank.hauenschild%40uni-leipzig.de?subject= mailto:muellner-riehl%40uni-leipzig.de?subject= https://doi.org/10.5852/ejt.2020.676 european journal of taxonomy 676: 1–15 (2020) 2 continuous distributional range, the genus shows high levels of local endemism exhibiting high degrees of ecological specialization, making it an ideal candidate to address hypotheses on speciation, patterns of diversification, and community assembly (atkins et al. 2001; bramley et al. 2004; clark et al. 2009; johnson et al. 2019). in recent years, several studies (wagner et al. 2001; bramley et al. 2003; atkins 2004; bramley 2005; lorence & perlman 2007; bone & atkins 2013; johnson 2017; kartonegoro et al. 2018; atkins et al. 2019; nishii et al. 2019) have led to an increase in numbers of species of cyrtandra. these studies emphasized the urgency to document and understand the biodiversity of cyrtandras before they succumb to anthropogenic pressures. however, little is still known about the taxonomy of members in areas which are considered centers of biodiversity for the genus. atkins et al. (2013) estimated a high species richness on borneo, in the philippines and on new guinea. understanding species boundaries of members from these areas is becoming increasingly important for examining biological trends of adaptation and speciation, and facilitating ecosystem and species conservation assessments. for the philippines, a comprehensive account of cyrtandra spp. was published by merrill (1922) in his an enumeration of philippine flowering plants noting 83 species, of which only c. oblongifolia c.b.clarke (de candolle & de candolle 1883) was listed as not endemic. atkins & cronk (2001) revised philippine cyrtandras from the island of palawan, describing seven new species and indicating that c. elatostemoides elmer (elmer 1913) is also found on borneo. both accounts were considered in an updated checklist of philippine cyrtandras available through ‘co’s digital flora of the philippines’ (pelser et al. 2011 onwards). a taxonomic revision of philippine cyrtandras, however, addressing species boundaries, distributions, and descriptions is yet to be achieved. atkins & cronk (2001) noted the striking vegetative similarities between c. villosissima merr. (merrill 1906) from the island of mindanao and a 1922 collection by merrill from the island of palawan which he labeled c. woodii merr. ined. with no accompanying publication. similarities include an erect suffrutescent habit, and large leaves that are slightly falcate and densely hirsute. however, in merrill’s 1922 enumeration of philippine plants the name c. woodii did not appear, and the distribution of c. villosissima was extended to palawan, which according to atkins & cronk (2001) seems to provide evidence for merrill’s decision to ‘sink’ his c. woodii into c. villosissima. increased sampling in the locality of c. woodii led to collections with reproductive structures that show the species’ distinctness from c. villosissima. this ultimately led to the description of c. hirtigera h.j.atkins & cronk (atkins & cronk 2001), favoring a new name to prevent confusion with the bornean c. woodsii b.l.burtt (burtt 1970). in the course of an ongoing research project, aimed at the taxonomic revision of philippine cyrtandras, it was found that c. ferruginea merr. (merrill 1915) and a series of collections from mindoro and the aurora province housed at the royal botanic garden edinburgh (rbge) share similar vegetative characters with c. villosissima. as a result, identification of several herbarium specimens was found to be intermediate between c. villosissima and c. ferruginea, without a clear distinction being possible between the two when no additional reproductive characters were present. in this paper, species sharing the character combination of an erect suffrutescent habit and large leaves that are slightly falcate and densely hirsute, are referred to as the c. villosissima group. members of this group are c. ferruginea, c. hirtigera, c. villosissima, and c. argentii sp. nov. our study aims at clarifying the differences between these species through keys and photographs, and provides a description and diagnosis for a new species often misidentified as either c. ferruginea, c. villosissima, or c. hirtigera. material and methods data for this study were derived from herbarium specimens including their corresponding field notes, photographs, and field observations. whenever available, living collections housed at the royal botanic garden edinburgh (rbge) were consulted and reproductive characters and measurements were olivar j.e.c. et al., cyrtandra argentii sp. nov. from the philippines 3 recorded from material preserved in alcohol. all philippine cyrtandra deposited at aah, bm, bo, e, gh, k, l, ny, p, pnh, and us were consulted through visits to these herbaria and access to digital images. herbaria acronyms follow index herbariorum (thiers, continuously updated). descriptions follow schemes of recently published accounts of new species (atkins & cronk 2001; johnson 2017; kartonegoro et al. 2018; atkins et al. 2019). assessment of conservation status was implemented using geocat (bachman et al. 2011), following the iucn red list category criteria (iucn standards and petitions subcommittee 2017). results cyrtandra ferruginea, c. villosissima, c. hirtigera, and the new species described here share the following characteristics: erect suffrutescent habit and large slightly falcate and densely hirsute opposite leaves. there exist minute differences in their vegetative characters and they are presented in the key. table 1 details more differences between the studied species. the studied species primarily differ, vegetatively, in color of indumentum and leaf symmetry. cyrtandra argentti sp. nov. is distinct among the three species by having a white indumentum, and c. ferruginea is distinct by having pronouncedly anisophyllous leaves. ultimately, the species are distinguishable by calyx and inflorescence type, corolla color and, to some degree, by geographic distribution (fig. 1). cyrtandra hirtigera is restricted c. argentii c. ferruginea c. hirtigera c. villosissima sp. nov. distribution sierra madre catanduanes, mt palawan negros island to mountain range to bulusan, mt isarog, mindanao island mindoro island mt mayon indumentum color white all throughout ferruginous crimson or pale white ferruginous leaf symmetry subequal anisophyllous subequal subequal inflorescence type compound cyme cyme cyme cyme inflorescence pedunculate pedunculate subsessile subsessile attachment no. of flowers per 10–15 1–3 4–many-flowered 4–many-flowered inflorescence calyx hirsuteness densely hirsute densely hirsute densely hirsute densely hirsute externally, glabrous externally, with externally, with internally glandular hairs scattered glandular internally at base hairs internally and of lobes prominent tufts of hairs at the base calyx lobes shape acuminate acuminate acute or narrowly linear acuminate corolla color white white dull reddish orange red or yellowish green corolla hirsuteness glabrous externally densely hirsute, glandular hairs densely hirsute and internally with glandular hairs externally and internally internally calyx persistence persistent entirely persistent entirely not persistent persistent enclosing the fruit enclosing the fruit style persistence persistent persistent not persistent persistent table 1. diagnostic characters separating the four studied species. european journal of taxonomy 676: 1–15 (2020) 4 to the island of palawan and c. villosissima is found in mindanao extending to negros island. only c. ferruginea and c. argentii sp. nov. occur on luzon island. class magnoliopsida brongn. (brongniart 1843) order lamiales bromhead (bromhead 1838) family gesneriaceae rich. & juss. (de candolle 1816) genus cyrtandra j.r.forst. & g.forst. (forster & forster 1775) key to the studied species 1. mature leaves anisophyllous (i.e., smaller leaves less than half the length of the larger leaves in a pair) ..................................................................................................................... c. ferruginea merr. – mature leaves subequal ..................................................................................................................... 2 2. indumentum white; inflorescences pendulous and pedunculate ......................................................... ........................................................................... c. argentii olivar, h.j. atkins & muellner sp. nov. – indumentum ferruginous; inflorescences erect and subsessile ......................................................... 3 3. calyx divided almost to the base; corolla red .................................................... c. villosissima merr. – calyx fused for half or more of its length; corolla yellowish green or dull reddish orange ............... ......................................................................................................... c. hirtigera h.j.atkins & cronk cyrtandra argentii olivar, h.j.atkins & muellner sp. nov. urn:lsid:ipni.org:names:77209562-1 figs 1, 2, 3a, 4a diagnosis the species’ pendulous compound cymose inflorescences (10–15 flowers) distinguish it from all other members of the genus in the philippines. the combination of subequal leaves, white woolly indumentum, glabrous corolla, and ovoid fruit separates this species from the rest of the c. villosissima group. etymology this species is named after george argent who was part of the team that collected specimens at the type locality. george’s contribution to our knowledge of the philippine flora is undisputed. his extensive fieldwork in the country has led to the discovery of several new species, recognition of important conservation areas, and promotion of biodiversity studies. material examined type philippines • mindoro island, oriental province, mt halcon; 600 m; 13 mar. 1997; mendum, m., argent, c.g.c., pennington, r.t., wilkie, p., reynoso, e.j., gaerlan, f. 29053 (holotype: e!; isotype: pnh). additional material philippines • mt halcon, mindoro; 12 may 1986; c.e. ridsdale 1762 (k000223279, l.2822762) • ibid.; 1 apr. 1991; stone, reynoso, sagcal 504 (k000223280, l.2822797, us00737625) • ibid.; 13 mar. 1997; argent, gaerlan, reynoso 20053 (l.3805692) • sierra madre mountains, baler aurora; 25 mar. 1968; jacobs, m. 8002 (l.2822694) • aurora national park; 8 mar. 1993; barbon, garcia, fernando http://www.ipni.org/urn:lsid:ipni.org:names:77209562-1 olivar j.e.c. et al., cyrtandra argentii sp. nov. from the philippines 5 9121 (k000223281, l.3794225) • llavac, infanta, quezon province; 25 jun. 1955; lagrimas, m. 521 (l.2822646). description an erect suffrutescent plant up to 3 m in height. stems terete or slightly grooved, with white woolly hairs throughout. leaves opposite, subequal; petioles 4–7 cm long, densely hirsute; blades 13–30 × 7–15 cm, oblong to oblong-elliptic, slightly falcate, apex attenuate, base rounded to oblique, pronouncedly asymmetrical, not decurrent, margins denticulate, 10–12 pairs of lateral veins, curving and uniting at the margins, densely hirsute on both sides. inflorescences compound cymes, axillary, pendulous, pedunculate, with 10–15 flowers; peduncle 5–6 cm, densely hirsute; bracts green, ca 9 × 2 mm, lanceolate, densely hirsute on both surfaces, persistent; bracteoles on every point of branching, lanceolate, green, ca 5 × 2 mm, densely hirsute on both surfaces; pedicels 3–5 cm long, densely hirsute. calyx tubular, pale green, ca 15 mm long, upper lobes ca 4 mm long, lower lobes ca 6 mm long, acuminate, densely hirsute externally, glabrous internally. corolla white, ca 30 mm long, funnel-shaped, lobes suborbicular, upper lobes, 10–12 × 5 mm, lateral lobes 7 × 7 mm, lower lobes 8 × 8 mm, 3–4 × 1–2 mm; glabrous externally and internally, lobes slightly recurved. stamens 2; filaments ca 12 mm long, attached ca 17 mm from base of corolla, glabrous; anthers ca 2.5 mm long, thecae parallel, coherent at apices; staminodes 3, lateral staminodes ca 3 mm long, central staminode ca 0.5 mm long. gynoecium ca 20 mm long overall; disc cupular with entire margin, ca 1.5 mm long, glabrous; ovary ca 6 mm long, glabrous, with some eglandular hairs towards base of style; style ca 14 mm long, with eglandular hairs throughout; stigma fig. 1. map of the philippines showing known distributions of cyrtandra argentii olivar, h.j.atkins & muellner sp. nov. (★), c. ferruginea merr. (■), c. hirtigera h.j.atkins & cronk (◆), and c. villosissima merr. (●) based on collection localities. a single point may represent more than one collection. european journal of taxonomy 676: 1–15 (2020) 6 fig. 2. cyrtandra argentii olivar, h.j.atkins & muellner sp. nov. a. flower, lateral view. b. detail of upper leaf surface. c. habit. d. inflorescence. e. calyx, longitudinal section. f. corolla, longitudinal section. g. gynoecium. h. fruit enclosed by the persistent calyx. drawn from mendum et al. 29053 deposited at e. habit, inflorescence, fruit and leaf indumentum drawn from dried material. flower parts all from material preserved in alcohol. drawing by claire banks. olivar j.e.c. et al., cyrtandra argentii sp. nov. from the philippines 7 bilobed, ca 2.5 mm across. fruits ovoid, green, glabrous, verrucose, ca 12 × 5 mm, excluding the style; calyx persistent and entirely enclosing the fruit, style persistent. distribution and habitat cyrtandra argentii sp. nov. is found growing on slopes near streams in primary forests. this species is distributed from the north of luzon to the island of mindoro. conservation status cyrtandra argentii sp. nov. occurs at an elevation of 600–800 m a.s.l. which corresponds to the forest land use zone (villanueva & buot jr 2018). using the online geocat conservation assessment tool (http://geocat.kew.org/), the proposed conservation category based on extent of occurrence (eoo) is near threatened (nt), and the category based on the estimated area of occupancy (aoo) calculated using the default 2 × 2 km grid is endangered (en). here, we consider this species’ status as nt due to: i) its occurrence in close proximity to the agroforest land use zone, the latter at approximately 100–400 m a.s.l. (villanueva & buot jr 2018); and ii) the fact that the forest areas wherein the species occurs are not declared protected by law (biodiversity management bureau 2015), making it highly susceptible to population decline through deforestation and other anthropogenic activities. notes like many species of cyrtandra, filaments of c. argentii sp. nov. recoil into the corolla tube after anther dehiscence. this is hypothesized as constituting a mechanism against self-pollination (bramley et al. 2003). the length of the style also varies developmentally, the style can be either exserted or inserted depending on the stage of maturity of the flower. cyrtandra ferruginea merr. (merrill 1915) figs 1, 3b, 4b material examined type philippines • luzon, camarines, mt cauayan; 9 dec. 1913; phil. pl. ramos 1548 (syntypes: bm!, gh!, ny!, p!, us!). additional material philippines • mt isarog, camarines sur; aug. 1915; ramos 23554 (us00081328) • ibid.; 22 mar. 1997; mendum et al. 29182 (e00057041) • ibid.; 23 mar. 1997; argent et al. 20182 (l.3805694) • mt bulusan, sorsogon; dec. 1915; elmer 16074 (l.2818244, us00081329, u.1341267, p03884333) • ibid.; 19 jun. 1958; sinclair 9624 (e00631523) • mt mayon, albay; 15 nov. 1991; reynoso, romero & fuentes 3584 (e00316099) • catanduanes; 11 dec. 1917; ramos 30288 (us00081330, p03884332) • mt malinao, albay; 29 oct. 1995; reynoso, sagcal & fernando 21406 (l.3805666). description an erect suffrutescent plant up to 1 m in height. stems terete with ferruginous hairs all throughout. leaves opposite, anisophyllous; petioles 4–7 cm long, densely hirsute; blades ca 20 × 12 cm, oblong to oblong-elliptic slightly falcate, apex acute or slightly acuminate, base acute or rounded, pronouncedly asymmetrical, not decurrent, margins denticulate, 10 pairs of lateral veins, curving and uniting at the margin, densely hirsute on both sides; blades of smaller leaves of a pair 6.5–9 × 2–2.5 cm, resembling the major leaves in all other respects. inflorescences cymose, axillary, erect, pedunculate, with 1–3 flowers; peduncle 2–3 cm long, densely hirsute; bracts green, ca 10 × 1 mm, linear lanceolate, densely hirsute http://geocat.kew.org/ european journal of taxonomy 676: 1–15 (2020) 8 fig. 3. leaf similarities. a. c. argentii olivar, h.j.atkins & muellner sp. nov. b. c. ferruginea merr. c. c. hirtigera h.j.atkins & cronk. d. c. villosissima merr. photos taken from co’s digital flora with permission (pelser et al. 2011 onwards). olivar j.e.c. et al., cyrtandra argentii sp. nov. from the philippines 9 fig. 4. inflorescence types. a. c. argentii olivar, h.j.atkins & muellner sp. nov. b. c. ferruginea merr. c. c. hirtigera h.j.atkins & cronk. d. c. villosissima merr. a, b & d from co’s digital flora with permission (pelser et al. 2011 onwards). c from a living collection in rbge. on both surfaces, persistent. calyx tubular, pale green, 20–30 mm long, upper lobes ca 3 mm long, lower lobes ca 7 mm long, acuminate, densely hirsute externally, with glandular hairs internally at base of lobes. corolla white, 50–65 mm long, funnel-shaped, upper lobes rounded, ca 8 × 9 mm, lower and lateral lobes rounded, ca 5 × 6 mm, densely hirsute, with glandular hairs internally, lobes slightly recurved. stamens 2; filaments ca 10 mm long, attached ca 18 mm from base of corolla, sparsely covered with glandular hairs; anthers ca 1.5 mm long, thecae parallel, coherent at apices; staminodes 3, lateral staminodes ca 4 mm long, central staminode ca 1 mm long. gynoecium ca 25 mm long overall; disc cupular with undulate margin, ca 2 mm long, glabrous; ovary 8–9 mm long, with glandular hairs throughout; style ca 12 mm long, with glandular hairs throughout; stigma bilobed, ca 1 mm across. fruits lanceolate, green, hirsute, verrucose, 30–40 × 6 mm; calyx persistent and entirely enclosing the fruit, style persistent. distribution and habitat cyrtandra ferruginea is found growing in damp forests at approximately 500–800 m a.s.l. and can be found on catanduanes, mt isarog, mt mayon, mt malinao and mt bulusan (fig. 1). notes cyrtandra ferruginea is morphologically most similar to c. argentii sp. nov., but can be separated by the following characters: ferruginous anisophyllous leaves, 1–3 flowered simple cymes, and hirsute corolla. european journal of taxonomy 676: 1–15 (2020) 10 cyrtandra hirtigera h.j.atkins & cronk (atkins & cronk 2001) figs 1, 3c, 4c material examined type philippines • palawan, cleopatra’s needle; 29 jan. 1998; cronk et al. 25433 (holotype: pnh!; isotypes: e!, k!, l!). additional material philippines • san vicente, palawan; 4 aug. 1988; soejarto & madulid 6353 (l.2818243) • mt beaufort, palawan; 12 mar. 1984; ridsdale smhi 23 (l.2818051) • pagdanan range, palawan; 22 apr. 1984; podzorski smhi 934 (l.2818049) • mt mantalingahan, palawan; 5 may 1948; edaño 122 (l.2818048) • ibid.; 4 mar. 1992; argent & romero 9666 (l.3805814) • malampaya bay, palawan; oct. 1922; merrill 11573 (us00081506) • mt capoas, palawan; apr. 1913; merrill 9500 (us00081487) • palawan; apr. 1906; foxworthy 581 (us00081485, p03899657) • taytay, palawan; 31 jan. 1991; stone 327 (l.2822680). description an erect suffrutescent plant, up to 2–2.5 m in height. stems terete, with crimson or white hairs throughout. leaves opposite, subequal; petioles 5–8 cm long, densely hirsute; blades 22–30 × 15–18 cm, broadly elliptic, slightly falcate, apex acuminate, base cuneate, not decurrent, margins denticulate, 12–14 pairs of lateral veins, curving and uniting at the margins, densely hirsute on both sides. inflorescences cymous, axillary, erect, subsessile, with 4–many flowers; peduncle 4–5 mm, densely hirsute; bracts green, ca 1 cm × 5 mm, lanceolate, densely hirsute on both surfaces, persistent; bracteoles up to 5 mm long, densely hirsute on both surfaces; pedicels ca 5 mm long, densely hirsute. calyx tubular, red or green, ca 1–1.5 cm long, lobes ca 5 mm long, lower lobes ca 6 mm long, acute or narrowly acuminate, densely hirsute externally, with scattered glandular hairs internally and has prominent tufts of hairs at the base. corolla dull reddish orange or yellowish green, ca 2 cm long, funnel-shaped, lobes slightly bilabiate or subequal, upper lobes, 1–1.5 × 2 mm, lateral lobes 1 × 1.5 mm, lower lobes 2 × 1.5 mm; glandular hairs externally and internally. stamens 2; filaments ca 1–1.5 cm long, attached ca 12–13 mm from base of corolla, glabrous; anthers ca 2 mm long, thecae parallel, coherent at apices; staminodes 2, 5–8 mm long. gynoecium ca 20 mm long overall; disc cupular with undulate margin, ca 1.5 mm long, glabrous; ovary ca 4–5 mm long, glabrous; style ca 10 mm long, with eglandular hairs throughout; stigma bilobed, ca 2.5 mm across. fruits ovoid, green, glabrous, ca 1 cm × 5 mm; inflorescence bracts persistent, calyx and style not persistent. distribution and habitat cyrtandra hirtigera is distributed throughout the island of palawan and is usually found on slopes near gullies at 30–900 m a.s.l. notes atkins & cronk (2001) described two varieties of this species, c. hirtigera var. hirtigera and c. hirtigera var. chlorina, distinguishable by color and shape of their calyces and corolla limbs. cyrtandra hirtigera var. hirtigera has a crimson indumentum, red calyces with acute lobe apices, and reddish orange corollas with slightly bilabiate limbs. cyrtandra hirtigera var. chlorina has pale indumentum, green calyces with acuminate lobe apices, and yellowish green corollas with subequal lobes. olivar j.e.c. et al., cyrtandra argentii sp. nov. from the philippines 11 cyrtandra villosissima merr. (merrill 1906) figs 1, 3d, 4d material examined type philippines • mindanao, camp keithley, lake lanao; jan. 1906; mrs clemens 51 (holotype: aah!; isotypes: f!, us!). additional material philippines • mt malindang, misamis occidental; 16 mar. 2004; opiso et al. 2088 (l.3794253) • ibid.; may 1993; gaerlan et al. 10925 (k000184579, l.3794112) • mt hibok-hibok, camiguin; 1999; rbge & pnh 48 (e00743749) • lake balunsasayao, negros oriental; 11 sep. 1953; britton 357 (l.2826671) • cuernos mountains, negros oriental; 13 may 1948; edaño 7393 (aah00092001) • ibid.; mar. 1908; elmer 9511 (l.2826672, us00081486) • mahilucot river, bukidnon; jul. 1920; ramos & edaño 38649 (l.2826670, us00081488) • mt daho, jolo; sep. 1924; ramos & edaño 43913 (p03899658). description an erect suffrutescent plant up to 7 m in height. stems terete or slightly grooved, with ferruginous woolly hairs throughout. leaves opposite, subequal; petioles 3–5 cm long, densely hirsute; blades 11–20 × 3.5–8 cm, oblong-ovate to ovate-lanceolate, slightly falcate, apex acuminate, base acute or acuminate, pronouncedly asymmetrical, not decurrent, margins denticulate, 12–14 pairs of lateral veins, curving and uniting at the margins, densely hirsute on both sides. inflorescences cymous, axillary, erect, subsessile, with 4–many flowers; peduncle 4–5 mm, densely hirsute; bracts green, ca 1 cm, linear, densely hirsute on both surfaces, persistent; bracteoles up to 5 mm long, densely hirsute on both surfaces; pedicels ca 5 mm long, densely hirsute. calyx tubular, pale green, lobes linear ca 1.5 cm × 1 mm, densely hirsute. corolla red, ca 18 mm long, funnel-shaped, lobes orbicular-ovate ca 4 mm long, densely hirsute. stamens 2; filaments ca 2 mm long; anthers ca 2.5 mm long, thecae parallel, coherent at apices. gynoecium ca 20 mm long overall, densely hirsute; disc cupular, glabrous; style densely hirsute. fruits oblong, green, densely hirsute, ca 1 cm × 4.5 mm; calyx and style persistent. distribution and habitat cyrtandra villosissima is distributed throughout the island of mindanao and extends to the island of negros in the visayas and is usually found in well-shaded areas near ravines. notes cyrtandra villosissima is vegetatively similar to c. hirtigera. it is distinguishable by its red corolla and green calyces with distinctly linear lobes. based on available distribution data, c. hirtigera appears to be restricted to the island of palawan while c. villosissima can be found from negros island to the island of mindanao. discussion the recognition of c. argentii sp. nov. as a new species was here aided by increased availability of collections with reproductive parts and continued alpha-taxonomic work. this highlights the importance of reproductive characters in establishing species boundaries among cyrtandra species. in phylogenetic analyses of the southeast asian cyrtandra (atkins et al. 2020), the c. villosissima group was not resolved as monophyletic; only c. villosissima and c. hirtigera belonged to the same subclade, but were not resolved as exclusive sister taxa. the character combination of erect suffrutescent habit with large leaves that are slightly falcate and densely hirsute has evolved at least three times independently (atkins european journal of taxonomy 676: 1–15 (2020) 12 et al. 2020). figure 3 shows this shared character combination and fig. 4 shows the inflorescence type that ultimately distinguishes the species from each other. the genus cyrtandra is the most taxonomically challenging in the gesneriaceae due to its large number and high proportion of poorly known and undescribed species (burtt 2001; atkins et al. 2013; clark et al. 2013). atkins et al. (2013) estimated 800 species of cyrtandra. since then, several authors (bone & atkins 2013; johnson 2017; kartonegoro et al. 2018; atkins et al. 2019) have described additional species. the number of species is expected to increase further as more alpha-taxonomic work and field collection are carried out. large genera can be systematically addressed by following a phylogenetically informed taxonomic approach on a region-by-region basis (atkins et al. 2013; clark et al. 2013). this has been applied effectively by bramley (2005) in a revision of cyrtandra section dissimiles in borneo. the approach involves a robustly sampled phylogenetic tree wherein monophyletic clades can be characterized morphologically by one or more salient characters (atkins et al. 2013; clark et al. 2013). taxa with molecular data can be assigned to the clades while taxa lacking molecular data can be tentatively assigned based on morphological similarities. upon assignment to a clade, taxonomic assessment can be streamlined by focusing on related taxa identified through both morphological and molecular data, therefore limiting the number of potential conspecifics for comparison. areas, particularly archipelagos, with high diversity can benefit from this approach since it provides a systematic way of prioritizing areas where additional fieldwork and alpha-taxonomic work are most needed. clark et al. (2013) suggest that phylogenetically defined areas can be addressed taxonomically first, followed by increased sampling in lesser-resolved areas. this strategy is currently being applied to study the genus in the philippines with the aim of producing a complete revision of cyrtandra in the archipelago (olivar et al., in preparation). acknowledgements this research received support from the synthesys project http://www.synthesys.info/ which is financed by european community research infrastructure action under the fp7 ‘capacities’ program. financial support for this study was also provided by the deutscher akademischer austauschdienst (daad project no. 91690870) to jeco and anm-r, and the federal ministry of education and research (bmbf project no. 16gw0120k) to anm-r. mr danilo tandang from pnh is acknowledged for his help with the types. curators of aah, bm, bo, e, gh, k, l, ny, p, pnh, and us are acknowledged for their assistance during visits, loans, and digital images. nathan 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https://doi.org/10.13057/biodiv/d190624 wagner w.l., wood k.r. & lorence d.h. 2001. a new species of cyrtandra (gesneriaceae) from kaua’i, hawaiian islands. novon 11 (1): 146–152. https://doi.org/10.2307/3393224 https://doi.org/10.3897/phytokeys.91.21623 https://doi.org/10.1016/j.ympev.2017.07.004 https://doi.org/10.1111/jbi.13567 https://doi.org/10.1017/s0960428618000045 https://doi.org/10.3417/1055-3177(2007)17[357:ansocg]2.0.co;2 https://doi.org/10.1017/s0960428619000106 http://www.philippineplants.org http://sweetgum.nybg.org/science/ih/ https://doi.org/10.13057/biodiv/d190624 https://doi.org/10.2307/3393224 olivar j.e.c. et al., cyrtandra argentii sp. nov. from the philippines 15 manuscript received: 2 september 2019 manuscript accepted: 6 january 2020 published on: 30 june 2020 topic editor: frederik leliaert desk editor: connie baak printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. calidolipeurus, new genus for lipeurus megalops piaget, 1880 (phthiraptera: ischnocera: oxylipeurus-complex), with a redescription of the type species and a preliminary key to the oxylipeurus-complex daniel r. gustafsson 1,*, lujia lei 2 & fasheng zou 3 1,2,3 guangdong key laboratory of animal conservation and resource utilization, guangdong public laboratory of wild animal conservation and utilization, guangdong institute of applied biological resources, guangzhou, guangdong, china. * corresponding author: kotatsu@fripost.org 2 email: leilujia456@163.com 3 email: pzoufs@giabr.gd.cn 1 urn:lsid:zoobank.org:author:8d918e7d-07d5-49f4-a8d2-85682f00200c 2 urn:lsid:zoobank.org:author:8f6d34b8-cd2d-42ee-8c17-695a22016fd8 3 urn:lsid:zoobank.org:author:a0e4f4a7-cf40-4524-aaae-60d0ad845479 abstract. the chewing louse species lipeurus megalops piaget, 1880, is redescribed and illustrated. this species has previously been placed in the genus oxylipeurus mjöberg, 1910, but marked differences in preantennal structure, male and female genitalia, abdominal chaetotaxy, and structure of abdominal plates indicate that this species is not closely related to other species in this genus. we therefore erect a new genus, calidolipeurus gen. nov. for this species. calidolipeurus is presently monotypic, containing only calidolipeurus megalops gen. et comb. nov. we also provide a preliminary key to the oxylipeuruscomplex. keywords. phthiraptera, ischnocera, oxylipeurus-complex, new genus, redescription. gustafsson d.r., lei l. & zou f. 2020. calidolipeurus, new genus for lipeurus megalops piaget, 1880 (phthiraptera: ischnocera: oxylipeurus-complex), with a redescription of the type species and a preliminary key to the oxylipeurus-complex. european journal of taxonomy 686: 1–15. https://doi.org/10.5852/ejt.2020.686 introduction the first comprehensive review of the oxylipeurus-complex was by clay (1938), who considered all species to belong to the same, morphologically variable, genus, oxylipeurus mjöberg, 1910. within this genus, she designated six species groups, based on rough similarities in morphology. the third group comprised four species parasitizing small-bodied partridges in southeast asia, including lipeurus megalops piaget, 1880, parasitizing the crested partridge, rollulus roulroul (scopoli, 1786). kéler (1958) considered this group distinct from oxylipeurus s. str. and proposed the subgeneric name megalipeurus kéler, 1958. most subsequent treatments of the oxylipeurus-complex have followed clay (1938) and hopkins & clay (1952), and considered megalipeurus and most other proposed genera in the oxylipeurus-complex to be synonyms of oxylipeurus (e.g., ledger 1980; price et al. 2003) european journal of taxonomy 686: 1–15 issn 2118-9773 https://doi.org/10.5852/ejt.2020.686 www.europeanjournaloftaxonomy.eu 2020 · gustafsson d.r. et al. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:ac52c43b-deb7-414b-b905-18d68f1d9dd9 1 mailto:kotatsu%40fripost.org?subject= mailto:leilujia456%40163.com?subject= mailto:zoufs%40giabr.gd.cn?subject= http://zoobank.org/urn:lsid:zoobank.org:author:8d918e7d-07d5-49f4-a8d2-85682f00200c http://zoobank.org/urn:lsid:zoobank.org:author:8f6d34b8-cd2d-42ee-8c17-695a22016fd8 http://zoobank.org/urn:lsid:zoobank.org:author:a0e4f4a7-cf40-4524-aaae-60d0ad845479 https://doi.org/10.5852/ejt.2020.686 https://doi.org/10.5852/ejt.2020.686 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:ac52c43b-deb7-414b-b905-18d68f1d9dd9 however, in recent decades, redescriptions of several species in the oxylipeurus-complex have been published (mey 1990, 2010), including that of the type species oxylipeurus inaequalis (piaget, 1880). these indicate that oxylipeurus s. str. is morphologically very different from most other species in the oxylipeurus-complex, and that several genera commonly synonymized with oxylipeurus (e.g., price et al. 2003) should be accepted as valid. mey (2009, 2010) has recently considered the genus megalipeurus as a valid genus, including the species l. megalops. we agree that all of the genera considered synonyms of oxylipeurus in the recent checklist of price et al. (2003) should be accepted as valid, following mey (2009, 2010) and gustafsson et al. (2020). this includes the recognition of the following genera considered synonymous with oxylipeurus in the checklist of price et al. (2003): eiconolipeurus carriker, 1945, epicolinus carriker, 1945, megalipeurus kéler, 1958, reticulipeurus kéler, 1958 and talegallipeurus mey, 1982. however, mey (2009) includes the species l. megalops in megalipeurus, following its placement in group iii by clay (1938) and in megalipeurus by kéler (1958). oxylipeurus megalops is similar to other species in megalipeurus in the presence of elongated and strongly sclerotized coni and in the general shape of the female subgenital plate, but it differs from all other species in the oxylipeurus-complex in many significant characters (see below). we therefore here propose a new genus, calidolipeurus gen. nov. for l. megalops. material and methods all examined specimens are deposited at the natural history museum, london, united kingdom (nhmuk). drawings were made using a drawing tube attached to a nikon eclipse ni (nikon corporation, tokyo, japan) and edited in gimp (www.gimp.org). measurements (all in mm) were made from live images in nis-elements (nikon corporation, tokyo, japan) for the following dimensions: hl = head length (along midline); hw = head width (at temples); pow = preocular width (at preantennal nodi); prw = prothoracic width; ptw = pterothoracic width; aw = abdominal width (at segment v); tl = total length (along midline, including stylus). terminology of morphological and setal characters and abbreviations thereof follow gustafsson & bush (2017). abbreviations used (following gustafsson & bush 2017 and gustafsson et al. 2020): al = anterior lobes; as1–2 = anterior setae 1–2; avs3 = anterior ventral seta 3; c = conus; cl = “claspers”; cls = clypeo-labral suture; dps = dorsal preantennal suture; dpas = dorsal post-antennal suture; e = eye; es = epistomal suture; hm = hyaline margin; lavp = lateral accessory vulval plates; mds = mandibular seta; mts3 = marginal temporal seta 3; os = ocular seta; pos = preocular seta; psps = principal postspiracular setae; pst1–2 = parameral setae 1–2; s1–7 = sensilla 1–7; sgp = subgenital plate; smc = secondary marginal carina; ss = sutural setae; sty = stylus; vm = vulval margin; vsms1–2 = ventral submarginal setae 1–2; y = y-shaped thickening. results systematics order phthiraptera haeckel, 1896 suborder ischnocera kellogg, 1896 family philopteridae burmeister, 1838 oxylipeurus-complex calidolipeurus gen. nov. urn:lsid:zoobank.org:act:e0395b9a-62ad-4a56-be5b-c99cd210e7d6 type species lipeurus megalops piaget, 1880. european journal of taxonomy 686: 1–15 (2020) 2 www.gimp.org http://zoobank.org/urn:lsid:zoobank.org:act:e0395b9a-62ad-4a56-be5b-c99cd210e7d6 diagnosis based on morphological similarities, calidolipeurus is likely most closely related to megalipeurus, reflecting the close relationship between the hosts of the two louse genera (wang et al. 2013). these two genera share the following characters: coni elongated in both sexes (figs 5, 7); female subgenital plate with deep median indentation on anterior margin (fig. 4); female vulval margin convex (fig. 4); dorsal preantennal suture present (fig. 5); tergopleurites ii–vii medianly separated in both sexes (figs 1–2); tergopleurites ix + x and xi fused (figs 1–2). calidolipeurus can be separated from megalipeurus by the following characters: marginal carina interrupted only near as1 in megalipeurus, but interrupted near as1 and as2 in calidolipeurus (fig. 5); dorsal preantennal suture transversal in megalipeurus, but longitudinal in calidolipeurus (fig. 5); figs 1–2. calidolipeurus megalops (piaget, 1880) gen. et comb. nov., based on specimens on slide nhmuk010682491 (nhmuk). 1. habitus, ♂, dorsal and ventral views. 2. habitus, ♀, dorsal and ventral views. gustafsson d.r. et al., new genus calidolipeurus (phthiraptera) 3 postmarginal carina extended medianly in calidolipeurus (fig. 5), but not in megalipeurus; eyes gigantic in calidolipeurus (fig. 5), but not in megalipeurus; preocular nodus present in megalipeurus but absent in calidolipeurus (fig. 5); stylus reaches beyond distal margin of abdomen in calidolipeurus (fig. 3), but not in megalipeurus; ss present on male abdominal segments vi–vii in megalipeurus, but absent in calidolipeurus (fig. 1); psps present and of equal size on male tergopleurites iv–vi in megalipeurus, but present on tergopleurites iv–vii in calidolipeurus (fig. 1) with those of tergopleurites vi–vii microsetae; male subgenital plate with lateral extensions following genital opening in calidolipeurus (fig. 3), but without such extensions in megalipeurus; parameres very broad proximally, with overall shape roughly triangular in calidolipeurus (fig. 12), but with slender heads in megalipeurus; proximal mesosome with large rounded lobes on each side in megalipeurus, but without such lobes in calidolipeurus (fig. 9); gonopore with lateral hook-shaped elongations in calidolipeurus (fig. 11), but roughly rounded in megalipeurus. figs 3–4. calidolipeurus megalops (piaget, 1880) gen. et comb. nov., based on specimens on slide nhmuk010682491 (nhmuk). 3. male subgenital plate and terminalia, ventral view. 4. female subgenital plate and terminalia, ventral view. european journal of taxonomy 686: 1–15 (2020) 4 etymology calidolipeurus is constructed from the latin ‘calidus’ for ‘white spot in the forehead’ and the genus name lipeurus nitzsch, 1818, commonly used for long and slender ischnocerans. this refers to the small, elongated dorsal preantennal suture of the type species, which appears white in many specimens in contrast to the otherwise brownish head. gender: masculine. figs 5–12. calidolipeurus megalops (piaget, 1880) gen. et comb. nov., based on specimens on slide nhmuk010682491 (nhmuk). 5. male head, dorsal and ventral views. 6. male antennae, ventral view. 7. female antenna, ventral view. 8. male genitalia (mesosome everted), dorsal view. 9. male mesosome (everted), ventral view. 10. male mesosome (not everted), dorsal view. 11. male mesosome (not everted), ventral view. 12. male paramere, dorsal view. male and female antenna at same scale as male head; all genitalic elements at same scale. gustafsson d.r. et al., new genus calidolipeurus (phthiraptera) 5 description both sexes overall body shape elongated, ‘lipeuroid’ (figs 1–2). head longer than wide (fig. 5). marginal carina widening considerably anteriorly, interrupted laterally near sites of as1 and as2. frons slightly protruding; marginal carina at frons with serrated posterior margin. dorsal preantennal suture narrow, median, longitudinal. ventral carina uninterrupted, indistinct. head chaetotaxy as in fig. 5; avs3 anterior, near vsms1–2; mds not visible in all examined specimens; s1–3, s5 and s7 present; pos posterior to eye; mts3 only macroseta. coni elongated. antennae sexually dimorphic (figs 5–7). eyes large. preocular nodus absent. marginal temporal carina slender. dorsal postantennal suture absent. thoracic and abdominal segments as in figs 1–2. legs as in figs 13–18; similar between sexes but some setae more slender in female than in male. mesoand metasterna fused. metepisterna long, with extensive striation. tergopleurites ii–vii medianly separated; tergopleurite viii medianly continuous in posterior end; tergopleurites ix + x and xi fused. sternites medianly continuous, reticulated in both sexes. subgenital plates differ between sexes, formed by fusion of sternites viii–ix. figs 13–18. calidolipeurus megalops (piaget, 1880) gen. et comb. nov., based on specimens on slide nhmuk010682494 (nhmuk). 13. male leg i, dorsal view. 14. male leg i, ventral view. 15. male leg ii, dorsal view. 16. male leg ii, ventral view. 17. male leg iii, dorsal view. 18. male leg, iii, ventral view. european journal of taxonomy 686: 1–15 (2020) 6 male pedicel and flagellomere i with rugose antero-dorsal surface; flagellomere with antero-dorsal claw-like elongation. thoracic and abdominal chaetotaxy as in fig. 1; ss present on segments ii–v; psps present on segments iv–vii, those of vi–vii microsetae. tergopleurite ix–xi with variable lateral incision reaching apertures of setae. subgenital plate with extensive reticulation (fig. 3), distally elongated into protruding stylus; lateral extensions on each side of base of stylus associated with anterior margin of ventral genital opening. basal apodeme elongated, of roughly equal width, but in some specimens with concave lateral margins (fig. 8). mesosome largely unsclerotized, and looks different in specimens with everted or non-everted genitalia. proximal mesosome flattened to concave, but diffuse and here illustrated approximately (figs 8–11); on each side sinusoid thickening articulating with parameral heads. distal mesosome roughly quadratic, with extensive folds and serrations on dorsal and ventral surfaces. internal sclerite roughly heart-shaped (fig. 10). gonopore longer than wide, with lateral hook-shaped in distal end; in anterior end a dark central sclerite may be proximal part of endophallus; one small aperture on each side of presumed endophallus may be microsetae, but setae not visible in examined specimens. in everted genitalia, the gonopore extends beyond distal margin of mesosome, and rugose areas of mesosome contracted (figs 8–9). parameres roughly triangular (fig. 12), median corners of heads almost touching anteriorly; median margin of anterior parameres weakly sclerotized; pst1 not visible; pst2 microsetae near distal end of paramere. female pedicel and flagellomere i without rugose areas and without claw-like elongation. head chaetotaxy similar to male, but s7 shorter. thoracic and abdominal chaetotaxy as in fig. 2; ss present on segments ii–v; psps present on segments iv–vii; those on vi–vii microsetae. no lateral incision of tergopleurite ix–xi in examined specimens. subgenital plate formed by fused sternites viii–ix (fig. 4); proximal margin deeply indented medianly. vulval margin strongly convex; setae situated anterior to margin and do not form distinct sets. subvulval sclerites absent. lateral and posterior margins of abdominal segment ix–xi with more or less equally spaced and equal-sized setae. host distribution only known from the type host of the type species, rollulus roulroul (scopoli, 1786) (galliformes: phasianidae). geographical range indo-malayan region, both mainland and the indonesian archipelago. calidolipeurus megalops (piaget, 1880) gen. et comb. nov. figs 1–18 lipeurus megalops piaget, 1880: 675. esthiopterum megalops – harrison 1916: 138. oxylipeurus megalops – clay 1938: 166. oxylipeurus (megalipeurus) megalops – kéler 1958: 327 [inferred]. megalipeurus megalops – mey 2009: 162 [inferred]. type host rollulus roulroul (scopoli, 1786) – crested partridge (phasianidae). gustafsson d.r. et al., new genus calidolipeurus (phthiraptera) 7 type locality madagascar [= in error]. known from southeast asia (thailand, peninsular malaysia, borneo). material examined non-type material borneo • 4 ♂♂, 9 ♀♀; r. meinertzhagen, 10905; nhmuk-010682491; nhmuk • 6 ♂♂, 10 ♀♀; r. meinertzhagen, 10891; nhmuk-010682483; nhmuk • 2 ♂♂, 1 ♀♀; jan. 1901; r. meinertzhagen, 3655; nhmuk-010682490; nhmuk. malaysia • 1 ♂, 1 ♀; terengganu [as trengganu]; 140 ft a.s.l.; 102°0′ e, 5°28′ n; 26 feb. 1974; gn. lawit expedition, brit. mus. 1974-2; nhmuk-010682494; nhmuk. thailand • 1 ♂♂, 4 ♀♀; 1939; r. meinertzhagen, 17661; nhmuk-010682866; nhmuk. description see genus description. male lobes of genital opening with 3–5 mesosetae and 1–3 short setae on each side. stylus with 12–16 microsetae ventrally or laterally (some situated near base of stylus). measurements (n = 14, except tl where n = 13): tl = 1.84–2.01 (1.93); hl = 0.44–0.50 (0.47); pow = 0.25–0.28 (0.27); hw = 0.26– 0.29 (0.28); prw = 0.21–0.24 (0.22); ptw = 0.35–0.41 (0.38); aw = 0.34–0.41 (0.37). female proximal margin of subgenital plate typically with two setae on each side, but placement asymmetrical and 1–3 setae may be present on each side; lateral setae about twice as long as median setae. vulval margin with 10–15 slender setae on each side (fig. 4). measurements (n = 25, except aw where n = 24): tl = 2.11–2.31 (2.21); hl = 0.49–0.53 (0.51); pow = 0.25–0.29 (0.27); hw = 0.28–0.32 (0.30); prw = 0.22–0.24 (0.23); ptw = 0.37–0.43 (0.40); aw = 0.40–0.46 (0.43). remarks piaget (1880) gives as type locality madagascar, which is well outside the range of the host species (madge & mcgowan 2002). clay (1938) examined piaget’s types, which she found to be identical to material from borneo. piaget’s type locality designation is therefore here considered erroneous. we have seen photos of the lectotype and paralectotype (at nhmuk), but not examined these specimens in person. the photo of the female lectotype does not differ from the non-type specimens we have examined. preliminary key to the genera of the oxylipeurus-complex characters taken primarily from clay (1938), clay & meinertzhagen (1941), carriker (1945, 1967), emerson & ward (1958), kéler (1958), elbel & price (1970), mey (1982, 1990, 2006, 2010), gustafsson et al. (2020), and examinations of specimens. additional groups deserving recognition at the genus level may exist, and many species of the complex are in need of further study and redescription. the genus labicotes kéler, 1939, may also belong to this complex, based on similarities in male and female terminalia, male genitalia, and temporal chaetotaxy between this genus and chelopistes. this needs to be confirmed by additional studies of species of labicotes, and the genus is not included here. european journal of taxonomy 686: 1–15 (2020) 8 1. broad-headed, with width of head similar to, or wider than, length of head (figs 19–20); temples with elongated “horns” (fig. 19) or with prominent lateral bulges (fig. 20) .................................... 2 – slender-headed, with head clearly longer than wide (fig. 5); temples generally rounded, never with prominent bulging .............................................................................................................................. 3 2. temporal setae mts1–2 macrosetae (fig. 20) ......................................trichodomedea carriker, 1946 – temporal setae mts1–2 microsetae (fig. 19) ................................................. chelopistes kéler, 1939 3. dorsal preantennal suture present (figs 5, 21) .................................................................................. 4 – dorsal preantennal suture absent or if present only visible around aperture of ads and not extending medianly (fig. 22) ........................................................................................................................... 10 4. dorsal preantennal suture as median, elongated oval, not expanded laterally (fig. 5); female terminalia with marginal mesosetae distributed more or less equally around distal margin (fig. 4); eye very large and preocular nodus absent (fig. 5) ..................................... calidolipeurus gen. nov. – dorsal preantennal suture transversal, normally reaching apertures of ads (fig. 21); female terminalia with marginal setae gathered in the same area (fig. 23); eye not very large (fig. 21), and preocular nodus present ..................................................................................................................................... 5 5. clypeo-labral suture present (fig. 24); stylus expanded distally, with small “hooks” on lateral margins (fig. 25) ........................................................................gallancyra gustafsson & zou, 2020 – clypeo-labral suture absent (fig. 5); stylus differing in shape, but never with lateral ‘hooks’ ......... 6 6. dorsal preantennal suture with postero-lateral elongations (“epistomal suture” sensu kéler 1958) extending towards preantennal nodi (fig. 26); hyaline margin present (fig. 26) ....................................................................... splendoroffula clay & meinertzhagen, 1941 – dorsal preantennal suture without such extensions (fig. 21); hyaline margin absent (fig. 21) ....... 7 7. dorsal postantennal suture present (fig. 27); male genitalia asymmetrical, with mesosome much reduced (fig. 28) ......................................................................................oxylipeurus mjöberg, 1910 – dorsal postantennal suture absent (fig. 21); male genitalia symmetrical, with prominent mesosome (variable, but similar to figs 8–11) .................................................................................................... 8 8. coni elongated (similar to fig. 5); male mesosome with prominent vor y-shaped thickening in distal half (fig. 29); proximal margin of mesosome with rounded lateral lobes (fig. 29); frons convergent to median point in most species (similar to fig. 27) .............. megalipeurus kéler, 1958. – coni short (fig. 21); male mesosome without thickening in distal half; proximal margin variable, but never with rounded lateral lobes; frons rounded ............................................................................... 9 9. male abdominal segments ix and ix + x with prominent postero-lateral extensions (“claspers” sensu carriker 1945) (fig. 30) ............................................................ eiconolipeurus carriker, 1945 – male abdomen without such structures ....................................................reticulipeurus kéler, 1958 10. frons convergent to median point (fig. 27) .....................................................................................11 – frons rounded (fig. 21) ................................................................................................................... 12 11. male tergopleurites ii–vii medianly continuous with no median indentations of anterior margin; male abdominal segments ix + x and xi fused into roughly triangular cone (fig. 31); stylus elongated and slender, in the shape of a posterior extension of the male subgenital plate (fig. 31); female terminalia without “claspers”, vulval margin more or less straight ............................... afrilipeurus mey, 2010 – male tergopleurites ii–vii either divided medianly, or with median indentation of anterior margin; male tergopleurites ix + x and xi separate, posterior margin concave (similar to fig. 1); stylus short and blunt (fig. 32); female terminalia with “claspers”, vulval margin deeply concave (fig. 33) ....................................................................................................talegallipeurus mey, 1982 gustafsson d.r. et al., new genus calidolipeurus (phthiraptera) 9 figs 19–26. 19. outline of head and temporal macrosetae (cut off distally) of male chelopistes meleagridis (linnaeus, 1758), redrawn from kéler (1939). 20. outline of head and temporal macrosetae (cut off distally) of female trichodomedea setosus carriker, 1946, redrawn from original description. 21. outline of head and dorsal preantennal suture of male reticulipeurus mesopelios (nitzsch, 1866), redrawn from gustafsson et al. (2020). 22. outline of head and dorsal preantennal suture of male cataphractomimus junae gustafsson et al., 2020, redrawn from original description. 23. female terminalia of reticulipeurus mesopelios (nitzsch, 1866), redrawn from gustafsson et al. (2020); vulval margin, lateral macrosetae, and subvulval plates not illustrated. 24. ventral view of preantennal area in gallancyra dentata (sugimoto, 1934) (redrawn from gustafsson & zou 2020a). 25. outline of stylus in gallancyra dentata (sugimoto, 1934) (redrawn from gustafsson & zou 2020a). 26. outline of preantennal area and dorsal preantennal suture of splendoroffula ampullaceal kéler, 1955, redrawn from kéler (1958). antennae not included in any illustration. abbreviations used: c = conus; cls = clypeo-labral suture; dps = dorsal preantennal suture; e = eye; es = epistomal suture; hm = hyaline margin; mts3 = marginal temporal seta 3; os = ocular seta. illustrations are not to scale. european journal of taxonomy 686: 1–15 (2020) 10 figs 27–37. 27. outline of head and dorsal post-antennal suture of oxylipeurus inaequalis (piaget, 1880), redrawn from mey (1990); original drawing asymmetrical. 28. male genitalia of oxylipeurus inaequalis (piaget, 1880), redrawn from mey (1990); some details left out for clarity. 29. ventral view of mesosome of megalipeurus sinensis gustafsson et al., 2020, redrawn from original description. 30. dorsal view of male terminalia of eiconolipeurus melanotis carriker, 1945, redrawn from original description; setae not illustrated. 31. dorsal and ventral views of male terminalia of afrilipeurus vincentei kéler, 1953, redrawn from mey (2010); seta not illustrated. 32. ventral view of male terminalia and stylus of talegallipeurus tenuis mey, 1982, redrawn from original description; setae not illustrated; original illustration asymmetrical (mey 1982). 33. ventral view of female terminalia and vulval margin of talegallipeurus tenuis mey, 1982, redrawn from original description; setae not illustrated. 34. outline of male paramere of sinolipeurus tetraophasis (clay, 1938), redrawn and simplified from gustafsson et al. (2020). 35. outline of male terminalia and stylus of sinolipeurus tetraophasis (clay, 1938), redrawn and simplified from gustafsson et al. (2020). 36. outline of male paramere of reticulipeurus ithaginis (clay, 1938), redrawn and simplified from gustafsson et al. (2020). 37. distal section of male genitalia of epicolinus clavatus (mcgregor, 1917), redrawn from carriker (1945). abbreviations used: al = anterior lobes; cl = “claspers”; dpas = dorsal post-antennal suture; ix–xi = tergopleurites ix–xi; sty = stylus; vm = vulval margin; y = y-shaped thickening. antennae not included in any illustration. illustrations are not to scale. gustafsson d.r. et al., new genus calidolipeurus (phthiraptera) 11 12. male parameres strongly s-curved (fig. 34); stylus arising centrally on abdominal segment ix + x (fig. 35) .....................................................................................sinolipeurus gustafsson et al., 2020 – male parameres not s-curved (fig. 36); stylus varying in shape, but always arising terminally or subterminally on subgenital plate (similar to fig. 3) ....................................................................... 13 13. male genitalia simple, with parameres fused to basal apodeme and mesosome much reduced (fig. 37) ......................................................................................................epicolinus carriker, 1945 – male genitalia with parameres articulating with basal apodeme, and mesosome not reduced (similar in structure but not shape to fig. 8) ................................................................................................. 14 14. lateral margins of postantennal head with secondary, ventral carina between antennal socket and site of mts2 or mts3 (fig. 38); area between margin of head and secondary carina, densely reticulated, including ventral surface of eye (fig. 38); male parameres with pst1–2 situated close together apically; male gonoporal complex does not reach distal margin of mesosome; female subgenital plate divided medianly (fig. 39) ..................................................... valimia gustafsson & zou, 2020 – lateral margins of postantennal head without secondary carina and without extensive ventral reticulation (similar to fig. 5); male parameres with pst1–2 separated, and only pst2 apical; male gonoporal complex reached to or beyond distal margin of mesosome; female subgenital plate medianly continuous (fig. 40) ........................................ cataphractomimus gustafsson et al., 2020 figs 38–40. 38. outline of male head of valimia polytrapezius (burmeister, 1838), with post-antennal ventral carina and densely reticulated area marked with grey dots; other characters omitted. 39. outline of ventral view of female terminalia of “oxylipeurus polytrapezius” (burmeister, 1838). setae not illustrated. (38–39 redrawn from gustafsson & zou 2020b) 40. outline of ventral view of female terminalia of cataphractomimus impervius gustafsson et al., 2020, redrawn and simplified from the original description. antennae not included in any illustration. abbreviations used: lavp = lateral accessory vulval plates; sgp = subgenital plate; smc = secondary marginal carina. illustrations are not to scale. european journal of taxonomy 686: 1–15 (2020) 12 discussion clay (1938) included illustrations of only a few species of oxylipeurus and most of these illustrations are partial. as clay (1938) is the only comprehensive revision of the genus, particularly for the old world species, the systematics of this group has been confused, and most recent authors have adopted an overly conservative approach (e.g., price et al. 2003). the complex includes a large number of morphologically homogeneous groups, some of which clay (1938) considered species groups. most of these species groups are distinct enough to warrant recognition at the genus level and a thorough revision of the complex is needed. within this complex, the following genera should be recognized as valid: afrilipeurus mey, 2010; calidolipeurus gen. nov.; cataphractomimus gustafsson et al., 2020; chelopistes kéler, 1939; eiconolipeurus carriker, 1945; epicolinus carriker, 1945; gallancyra gustafsson & zou, 2020; megalipeurus kéler, 1958; oxylipeurus mjöberg, 1910; reticulipeurus kéler, 1958; sinolipeurus gustafsson et al., 2020; splendoroffula clay & meinertzhagen, 1941; talegallipeurus mey, 1982; trichodomedea carriker, 1946 and valimia gustafsson & zou, 2020. additional morphologically distinct groups exist within this complex, which represent undescribed genera. acknowledgements work was funded by the introduction of full-time high-level talent fund of the guangdong academy of sciences grant 2018gdascx-0809, gdas special project of science and technology development grants 2018gdascx-0107 and giabr-gjrc201701, and the national natural science foundation of china grant 31961123003. these agencies had no hand in the design or execution of this study and we declare no conflict of interest. we would like to thank two anonymous reviewers for helpful comments on this manuscript. we would like to thank dr vince smith and paul brown (nhmuk) who hosted drg in london and enabled the loan of specimens that this manuscript is based on. references carriker m.a., jr. 1945. studies in neotropical mallophaga (v) the lipeuroid forms of the new world “galliformes”. part 1. revista brasileira de biologia 4: 537–585. carriker m.a., jr. 1946. studies in neotropical mallophaga (vii) goniodes and allied genera from gallinaceous hosts. revista de le academia colombiana de sciencias exactas (bogotá) 6: 355–399. carriker m.a., jr. 1967. new species of colinicola, passonomedea, eiconolipeurus, and oxylipeurus (mallophaga: philopteridae) from neotropical gallinaceous birds. united states national museum bulletin 248: 46–55. clay t. 1938. a revision of the genera and species of mallophaga occurring on gallinaceous hosts. — part i. lipeurus and related genera. proceedings of the zoological society of london 108: 108–204. clay t. & meinertzhagen r. 1941. mallophaga miscellany. — no. 2. annals and magazine of natural history 40: 329–346. https://doi.org/10.1080/00222934108527162 elbel r.e. & price r.d. 1970. two new species of ischnoceran mallophaga from an oriental partridge (mallophaga: philopteridae). journal of the kansas entomological society 43: 238–242. emerson k.c. & ward r.a. 1958. notes on philippine mallophaga, i. species from ciconiiformes, anseriformes, falconiformes, galliformes, gruiformes and charadriiformes. fieldiana – zoology 42: 49–61. gustafsson d.r. & bush s.e. 2017. morphological revision of the hyperdiverse brueelia-complex (insecta: phthiraptera: ischnocera: philopteridae) with new taxa, checklists and generic key. zootaxa 4313: 1–443. https://doi.org/10.11646/zootaxa.4313.1.1 gustafsson d.r. et al., new genus calidolipeurus (phthiraptera) 13 https://doi.org/10.1080/00222934108527162 https://doi.org/10.11646/zootaxa.4313.1.1 gustafsson d.r. & zou f. 2020a. gallancyra gen. nov. (phthiraptera: ischnocera), with an overview of the geographical distribution of chewing lice parasitizing chicken. european journal of taxonomy 685: 1–36. https://doi.org/10.5852/ejt.2020.685 gustafsson d.r. & zou f. 2020b. descriptions of three congeneric species of chewing lice of the oxylipeurus-complex (insecta: phthiraptera: philopteridae) from the turkey, meleagris gallopavo, including a new genus and a new species. zootaxa 4801: 488–512. https://doi.org/10.11646/zootaxa.4801.3.4 gustafsson d.r., lei l., chu x. & zou f. 2020. review of the chinese species of the oxylipeuruscomplex (phthiraptera: philopteridae), with descriptions of two new genera and five new species. zootaxa 4742: 201–255. https://doi.org/10.11646/zootaxa.4742.2.1 harrison l. 1916. the genera and species of mallophaga. parasitology 9: 1–156. https://doi.org/10.1017/s0031182000005989 hopkins g.h. & clay t. 1952. a check list of the genera and species of mallophaga. british museum (natural history), london. kéler s. von. 1939. baustoffe zu einer monographie der mallophagen. ii. teil: überfamilie der nirmoidea (1). die familien trichophilopteridae, goniodidae, heptapsogastridae. nova acta leopoldina. abhandlungen der kaiserlischen leopoldinisch-carolinisch deutschen akademie der naturforscher, neue folge 8: 1–254. kéler s. von. 1958. the genera oxylipeurus mjöberg and splendoroffula clay and meinertzhagen (mallophaga). deutsche entomologische zeitschrift, neue folge 5: 300–347. ledger j.a. 1980. phthiraptera (insecta). volume iv of the arthropod parasites of vertebrates in africa south of the sahara. publications of the south african institute for medical research 56. south african institute for medical research, johannesburg. madge s. & mcgowan p. 2002. pheasants, partridges and grouse. a guide to the pheasants, partridges, quails, grouse, guineafowl, buttonquails, and sandgrouse of the world. christopher helm, london. mey e. 1982. zur taxonomie und biologie der mallophagen von talegalla jobiensis longicaudus a.b. meyer, 1891 (aves, megapodiidae). reichenbachiana, staatliches museum für tierkunde in dresden 20: 223–246. mey e. 1990. zur taxonomie der auf großfußhühnern (megapodiidae) schmarotzenden oxylipeurusarten (insecta, phthiraptera, ischnocera: lipeuridae). zoologische abhandlungen staatliches museum für tierkunde dresden 46: 103–116. mey e. 2006. rätselhaftes vorkommen zweier federlingsarten (insecta, phthiraptera, ischnocera) auf dem haldenhuhn lerwa lerwa (galliformes, phasianidae)? in: hartmann m. & weipert j. (eds) biodiversität und naturausstattung im himalaya. naturkundemuseum erfurt, germany. mey e. 2009. die mallophagen (insecta, phthiraptera: amblycera & ischnocera) der galloanseres (aves) – ein überblick. beiträge zur jagd und wild forschung 34: 151–187. mey e. 2010. afrilipeurus novum genus pro oxylipeurus vincentei v. kéler (insecta, phthiraptera, ischnocera, philopteridae s. l.) vom kräuselhauben-perlhuhn guttera pucherani (hartlaub). rudolstädter naturhistorische schriften 16: 99–110. mjöberg, e. 1910. studien über mallophagen und anopluren. arkiv för zoologi 6: 1–150. piaget e. 1880. les pédiculines. essai monographique. e.j. brill, leiden. price r.d., hellenthal r.a. & palma r.l. 2003. world checklist of chewing lice with host associations and keys to families and genera. in: price r.d., hellenthal r.a., palma r.l., johnson k.p. & clayton d.h. european journal of taxonomy 686: 1–15 (2020) 14 https://doi.org/10.5852/ejt.2020.685 https://doi.org/10.11646/zootaxa.4801.3.4 https://doi.org/10.11646/zootaxa.4742.2.1 https://doi.org/10.1017/s0031182000005989 (eds) the chewing lice: world checklist and biological overview. illinois natural history survey special publication 24. illinois natural history survey, champaign, illinois. wang n., kimball r.t., braun e.l., liang b. & zhang z. 2013. assessing phylogenetic relationships among galliformes: a multigene phylogeny with expanded taxon sampling in phasianidae. plos one: e64312. https://doi.org/10.1371/journal.pone.0064312 manuscript received: 18 november 2019 manuscript accepted: 4 may 2020 published on: 15 july 2020 topic editor: nesrine akkari desk editor: kristiaan hoedemakers printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. gustafsson d.r. et al., new genus calidolipeurus (phthiraptera) 15 https://doi.org/10.1371/journal.pone.0064312 1 european journal of taxonomy 638: 1–7 issn 2118-9773 https://doi.org/10.5852/ejt.2020.638 www.europeanjournaloftaxonomy.eu 2020 · iczn this work is licensed under a creative commons attribution license (cc by 4.0). o p i n i o n p a p e r urn:lsid:zoobank.org:pub:34ea0e72-90b8-495f-b049-3b56319fb77d proposed amendments to the constitution of the international commission on zoological nomenclature international commission on zoological nomenclature c/o lee kong chian natural history museum, 2 conservatory drive, singapore 117377, republic of singapore. email: iczn@nus.edu.sg urn:lsid:zoobank.org:author:a6302ea9-154f-4a21-b1b0-bc2d1b2d0bce abstract. the international commission on zoological nomenclature (iczn) proposes amendments to its constitution and solicits constructive feedback from the zoological community. in compliance with the constitution, the proposed amendments are made broadly available, and there will be a one-year period for submission of comments starting on 30 april 2020. the amendments may be modified in the light of the comments, before the final version is voted on by the commissioners. iczn. 2020. proposed amendments to the constitution of the international commission on zoological nomenclature. european journal of taxonomy 638: 1–7. https://doi.org/10.5852/ejt.2020.638 introduction the international commission on zoological nomenclature (iczn, or commission) considered amendments to articles of its constitution (iczn 1999a) at a special session in singapore, convened on june 3–7, 2019. during this meeting, commissioners also planned revisions to the bylaws, the current international code of zoological nomenclature (iczn 1999b, 2003, 2012, 2017) and zoobank user policies. the commissioners voted to begin the process required to amend the constitution. notably, revisions were proposed and approved in principle by majority vote to clarify and update constitutional articles pertaining to: 1. commissioners’ terms of service (article 3); 2. inclusion of the iczn web site as a required venue for information dissemination (articles 2.1, 4.2 and 12.2); 3. outdated concepts (such as the itzn, article 13, or postal voting, articles 4.6, 12.2, 16.1.4 and 16.1.5), and new responsibilities (such as zoobank, article 14.5); 4. duration of votes on amendments to the code and constitution (article 16.1.4); and 5. separation of procedures of the commission from its bylaws (article 17). http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:34ea0e72-90b8-495f-b049-3b56319fb77d mailto:iczn%40nus.edu.sg?subject= http://zoobank.org/urn:lsid:zoobank.org:author:a6302ea9-154f-4a21-b1b0-bc2d1b2d0bce https://doi.org/10.5852/ejt.2020.638 european journal of taxonomy 638: 1–7 (2020) 2 some constitutional articles were also proposed to be revised for consistency and accuracy, including: 1. article 2.1 (“contact information” instead of “addresses”); 2. article 4.1.1 (“countries of residence” instead of “nationalities”); and 3. adding “executive” before “secretary” in articles 5.2, 7.1 and 14 to bring these articles into accord with articles 4.3, 5.3 and 9. procedure for amendment the constitution of the iczn (article 12.2) requires publication of notice of a proposed amendment of the constitution in the bulletin of zoological nomenclature (bzn), and submission to at least three other journals, for comment from the zoological community for at least one year prior to the commissioners’ voting on the proposed amendment (article 16.1.2). the one-year period for comment on the current proposal, prior to any vote, will start on 30 april 2020, with the publication of notice in the bulletin of zoological nomenclature. a copy of the proposed amendment will also be posted on the iczn web site (www.iczn.org). the iczn requests comments from the community of zoologists on various parts of this amendment, either opposing, supporting, or suggesting improvements and alternatives. the commission will revise the amendment in light of comments received from zoologists (constitution article 16.1.4). comments that have been vetted for content and language will be published in bzn with open access and posted on the iczn web site (www.iczn.org). formal comments should be sent to dr gwynne lim, executive secretary of the iczn (iczn@nus.edu.sg). zoologists may also contact iczn commissioners directly for informal discussions (https://www.iczn.org/about-the-iczn/commissioners/). assuming that sufficient consensus is reached in the zoological community, the final-worded amendment will be presented to the international union of biological sciences (iubs) for provisional ratification (constitution article 16.1.5.1). effective ratification will be contingent on a subsequent vote by the commission (constitution article 16.1.5.1). the decision and date of effective ratification will be published in bzn (constitution article 16.1.6). proposed amendment in the proposed amendment, normal font represents existing text of the constitution that is retained. strikethrough text is existing text that is to be deleted in the amended articles. underlined text is new text. indented text in square brackets describes the changes. the amendment affects articles 2, 3, 4, 5, 7, 9, 12, 13, 14, 16, and 17. article 2. membership of the commission. [article 2.1 is to be revised to specify “contact details” instead of “addresses” and to include the iczn web site as a source of information] 2.1. number. the commission shall ordinarily consist of 18 members or such larger number as the commission may decide. the names and contact details addresses of the members at any given time shall be published in the bulletin of zoological nomenclature and on the commission web site. [articles 2.2 and 2.3 are unchanged.] [article 3 is to be revised to include the criteria for eligibility of re-election specified by article 3.2] http://www.iczn.org http://www.iczn.org mailto:iczn%40nus.edu.sg?subject= https://www.iczn.org/about-the-iczn/commissioners/ iczn, amendments to iczn constitution 3 article 3. term of service and eligibility of members of the commission. [article 3.1 is to be reformulated to simplify the rules for commissioners’ terms of service and to eliminate the class system. articles 3.1.1, 3.1.2 and 3.1.3 are to be deleted, and elements of article 3.2 are to be simplified and merged into revised articles 3.1.1 (length of term of service) and 3.1.2 (term of service for the president of the commission).] 3.1. normal term of service. 3.1.1. the normal term of service of a member of the commission shall be reckoned as follows: is eighteen years. 3.1.2. the term of service of the president of the commission shall end at the end of the term indicated in the bylaws even if it exceeds the eighteen years term of service above. [article 3.2 is to be revised to simplify the rules for re-election of existing commissioners. article 3.2.1 is to be deleted, having been subsumed within revised article 3.1.1. article 3.2.2 is to be reformulated as part of revised article 3.2.1. article 3.2.2.1 is to be reformulated as part of revised article 3.2.2. article 3.2.3 is to be deleted.] 3.2. maximum term of service and eligibility for re-election. a member whose normal term of service has terminated terminates may be re-elected but: 3.2.1. on completion of the period specified in article 3.1 three years must elapse before a former member of the commission is eligible for re-election; 3.2.2. this provision [art. 3.2.1] shall not apply when a retiring or former member is pre-elected by the commission to continue as or to become its president if re-elected as a member. [article 3.3 is to be revised to exclude article 3.3.1, which is to be deleted. article 3.3.2 is to be renumbered as article 3.3.1. article 3.3.3 is to be revised and renumbered as article 3.3.2] 3.3. prior termination of membership. the membership of any member of the commission shall terminate prior to the expiration of the term of service under art. 3.1 above: 3.3.1. on acceptance by the council of notice of resignation tendered in writing to the executive secretary; 3.3.2. if, not being on leave of absence, he or she fails on five consecutive occasions to return the ballot, or, when no ballot is provided, fails to record a vote for or against or an abstention to record a vote or an abstention on questions put to the commission for decision, provided that within a period of three months following such failure no written explanation has been made which the council finds adequate. article 4. election of members of the commission. [article 4.1.1 is to be revised to specify “countries of residence” instead of “nationalities” in recognition of the fact that some commissioners are expatriates. article 4.1.2. is to be revised to specify “refers” instead of “quotes”.] 4.1. notice. the commission shall publish, not less than one year before a general session of the commission [art. 11.1], a notice which: european journal of taxonomy 638: 1–7 (2020) 4 4.1.1. gives the names, countries of residence nationalities and fields of specialization of the members whose terms of service will end at the close of that session in accordance with article 3; 4.1.2. quotes refers to article 2.2 and invites nominations for membership of the commission; 4.1.3. gives a date, not more than three months before the forthcoming general session, by which nominations must be received. [article 4.2 is to be revised to include web sites as, depending on circumstances, either mandated or optional outlets for information] 4.2. circulation. the notice specified in article 4.1 shall be published on the commission web site and submitted to iubs (or to its successor body, if any), to the organizers of the congress where the general session is to be held, and to appropriate journals and/or web sites in different parts of the world, with a request for its dissemination. [article 4.3 is revised to add “executive” before “secretary” for consistency.] 4.3. nominations. nominations, accompanied by a statement of the fields of specialization and qualification under article 2.2 of each nominee, are to be sent to the executive secretary of the commission. unless the nomination contains the information, the executive secretary shall require each nominee to give consent to the nomination and to provide a curriculum vitae, a list of publications and a statement of his or her nomenclatural experience. [article 4.6 is to be revised to delete “postal”.] 4.6. by-elections. the commission may by a postal ballot fill vacancies arising from prior termination of membership [art. 3.3], or which have not been filled by election at a session of the section of zoological nomenclature [art. 4.4.1], or which result from an increase in the number of members decided by the commission in accordance with article 2.1. [articles 5.2 and 5.3 are to be revised to add “executive” before “secretary”] 5.2. between sessions. it shall be the duty of a member of the commission to vote, within the prescribed period, upon each question submitted to him or her for that purpose by the executive secretary. 5.3. leave of absence. a member of the commission who is temporarily unable to perform his or her duties should apply through the executive secretary (if possible in advance) for leave of absence for a specified period. [article 6 is to be unchanged. article 7.1 is to be revised to include “executive” before “secretary” and to include “or she” after “he”.] 7.1. the executive secretary to the commission is also the secretary to the council but neither he or she nor any other member of the secretariat shall vote in its deliberations. iczn, amendments to iczn constitution 5 [article 8 is to be unchanged. article 9 is to be revised to remove references to a secretary-general and the international trust for zoological nomenclature. the appointment of the secretary-general is to be covered more generally in revised article 14.] article 9. secretariat. the council may appoint an executive secretary for such a term and with such duties as may be fixed in the bylaws. a member of the commission may be appointed similarly as secretary-general. the executive secretary may be an employee of an appropriate body such as the international trust for zoological nomenclature. [articles 10 and 11 are to be unchanged. article 12.2 is to be revised to remove “postal”, to include the commission web site and to remove the necessity of submitting proposed amendments to the constitution to multiple journals before voting on them. article 12.2 is also to be revised to reduce the voting period from three months to two, in recognition of the faster transmission speed of electronic mail compared to postal mail.] 12.2. in cases involving the use of the plenary power or amendments to the code or constitution. in such cases (see articles 78 to 81 of the code for the use of the plenary power and article 16 of this constitution for amendments to the code or constitution) an affirmative decision shall be deemed to have been taken only when two thirds of the votes validly cast in a postal vote lasting three two months are in favour of the proposal, and provided that notice of the proposal had been published in the bulletin of zoological nomenclature, on the commission web site and, only for the amendments to the code, submitted for publication to at least three two appropriate journals at least six months (in the case of amendments to the code or constitution, twelve months) prior to the vote. [article 13 is to be revised to remove the reference to the international trust for zoological nomenclature.] article 13. financial arrangements. the commission when not prepared to raise or administer its own funds is empowered for such purposes to enter into a beneficent relationship with one or more bodies such as the international trust for zoological nomenclature, that undertake to act in accordance with the policy of the commission and iubs (or its successor body, if any). the commission may terminate such a relationship at its discretion. [article 14 is to be revised to include “executive” before “secretary” and to incorporate elements from article 9.] article 14. editorial duties of the commission. the commission shall issue and, finances permitting, may itself publish various communications, to be prepared and edited in the office of the executive secretary, or by another person appointed for that purpose, under the guidance of the council. [article 14.3 is to be revised to delete redundant language.] 14.3. maintenance of official lists and indexes. the commission shall compile and maintain the undermentioned lists and indexes: 14.3.1. official list of family-group names in zoology; european journal of taxonomy 638: 1–7 (2020) 6 14.3.2. official list of generic names in zoology; 14.3.3. official list of specific names in zoology; 14.3.4. official index of rejected and invalid family-group names in zoology; 14.3.5. official index of rejected and invalid generic names in zoology; 14.3.6. official index of rejected and invalid specific names in zoology; 14.3.7. official list of works approved as available for zoological nomenclature; 14.3.8. official index of rejected and invalid works in zoological nomenclature. [a new article 14.5 is to be added to specify the commission’s duties with regards to the official register of zoological nomenclature.] 14.5. official register of zoological nomenclature (zoobank). the commission shall maintain zoobank, the official register of zoological nomenclature. [article 15 is to be unchanged. article 16.1.2 is to be revised to specify the time span as “twelve months” instead of “one year”. articles 16.1.4 and 16.1.5 are to be revised to remove “postal”. article 16.1.4 is to be revised to reduce the voting period from three months to two, in recognition of the faster transmission speed of electronic mail compared to postal mail.] 16.1.2. receive and consider comments from zoologists that are received within twelve months one year of the publication of the proposals; 16.1.4. vote on the proposed amendments (which may be modified in the light of the comments by zoologists and the section) in a postal vote lasting two three months [art. 12.2]; 16.1.5. submit the amendments subject to their approval by two thirds of the votes validly cast in the postal vote, and with the support of the section for the major principles, to iubs (or to its successor body, if any) for ratification [art. 90 of the code]; [article 17 is to be revised to distinguish between bylaws and procedures.] article 17. bylaws and procedures. the commission is empowered to adopt a set of bylaws and of procedures governing those of its regulations and activities not covered by the constitution. the commission has authority to modify these bylaws and procedures by a majority vote as the occasion demands. these bylaws will deal with such matters as the duties of the officers, the methods by which nominations are to be obtained for vacancies on the commission, the relations between the commission and the secretariat, and with other business matters of the commission. the procedures will deal with regulations concerning the treatment to be given to applications and the adoption of time schedules and priorities, and with other business matters of the commission. [article 18 is to be unchanged.] iczn, amendments to iczn constitution 7 references iczn 1999a. the constitution of the international commission on zoological nomenclature. in: international code of zoological nomenclature: 264–271. fourth edition. the international trust for zoological nomenclature, london. iczn 1999b. international code of zoological nomenclature. fourth edition. the international trust for zoological nomenclature, london. iczn 2003. declaration 44. amendment of article 74.7.3. bulletin of zoological nomenclature 60 (4): 263. iczn 2012. amendment of articles 8, 9, 10, 21 and 78 of the international code of zoological nomenclature to expand and refine methods of publication. zookeys 219: 1–10. https://doi.org/10.3897/zookeys.219.3944 iczn 2017. declaration 45 – addition of recommendations to article 73 and of the term “specimen, preserved” to the glossary. bulletin of zoological nomenclature 73 (2–4): 96–97. https://doi.org/10.21805/bzn.v73i2.a2 manuscript received: 14 february 2020 manuscript accepted: 26 february 2020 published on: 30 april 2020 topic editor: koen martens desk editor: chloë chester printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. http://https://doi.org/10.3897/zookeys.219.3944 https://doi.org/10.21805/bzn.v73i2.a2 1 european journal of taxonomy 828: 1–15 issn 2118-9773 https://doi.org/10.5852/ejt.2022.828.1847 www.europeanjournaloftaxonomy.eu 2022 · van valkenburg j.l.c.h. et al. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e myriophyllum rubricaule sp. nov., a m. aquaticum look-alike only known in cultivation johan l.c.h. van valkenburg  1,*, leni (h.) duistermaat  2, edu boer 3 & tom m. raaymakers  4 1,3,4 national reference centre for plant health, nvwa, geertjesweg 15, 6706 ea wageningen/postbus 9102, 6700 hc wageningen, the netherlands. 2,3 naturalis biodiversity center, darwinweg 2, 2333 cr leiden/postbus 9517, 2300 ra leiden, the netherlands. * corresponding author: j.l.c.h.vanvalkenburg@nvwa.nl 2 email: leni.duistermaat@naturalis.nl 3 email: e.boer2@nvwa.nl 4 email: t.m.raaymakers@nvwa.nl abstract. a confusingly labeled water-milfoil of obscure status, known only in cultivation, is here formally described as a new species, myriophyllum rubricaule valk. & duist. sp. nov. this species has fully replaced m. aquaticum in the horticultural trade in europe since the addition of m. aquaticum to the list of invasive alien species of union concern (eu regulation no. 1143/2014) in 2016. this manuscript provides a morphological description of m. rubricaule sp. nov., and its distinction from m. aquaticum is further supported by molecular data (chloroplast and nuclear loci). keywords. myriophyllum, water-milfoil, eu regulation no. 1143/2014, invasive plant, horticulture. van valkenburg j.l.c.h., duistermaat l.(h.), boer e. & raaymakers t.m. 2022. myriophyllum rubricaule sp. nov., a m. aquaticum look-alike only known in cultivation. european journal of taxonomy 828: 1–15. https://doi.org/10.5852/ejt.2022.828.1847 introduction incorrect labeling of plants in trade and misidentification are widespread and may be caused by negligence or willful disrespect of regulations (brunel 2009; thum et al. 2012; verbrugge et al. 2014; hulme et al. 2018). mislabeling may consist of simple misspelling of names, or treating a variety as a true species, using synonyms or just preferring a name that sounds nice or a name that customers are familiar with. the latter case actually equals misidentification of the plant in trade. from observations of species of myriophyllum ponted. ex l. in trade in the netherlands, it is clear that proper labeling is generally neglected, causing confusion about the proper identity of the species in trade (van valkenburg & boer 2015; van valkenburg et al. 2015). plants labelled as myriophyllum ‘brasiliensis’ or m. ‘brasiliense’ have been in the horticultural trade in north-western europe for several decades, though their taxonomy has caused some confusion among botanists, plant growers and regulators. when looking https://doi.org/10.5852/ejt.2022.828.1847 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ https://orcid.org/0000-0001-7281-7819 https://orcid.org/0000-0002-0931-030x https://orcid.org/0000-0003-1056-7177 mailto:j.l.c.h.vanvalkenburg%40nvwa.nl?subject= mailto:leni.duistermaat%40naturalis.nl?subject= mailto:e.boer2%40nvwa.nl?subject= mailto:t.m.raaymakers%40nvwa.nl?subject= https://doi.org/10.5852/ejt.2022.828.1847 european journal of taxonomy 828: 1–15 (2022) 2 at species of myriophyllum, moody & les (2010) refer to sterile plants of unknown origin in horticulture as myriophyllum ‘red 1’ and ‘red 2’, respectively. one of these taxa may well be identical to the taxon we found in trade in north-western europe. the european material in trade was included in our previous study on barcoding invasive aquatic plants as m. ‘brasiliense’ trade material and erroneously referred to as m. robustum hook.f. (ghahramanzadeh et al. 2013). after receiving the true m. robustum from new zealand, and observing it flowering in 2013 (valkenburg 3739 wagpd; only female flowers, however, m. robustum is described as monoecious by orchard (1980)), the identification of the trade material was adjusted and the designation in q-bank plants updated to myriophyllum sp. trade name ‘brasiliensis’. the name myriophyllum brasiliense cambess. is a synonym of m. aquaticum (vell.) verdc. in the absence of any knowledge of its occurrence in the wild, we refrained from formally describing it as a new species. in the meantime, plants have proved to be hardy in outdoor mesocosms at the nvwa (nederlandse voedsel en warenautoriteit) in wageningen (the netherlands) since 2007. after m. aquaticum was included in the list of invasive alien species of union concern (‘the union list’) under eu regulation no. 1143/2014 in august 2016, it has been fully replaced by myriophyllum sp. trade name ‘brasiliensis’ in the horticultural trade in europe. since then, there has been an increase in the number of recordings of myriophyllum sp. trade name ‘brasiliensis’ in urban waters as well as in more natural habitats. this is likely a result of inappropriate disposal of garden waste or the deliberate planting of the species in urban waterways and residential areas. to-date, there are records of establishment for the netherlands, belgium and hungary. the ongoing confusion with respect to the epithet ‘brasiliensis’ has prompted the authors to formally describe this taxon known only in cultivation. material and methods plant material of myriophyllum has been collected since 2007, and deposited at the herbaria l, wag and wagpd (appendix 1). acronyms of herbaria follow index herbariorum (thiers continuously updated). in addition to the above mentioned specimens used for the present study, all myriophyllum collections from south america at naturalis (ams, l, u, wag) were consulted as well as most relevant revisions and new taxonomical publications on the genus myriophyllum to ascertain the novelty of the new species (meijden & caspers 1971; orchard 1980, 1981, 1986; orchard & kasselmann 1992; wang et al. 2002; yu et al. 2002; wang & yu 2007). morphological description is based on both living and herbarium material and follows meijden & caspers (1971) and crow & ritter (1999) for characters considered important in species recognition. no plant material was consulted of the myriophyllum sp. ‘red 1’ and ‘red 2’ that were used in the molecular analysis as they were reported to be sterile (moody & les 2010). an annotated collection of m. aquaticum (pedersen 3977 in orchard 1981) from argentina, as well as a diminutive collection of m. aquaticum from brazil (lindeman 8405) were included to ascertain the molecular novelty of the new species. a laurembergia tetrandra (schott) kanitz collection (van proosdij 33) was used as an outgroup (appendix 1). dna extractions genomic dna was isolated from approximately 100 mg of plant material with the dneasy plant mini kit (qiagen, venlo, the netherlands) using the tissuelyser procedure and eluted with 50 μl of prewarmed (65ºc) ae buffer. dna was stored at -20°c until use. pcr amplification and sanger sequencing pcr reactions for the chloroplast loci rbcl and trnh-psba intergenic spacer and the nuclear locus its (partial 18s, its1, 5.8s, its2, partial 28s) were performed in 25 μl reaction mixes containing 1 × myfitm mix (bio-line, taunton, usa), 2 µl of genomic dna and 200 nm of either primer combinations rbcl-a f and rbcla si_rev, trnh2 and psbaf, or its5 and its4 (table 1), respectively. http://q-bank plants van valkenburg j.l.c.h. et al., on a confusingly labeled water-milfoil (myriophyllum) 3 the cycle conditions for rbcl and trnh-psba loci were as follows: 5 min at 95ºc, followed by 5 cycles of 30 s at 94ºc, 30 s at 45ºc, 30 s at 72ºc and 35 cycles of 30 s at 94ºc, 30 s at 50ºc, 30 s at 72ºc and a final extension for 10 min at 72ºc. for the its locus we used the following conditions: 5 min at 95ºc, followed by 40 cycles of 30 s at 94ºc, 30 s at 52ºc, 100 s at 72ºc and a final extension for 10 min at 72ºc. the obtained pcr products were purified using the qiaquick pcr purification kit (qiagen, venlo, the netherlands) preceding bidirectional cycle sequencing with the bigdye terminator ver. 1.1 cycle sequencing kit (thermo fisher scientific, bleiswijk, the netherlands) using the aforementioned amplification primers as sequencing primers in separate reactions according to the manufacturer’s instructions. cycle sequence products were purified with the dyeex 2.0 spin kit (qiagen,venlo, the netherlands) and subsequently sequenced using a 3500 genetic analyzer (thermo fisher scientific). consensus sequences were generated from an assembly with trace files from both sanger sequencing runs in geneious prime® 2021.1.1 (biomatters auckland, new zealand). amplification primer sequences were trimmed in the assembly, and when needed, additional trimming was performed to obtain high-quality (phred > 30) consensus sequences. illumina sequencing dna extracts were sent to genomescan (leiden, the netherlands) for illumina 150pe (paired-end) sequencing using the novaseq 6000 (myriophyllum aquaticum jl 8405, jvv 3494 and tp 3977, myriophyllum rubricaule jvv 3298, 3342, 3495, 4118 and 4119) or 100pe sequencing with the hiseq 2500 platform (laurembergia tetrandra ap 33, myriophyllum heterophyllum jvv 3651 and m. rubricaule jvv 3648) with at least 2 gb output per sample. samples that were sequenced with the novaseq 6000 were processed using the nebnext® ultra ii fs dna module and the nebnext® ultra ii ligation module. fragmentation, a-tailing and ligation of sequencing adapters and pcr using nebnext® ultra ii q5 master mix of the resulting product was performed according to the procedure described in the nebnext ultra ii fs dna module and nebnext ultra ii ligation module instruction manual. hiseq 2500 samples were processed using the nebnext® ultra dna library prep kit for illumina (new england biolabs, ipswich, usa). fragmentation of the dna using the covaris, ligation of the sequencing adapters and pcr amplification of the resulting product were performed according to the procedure described in the nebnext ultra dna library prep kit for illumina instruction manual. after preparation, the quality and yield for all samples was measured with the fragment analyzer (agilent technologies, usa). the size of the resulting products was consistent with the expected size of approximately 500–700 bp. loci primer name primer sequence reference rbcl rbcl-a f atgtcaccacaaacagagactaaagc kress & erickson 2007 rbcla si_rev gtaaaatcaagtccaccrcg kress et al. 2009 trnh-psba trnh2 cgcgcatggtggattcacaatcc tate 2002 psbaf gttatgcatgaacgtaatgctc sang et al. 1997 its its5 ggaagtaaaagtcgtaacaagg white et al. 1990 its4 tcctccgcttattgatatgc white et al. 1990 table 1. primers used in this study. european journal of taxonomy 828: 1–15 (2022) 4 de novo assembly and extraction of loci reads of samples were trimmed and de novo assembled in clc genomics workbench 21.0.3 (qiagen, aarhus, denmark) using default settings. loci of interest were extracted with blastn (standard settings) from assembled contigs using a custom blast database in geneious prime® 2021.1.1 (biomatters auckland, new zealand) containing the reference sequences of m. aquaticum its (genbank no. ky767734; lü et al. 2017), myriophyllum sp. ‘red 1’ matk and trnk3 (genbank nos fj870932 and fj861339, respectively; moody & les 2010) and m. rubricaule trnh-psba and rbcl (this paper, genbank nos mz399141 and mz399133). phylogenetic analyses sequences were aligned using muscle ver. 3.8.425 (edgar 2004) within geneious prime® 2021.1.1 (biomatters auckland, new zealand) with standard settings. we performed model testing on the alignments to select the most appropriate model for constructing a maximum likelihood tree using the model selection tool in mega x with standard settings (kumar et al. 2018). selected models were used to generate the phylogenetic trees in mega x. data accessibility the annotated dna sequences are available from the ncbi genbank database under the following accession numbers (supp. file 1): mz401372–mz401382 and ol806572–ol806573 (its), mz399118– mz399126 and ol827545–ol827546 (matk), mz399127–mz399134, mz399152–mz399156 and ol827547–ol827548 (rbcl), mz399135–mz399142, mz399157–mz399161 and ol827549– ol827550 (trnh-psba intergenic spacer) and mz399143–mz399151 and ol827551–ol827552 (trnk3). illumina data are deposited in the sequence read archive (sra) under accession numbers err6000187– err6000195, err7645488 and err7645489. see supp. file 1 for an overview. sequences are also available at https://qbank.eppo.int/plants/. results taxonomic treatment class magnoliopsida brongn. order saxifragales bercht. & j.presl. family haloragaceae r.br. genus myriophyllum ponted. ex l. myriophyllum rubricaule valk. & duist. sp. nov. urn:lsid:ipni.org:names:77299815-1 fig. 1 myriophyllum robustum auct. non hook.f. molecular ecology resources 13: 21–31 (ghahramanzadeh et al. 2013). myriophyllum brasiliense auct. non cambess.: plants in trade. myriophyllum sp. trade name ‘brasiliensis’: beringen r. 2020. — pot r. et al. 2021. — q-bank invasive plants. 2019–2021. — eppo-q-bank. 2022. diagnosis herba perennis aquatica vel paludigena. folia omnia verticillata pectinata. flores unisexuales, solae plantae femininae culturaeque cognitus. flores intra axillas foliorum solitarii. planta valde similaris myriophyllo aquatico, sed caules foliaque modice, caules purpurei et folia emergentia virida. flores feminei subrosei. https://doi.org/10.5852/ejt.2022.828.1847.7211 https://doi.org/10.5852/ejt.2022.828.1847.7211 https://qbank.eppo.int/plants/ http://www.ipni.org/urn:lsid:ipni.org:names:77299815-1 https://www.verspreidingsatlas.nl/10588# https://keys.lucidcentral.org/keys/v3/aquatic_plants/en/index.html https://q-bankplants.eu/page/organisms%20included%20display/1129 https://qbank.eppo.int/plants/taxon/z00259/ van valkenburg j.l.c.h. et al., on a confusingly labeled water-milfoil (myriophyllum) 5 etymology the species epithet is based on the purplish red color of the stem. type material netherlands • grashoek, roomweg 85, 5985 ns, golf course kapelkeshof; 51°22.0′ n, 5°56.5′ e; herbarium specimen; 13 jul. 2009; fl; j.l.c.h. van valkenburg & j. hoogveld 3495; holotype: l!; isotypes: wagpd[wag0453615]!, a!, bm!, br!, mo!, nsw!; genbank nos: mz399148 (trnk3), mz399140 (trnh-psba), mz399132 (rbcl), mz399123 (matk), mz401380 (its); sra: err6000190. additional specimens examined netherlands • wageningen, open air pond at pd; herbarium specimen; originally from wageningen, garden centre d’oude tol; 14 jun. 2007; fl; j.l.c.h. van valkenburg 3298; j.l.c.h. van valkenburg det.; wagpd[wag0453612]; genbank nos: mz399146 (trnk3), mz399138 (trnh-psba), mz399130 (rbcl), mz399121 (matk), mz401378 (its); sra: err6000191 • vinkeveen, aquaflora vinkeveen, ter aase zuwe, near junction main road vinkeveen to wilnis; herbarium specimen; cultivated at wageningen, open air pond at pd, originally from vinkeveen, collected 5 apr. 2007; 9 jul. 2007; fl; j.l.c.h. van valkenburg 3314; j.l.c.h. van valkenburg det.; wagpd[wag0453613] • herbarium specimen; grown by ‘simon van der velde waterplanten bv’, albert van ‘t hartweg 1, bleiswijk; 30 jul. 2007; j.l.c.h. van valkenburg 3342; j.l.c.h. van valkenburg det.; wagpd[wag0453614]; genbank nos: mz399147 (trnk3), mz399139 (trnh-psba), mz399131 (rbcl), mz399122 (matk), mz401377 (its); sra: err6000192 • wageningen, garden centre d’oude tol; herbarium specimen; culta the netherlands; 11 apr. 2009; j.l.c.h. van valkenburg 3472; j.l.c.h. van valkenburg det.; l[l 0767219; l 0909277]; genbank no.: jx100590 (trnh-psba) • wageningen, geertjesweg 15, plant protection service; herbarium specimen; culta, first received 1 may 2009 from stoffels international bv, maalbekerweg 14, 5951 nt, belfeld, the netherlands; 11 sep. 2009; fl; j.l.c.h. van valkenburg 3510; j.l.c.h. van valkenburg det.; l[l 0767220; l 0909323]; genbank nos: jx100591 (trnh-psba), jx100753 (rbcl) • wageningen, greenhouse plant protection service; herbarium specimen; culta from plant obtained at garden centre; 28 jul. 2011; j.l.c.h. van valkenburg 3648; j.l.c.h. van valkenburg det.; wagpd[wag0453609]; genbank nos: mz399149 (trnk3), mz399158 (trnh-psba), mz399153 (rbcl), mz399124 (matk), mz401372 (its); sra: err6000195 • hoogeveen, kinholt; 52°43.34′ n, 6°26.00′ e; herbarium specimen; 5 nov. 2018; j.l.c.h. van valkenburg 3971; j.l.c.h. van valkenburg det.; wagpd[wag0453559]; genbank nos: mz399159 (trnh-psba), mz399154 (rbcl) • steenwijk, het eemter; 52°47.415′ n, 6°08.840′ e; herbarium specimen; 8 sep. 2020; j.l.c.h. van valkenburg 4116; j.l.c.h. van valkenburg det.; wag[wag.1971556], wagpd[wag0452369]; genbank nos: mz399160 (trnh-psba), mz399155 (rbcl), mz401381 (its) • hattem, palmstraat; 52°27.768′ n, 6°04.428′ e; herbarium specimen; 8 sep. 2020; j.l.c.h. van valkenburg 4117; j.l.c.h. van valkenburg det.; wag[wag.1971558], wagpd[wag0452378]; genbank nos: mz399161 (trnh-psba), mz399156 (rbcl), mz401382 (its) • klazienaveen; 52°44.580′ n, 7°00.615′ e; herbarium specimen; 7 oct. 2020; j.l.c.h. van valkenburg 4118; j.l.c.h. van valkenburg det.; wagpd[wag0452243]; genbank nos: mz399150 (trnk3), mz399141 (trnh-psba), mz399133 (rbcl), mz399125 (matk), mz401375 (its); sra: err6000193 • wageningen, greenhouse nvwa; herbarium specimen; culta, origin garden center more than 8 years ago; 8 oct. 2020; j.l.c.h. van valkenburg 4119; j.l.c.h. van valkenburg det.; wagpd[wag0452240]; genbank nos: mz399151 (trnk3), mz399142 (trnh-psba), mz399134 (rbcl), mz399126 (matk), mz401376 (its); sra: err6000194. hungary • borsod-abaúj-zemplén, kács; 47°57.44′ n, 20°37.00′ e; herbarium specimen; 20 sep. 2019; a. mesterházy s.n.; j.l.c.h. van valkenburg det.; wagpd[wag0452409]; genbank nos: mz399157 (trnh-psba), mz399152 (rbcl). european journal of taxonomy 828: 1–15 (2022) 6 description dioecious amphibic or aquatic herb. stem unbranched or with up to 6 branches per 20 cm, often rooting at submerged and lower emerged nodes; submerged part green to tinged red-brown and internodes 10–50 mm long; emerged part red or purplish red and internodes 3–25 mm long. hydathodes few at base of submerged leaves, many at base of emerged leaves, ca 1 mm long, filiform, pale or reddish brown. leaves in whorls of 4 or 5, pinnatifid with pinnae placed opposite and/or alternate. submerged leaves 10–25(–50) × 3–10 mm, olive green or turning pale to dark reddish brown; pinnae 12–21, 3–14(–30) × 0.1–0.2 mm. emerged leaves (7–)10–25 × 3–8 mm, bright green to bluish green, not glaucous, sometimes tinged red brown or pinnae red-tipped; pinnae (5–)7–21, 2.3–9 × 0.15–0.3(–0.5) mm. flowers solitary in the axils of the emerged leaves, only female known, tinged pink. pedicel 0.3–0.5 mm long. bracteoles 2 at basis of pedicel, 0.5–1 × 0.1 mm, with 1–3 alternate and filiform lobes. sepals 4, erecto-patent to reflexed, 0.5–0.7 × 0.2–0.3 mm, margin remotely fimbriate. petals absent. stamens absent. ovary 4-sulcate, with 4 styles with feathery and more or less inrolled stigmas. fruits unknown. amphibic herb with pinnate leaves in whorls of 4 or 5, only known from female plants and (escapes from) horticulture. differs from m. aquaticum in its generally more modest dimensions, the stems being purplish red, the emergent leaves being green to bluish green and not glaucous (i.e., leaves without waxy coating) and the female flowers being pinkish (fig. 1). fig. 1. plants, prior to collection, of m. rubricaule valk. & duist. sp. nov. (valkenburg 3495; red stem) growing amidst m. aquaticum (valkenburg 3494; green stem) in an artificial pond at a golf course, illustrating the more modest dimensions and color differences (13 july 2009, grashoek, the netherlands). van valkenburg j.l.c.h. et al., on a confusingly labeled water-milfoil (myriophyllum) 7 distribution origin unknown. known from cultivation (netherlands, belgium) and as escapes from cultivation since 2018 in the netherlands (hoogeveen, steenwijk, hattem, klazienaveen) and belgium (maasmechelen, houthalen-helchteren, brugge, gent, beauraing, waimes; all confirmed from photographs at waarnemingen.be), and at least since 2019 in hungary (kács). it is unclear whether the observations in belgium before 2018 (in retrospect the first observation dates from 2012; see waarnemingen.be records 70176185 and 95286205) refer to escapes or planted material. note if grown in particularly nutrient rich and/or high light conditions plants develop much bigger submerged leaves (e.g., valkenburg 3298 and 4116 wagpd). similar species because both the submerged and emergent leaves are whorled, pectinate and not much different in length, and the inflorescence is emergent, the species fits in section pectinatum m.l.moody & d.h.les. this section includes m. aquaticum and m. mattogrossense hoehne (moody & les 2010). myriophyllum aquaticum differs from m. rubricaule sp. nov. in having green stems only turning red when grown as a potted plant or in unfavorable conditions, but never purplish red, with leaves in whorls of 4–6, submerged leaves 25–45 mm long and green or red brown, emerged leaves 25–35 mm long and bluish green glaucous (i.e., with a waxy coating that can be rubbed of), and white flowers. myriophyllum mattogrossense has green stems, small, globular, sessile glands on leaves and stems, submerged leaves 10–35 mm wide, and bisexual flowers (crow & ritter 1999). while performing the dna barcoding study of m. aquaticum (ghahramanzadeh et al. 2013), we initially thought that the plants described here as a new species belonged to m. robustum. this is a species resembling m. aquaticum with similar emergent pectinate leaves and solitary axillary flowers. however, m. robustum is described as having hermaphrodite flowers whereas m. aquaticum is dioecious (orchard 1980). shortly after our paper was published, plants of m. robustum were received from new zealand and were grown in a greenhouse. the cultivated plants had more robust erect emergent stems that were pink in the upper part, and subglaucous leaves that were oblong in outline with an acute tip. surprisingly, the plants produced only female flowers in the greenhouse (valkenburg 3739 wagpd). grown outdoors in a mesocosm in later years, the plants first produced female flowers, a row of hermaphrodite flowers and then, distally, male flowers (valkenburg 3853 wagpd). distinguishing molecular features of myriophyllum rubricaule sp. nov. as described above, m. rubricaule sp. nov. is morphologically distinguishable from m. aquaticum. moody & les (2010) described two specimens, m. sp. ‘red 1’ and m. sp. ‘red 2’, that were morphologically and genetically related to m. aquaticum. we made a molecular comparison of m. rubricaule, m. aquaticum, m. sp. ‘red 1’ and m. sp. ‘red 2’ using the same molecular markers (moody & les 2010), namely the nuclear locus its (partial 18s, its1, 5.8s, its2, partial 28s), and the chloroplast loci trnk3 and matk to find out how m. sp. ‘red 1’ and m. sp. ‘red 2’ relate to m. rubricaule (figs 2–3). we included myriophyllum heterophyllum michx. and laurembergia tetrandra (schott) kanitz as more distantly related species within the haloragaceae in this comparison. based on its (fig. 2), all m. rubricaule sp. nov. specimens showed 100% sequence similarity. myriophyllum sp. ‘red 1’ was most similar to m. rubricaule sharing 98.62% sequence identity. we only observed differences in the its1 sequence (38–42delccccg and 87delg). interestingly, the specimens of m. aquaticum group with m. sp. ‘red 2’, albeit with a relatively low confidence value, sharing 96.27% and 93.65% identity with m. rubricaule, respectively. https://waarnemingen.be/ https://waarnemingen.be/ european journal of taxonomy 828: 1–15 (2022) 8 the trnk3 and matk loci were extracted from the sequence data and subsequently concatenated, before tree generation. similarly to its, the trnk3-matk sequences were identical for all samples of m. rubricaule sp. nov. (fig. 3). intriguingly, the m. sp. ‘red 2’ and m. aquaticum jl 8405 trnk3-matk loci were identical to m. rubricaule. myriophyllum aquaticum jvv 3494 and tp 3977 showed 99.88% similarity with m. rubricaule and m. sp. ‘red 2’ with only two single nucleotide polymorphisms (snps; t32a and t1243a). myriophyllum sp. ‘red 1’ differs on 10 positions with m. aquaticum, m. rubricaule and m. sp. ‘red 2’ sharing 99.63% similarity. finally, we compared two other well-known chloroplast loci, rbcl and trnh-psba, to identify more distinguishing molecular features between m. aquaticum and m. rubricaule sp. nov. we incorporated previously described sanger sequences (ghahramanzadeh et al. 2013) as well as newly generated sanger and illumina sequences in the comparison (supp. file 1 and 2). for trnh-psba, all 12 specimens of m. rubricaule were identical to each other. seven out of eight m. aquaticum were identical, for m. aquaticum jl 8405 we observed one snp (g230a) (supp. file 3). however, for two samples of m. aquaticum not all sanger data could be resolved resulting in two ambiguities: m188a for m. aquaticum jvv 3329 and k255t for m. aquaticum hd 426. a clear distinction was observed towards the 5’-end of the trnh-psba intergenic spacer (nucleotides #237–292) where 13 snps were identified between the specimens of m. aquaticum and m. rubricaule resulting in approximately 96% similarity. interestingly, we did not find any differences between m. aquaticum and m. rubricaule rbcl sequences (supp. file 4). even the distantly related plant species l. tetrandra shared 97.29% of rbcl identity. in contrast, for trnh-psba, only a ~ 69–73% identity score was observed between l. tetrandra and m. rubricaule and m. aquaticum. discussion during our study of the confusingly labeled water-milfoil in trade, we consulted the most relevant revisions and new taxonomical publications on the genus myriophyllum (meijden & caspers 1971; orchard 1980, 1981, 1986; orchard & kasselmann 1992; wang et al. 2002; yu et al. 2002; wang & yu 2007). after attempting to identify the plants using the revised key for new zealand (orchard 1980), the two choices where m. aquaticum and m. robustum. firstly, our taxon cannot be m. aquaticum because of the color of the leaves in the field, and secondly it is not m. robustum because the flowers are not hermaphroditic. the revision for south america (orchard 1981) likewise does not yield a result as m. aquaticum has glaucous leaves and m. mattogrossense is disqualified by the plants being weak and having hermaphrodite flowers. in the revision for australia (orchard 1985), m. aquaticum is also the only realistic option. none of the species covered by the myriophylllum treatment for malesia (meijden & caspers 1971), nor recent publications for new species in asia (wang et al. 2002; yu et al. 2002; wang & yu 2007) match our material as all of those species have dimorphic leaves. the new species fits in section pectinatum m.l.moody & les (2010: 136). they formally described the section, with m. aquaticum (vell.) verdc. as the type species, as follows: “folia omnia verticillata; difert a m. subsect. myriophyllum folia omnia emersa pectinata et difert a m. subsect. isophylleae folia emersa comparate non redacta de foliis submersis. diagnosis: submerged and emergent leaves whorled and pectinate. emergent leaves pectinate and not highly reduced in relation to submerged leaves. plants dioecious or flowers all hermaphrodite.” species within this section have in common the emergent inflorescences and all leaves whorled and pectinate; the emergent leaves are not highly reduced. the definition as such would also fit for m. robustum, that is placed in m. subsection myriophyllum but is the exception in the subsection for having all pectinate emergent leaves that are not highly reduced in relation to submerged leaves. https://doi.org/10.5852/ejt.2022.828.1847.7211 https://doi.org/10.5852/ejt.2022.828.1847.7213 https://doi.org/10.5852/ejt.2022.828.1847.7215 https://doi.org/10.5852/ejt.2022.828.1847.7217 van valkenburg j.l.c.h. et al., on a confusingly labeled water-milfoil (myriophyllum) 9 fig. 2. maximum likelihood tree based on its (partial 18s, its1, 5.8s, its2, partial 28s) sequences. the tamura 3 parameter model was used (tamura 1992). the tree with the highest log likelihood (-1398.71) is shown. bootstrap values after 1000 replicates are expressed as percentages. a discrete gamma distribution was used to model evolutionary rate differences among sites (5 categories (+g, parameter = 0.4031)). the tree is drawn to scale, with branch lengths measured in the number of substitutions per site. laurembergia tetrandra ap 33 was used as an outgroup. european journal of taxonomy 828: 1–15 (2022) 10 fig. 3. maximum likelihood tree based on concatenated trnk3-matk sequences. this tree was inferred by using the maximum likelihood method and tamura 3 parameter model (tamura 1992). the tree with the highest log likelihood (-2853.86) is shown. bootstrap values after 1000 replicates are expressed as percentages. the tree is drawn to scale, with branch lengths measured in the number of substitutions per site. laurembergia tetrandra ap 33 was used as an outgroup. van valkenburg j.l.c.h. et al., on a confusingly labeled water-milfoil (myriophyllum) 11 its, and the chloroplast locus trnh-psba demonstrate that m. aquaticum markedly differs from m. rubricaule sp. nov. (fig. 2 and supp. file 3). the rbcl locus is not suitable to distinguish between the species, little variation was observed even with the more distantly related l. tetrandra (supp. file 4). intriguingly, we observed that the concatenated loci trnk3-matk of m. aquaticum jl 8405 were identical to m. rubricaule. however, only two snps were observed in comparison with the other m. aquaticum specimens (fig. 3). furthermore, stark genetic differences between m. aquaticum jl 8405 and m. rubricaule were found in comparison with the other loci. moody & les (2010) described two specimens, m. sp. ‘red 1’ and ‘red 2’, that were closely related to m. aquaticum and showed intriguing morphological similarities with the specimens of m. rubricaule sp. nov. described here. based on the chloroplast loci, m. sp. ‘red 2’ is identical to m. rubricaule. however, its sequences of m. rubricaule are more similar to m. sp. ‘red 1’. based on these data, we cannot conclude whether either m. sp. ‘red 1’, m. sp. ‘red 2’ or both are m. rubricaule. the elucidation of additional dna sequences of the other loci described herein for m. sp. ‘red 1’ and 2 may help in resolving this question. interestingly, our comparison between chloroplast loci yielded results that deviated from the analyses of moody & les (2010). in our hands, m. aquaticum seems to be more closely related to m. sp. ‘red 2’ than to m. sp. ‘red 1’, where the former only had 2 snps compared to m. aquaticum, whereas the latter showed 10. possibly, the its or chloroplast sequences of m. sp. ‘red 1’ and m. sp. ‘red 2’ were switched when they were deposited into genbank. if we assume this to be the case either m. sp. ‘red 1’ or m. sp. ‘red 2’ is likely a m. rubricaule specimen. in addition to species in trade of unknown origin as further mentioned by thum et al. (2012) incorrect labeling of plants is widespread and may be caused by negligence or willful disrespect of regulations. what we have noticed so far with respect to species of myriophyllum in trade in the netherlands is that, in general, negligence of proper labeling causes confusion in the proper identity of the species in trade (van valkenburg & boer 2015; van valkenburg et al. 2015). concerning the new species described, the only mislabeling observed was m. proserpinacoides gillies ex hook. & arn. 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(ed.) abstracts 47th robson meeting, reading, england: 16–17, waterland management, united kingdom. van valkenburg j., duistermaat l., westenberg m. & van de vossenberg b. 2015. myriophyllum in trade in western europe, what species are we really talking about? in: aquatic plants 2015, 14th international symposium on aquatic plants: 57. the centre for ecology & hydrology, edinburgh, united kingdom. verbrugge l.n.h., leuven r.s.e.w., van valkenburg j.l.c.h. & van den born r.j.g. 2014. evaluating stakeholder awareness and involvement in risk prevention of aquatic invasive plant species by a national code of conduct. aquatic invasions 9: 369–381. https://doi.org/10.3391/ai.2014.9.3.11 wang d., yu d. & li z.-y. 2002. myriophyllum exasperatum (haloragaceae), a new species from china. annales botanici fennici 39: 267–271. wang d. & yu d. 2007. a new subspecies of myriophyllum oguraense (haloragaceae) from china. annales botanici fennici 44: 227–230. white t.j., bruns t., lee s. & taylor j. 1990. amplification and direct sequencing of fungal ribosomal rna genes for phylogenetics. in: innis m.a., gelfand d.h., snisnky j.j. & white t.j. (eds) pcr protocols: a guide to methods and applications: 315–332. academic press, san diego. yu d., wang d. & li z.-y. 2002. taxonomic revision of the genus myriophyllum (haloragaceae) in china. rhodora 104: 396–421. manuscript received: 23 september 2021 manuscript accepted: 19 april 2022 published on: 5 july 2022 topic editor: frederik leliaert desk editor: radka rosenbaumová printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. https://doi.org/10.1093/oxfordjournals.molbev.a040752 http://sweetgum.nybg.org/science/ih/ https://doi.org/10.1007/s10530-011-0130-8 https://doi.org/10.3391/ai.2014.9.3.11 european journal of taxonomy 828: 1–15 (2022) 14 appendix 1 specimens of other species used in the present study. myriophyllum aquaticum argentina • estancia ‘santa teresa’, dep. mburucuyá, prov. corrientes; herbarium specimen; 7 sep. 1956; fl; t.m. pedersen 3977; t.m. pedersen det.; l[l.2571172]; genbank nos: ol827552 (trnk3), ol827550 (trnh-psba), ol827548 (rbcl), ol827546 (matk), ol806573 (its); sra: err7645488. brazil • rio grande do sul, 16 km n da br 290 km 380, ne de rosario; herbarium specimen; 16 oct. 1971; j.c. lindeman, b.e. irgang & j.f.m. valls 8405; j.c. lindeman det.; u[u.1329313]; genbank nos: ol827551 (trnk3), ol827549 (trnh-psba), ol827547 (rbcl), ol827545 (matk), ol806572 (its); sra: err7645489. netherlands • eindhoven, jan van eijckgracht; 51°25.73′ n, 5°30.80′ e; herbarium specimen; 2 jul. 2007; h. duistermaat 418; h. duistermaat det.; l[l 0768064 ]; genbank nos: jx100595 (trnhpsba), jx100747 (rbcl) • wageningen, open air pond at plant protection service; herbarium specimen; plants raised in pond, obtained as potplant (cultivated above ground) on 15 may 2007 from wageningen, garden centre d’oude tol.; 8 jun. 2007; fl; j.l.c.h. van valkenburg 3294; j.l.c.h. van valkenburg det.; wagpd[wag0453553]; genbank nos: jx100592 (trnh-psba), jx100749 (rbcl) • emmer erfscheidenveen, kanaal e, middenweg zw; 52°48.30′ n, 6°59.38′ e; herbarium specimen; 20 jul. 2007; fl; j.l.c.h. van valkenburg 3329; j.l.c.h. van valkenburg det.; wagpd[wag0453554]; genbank nos: jx100589 (trnh-psba), jx100752 (rbcl) • grashoek, roomweg 85, 5985 ns, golf course kapelkeshof; 51°22.0′ n, 5°56.5′ e; herbarium specimen; 13 jul. 2009; fl; j.l.c.h. van valkenburg 3494; j.l.c.h. van valkenburg det.; wagpd[wag0453552]; genbank nos: mz399144 (trnk3), mz399136 (trnhpsba), mz399128 (rbcl), mz399119 (matk), mz401379 (its); sra: err6000188 • wageningen, geertjesweg 15, plant protection service; herbarium specimen; culta, first received 1 may 2009 from stoffels internationaal bv, maalbekerweg 14, 5951 nt, belfeld, the netherlands; 11 sep. 2009; fl; j.l.c.h. van valkenburg 3511; j.l.c.h. van valkenburg det.; l[l 0909322]; genbank nos: jx100594 (trnh-psba), jx100754 (rbcl). france • bretagne, plouay; 47°55′ n, 3°20′ e; herbarium specimen; 25 jul. 2007; h. duistermaat 426; h. duistermaat det.; l[l 0768057]; genbank nos: jx100596 (trnh-psba), jx100748 (rbcl). myriophyllum heterophyllum netherlands • wageningen, greenhouse plant protection service; herbarium specimen; culta; 28 jul. 2011; j.l.c.h. van valkenburg 3651; j.l.c.h. van valkenburg det.; wagpd[wag0453530]; genbank nos: mz399145 (trnk3), mz399137 (trnh-psba), mz399129 (rbcl), mz399120 (matk), mz401373 (its); sra: err6000189. laurembergia tetrandra gabon • ogooué-maritime, gamba, just w of the shell terminal; 2.47300° s, 10.00600° e; herbarium specimen; 10 jan. 1998; a.s.j. van proosdij 33; j.j. wieringa det.; wag[wag.1550989]; genbank nos: mz399143 (trnk3), mz399135 (trnh-psba), mz399127 (rbcl), mz399118 (matk), mz401374 (its); sra: err6000187. van valkenburg j.l.c.h. et al., on a confusingly labeled water-milfoil (myriophyllum) 15 supplementary material supp. file 1. sra and genbank accession numbers of sequences used in this study. https://doi.org/10.5852/ejt.2022.828.1847.7211 supp. file 2. length of sequences in nucleotides that were used in this study. https://doi.org/10.5852/ejt.2022.828.1847.7213 supp. file 3. maximum likelihood tree based on trnh-psba sequences. the tamura 3 parameter model was used (tamura 1992). the tree with the highest log likelihood (-898.13) is shown. bootstrap values after 1000 replicates are expressed as percentages. the tree is drawn to scale, with branch lengths measured in the number of substitutions per site. laurembergia tetrandra ap 33 was used as an outgroup. https://doi.org/10.5852/ejt.2022.828.1847.7215 supp. file 4. maximum likelihood tree based on rbcl sequences. this tree was inferred by using the maximum likelihood method and jukes-cantor model (jukes & cantor 1969).the tree with the highest log likelihood (-883.52) is shown. bootstrap values after 1000 replicates are expressed as percentages. the tree is drawn to scale, with branch lengths measured in the number of substitutions per site. laurembergia tetrandra ap 33 was used as an outgroup. https://doi.org/10.5852/ejt.2022.828.1847.7217 https://doi.org/10.5852/ejt.2022.828.1847.7211 https://doi.org/10.5852/ejt.2022.828.1847.7213 https://doi.org/10.5852/ejt.2022.828.1847.7215 https://doi.org/10.5852/ejt.2022.828.1847.7217 121 european journal of taxonomy 833: 121–142 issn 2118-9773 https://doi.org/10.5852/ejt.2022.833.1889 www.europeanjournaloftaxonomy.eu 2022 · zhang y.-h. et al. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:c82a34e8-7a30-4581-8c14-269c178a66fd new species of genus atractides koch, 1837 (acari: hydrachnidiae, hygrobatidae) from qinghai, china yu-hao zhang 1,#, hai-tao li 2,#, cai-yun li 3 & jian-jun guo 4,* 1,2,3,4 institute of entomology, guizhou university, scientific observing and experimental station of crop pest in guiyang, ministry of agriculture and rural affairs of the p.r. china, guiyang 550025, p.r. china. * corresponding author: jjguo@gzu.edu.cn 1 email:1289568720@qq.com 2 email: lhtzlp424666@outlook.com 3 email: 1377195309@qq.com # yu-hao zhang and hai-tao li contributed equally to this study 1 urn:lsid:zoobank.org:author:9413e3a0-c9fb-4ccb-8f60-de9454cf856d 2 urn:lsid:zoobank.org:author:5b5d8a0d-55b4-4ec7-a6b5-a3b4b7a8a475 3 urn:lsid:zoobank.org:author:5e4c3dc0-812d-4822-9b1f-119eee651c4c 4 urn:lsid:zoobank.org:author:60680105-e469-4f51-af3d-094bed1c361e abstract. the paper deals with five new species of the genus atractides koch, 1837 (acari: hydrachnidiae, hygrobatidae) collected from qinghai province, p.r. china, atractides (atractides) biprojectus zhang, li & guo sp. nov., atractides (atractides) smiti zhang, li & guo sp. nov., atractides (atractides) menyuanensis zhang, li & guo sp. nov., atractides (atractides) longiprojectus zhang, li & guo sp. nov. and atractides (atractides) xianmiensis zhang, li & guo sp. nov. all the new species are described and illustrated in detail, and all the type specimens are deposited in the institute of entomology, guizhou university, guiyang, china (gugc). keywords. water mite, atractides, new species, taxonomy, qinghai-tibet plateau. zhang y.-h., li h.-t., li c.-y. & guo j.-j. 2022. new species of genus atractides koch, 1837 (acari: hydrachnidiae, hygrobatidae) from qinghai, china. european journal of taxonomy 833: 121–142. https://doi.org/10.5852/ejt.2022.833.1889 introduction atractides koch, 1837 are reported in all biogeographical areas except for australia and antarctica (cook 1974; gerecke 2003). most species are known from the northern hemisphere, and only found in clean waters where the substratum is well preserved, so they are probably sensitive to many forms of human impact (gerecke 2003; pešić & smit 2011; gülle et al. 2015). https://doi.org/10.5852/ejt.2022.833.1889 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:c82a34e8-7a30-4581-8c14-269c178a66fd https://orcid.org/0000-0002-0523-6704 mailto:jjguo@gzu.edu.cn mailto:jjguo@gzu.edu.cn mailto:jjguo@gzu.edu.cn mailto:jjguo@gzu.edu.cn mailto:jjguo@gzu.edu.cn mailto:1289568720@qq.com mailto:lhtzlp424666@outlook.com mailto:1377195309@qq.com http://zoobank.org/urn:lsid:zoobank.org:author:9413e3a0-c9fb-4ccb-8f60-de9454cf856d http://zoobank.org/urn:lsid:zoobank.org:author:5b5d8a0d-55b4-4ec7-a6b5-a3b4b7a8a475 http://zoobank.org/urn:lsid:zoobank.org:author:5e4c3dc0-812d-4822-9b1f-119eee651c4c http://zoobank.org/urn:lsid:zoobank.org:author:60680105-e469-4f51-af3d-094bed1c361e https://doi.org/10.5852/ejt.2022.833.1889 european journal of taxonomy 833: 121–142 (2022) 122 currently, the genus atractides has four subgenera (atractides koch, 1837; tympanomegapus thor, 1923; polymegapus k. viets, 1926; maderomegapus lundblad, 1941), and about 405 species have been described from all over the world (k.o. viets 1987; gerecke 2003; pešić et al. 2018; pešić & smit 2018, 2021a, 2021b; smit 2020; gülle et al. 2021; http://www.watermite.org/). based on previous information, two subgenera and 18 species of atractides have been reported for the chinese fauna: atractides s. str. with 14 species and tympanomegapus thor, 1923 with 4 species (jin 1997; yi et al. 2010; wang & jin 2012, 2013; wang et al. 2015). the qinghai province lies in the west of china, on the qinghai-tibet plateau. the qinghai-tibet plateau is the highest and largest plateau in the world, which has a wide range of geological, topographical and climatic gradients. therefore, there is a greater biodiversity than in the surrounding lowlands and other high elevation regions (sun et al. 2014). through a joint investigation of water mites in qinghai province, five new species are found: atractides (atractides) biprojectus zhang, li & guo sp. nov., atractides (atractides) smiti zhang, li & guo sp. nov., atractides (atractides) menyuanensis zhang, li & guo sp. nov., atractides (atractides) longiprojectus zhang, li & guo sp. nov. and atractides (atractides) xianmiensis zhang, li & guo sp. nov. all the new species are described and illustrated in detail. material and methods specimens in this study were all collected by hai-tao li from qinghai province, p.r. china. water mites were collected by hand netting, sorted on the spot from the living material, preserved in koenike’s fluid and dissected following jin (1997) under a motic smz-168 stereo microscope. specimens were observed under a nikon ni-e microscope (with a nikon ds-ri2 camera). all illustrations were edited with adobe photoshop cc2018. all measurements are given in micrometers (μm) following gerecke (2003). terms are modified from jin (1997) and gerecke (2003). the following abbreviations are used: a.s.l. = above sea level a1 = preantennal glandularia a2 = postantennal glandularia ac = acetabulum (pl. acetabula, numbered 1 to 3) acg = anterior coxal group (cx-i+cx-ii) ap = acetabular plates c1–c4 = coxoglandularia 1–4 cx-i–iv = coxae i–iv d1–d4 = dorsoglandularia 1–4 dl = dorsal length hb = central height il = lateral length i-l-1–6, etc. = the first–sixth segment of the first leg, etc. l = length l1–l4 = lateroglandularia 1–4 ml = median length o1 = preocularia o2 = postocularia p-1–p-5 = the first–fifth segment of palp pcg = posterior coxal group (cx-iii+cx-iv) http://www.watermite.org/ zhang y.-h. et al., new species of atractides from china 123 s-1 = proximal large ventral seta at i-l-5 s-2 = distal large ventral seta at i-l-5 so1–so5 = slit organs 1–5 v1–v4 = ventroglandularia 1–4 w = width holotype and paratypes are deposited in the institute of entomology, guizhou university, guiyang, china (gugc). results class arachnida lamarck, 1801 order trombidiformes reuter, 1909 superfamily hygrobatoidea koch, 1842 family hygrobatidae koch, 1842 genus atractides koch, 1837 subgenus atractides koch, 1837 atractides (atractides) biprojectus zhang, li & guo sp. nov. urn:lsid:zoobank.org:act:06ca408f-1fc6-4fd0-aa2a-e94b784621c0 figs 1–3 diagnosis male dorsal muscle attachment unsclerotized. i-l-5 longish, s-1 and s-2 with blunt tips and with a narrow setal interspace between them; i-l-6 curved. ac in an obtuse triangle, v1 separated from v2. p-2 and p-3 with a ventral projection respectively; p-4 divided by two long ventral hairs in sectors 2:3:1, sword seta between two ventral hair insertions and near the terminal. female similar to male. ventral projection of p-2 not obvious, and p-3 ventral margin nearly straight. etymology the latin prefix ‘bi-’ means two, in the male of the new species p-2 and p-3 are with a ventral projection respectively. type material holotype china • ♂; qinghai province, huangnan tibetan autonomous prefecture, zeku county, maixiu town; 35°18′64′′ n, 101°52′32′′ e; 3201 m a.s.l.; 17 jul. 2020; hai-tao li leg.; running water; gugc, slide no. qh-hy-2020071701. paratypes china • 3 ♂♂; same collection data as for holotype; gugc, slides no. qh-hy-2020071702 to 2020071704 • 1 ♀; same collection data as for holotype; gugc, slide no. qh-hy-2020071705. description male (n = 4) idiosoma oval; dorsal muscle attachment unsclerotized, o2 and d1 at the same level; setae of d1 and d2 longer than others, setae of d2 reaching to d3; all slit organs visible, so1 near the eye capsule and at http://zoobank.org/urn:lsid:zoobank.org:act:06ca408f-1fc6-4fd0-aa2a-e94b784621c0 european journal of taxonomy 833: 121–142 (2022) 124 fig. 1. atractides (atractides) biprojectus zhang, li & guo sp. nov. a–e. holotype, ♂ (gugc, slide no. qh-hy-2020071701). f. paratype, ♀ (gugc, slide no. qh-hy-2020071705). a. idiosoma, dorsal view. b. idiosoma, ventral view. c. gnathosoma. d. chelicera. e. palp. f. palp. scale bars = 100 μm. zhang y.-h. et al., new species of atractides from china 125 the level of o1, so2 in front of l2, so3 near d2, so4 at the middle of d3 and l4; so5 behind d4 (fig. 1a). acg fused together and with a suture, pcg separated; apodemes from acg not reaching to cx-iv. acetabula three pairs and in an obtuse triangle; v1 separated from v2; v3 and v4 forming a trapezoid, v4 at the same level as the middle of ap (fig. 1b). palp five-segmented; p-2 and p-3 with a ventral projection respectively; p-4 with numerous dorsal hairs, and divided by two long ventral hairs in sectors 2:3:1, sword seta between two ventral hair insertions and near the terminal (fig. 1e). i-l-5 longish, s-1 and s-2 both with blunt tips and with a narrow setal interspace between them; i-l-6 curved (fig. 3a). female (n = 1) similar to male (fig. 2). setae of d1 and d2 shorter than that in male (fig. 2a); ventral projection of p-2 not obvious, and p-3 ventral margin nearly straight (fig. 1f). measurements male (n = 4) idiosoma l 641 (585–645), w 546 (443–546); coxal field l 316 (314–325), cx-iii w 333 (333–347), acg il 237 (234–239), ml 125 (107–125), w 268 (268–284); infracapitular bay l 123 (118–127); genital field l 120 (119–120), ac1–3 l 34 (33–38), 34 (34–42), 37(37–40); chelicera l 205 (204–207); infracapitulum l 159 (140–177); palp dl: p-1 29 (29–32), p-2 74 (68–74), p-3 62(59–66), p-4 95(95– 99), p-5 27 (27–29); legs segments: i-l-1 dl 48 (46–52), i-l-2 dl 93 (89–93), i-l-3 dl 80 (78–80), i-l-4 dl 119 (119–125), i-l-5 dl 163 (163–176), hb 53 (53–60), i-l-6 dl 119 (119–122), hb 22 (21–22), s-1 l 90 (78–90), s-2 l 74 (74–75); dl: ii-l-1 47 (47–50), ii-l-2 82 (74–82), ii-l-3 71 (70–74), ii-l-4 92 (92–102), ii-l-5 116 (116–120), ii-l-6 124 (124–127); dl: iii-l-1 56 (53–57), iii-l-2 90 (81–90), iii-l-3 75 (73–77), iii-l-4 124 (124–125), iii-l-5 146 (146–154), iii-l-6 141 (133–147); dl: iv-l-1 117 (117–120), iv-l-2 105 (102–113), iv-l-3 138 (135–138), iv-l-4 174 (174–180), iv-l-5 199 (199–208), iv-l-6 163 (162–170). fig. 2. atractides (atractides) biprojectus zhang, li & guo sp. nov. a–b. paratype, ♀ (gugc, slide no. qh-hy-2020071705). a. idiosoma, dorsal view. b. idiosoma, ventral view. scale bar = 100 μm. european journal of taxonomy 833: 121–142 (2022) 126 female (n = 1) idiosoma l 902, w 721; coxal field l 390, cx-iii w 477, acg il 272, ml 137, w 355; infracapitular bay l 165; gonopore l 146, ap l 140, ac1–3 l 43, 47, 47; chelicera l 267; infracapitulum l 222; palp dl: p-1 38, p-2 84, p-3 84, p-4 123, p-5 34; legs segments: i-l-1 dl 64, i-l-2 dl 113, i-l-3 dl 114, i-l-4 dl 176, i-l-5 dl 242, hb 77, i-l-6 dl171, hb 26, s-1 l 120, s-2 l 106; dl: ii-l-1 72, ii-l-2 fig. 3. atractides (atractides) biprojectus zhang, li & guo sp. nov., holotype, ♂ (gugc, slide no. qh-hy-2020071701). a–d. i-l–iv-l. scale bar = 100 μm. zhang y.-h. et al., new species of atractides from china 127 98, ii-l-3 96, ii-l-4 135, ii-l-5 153, ii-l-6 162; dl: iii-l-1 69, iii-l-2 100, iii-l-3 107, iii-l-4 168, iii-l-5 193, iii-l-6 181; dl: iv-l-1 153, iv-l-2 141, iv-l-3 184, iv-l-4 231, iv-l-5 267, iv-l-6 211. remarks the new species atractides (atractides) biprojectus sp. nov. is similar to atractides yazdensis pešić, smit & saboori, 2021 in the following points: (1) male p-2 and p-3 with ventral projections; (2) setae s-1 and s-2 separated, with a narrow setal interspace; (3) v1 separated from v2. however, a. (a.) biprojectus differs from a. yazdensis in following aspects: (1) p-4 sword seta between two ventral hair insertions in a. (a.) biprojectus, but slightly proximal to posteroventral hair in a. yazdensis; (2) apodemes from acg not reaching to cx-iv in a. (a.) biprojectus, but reaching to cx-iv in a. yazdensis; (3) genital field of a. (a.) biprojectus much rounder than that of in a. yazdensis (pešić et al. 2021). atractides (atractides) smiti zhang, li & guo sp. nov. urn:lsid:zoobank.org:act:d2da763a-88e0-4432-810f-76bc5fbf1f3b figs 4–5 diagnosis muscle attachments between d3 sclerotized. i-l-5 longish, s-1 and s-2 both with blunt tips and with a narrow setal interspace between them; i-l-6 curved. ac in an obtuse triangle; v1 fused to v2, excretory pore surrounded by sclerotized ring. p-2 and p-3 ventral margin slightly straight; p-4 divided by two ventral hairs in sectors 3:3:2, sword seta at the middle of p-4. etymology the species is named after dr harry smit in appreciation of his contributions to the taxonomy of water mites. type material holotype china • ♂; qinghai province, huangnan tibetan autonomous prefecture, zeku county, maixiu town; 35°18′64′′ n, 101°52′32′′ e; 3201 m a.s.l.; 17 jul. 2020; hai-tao li leg.; running water; gugc, slide no. qh-hy-2020071706. description male idiosoma oval; o2 and l1 at the same level; setae of d1 about two thirds of the distance from d1 to d2; setae of d2 reaching to the level of d3; muscle attachments between d3 sclerotized; all setae surrounded by sclerites; so1 in front of a2, so2 at the same level as d1, so3 near d2, so4 at the same level as d3, so5 behind d4 (fig. 4a). acg fused together and with a suture, apodemes of acg reaching to cx-iii; genital field with a development sclerotization, the anterior part of ap with a projection, ac in an obtuse triangle, ac2 near ac3 rather than ac1, ac3 the biggest; v1 fused to v2, v3 and v4 forming a trapezoid, v4 at the same level as the anterior of acetabular plate; excretory pore surrounded by sclerotized ring (fig. 4b). palp five-segmented; p-2 and p-3 ventral margins slightly straight; p-4 with numerous dorsal hairs, and divided by two ventral hairs in sectors 3:3:2, sword seta at the middle of p-4 (fig. 4e). i-l-5 longish, s-1 and s-2 both with blunt tips and with a narrow setal interspace between them; i-l-6 curved (fig. 5a). measurements male (n = 1) idiosoma l 513, w 369; coxal field l 264, cx-iii w 164, acg il 175, ml 103, w 216; infracapitular bay l 97; genital field l 131, ac1–3 l 26, 28, 27; chelicera l 145; infracapitulum l 125; palp dl: p-1 23, p-2 49, p-3 49, p-4 70, p-5 26; legs segments: i-l-1 dl 38, i-l-2 dl 74, i-l-3 dl 67, i-l-4 dl 107, http://zoobank.org/urn:lsid:zoobank.org:act:d2da763a-88e0-4432-810f-76bc5fbf1f3b european journal of taxonomy 833: 121–142 (2022) 128 fig. 4. atractides (atractides) smiti zhang, li & guo sp. nov., holotype, ♂ (gugc, slide no. qhhy-2020071706). a. idiosoma, dorsal view. b. idiosoma, ventral view. c. gnathosoma. d. chelicera. e. palp. f. ejaculatory complex. scale bars = 100 μm. zhang y.-h. et al., new species of atractides from china 129 i-l-5 dl 122, hb 37, i-l-6 dl 72, hb 19, s-1 l 54, s-2 l 60; dl: ii-l-1 32, ii-l-2 61, ii-l-3 57, ii-l-4 82, ii-l-5 85, ii-l-6 91; dl: iii-l-1 36, iii-l-2 64, iii-l-3 61, iii-l-4 95, iii-l-5 106, iii-l-6 103; dl: iv-l-1 70, iv-l-2 83, iv-l-3 107, iv-l-4 138, iv-l-5 148, iv-l-6 129. female unknown. remarks the new species atractides (atractides) smiti sp. nov. is similar to atractides protendens k.o. viets, 1955 in the following points: (1) p-2 and p-3 ventral margins slightly straight; (2) s-1 and s-2 with a narrow setal interspace between them; (3) the anterior part of ap with a projection; (4) apodemes of acg reaching to cx-iii. however, a. (a.) smiti differs from a. protendens in the following aspects: (1) v1 fused to v2 in a. (a.) smiti, but not fused in a. protendens; (2) excretory pore surrounded by sclerotized ring in a. (a.) smiti, but smooth in a. protendens; (3) the i-l-6 of a. (a.) smiti more curved than that of a. protendens (gerecke 2003). fig. 5. atractides (atractides) smiti zhang, li & guo sp. nov., holotype, ♂ (gugc, slide no. qhhy-2020071706). a–d. i-l–iv-l. scale bar = 100 μm. european journal of taxonomy 833: 121–142 (2022) 130 fig. 6. atractides (atractides) menyuanensis zhang, li & guo sp. nov. a–e. holotype, ♂ (gugc, slide no. qh-hy-2020072901). f. paratype, ♀ (gugc, slide no. qh-hy-2020072902). a. idiosoma, dorsal view. b. idiosoma, ventral view. c. gnathosoma. d. chelicera. e. palp. f. palp. scale bars = 100 μm. zhang y.-h. et al., new species of atractides from china 131 atractides (atractides) menyuanensis zhang, li & guo sp. nov. urn:lsid:zoobank.org:act:c836e7cf-0a50-4334-aa1d-dde500ddbcf8 figs 6–8 diagnosis dorsal muscle attachment unsclerotized. i-l-5 longish, s-1 close to s-2. i-l-6 curved. ac in an obtuse triangle; v1 separated from v2. p-2 ventral margin slightly convex; p-3 ventral margin nearly straight; two long hairs on the ventral surface of p-4 near the base, sword seta near the base. ac in a weakly curved line in female. etymology the new species is named after the name of menyuan hui autonomous county where the specimens were collected. type material holotype china • ♂; qinghai province, haibei tibetan autonomous prefecture, menyuan hui autonomous county; 37°31′31′′ n, 101°21′09′′ e; 2427 m a.s.l.; 29 jul. 2020; hai-tao li leg.; running water; gugc, slide no. qh-hy-2020072901. paratypes china • 5 ♀♀; same collection data as for holotype; gugc, slides no. qh-hy-2020072902 to 2020072906. fig. 7. atractides (atractides) menyuanensis zhang, li & guo sp. nov., paratype, ♀ (gugc, slide no. qh-hy-2020072902). a. idiosoma, dorsal view. b. idiosoma, ventral view. scale bars = 100 μm. http://zoobank.org/urn:lsid:zoobank.org:act:c836e7cf-0a50-4334-aa1d-dde500ddbcf8 european journal of taxonomy 833: 121–142 (2022) 132 description male (n = 1) idiosoma oval, dorsal muscle attachment unsclerotized; all slit organs visible, so1 near the eye capsule, so2 near l2, so3 at the level of d2, so4 at the level of d3, so5 on the outside of d4 (fig. 6a). acg fused together and with a suture, pcg separated; apodemes from acg not reaching to pcg; ac in an obtuse fig. 8. atractides (atractides) menyuanensis zhang, li & guo sp. nov., holotype, ♂ (gugc, slide no. qh-hy-2020072901). a–d. i-l–iv-l. scale bar = 100 μm. zhang y.-h. et al., new species of atractides from china 133 triangle; v1 separated from v2; v3 and v4 forming a trapezoid, v4 at the same level as the anterior of acetabular plate (fig. 6b). palp five-segmented; p-2 ventral margin slightly convex; p-3 ventral margin nearly straight; p-4 with numerous dorsal hairs, two long hairs on the ventral surface of p-4 near the base, sword seta near the base (fig. 6e). i-l-5 longish, s-1 close to s-2; i-l-6 curved (fig. 8a). female (n = 5) similar to male (figs 6f, 7). idiosoma oval; acg and pcg significantly smaller than the male; ac in a weakly curved line (fig. 7b). measurements male (n = 1) idiosoma l 708, w 582; coxal field l 352, cx-iii w 415, acg il 232, ml 92, w 323; infracapitular bay l 146; genital field l 141, ac1–3 l 35, 35, 38; chelicera l 223; infracapitulum l 210; palp dl: p-1 37, p-2 70, p-3 63, p-4 122, p-5 31; legs segments: i-l-1 dl 57, i-l-2 dl 106, i-l-3 dl 101, i-l-4 dl 139, i-l-5 dl 182, hb 50, i-l-6 dl 102, hb 32, s-1 l 58, s-2 l 58; dl: ii-l-1 57, ii-l-2 95, ii-l-3 93, ii-l-4 135, ii-l-5 158, ii-l-6 157; dl: iii-l-1 55, iii-l-2 102, iii-l-3 114, iii-l-4 165, iii-l-5 191, iii-l-6 180; dl: iv-l-1 118, iv-l-2 138, iv-l-3 181, iv-l-4 240, iv-l-5 272, iv-l-6 217. female (n = 5) idiosoma l 1074 (1050–1113), w 935 (868–935); coxal field l 385 (385–432), cx-iii w 575 (550– 585), acg il 248 (248–299), ml 111 (111–130), w 373 (373–417); infracapitular bay l 130 (105– 163); gonopore l 168 (161–190), ap l 157 (145–171), ac1–3 l 38 (34–48), 39 (39–49), 37 (32–48); chelicera l 264 (253–283); infracapitulum l 215 (214–244); palp dl: p-1 46 (40–46), p-2 75 (75–86), p-3 66 (66–80), p-4 126 (121–145), p-5 37 (36–37); legs segments: i-l-1 dl 62 (62–77), i-l-2 dl 106 (106–136), i-l-3 dl 107 (107–126), i-l-4 dl 153 (151–176), i-l-5 dl 200 (200–233), hb 53 (53–59), i-l-6 dl 120 (120–132), hb 31 (31–36), s-1 l 63 (62–68), s-2 l 63 (62–69); dl: ii-l-1 61 (61–72), ii-l-2 93 (93–109), ii-l-3 109 (98–124), ii-l-4 144 (142–163), ii-l-5 168 (168–204), ii-l-6 170 (170– 191); dl: iii-l-1 67 (65–84), iii-l-2 102 (102–116), iii-l-3 119 (115–137), iii-l-4 178 (178–203), iii-l-5 208 (206–233), iii-l-6 200 (190–212); dl: iv-l-1 149 (129–149), iv-l-2 159 (156–179), iv-l-3 202 (191–234), iv-l-4 261 (252–294), iv-l-5 286 (276–321), iv-l-6 232 (223–255). remarks the new species atractides (atractides) menyuanensis sp. nov. is similar to atractides algeriensis lundblad, 1942 in the following points: (1) apodemes from acg not reaching to cx-iv; (2) p-2 ventral margin slightly convex, p-3 ventral margin nearly straight; (3) ac in a weakly curved line in female. however, a. (a.) menyuanensis differs from a. algeriensis in following aspects: (1) two long hairs on the ventral surface of p-4 near the base, sword seta near the base in a. (a.) menyuanensis, but the ventral surface of p-4 divided by two long ventral hairs in 1:1:1, sword seta at the middle of p-4 in a. algeriensis; (2) s-1 much more close to s-2 in a. (a.) menyuanensis than in a. algeriensis; (3) i-l-6 of a. algeriensis much longer and more slender than that of a. (a.) menyuanensis (gerecke 2003). atractides (atractides) longiprojectus zhang, li & guo sp. nov. urn:lsid:zoobank.org:act:a043146a-40fb-45e2-888a-3613e5a14165 figs 9–10 diagnosis dorsal muscle attachment unsclerotized, setae of d1 and d2 significantly longer than others, setae of d1 nearly reaching to d2, setae of d2 extending beyond d3. i-l-5 longish, s-1 and s-2 with a narrow setal interspace between them, s-1 longer than s-2. i-l-6 straight. ac in a weakly curved line; v1 separated http://zoobank.org/urn:lsid:zoobank.org:act:a043146a-40fb-45e2-888a-3613e5a14165 european journal of taxonomy 833: 121–142 (2022) 134 from v2. p-2 with an unobvious ventral projection; p-3 ventral margin slightly convex; the ventral edge of p-4 divided by two long hairs in 2:1:2, sword seta at two thirds of p-4. etymology the latin prefix ‘longi-’ means long, acg of the female of this new species is with a particularly long hind projection. type material holotype china • ♀; qinghai province, haibei tibetan autonomous prefecture, menyuan hui autonomous county, quankou town; 37°31′31′′ n, 101°81′12′′ e; 2691 m a.s.l.; 29 jul. 2020; hai-tao li leg.; running water; gugc, slide no. qh-hy-2020072907. paratypes china • 2 ♀♀; same collection data as for holotype; gugc, slides no. qh-hy-2020072908, 2020072909. description female (n = 3) idiosoma oval, dorsal muscle attachment unsclerotized; o2 and d1 at the same level; setae of d1 and d2 significantly longer than others, setae of d1 nearly reaching to d2, setae of d2 extending beyond d3; all slit organs visible, so1 near the eye capsule, so2 at the same level as d1, so3 near d2, so4 in front of l4, so5 behind of d4 (fig. 9a). coxal group occupying a half of ventral surface, projections from acg extending to cx-iv; ac in a weakly curved line; v1 separated from v2; v3 and v4 forming a rectangle, v4 at the same level as the anterior of acetabular plate (fig. 9b). palp five-segmented; p-2 with an unobvious ventral projection; p-3 ventral margin slightly convex; p-4 with numerous dorsal hairs, the ventral edge of p-4 divided by two long hairs in 2:1:2, sword seta at two thirds of p-4 (fig. 9e). i-l-5 longish, s-1 and s-2 with a narrow setal interspace, s-1 longer than s-2. i-l-6 straight (fig. 10a). measurements female (n = 3) idiosoma l 530 (530–663), w 419 (419–547); coxal field l 320 (320–361), cx-iii w 353 (353–406), acg il 244 (244–295), ml 126 (126–163), w 297 (296–336); infracapitular bay l 143 (133–151); gonopore l 98 (98–118), ap l 120 (118–128), ac1–3 l 35 (35–41), 39 (39–45), 30 (30–38); chelicera l 251 (251–275); infracapitulum l 227 (226–239); palp dl: p-1 35 (32–35), p-2 68 (68–79), p-3 75 (75–82), p-4 106 (106–110), p-5 27 (27–31); legs segments: i-l-1 dl 45 (45–53), i-l-2 dl 103 (100– 113), i-l-3 dl 103 (96–112), i-l-4 dl 153 (152–163), i-l-5 dl 166 (166–180), hb 37 (37–39), i-l-6 dl 134 (134–144), hb 34 (34–37), s-1 l 57 (57–72), s-2 l 47 (46–54); dl: ii-l-1 53 (53–56), ii-l-2 96 (89–104), ii-l-3 96 (92–102), ii-l-4 130 (130–141), ii-l-5 140 (140–153), ii-l-6 141 (140–152); dl: iii-l-1 53 (51–59), iii-l-2 87 (79–106), iii-l-3 101 (101–115), iii-l-4 154 (153–168), iii-l-5 178 (170–189), iii-l-6 163 (160–178); dl: iv-l-1 108 (108–120), iv-l-2 141 (135–141), iv-l-3 182 (178–195), iv-l-4 219 (219–239), iv-l-5 246 (234–255), iv-l-6 204 (197–218). male unknown. remarks the new species atractides (atractides) longiprojectus sp. nov. is similar to atractides inflatipalpis k. viets, 1950 in the following points: (1) v1 separated from v2; (2) s-1 longer than s-2; (3) p-2 zhang y.-h. et al., new species of atractides from china 135 with an unobvious ventral projection. however, a. (a.) longiprojectus differs from a. inflatipalpis in the following aspects: (1) p-3 ventral margin slightly convex in a. (a.) longiprojectus, but straight in a. inflatipalpis; (2) i-l-6 straight in a. (a.) longiprojectus, but curved in a. inflatipalpis; (3) the pregenital sclerite of a. inflatipalpis in female much bigger than a. (a.) longiprojectus (gerecke 2003). fig. 9. atractides (atractides) longiprojectus zhang, li & guo sp. nov., holotype, ♀ (gugc, slide no. qh-hy-2020072907). a. idiosoma, dorsal view. b. idiosoma, ventral view. c. gnathosoma. d. chelicera. e. palp. scale bars = 100 μm. european journal of taxonomy 833: 121–142 (2022) 136 fig. 10. atractides (atractides) longiprojectus zhang, li & guo sp. nov., holotype, ♀ (gugc, slide no. qh-hy-2020072907). a–d. i-l–iv-l. scale bar = 100 μm. zhang y.-h. et al., new species of atractides from china 137 atractides (atractides) xianmiensis zhang, li & guo sp. nov. urn:lsid:zoobank.org:act:fb29b7eb-1a14-450e-80e2-4795c3b299ea figs 11–13 diagnosis male dorsal muscle attachment unsclerotized. i-l-5 longish, s-1 and s-2 both with blunt tips and with a setal interspace between them; i-l-6 straight. ac in an obtuse triangle; v1 separated from v2. p-2 and p-3 with a ventral projection respectively, p-4 with numerous dorsal hairs, ventral hairs long, one at the middle of the surface, and the other one at the terminal of lateral edge, sword seta at the middle of p-4. female ac in a weakly curved line. p-2 with a ventral projection, p-3 ventral margin slightly convex, p-4 divided by two ventral hairs in sectors 1:1:1. etymology the new species is named after the name of the xianmi national nature reserve where the specimens were collected. type material holotype china • ♂; qinghai province, xianmi national nature reserve; 37°10′56′′ n, 102°20′03′′ e; 2949 m a.s.l.; 29 jul. 2020; hai-tao li leg.; running water; gugc, slide no. qh-hy-2020072910. paratypes china • 3 ♂♂; same collection data as for holotype; gugc, slides no. qh-hy-2020072911 to 2020072913 • 2 ♀♀; same collection data as for holotype; gugc, slides no. qh-hy-2020072914, 2020072915. description male (n = 4) idiosoma oval, dorsal muscle attachment unsclerotized; so1 near eye capsule; so2 at the same level as d1; so3 at the same level as d2; so4 in front of l4; so5 behind d4 (fig. 11a). acg fused together and with a suture, apodemes of acg well developed, and reaching to cx-iii; ac in an obtuse triangle, ac3 biggest; v1 separated far from v2, v3 and v4 forming a rectangle, v4 at the same level as the anterior part of acetabular plate (fig. 11b). palp five-segmented; p-2 and p-3 with a ventral projection respectively, p-4 with numerous dorsal hairs, ventral hairs long, one at the middle of the surface, and the other one at the terminal of lateral edge, sword seta at the middle of p-4 (fig. 11e). i-l-5 longish, s-1 and s-2 with blunt tips and with a setal interspace between them; i-l-6 straight (fig. 13a). female (n = 2) similar to male (fig. 12). ac in a weakly curved line (fig. 12b). p-2 with a ventral projection, p-3 ventral margin slightly convex, p-4 divided by two ventral hairs in sectors 1:1:1 (fig. 11f). measurements male (n = 4) idiosoma l 724 (724–896), w 605 (605–734); coxal field l 351 (351–395), cx-iii w 426 (426–483), acg il 251 (251–287), ml 131 (131–159), w 329 (329–372); infracapitular bay l 132 (132–160); genital field l 131 (131–144), ac1–3 l 39 (36–41), 32 (32–43), 36 (36–46); chelicera l 227 (227–267); infracapitulum l 207 (207–239); palp dl: p-1 30 (30–37), p-2 69 (69–80), p-3 76 (76–89), p-4 107 http://zoobank.org/urn:lsid:zoobank.org:act:fb29b7eb-1a14-450e-80e2-4795c3b299ea european journal of taxonomy 833: 121–142 (2022) 138 fig. 11. atractides (atractides) xianmiensis zhang, li & guo sp. nov. a–e. holotype, ♂ (gugc, slide no. qh-hy-2020072910). f. paratype, ♀ (gugc, slide no. qh-hy-2020072914). a. idiosoma, dorsal view. b. idiosoma, ventral view. c. gnathosoma. d. chelicera. e. palp. f. palp. scale bars = 100 μm. zhang y.-h. et al., new species of atractides from china 139 (107–121), p-5 34 (33–34); legs segments: i-l-1 dl 47 (47–56), i-l-2 dl 107 (107–126), i-l-3 dl 110 (110–125), i-l-4 dl 165 (165–191), i-l-5 dl 188 (188–214), hb 42 (42–46), i-l-6 dl 145 (145–160), hb 34 (32–34), s-1 l 67 (67–73), s-2 l 58 (58–65); dl: ii-l-1 55 (47–59), ii-l-2 94 (94–108), ii-l-3 100 (100–113), ii-l-4 141 (141–162), ii-l-5 161 (161–184), ii-l-6 154 (154–174); dl: iii-l-1 54 (51– 63), iii-l-2 93 (93–112), iii-l-3 109 (109–124), iii-l-4 166 (166–191), iii-l-5 194 (194–226), iii-l-6 178 (178–200); dl: iv-l-1 131 (129–137), iv-l-2 154 (154–161), iv-l-3 187 (187–211), iv-l-4 240 (240–274), iv-l-5 260 (260–303), iv-l-6 224 (224–246). female (n = 2) idiosoma l 925 (1215), w 754 (1042); coxal field l 403 (422), cx-iii w 515 (576), acg il 303 (323), ml 133 (131), w 405 (403); infracapitular bay l 180 (179); gonopore l 145 (190), ap l 156 (156), ac1–3 l 49 (48), 43 (45), 43 (46); chelicera l 291 (297); infracapitulum l 269 (268); palp dl: p-1 36 (46), p-2 84 (88), p-3 97 (103), p-4 124 (123), p-5 36 (39); legs segments: i-l-1 dl 59 (64), i-l-2 dl 122 (136), i-l-3 dl 131 (141), i-l-4 dl 193 (205), i-l-5 dl 220 (227), hb 47 (47), i-l-6 dl 165 (173), hb 39 (35), s-1 l 87 (78), s-2 l 74 (72); dl: ii-l-1 64 (50), ii-l-2 108 (106), ii-l-3 121 (133), ii-l-4 166 (180), ii-l-5 188 (200), ii-l-6 175 (186); dl: iii-l-1 54 (68), iii-l-2 114 (112), iii-l-3 131 (142), iii-l-4 198 (212), iii-l-5 233 (241), iii-l-6 208 (215); dl: iv-l-1 150 (160), iv-l-2 162 (174), iv-l-3 211 (228), iv-l-4 270 (293), iv-l-5 246 (314), iv-l-6 211 (265). remarks the new species atractides (atractides) xianmiensis sp. nov. is similar to atractides inflatus walter, 1925 in the following points: (1) p-2 and p-3 of the male both with a ventral projection respectively; (2) v1 not fused to v2; (3) s-1 and s-2 both with blunt tips and with a setal interspace between them; (4) ac in an obtuse triangle in the male and in a weakly curved line in the female. however, a. (a.) xianmiensis fig. 12. atractides (atractides) xianmiensis zhang, li & guo sp. nov., paratype, ♀ (gugc, slide no. qh-hy-2020072914). a. idiosoma, dorsal view. b. idiosoma, ventral view. scale bar = 100 μm. european journal of taxonomy 833: 121–142 (2022) 140 fig. 13. atractides (atractides) xianmiensis zhang, li & guo sp. nov., holotype, ♂ (gugc, slide no. qh-hy-2020072910). a–d. i-l–iv-l. scale bar = 100 μm. zhang y.-h. et al., new species of atractides from china 141 differs from a. inflatus in the following aspects: (1) apodemes of acg in female well developed and reaching to cx-iii in a. (a.) xianmiensis, but not reaching to cx-iii in a. inflatus; (2) i-l-6 straight in a. (a.) xianmiensis, but curved in a. inflatus (gerecke 2003). discussion due to the soft body wall and few specialized structures, there are only a few taxonomic features that can be used for species identification in atractides, which might cause confusion in the taxonomy of these species. so it is of great importance to search for new taxonomic features in atractides. jin (1997) mentioned that the relative position of glandularia is different in various groups of water mites, and ramírez-sánchez et al. (2016) also mentioned that glandularia in arrenuridae thor, 1900 could be potential taxonomic characters. so the relative position of glandularia may be among the new taxonomic features that can be used in the taxonomy of atractides. for example, the coxae group and palps of atractides (atractides) biprojectus sp. nov. and atractides (atractides) xianmiensis sp. nov. are similar in morphology, but the relative position of v3 in relation to v4 and v4 in relation to the genital field in a. (a.) biprojectus is significantly different from that in a. (a.) xianmiensis. therefore, it is suggested to mention the relative position of glandularia on the body wall when describing and drawing species of atractides. acknowledgements special thanks are due to professor zhen-ning chen (qinghai normal university, p.r. china) for providing the opportunity for water mite investigation in qinghai region. this research was supported by guizhou provincial science and technology projects (qiankehe pingtai rencai-gcc [2022]029-1) and commissioned project of the qilian mountain national park administration of qinghai province, p.r. china (qhtx-2021-006). references cook d.r. 1974. water mite genera and subgenera. memoirs of the american entomological institute 21: 1–860. gerecke r. 2003. water mites of the genus atractides koch, 1837 (acari: parasitengona: hygrobatidae) in the western palaearctic region: a revision. zoological journal of the linnean society 138 (2–3): 141–378. https://doi.org/10.1046/j.1096-3642.06-0.00051.x gülle p., gülle i. & boyaci y. ö. 2015. a new atractides koch (acari: hydrachnidia, hygrobatidae) species from southwestern turkey. zootaxa 3957 (2): 246–248. https://doi.org/10.11646/zootaxa.3957.2.11 gülle p., erman o. & boyaci y.ö. 2021. faunal diversity and distribution of hygrobatidae koch, 1842 species (acari: hydrachnidia) of turkey and three neighboring countries. acta aquatica turcica 17 (1): 25–33. https://doi.org/10.22392/actaquatr.747911 jin d.c. 1997. hydrachnellae – morphology systematics. a primary study of chinese fauna. science and technology publishing house, guiyang. [in chinese.] pešić v. & smit h. 2011. a new species of atractides koch, 1837 (acari, hydrachnidia, hygrobatidae) from ethiopia, with a discussion on the biodiversity of the genus atractides in the afrotropical region. zookeys 86: 1–10. https://doi.org/10.3897/zookeys.86.972 pešić v. & smit h. 2018. a checklist of the water mites of central asia with description of six new species (acari, hydrachnidia) from kyrgyzstan. acarologia 58 (1): 165–185. https://doi.org/10.24349/acarologia/20184236 pešić v. & smit h. 2021a. a new species of the genus atractides koch, 1837 from turkey (acari: hydrachnidia: hygrobatidae). ecologica montenegrina 43: 44–50. https://doi.org/10.37828/em.2021.43.6 https://doi.org/10.1046/j.1096-3642.06-0.00051.x https://doi.org/10.11646/zootaxa.3957.2.11 https://doi.org/10.22392/actaquatr.747911 https://dx.doi.org/10.3897%2fzookeys.86.972 https://doi.org/10.24349/acarologia/20184236 https://doi.org/10.37828/em.2021.43.6 european journal of taxonomy 833: 121–142 (2022) 142 pešić v. & smit h. 2021b. water mites of the genus atractides koch, 1837 from kyrgyzstan (acari: hydrachnidia: hygrobatidae) with the description of six new species. acarologia 61 (2): 332–355. https.//doi.org/10.24349/acarologia/20214434 pešić v., zawal a., smit h. & bańkowska a. 2018. new records of water mites from sri lanka (acari: hydrachnidia) with the description of four new species. systematic and applied acarology 23 (1): 178– 195. https://doi.org/10.11158/saa.23.1.14 pešić v., smit h. & saboori a. 2021. new records of the water mite genus atractides koch, 1837 from iran (acari: hydrachnidia: hygrobatidae). ecologica montenegrina 44: 1–10. https://doi.org/10.37828/em.2021.44.1 ramírez-sánchez m.m., de luna e. & cramer c. 2016. geometric and traditional morphometrics for the assessment of character state identity: multivariate statistical analyses of character variation in the genus arrenurus (acari, hydrachnidia, arrenuridae). zoological journal of the linnean society 177 (4): 720–749. https.//doi.org/10.1111/zoj.12384 smit h. 2020. water mites of the world, with keys to the families, subfamilies, genera and subgenera (acari: hydrachnidia). monografieën van de nederlandse entomologische vereniging, leiden. sun h., niu y., chen y.s., song b., liu c.q., peng d.l., chen j.g. & yang y. 2014. survival and reproduction of plant species in the qinghai–tibet plateau. journal of systematics and evolution 52 (3): 378–396. https://doi.org/10.1111/jse.12092 viets k.o. 1987. die milben des süßwassers (hydrachnellae und halacaridae (part.), acari). ii.: katalog. sonderbände des naturwisschaftlichen vereins in hamburg 9: 1–1012. wang z.y. & jin d.c. 2012. two new species of atractides koch, 1837 (acari: hydrachnida: hygrobatidae) from sichuan province, china. entomotaxonomia 34 (3): 567–573. wang z.y. & jin d.c. 2013. a new species of the water mite genus atractides koch, 1837 (acarina, hygrobatidae). journal of guangxi normal university (natural science edition) 31 (2): 93–95. wang z.y., yi t.c. & jin d.c. 2015. descriptions of atractides aprojectus sp. nov. and male of a. huaxiensis yi & jin (acari: hydrachnidia) from china. acta zootaxonomica sinica 40 (1): 53–57. https://doi.org/10.11865/zs.20150104 yi t.c., jin d.c. & wang x.j. 2010. water mites of the subgenus tympanomegapus thor (acari: hygrobatidae: atractides) from china. international journal of acarology 36 (5): 419–429. https://doi.org/10.1080/01647954.2010.483236 manuscript received: 23 february 2022 manuscript accepted: 1 june 2022 published on: 3 august 2022 topic editor: tony robillard section editors: rudy jocqué desk editor: pepe fernández printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; leibniz institute for the analysis of biodiversity change, bonn – hamburg, germany; national museum, prague, czech republic. https://dx.doi.org/10.24349/acarologia/20214434 https://doi.org/10.11158/saa.23.1.14 https://doi.org/10.37828/em.2021.44.1 http://dx.doi.org/10.1111/zoj.12384 https://doi.org/10.1111/jse.12092 https://doi.org/10.11865/zs.20150104 https://doi.org/10.1080/01647954.2010.483236 european journal of taxonomy 223: 1–38 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2016.223 www.europeanjournaloftaxonomy.eu 2016 · sinev a.y. & dumont h.j. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:aea6eaee-69d8-453f-9cf9-ce09d7aa393d 1 revision of the costata-group of alona s. lato (cladocera: anomopoda: chydoridae) confirms its generic status artem y. sinev 1,* & henri j. dumont 2 1 department of invertebrate zoology, biological faculty, lomonosov moscow state university, leninskie gory, moscow 119991, russia and a.n. severtsov institute of ecology and evolution, leninsky prospect 33, moscow 119071, russia. 2 department of ecology, jinan university, guangzhou 510632, china. 2 department of biology, ghent university, 9000 ghent, belgium. * corresponding author: artemsinev@gmail.com 2 email: henri.dumont@ugent.be 1 urn:lsid:zoobank.org:author:fbb6aaef-4f83-4f43-b6c3-e50c150ea9fe 2 urn:lsid:zoobank.org:author:647cc682-c915-4855-b447-74575898b4cb abstract. by taking flavalona gen. nov. out of alona s.l. (cladocera: anomopoda: chydoridae), the last major clade has now been removed from this polyphyletic assemblage. flavalona gen. nov. is a monophylum defined by having three, rarely two connected head pores and slit-shaped, rarely rounded lateral head pores. postabdomen rather long, distally narrowed, with robust marginal denticles and weakly developed lateral fascicles of setules. end-claw weakly curved and with short basal spine. male postabdomen with gonopores opening at the end of a penis-like outgrowth. trunk limbs: exopodite of p2 with seta; inner portion of p4 with flaming-torch shaped setae; p5 with filter plate of three setae; p6 a large simple lobe. the relationship of the new genus with other aloninae remains to be determined. a key to the 11 species of the genus is provided and a discussion of their geographic distribution and habitat type is given. keywords. cladocera, alona, taxonomy, morphology, revision. sinev a.y. & dumont h.j. 2016. revision of the costata-group of alona s. lato (cladocera: anomopoda: chydoridae) confirms its generic status. european journal of taxonomy 223: 1–38. http://dx.doi.org/10.5852/ ejt.2016.223 introduction species in the polyphyletic genus alona baird, 1843, historically the result of merging about a dozen of species groups, are similar in gross morphology by convergence (van damme & dumont 2008a, 2008b; van damme et al. 2010). recent revisions of these groups have led to their assignment to a long series of genera (dumont & silva-briano 2000; sinev 2004; sinev & shiel 2008; van damme & dumont 2008a, 2009; van damme et al. 2009; van damme et al. 2011; van damme & sinev 2011; van damme & maiphae 2013). some groups were transferred to new, others to previously described but reinstated http://dx.doi.org/10.5852/ejt.2016.223 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:aea6eaee-69d8-453f-9cf9-ce09d7aa393d mailto:artemsinev%40gmail.com?subject= mailto:henri.dumont%40ugent.be?subject= http://zoobank.org/urn:lsid:zoobank.org:author:fbb6aaef-4f83-4f43-b6c3-e50c150ea9fe http://zoobank.org/urn:lsid:zoobank.org:author:647cc682-c915-4855-b447-74575898b4cb http://dx.doi.org/10.5852/ejt.2016.223 http://dx.doi.org/10.5852/ejt.2016.223 european journal of taxonomy 223: 1–38 (2016) 2 and/or redefined genera (sinev et al. 2005; sinev & kotov 2012), while van damme & dumont (2008b) restricted true alona to the quadrangularis-group, now consisting of five species (sinev 2012). with the assignment of the pulchella-group to ovalona van damme & dumont, 2008 (sinev 2015b), today only few clades of alona s.l. remain unrevised. the largest of these clades is the costata-group. it was outlined by sinev (1999a, 2001, 2008, 2009) and is defined by the unique morphology of its lateral head pores (sinev 1999a). they are slit-shaped, with pocket-like cavities below, and the length of the lateral pores and size of their pocket vary between species. other distinctive features include a distally narrowing postabdomen with well-developed composite marginal denticles, armed with small spinules, and weakly developed lateral setule fascicles, idl with three setae, the endite of limb four with a row of robust marginal setae, shaped as flamingtorches, a large limb vi shaped as a rounded lobe, and а penis-like process on the male postabdomen. according to van damme & dumont (2008a), the costata-clade is part of the hexalona-branch of alona s.l., a supra-generic category characterized by having six thoracic limbs and a full set of setae on each limb. all species of hexalona have inner setae on endites 2–3 of limb i, seven setae on exopodite iii, and filter plate v with three setae. a generic status for the costata-group was therefore suggested but not formally instated by van damme & dumont (2008a) and by sinev (2008). in recent years, the group has been intensively studied (sinev 1999a, 2001, 2008, 2009; kotov & sinev 2004; van damme et al. 2011; van damme & eggermont 2011; sousa et al. 2015), and several new species have been described. currently, it includes 11 species. here, we formally raise it to the rank of a genus, and discuss the taxonomy and geographic ranges of all currently recognized species. material and methods animals and exuviae were selected from samples under a stereoscopic microscope, placed on slides (in a drop of a glycerol-formaldehyde mixture), and studied under a microscope in toto for identification and measurements. one to five specimens from each population were dissected for analysis of appendages. measurements were conducted using an eyepiece-micrometer; all drawings were made with a camera lucida. we here use the system of enumeration for different setae on thoracic limbs proposed by kotov et al. (2010) as for all anomopoda. abbreviations i–v = thoracic limbs i–v as = accessory seta of limb i cbs = copulatory brush seta of limb i ep = epipodite ex = exopodite gfp = gnathobase filter plates of limbs ii–v idl = inner distal lobe of limb i ip = interpore distance (distance between interior and posterior major head pores) ms = male seta of limb i mxp = maxillar process odl = outer distal lobe of limb i pep = preepipodite pp = postpore distance (distance between posterior head pore and posterior corner of head shield) s = sensillum sinev a.y. & dumont h.j., revision of the costata-group of alona (cladocera) 3 results order anomopoda sars, 1865 family chydoridae dybowski & grochowski, 1894 emend. frey 1967 subfamily aloninae dybowski & grochowski, 1894 emend. frey 1967 tribe alonini dybowski & grochowski, 1894 emend. kotov 2000 flavalona gen. nov. urn:lsid:zoobank.org:act:8cc51b78-c807-4a31-8808-f167b8e5095a type species alona costata sars, 1862. diagnosis female aloninae of moderate size, maximum length of adult female 0.5–0.7 mm. body oval, of moderate height, compressed laterally. maximum height usually at mid-body. valves and head shield without keel, valves from previous molts sometimes retained. dorsal, posterior and ventral margins of valves evenly curved, postero-dorsal, postero-ventral angles and anterior-ventral angles of valves broadly rounded. posteroventral angle of valves without denticles. valves with moderately to well-developed linear sculpture or tubercles. head small, triangular to round in lateral view, rostrum short, pointing downward. eye larger than ocellus. head shield broad, with broadly rounded rostrum. posterior margin of head shield usually broadly rounded. three, in one species two, main head pores, narrowly connected. lateral head pores transverse slits (rounded in f. sphagnophila (van damme & eggermont, 2011)), with flattened pocketlike cavities below. labrum with moderately broad keel, keel apex rounded, posterior margin of keel with two small clusters of setae. thorax 1.5 times longer than abdomen. abdomen without abdominal joint, middle abdominal segment not saddle-shaped. postabdomen moderately long, narrowing distally in postanal part. ventral margin straight. basis of claws bordered from distal margin by clear incision. distal margin straight or convex, distal angle usually acute, with rounded tip. dorsal margin concave in anal portion, straight in postanal portion. preanal angle well-defined, postanal angle weakly-defined. postanal margin with well-developed composite marginal denticles with spines at anterior margin. lateral fascicles of setules narrow, weakly developed, distalmost setules in fascicles not thicker than others, shorter than marginal denticles. postabdominal claw of moderate length, weakly curved. basal spine short. antennule not protruding beyond the tip of rostrum, with 3–4 transverse rows of short setules on anterior surface. antennular seta thin. nine terminal aesthetascs of variable length. antenna with antennal formula setae 0–0–3/1–1–3, spines 1–0–1/0–0–1. basal segment robust, branches elongated, all segments narrow, cylindrical, basal segment of each branch 1.5 times longer than two others. seta of basal segment of endopodite not reaching end of endopodite. seta of middle segment of endopodite and apical setae of same morphology, similar length and thickness. antennal spines well developed. six pairs of thoracic limbs. http://zoobank.org/urn:lsid:zoobank.org:act:8cc51b78-c807-4a31-8808-f167b8e5095a european journal of taxonomy 223: 1–38 (2016) 4 limb i. with accessory seta. idl with 3 setae, seta 1 short, setae 2 and 3 of similar size, armed with thin setules in distal portion. endite 3 with four setae. endite 2 with three outer setae and short inner seta. endite 1 with two 2-segmented setae, shortened, stub-like flat seta, and inner seta longer than inner seta of endite 2. ventral surface of limb with rows of long setules. two ejector hooks, of unequal size. maxillar process with single seta. limb ii. exopodite narrow, elongated, with seta. inner portion of limb with eight scraping spines, spines either of similar morphology, evenly decreasing in size basally, or spines 3 and 6 more robust, armed with thicker denticles. distal armature of gnathobase with four elements. filter plate ii with seven setae, posteriormost considerably shorter than others. limb iii. exopodite rounded, with seven setae, seta 3 being longest, setae 6–7 of moderate length, setae 1–2 and 4–5 very short. setae 1–5 plumose, setae 6–7 armed with short setules or naked. distal endite with three setae, two distalmost ones slender, sharp, with denticles; basalmost seta shorter, geniculated, with thin setules. basal endite with four stiff setae. four soft setae increasing in size basally. distal armature of gnathobase with four elements: elongated, cylindrical sensillum; thin, geniculated seta; and two short pointed spines. filter plate iii with seven setae. limb iv. exopodite rounded, with six setae, setae 1–4 plumose, setae 5–6 with short setules or naked. inner portion of limb iv with four setae and small sensillum. distalmost seta robust, sharp, first flamingtorch seta (2) wide, armed with thick setules, two other 2–3 times thinner, armed with thin setiules. three short soft setae increasing in size basally. gnathobase with one long 2-segmented seta and a blunt process near it. filter plate iv with five setae. limb v. exopodite v separated into two lobes, with 4 plumose setae, seta 1 longest, evenly decreasing in width basally. inner limb portion as elongated lobe. at inner face, two setae with wide bases, one 1.5–2 times longer than other. filter comb v with three setae. limb vi. large, of same size or larger than exopodite v, an oval lobe with setulated margin. male body lower than in female, similar to that of juveniles, with weakly convex dorsal margin. postabdomen moderately long, narrowing distally, with obtuse preanal angle, gonopores located at of the end of the genital process protruding above the postabdominal claws. clusters of setules in place of female marginal denticles, lateral fascicles of setules as in female. postabdominal claw much shorter than in female, weakly curved, with short basal spine or without spine. antennule shorter and wider than in female, with twelve terminal aesthetascs, male seta long. limb i with u-shaped copulatory hook. idl seta 1 absent, male seta large, of similar size with setae 2 and 3. etymology the name is formed by adding the prefix flavo-, from the latin adjective flavus, “yellow”, to the preexisting name alona, to express the fact that many of its species often are yellow-golden in color. differential diagnosis stands out from other aloninae by the morphology of the lateral head pores. also differs from all other alona s.l. by a male postabdomen with gonopores at the end of a penis-like process (similar structures only in leydigia kurz, 1875 and armatalona sinev, 2004). flavalona gen. nov. belongs to the hexalonabranch of alona s.l. and differs from alona s. str. (see van damme & dumont 2008b), ovalona (see sinev 2015b) and genera of the coronatella-branch (see van damme & dumont 2008a) by the presence of six trunk limbs and a filter plate v of three setae. within hexalona, flavalona gen. nov. also differs sinev a.y. & dumont h.j., revision of the costata-group of alona (cladocera) 5 from the affinis-group (see alonso 1996; sinev 2009) by its smaller size, shape of postabdomen, weakly developed lateral setules on the postabdomen, and weakly developed idl seta 1; from the intermediagroup (see alonso 1996) by the shape of the postabdomen, strong marginal denticles of the postabdomen, and by weakly developed lateral setules of the postabdomen; from the guttata-group (see alonso 1996; sinev & silva-briano 2012) by a more elongated postabdomen with weakly developed preanal angle. flavalona gen. nov. clearly differs from matralona van damme & dumont, 2009, another genus related to the hexalona-branch, in the shape and armament of the postabdomen, in a present idl seta 1, and in an exopodite iii with seven setae. key to the known species of flavalona gen. nov. 1. distal angle of postabdomen obtuse, with clearly defined tip ……………………………………… ……………………………………………f. weltneri (keilhack, 1905) comb. nov.; west palearctic – distal angle of postabdomen acute or with broadly rounded apex …………………………………2 2. lateral head pores long, length over 0.5 ip. distal angle of postabdomen acute ………………3 – lateral head pores short, length about less than 0.5 ip. distal angle of postabdomen rounded…7 3. two major head pores ……………………………………………………………………………… ……………………f. setigera (brehm, 1931) comb. nov.; australia and new zealand, neotropic – three major head pores ……………………………………………………………………………4 4. lateral head pore pockets deep, rounded, depth ca 1.5–2 lateral pore length …………………5 – lateral head pore pockets shallow, semicircular, depth ca 1/2–1/3 pore length …………………6 5. limb v strongly bilobed ………………………f. costata (sars, 1962) s.l. comb. nov.; holarctic – limb v only slightly bilobed ………………………………………………………f. margipluma (sousa, santos, güntzel, diniz, de melo junior & elmoor-loureiro, 2015) comb. nov.; neotropics 6. valve sculpture composed of thin longitudinal striae or hexagons ………………………………… ……………………………………………………f. cheni (sinev, 1999) comb. nov.; indo-malaya – valve sculpture composed of dense, broad longitudinal ridges …………………………………… …………………………………f. natalensis (sinev, 2008) comb. nov.; mountains of south africa 7. anterior margin of labrum with one or two denticles ……………………………………………… ……………………………………………………f. bicolor (frey, 1965) comb. nov.; east nearctic – anterior margin of labrum without denticles ………………………………………………………8 8. length of adult female up to 0.7 mm. seta 6 of exopodite iii about 2/3 lengths of seta 3 ……… ……………f. hudeci (sinev, 1999) comb. nov.; neotropics (syn. alona rusticoides hudec, 1998) – length of adult female not exceeding 0.52 mm. seta 6 of exopodite iii almost as long as seta 3 …9 9. lateral head pores dot-like with rounded pockets, widths of pockets over two lengths of pore …… …………f. sphagnophila (van damme & eggermont, 2011) comb. nov.; mountains of east africa – lateral head pores transverse with semicircular pockets …………………………………………10 10. central major head pore located closer to posterior one. exopodite v strongly bilobed. epipodites iv–v with finger-like processes longer than epipodite itself ……………………… ………………………………………f. iheringula (kotov & sinev, 2004) comb. nov.; neotropics – central major head pore located halfway between anterior and posterior pores. exopodite v not divided into two lobes. epipodites iv–v without processes ……………………………………… ……………………………………………………f. rustica (scott, 1901) s.l. comb. nov.; holarctic european journal of taxonomy 223: 1–38 (2016) 6 costata-clade characteristics of the costata-clade clade include: (1) valves with pronounced linear sculpture; (2) shortened anterior group of setae on valves, with longest setules in the middle of the group; (3) long lateral head pores with large pockets, length of pore over 0.5 ip. (4) postabdomen with acute or right distal angle; (5) lateral fascicles of setules on postabdomen with distalmost setae the longest; (6) male postabdomen with short genital process, its length about 1/3 or less of length of claw. the clade is rather uniform, with no marked differences in outer morphology. flavalona natalensis comb. nov. differs by the sculpture of its valves, and f. weltneri comb. nov. by the shape of its postabdomen. armament of postabdomen and morphology of head appendages similar within clade. differences in thoracic limb morphology minimal, the only variable feature being the shape of exopodite v, which can be bilobed or oval. species differ mostly by the morphology of the major and lateral head pores. male morphology known only in f. costata comb. nov. and f. setigera comb. nov., but no significant difference between them. flavalona costata (sars, 1862) comb. nov. figs 1–2 alona costata sars, 1862: 286. alona lineata schoedler, 1862: taf. i, fig. 23. phryxira rectirostris müller, 1867: 184–185, tab. iv, fig. 15. lynceus costatus norman & brady, 1867: 28, pl. xviii, fig. 2, pl. xxi, fig. 7. lynceus costata lilljeborg, 1901: 465–468, tab. lxviii, fig. 9–15. alona costata – hellich 1877: 90, figs 47–48. — behning 1941: 299–301, fig. 123. — šrámek-hušek et al. 1962: 336–338, fig. 124. — herbst 1962: 86, fig. 61. — frey 1965): 159–162, figs 1–6. — flössner & frey 1970: 327, figs 4, 6–7. — smirnov 1971: 371–373, figs 345–340. — flössner 1972: 299–301, figs 140–141. — negrea 1983: 291–292, fig. 118. — margaritora 1985: 313–315, fig. 124. — sars 1993: 135, pl. 98. — alonso 1996: 322–323, fig. 143. — flössner 2000: 315–322, fig. 117. — sinev 1999a: 132–138, figs 1–5. — sinev 2002: 394, figs 3в,м, 4в, 5в, з. — hudeč 2010: 319–322, fig. 77. — sinev et al. 2015: fig. 1a–c. alona lineata – müller 1867: 178, table iv, figs 2–3. non alona costata – idris 1983: 106–108, fig. 50. differential diagnosis flavalona costata comb. nov. differs from f. setigera comb. nov. by having three main head pores, and from f. natalensis comb. nov. and f. cheni comb. nov. by the deep pockets of the lateral head pores; in these species the depth of the pockets is less than half the depth of the pore. it also differs from f. natalensis comb. nov. by the costata-like sculpture of its valves. flavalona costata comb. nov. is similar in morphology to f. margipluma comb. nov. too but differs by the basal spine of its postabdomen exceeding the width of the claw base and by having a bilobed exopodite v. as flavalona costata comb. nov. is the type species of flavalona gen. nov., a detailed description follows. type material material in sars’ collection (zoological museum of oslo university, f12410) labeled “norway”; no exact locality or dates given. sinev a.y. & dumont h.j., revision of the costata-group of alona (cladocera) 7 fig. 1. flavalona costata (sars, 1862) comb. nov., randsfjorden lake, norway. a–k. parthenogenetic ♀. a. lateral view. b. ventral margin of valves. c. anterior setae of valves. d. posterior setae of valves. e. posteroventral corner and posterior margin of valves. f. head shield. g–h. head pores. i. postabdomen. j. antennule. k. antenna. l. ephippial ♀. m. ephippium. n–p. ♂. n. lateral view. o. postabdomen. p. antennule and outline of rostrum. european journal of taxonomy 223: 1–38 (2016) 8 fig. 2. flavalona costata (sars, 1862) comb. nov., randsfjorden lake, norway. a–l. labrum and thoraciс limbs of parthenogenetic ♀. a. labrum. b. limb i. c. odl of limb i. d. ejector hooks of limb i. e. limb ii. f. exopodite of limb iii. g–h. inner portion of limb iii. i. exopodite of limb iv. j. inner portion of limb iv. k. limb v. l. limb vi. m–n. thoracic limb i of adult ♂. sinev a.y. & dumont h.j., revision of the costata-group of alona (cladocera) 9 material studied earlier see sinev (1999a) for list of material from norway, poland, germany, russia and mongolia. material studied here norway: 11 parthenogenetic ♀♀, over 30 ephippial ♀♀, 13 ♂♂ from littoral zone of randsfjorden lake, 60º14.37ʹ n, 10º23.94ʹ e, 9 oct. 2015, coll. a.y. sinev & t. jensen. china: over 30 parthenogenetic ♀♀ from littoral zone of erhai lake, yunnan province, 25º49.887ʹ n, 100º13.166ʹ e, 19 jan. 2015, coll. a.y. sinev & y. gu; 3 parthenogenetic ♀♀ and exuviae from artificial lake in botanical garden of changsha city, hunan province, 28º06.253ʹ n, 113º01.783ʹ e, 10 apr. 2014, coll. a.y. sinev & y. gu; 8 parthenogenetic ♀♀ from artificial lake in botanical garden of guangzhou city, guangdong province, 23º11.225ʹ n, 113º21.834ʹ e, 5 apr. 2014, coll. a.y. sinev & y. gu; over 30 parthenogenetic ♀♀ from shuisheng reservoir, hainan island, 18°53.097ʹ n, 110°23.514ʹ e, 24 apr. 2014, coll. a.y. sinev & y. gu. type locality lake ostensjovand, oslo, norway. description parthenogenetic female length of adult up to 0.5 mm. in lateral view body oval or slightly ovoid (fig. 1a), maximum height at middle or somewhat behind it, height-length ratio ca 0.58–0.65. dorsal margin uniformly curved. postero-dorsal and postero-ventral angles broadly rounded. posterior margin uniformly curved. ventral margin almost straight, with about 50–60 setae (fig. 1b), about 10 first setae of moderate length, next 15 setae short, other setae of moderate length. setae of the anterior group (fig. 1c) not longer than those of the posterior group (fig. 1d), longest setae of anterior group located at the middle of the group. groups of small setules located between the bases of all ventral setae. posteroventral angle (fig. 1e) with only 10–15 short setules protruding beyond margin of valves. a row of about 150 setules along posterior margin on inner side of carapace. in ventral third of the row these setules uniform, in middle third, row formed by long setules and groups of shorter setules between them, in dorsal third by long setules only. carapace covered by moderately thick, sparsely spaced longitudinal lines, without intervening striae. head relatively small, triangular-round in lateral view, rostrum short, pointing downward. eye larger than ocellus. distance from tip of rostrum to ocellus greater than that between ocellus and eye. head shield with maximum width behind mandibular articulation, without any sculpture (fig. 1f). rostrum short, broadly rounded. posterior margin of head shield broadly rounded. three major head pores (fig. 1g–h), middle pore slightly smaller than others, located closer to posterior pore. connection between anterior and middle pores narrow, connection between middle and anterior pores varies from narrow to rather wide in some populations (see alonso 1996; sinev et al. 2015). pp a. 0.5–0.8 ip. transverse lateral head pores ca 0.5–0.8 ip lengths, located about 0.5 ip distance from midline, at a level between anterior and middle main head pores. lateral pore pockets large, rounded, with depth ca 1.5–2 pore lengths. labrum moderate size (fig. 2a). distal labral plate without setulation. labral keel of moderate width, with a blunt or rounded apex. anterior margin of keel convex, posterior margin with two clusters of setules. thorax and abdomen subequal in length, dorsal surface of abdominal segments not saddle-shaped. second abdominal segment with two transverse rows of setules. european journal of taxonomy 223: 1–38 (2016) 10 postabdomen (fig. 1i) wide, narrowing distally, with well defined, acute angle between distal and dorsal margins, length about 2.2–2.5 times height. ventral margin straight. inflated basis of claws bordered from distal margin by clear incision. distal margin weakly convex to straight. dorsal margin with distal part about 3 times longer than preanal one, with postanal portion about 1.5 times longer than anal one. postanal portion of distal margin straight, anal portion weakly concave. preanal and postanal angles weakly defined. preanal margin almost straight. postabdomen with 8–9–10 well-developed composite marginal denticles, each with several spinules on anterior margin, and with two groups of marginal spinules and 2–3 groups of short setules on anal margin. length of denticles decrease basally, length of longest denticles exceed the width of postabdominal claw base. ten–twelve lateral fascicles of thin setules; in postanal fascicles distalmost seta being longest, 1.5 times shorter than marginal denticles. postabdominal claw of moderate length, somewhat longer than preanal portion of postabdomen. basal spine ca 0.2 of length of claw. antennule (fig. 1j) moderate length, almost reaching the tip of rostrum, with four transverse rows ofshort setules at anterior face. antennular sensory seta slender, 1.5 times shorter than antennule, arising at ⅔ distance from base. nine aesthetascs, two of them longer than others, of about 2/3 length of antennule, others of similar length. . antenna (fig. 1k) moderate length. antennal formula, setae 0–0–3/1–1–3, spines 1–0–1/0–0–1. basal segment robust, branches elongated, slender, all segments cylindrical, with short setules around distal margin. seta arising from basal segment of endopodite short and thin, not reaching tip of distal segment. seta arising from middle segment of endopodite of similar size with apical setae. spine on basal segment of exopodite shorter then middle segment. apical spines slightly longer than respective apical segments. thoracic limbs six pairs. limb i (fig. 2b–d). epipodite oval, with finger–like process 2 times longer than epipodite itself. accessory seta about 1/3 length of odl seta. idl with three setae and several clusters of small setules on ventral face, idl seta 1 small, thin, setae 2 and 3 subequal in length to odl seta, both with thin setules in distal part. endite 3 with inner seta (1) shorter than outer setae (a–c). endite 2 with three outer setae (d–f) and short inner seta (2) and sensillum near its base, setae e–f short, about half length of limb itself, seta f slightly longer than seta e. endite 1 with inner seta (3) longer than inner seta of endite 2 and sensillum near its base, two 2-segmented setae (g–h), both setulated in distal part, and a flat seta (i) of characteristic shape, with reduced distal portion, armed with long setules. seven rows of thin long setules on ventral surface of limb. two ejector hooks, one of them 1.5 times longer than the other. maxillar process elongated, with single seta. limb ii (fig. 2e). exopodite oval, with seta almost as as long as exopodite. eight scraping spines, scrapers 1–5 long, scrapers 6–8 much shorter, armed with small setules. scrapers 3 and 6 of similar length with neighbors, but slightly thicker and armed with slightly more thick setules. distal armature of gnathobase with four elements. filter plate with seven setae, the posteriormost seta considerably shorter and thicker than other, armed with very long setules, next seta of same morphology with others, but two times shorter. limb iii (fig. 2f–h). еpipodite oval, without finger-like process. exopodite trapezium-shaped, with seven setae. seta 3 being longest, setae 6 and 7 of ⅔ and ½ length of seta 3, respectively, other setae short. setae 6 and 7 naked, all other plumose, setules on seta 3 considerably longer than on short setae. distal endite with three setae, two distalmost members slender, sharp, of similar length, with distal parts unilaterally armed small sharp denticles; basalmost seta thicker, bilaterally armed with long setules. basal endite with four plumose outer setae (a–d), increasing in size in basal direction. four inner setae (4–7) increasing in size basally, a small sensillum near the base of distalmost seta (4). distal armature sinev a.y. & dumont h.j., revision of the costata-group of alona (cladocera) 11 of gnathobase with four elements. the first one elongated, cylindrical sensillum, second thin, bent seta, others two blunt spines with fused bases. filter plate iii with seven setae. limb iv (fig. 2i–j). pre-epipodite setulated; epipodite with a finger-like process longer than epipodite itself. exopodite subquadrangular, with six setae. seta 3 the longest , seta 1–2 slightly shorter, setae 4–6 ca 2/3 length of seta 3, setae 5–6 slightly longer than seta 4. setae 1–4 flat, plumose, setae 5–6 slender, seta 5 with short setules at the middle, seta 6 naked. inner-distal portion of limb iv with four setae and small rounded sensillum. seta 1 slender, sharp, armed with short spinules. distal flaming-torch seta (2) of typical for aloninae shape, broad, with curved distal portion, armed with about 10 long thick setules. two other flaming-torch setae (3–4) thin and straight, with straight distal portion, armed with thin, hair-like setules, a small sensillum located near their bases. three inner setae (a–c) long, increasing in size basally. gnathobase with a short 2-segmented seta, only little longer than seta 1, and a hillock with sensilla distally. filter plate with five setae. limb v (fig. 2k). pre-epipodite setulated, epipodite with finger-like process longer than epipodite itself. exopodite bilobed, with 4 plumose setae. seta 1 being longest, setae 2–3 slightly shorter than seta 1, seta 4 two times shorter than seta 1. inner limb portion an elongated oval lobe, with setulated inner margin. at inner face, two setae, one three times longer than other, and a triangular sensillum. filter plate with three setae. limb vi (fig. 2l). an elongated lobe, its length about twice maximum width, its margin densely setulated in distal part. ephippial female outline similar to that of parthenogenetic female (fig. 1l), dorsal margin more highly arched in most specimens. ephippium of living specimens blackish, in preserved specimens yellow-brown. ephippium (fig. 1m) sculptured by anastomosing longitudinal lines, thicker than in ventral portion of valves. adult male length 0.36–0.42 mm. body (fig. 1n) low oval, dorsal margin weakly convex. maximum height at the middle of the body, height/length ratio ca 0.55. ocellus larger that in female. rostrum short, slightly truncated (fig. 1p). postabdomen (fig. 1o) moderately long, narrowing distally in postanal part. ventral margin straight or wavy. basis of claws bordered from distal margin by clear incision. genital process short, about ¼– ⅓ length of postabdominal claw. distal margin very short, distal angle obtuse. dorsal margin concave in anal portion, straight in postanal portion. preanal and postanal angles not defined. clusters of short setules in place of marginal denticles, distalmost cluster consisting of 3–5 thick short setules. lateral fascicles as in female. a cluster of short setules above the base of postabdominal claw. postabdominal claw as long as preanal portion of postabdomen, weakly curved. basal spine short and thin, visible only under oil immersion. antennule (fig. 1p) shorter and broader than in female, with 12 terminal aesthetascs, longest about ¾ length of antennule, shortest ca ½ length of antennule. male seta arising at ¼ length from tip, ca ⅓ length of antennule. thoracic limb i (fig. 2m–n) with u-shaped copulatory hook, half as long as limb itself. idl with three setae, idl seta 1 absent. idl setae 2 and 3 much thinner and two times shorter than in female, of similar length. male seta curved, slightly shorter than seta 2. copulatory brush seta ca 2/3 length of idl seta 2. ventral face of the limb under the copulatory brush with row of 20 long stiff stiff setules. inner seta (1) of endite 3 longer and thinner than in female, with long setules distally. european journal of taxonomy 223: 1–38 (2016) 12 for description of juvenile males and females, see sinev (1999a). our data agree with those of previous descriptions of the species, but add several features. thin setules between ventral setae of the valves were overlooked by sinev (1999), so sousa et al. (2015) used this feature as diagnostic for f. margipluma comb. nov. most authors also overlooked the basal spine of the male postabdomen. distribution presumed holarctic (flössner & frey 1970; smirnov 1971). flavalona costata comb. nov. is found in the whole of europe, european and asian russia, kazakhstan, mongolia, china, korea and japan but its southern limit is unclear (smirnov 1971; sinev 1999a). the southernmost certified record is from hainan island (sinev et al. 2015). it has been cited from mainland south-east asia (see korovchinsky 2013) and india (see chatterjee et al. 2013), but these records were not accompanied by descriptions, and the related f. cheni comb. nov. is present in these regions. flössner & frey (1970) found no significant differences between north american and european f. costata comb. nov., but the appendages were not studied, so possibly nearctic populations constitute a separate species. flavalona cheni (sinev, 1999) comb. nov. fig. 3a alona cheni sinev, 1999a: 142–146, figs 8–10. alona cheni – maiphae et al. 2008: 6, fig. 3. alona costata – idris 1983: 106–108, fig. 50. alona setigera – chen & peng 1993: 19, figs 1–5. diagnosis female length of adult up to 0.47 mm. body regular oval, maximum height at the middle of body, height/ length ratio ca 0.6. valves from previous molts not retained. posteroventral angle without setules protruding beyond valve margins. ventral margin with ca 45 setae. valves covered by thin, sparsely spaced longitudinal lines. eye larger than ocellus. rostrum short, broadly rounded. posterior margin of head shield broadly rounded. three major head pores (fig. 3a), middle pore slightly smaller than others, located near posterior pore. connection between middle and posterior pore wide, connection between middle and anterior pore narrow. pp ca 0.7–0.9 ip. lateral head pores of ca 0.9–1.2 ip length, lateral pore pockets semicircular, with depth ca 0.5 of pore length. labral keel without denticles. postabdomen truncated, length 2.2–2.4 heights. distal margin straight, distal angle prominent, acute with rounded tip. dorsal margin weakly convex in preanal part, concave in anal part, straight in postanal part. distal part of dorsal margin 3 times longer than preanal one, with preanal portion 1.3–1.5 times longer than anal one. preanal and postanal angles weakly defined. postanal margin of postabdomen provided with 10–14 marginal denticles. anal margin with 4–5 clusters of marginal setules. 10–12 lateral fascicles of setules, distal setules in fascicle being longest. postabdominal claw of moderate length, 1.5 times longer than preanal portion of postabdomen. basal spine ca 0.15 of length of claw. antennule with longest aesthetasc of ca 2/3 the ength of antennule. scrapers of limb ii of similar morphology. exopodite iii with seta 6 of ca 2/3 the length of seta 3. exopodite iv with seta 3 being sinev a.y. & dumont h.j., revision of the costata-group of alona (cladocera) 13 longest, setae 1–2 slightly shorter, setae 4–6 ca 0.4 length of seta 3. epipodites iv without process. exopodite v strongly bilobed. epipodite v with processes as long as epipodite itself. male unknown. differential diagnosis flavalona cheni comb. nov. differs from f. setigera comb. nov. in having three major head pores; from f. costata comb. nov. and f. margipluma comb. nov. by the morphology of lateral head pores, with semicircular pore pockets about half the depth of pore length; from f. natalensis comb. nov. in the typical costata-like sculpture of the valves; from f. weltneri comb. nov. in the shape of its postabdomen. type material holotype parthenogenetic ♀, zoological museum of moscow state university, ml-05. paratypes two dissected parthenogenetic ♀♀ on slides, zoological museum of moscow state university, ml-06, ml-07. fig.3. a. flavalona cheni (sinev, 1999) comb. nov., head pores of parthenogenetic ♀. b–c. flavalona natalensis (sinev, 2008) comb. nov., parthenogenetic ♀. b. lateral view. c. head pores. d. flavalona setigera (brehm, 1931) comb. nov., head pores of parthenogenetic ♀. e–f. flavalona weltneri (keilhack, 1905) comb. nov., parthenogenetic ♀. e. head pores. f. postabdomen. figures a and d from sinev (1999a), b–c from sinev (2008), e redrawn from flössner (2000) and f redrawn from van damme et al. (2010). european journal of taxonomy 223: 1–38 (2016) 14 material studied earlier see sinev (1999a) for list of material from india and maiphae et al. (2008) for a list of material from thailand. type locality india: aiwa reservoir, ahmedabad. distribution india, south-east asia and china (sinev 1999a; maiphae et al. 2008; idris 1983), northernmost record in china from wuhan province (chen & peng 1993). flavalona margipluma (sousa, santos, güntzel, diniz, de melo junior & elmoor-loureiro, 2015) comb. nov. alona margipluma sousa, santos, güntzel, diniz, de melo junior & elmoor-loureiro, 2015: 445–455, figs 1–4. alona iheringula – silva, abreu, orlando, wisniewski & santos-wisniewski 2014: 2–5, fig. 1. diagnosis (according to sousa et al. 2015) female length of adult 0.32–0.43 mm. body regular oval, maximum height at mid-body, height/length ratio about 0.65. valves from previous molts not retained. posteroventral angle without setules protruding beyond valve margins. ventral margin with 45–50 setae. valves covered by thin, sparsely spaced longitudinal lines. eye larger than ocellus. rostrum short, broadly rounded. posterior margin of head shield broadly rounded. three major head pores, middle pore slightly smaller than others, located near posterior pore. connection between middle and posterior pore wide, connection between middle and anterior pore narrow. pp ca 0.7–0.95 ip. lateral head pores of ca 0.7–0.85 ip length, lateral pore pockets rounded, with depth about 1.5 of pore length. labral keel without denticles. postabdomen truncated, length 2.4–2.6 heights. distal margin straight, distal angle prominent, acute with rounded tip. dorsal margin weakly convex in preanal part, concave in anal part, straight in postanal part. distal part of dorsal margin 3 times longer than preanal one, with preanal portion 1.3–1.5 times longer than anal one. preanal and postanal angles weakly defined. postanal margin of postabdomen provided with 11–13 marginal denticles. anal margin with 4–5 clusters of marginal setules. about 10 lateral fascicles of setules, distal setules in fascicle being longest. postabdominal claw of moderate length, longer than preanal portion of postabdomen. basal spine small, its length less than width of postabdominal claw base. antennule with longest aesthetasc of about 2/3 length of antennule. scrapers of limb ii of similar morphology. exopodite iii with seta 6 of ca 2/3 length of seta 3. exopodite iv with seta 3 being longest, setae 1–2 slightly shorter, setae 4–6 ca 0.5 length of seta 3. epipodite iv and v with processes as long as the epipodite itself. exopodite v not bilobed. male unknown. sinev a.y. & dumont h.j., revision of the costata-group of alona (cladocera) 15 differential diagnosis flavalona margipluma comb. nov. differs from f. setigera comb. nov. by three main head pores, from f. natalensis comb. nov. and f. cheni comb. nov. by deep pockets of lateral head pores (in these species the depth of the pockets is less than half the depth of the pore). it also differs from f. natalensis comb. nov. by its costata-like valve sculpture. flavalona margipluma comb. nov. is similar in morphology to f. costata comb. nov., but differs in exopodite v not separated into two lobes and in basal spine of postabdomen shorter than the width of claw base. unique to the species is the bottle-shaped main sensillum of limb iv. sousa et al. (2015) suggested that the presence of thin setules between the bases of setae on the ventral valve margin is unique to f. margipluma comb. nov., but such setules are present in all flavalona gen. nov., yet were overlooked in previous studies. type material holotype adult parthenogenetic ♀, the museum of zoology of the university of são paulo, mzusp 33196. paratypes see sousa et al. (2015) for the list of paratypes. type locality brazil: criminosa lake, porto murtinho, pantanal, mato grosso do sul, 21º40ʹ28.8″ s, 57º53ʹ28.5″ w. distribution so far known from mato grosso do sul state, pernambuco state and distrito federal of brazil (sousa et al. 2015). in littoral zone of lakes among macrophytes. flavalona natalensis (sinev, 2008) comb. nov. fig. 3b–c alona natalensis sinev, 2008: 24–32, fig. 1–4. diagnosis female length of adult 0.46–0.54 mm. body (fig. 3b) regular oval, maximum height at the middle of body, height/length ratio about 0.6. valves from previous molts not retained. posteroventral angle without setules protruding beyond valve margins. ventral margin with ca 50 setae. valves covered by dense, broad longitudinal lines. eye larger than ocellus. rostrum short, broadly rounded. posterior margin of head shield broadly rounded. three major head pores (fig. 3c), middle pore slightly smaller than others, located near the posterior pore. connection between middle and posterior pore wide, connection between middle and anterior pore narrow. pp ca 1.1–1.5 ip. lateral head pores of ca 1.0–1.2 ip length, lateral pore pockets shallow, with depth about 1/3 of pore length. labral keel without denticles. postabdomen truncated, length 2.2–2.4 heights. distal margin straight, distal angle prominent, acute, with rounded tip. dorsal margin weakly convex in preanal part, concave in anal part, straight in postanal part. distal part of dorsal margin 3 times longer than preanal one, with preanal portion 1.3–1.5 times longer than anal one. preanal and postanal angles weakly defined. postanal margin of postabdomen provided with 8–10 marginal denticles. anal margin with 4–5 clusters of marginal setules. 10–12 lateral european journal of taxonomy 223: 1–38 (2016) 16 fascicles of setules, distal setules in fascicle being longest. postabdominal claw of moderate length, 1.5 times longer than preanal portion of postabdomen. basal spine about 0.2 of length of claw. antennule with longest aesthetasc of about 2/3 length of antennule. scrapers of limb ii of similar morphology. exopodite iii with seta 6 of about 2/3 length of seta 3. exopodite iv with seta 3 being longest, setae 1–2 slightly shorter, setae 4 about half length of seta 3, setae 5–6 about 2/3 length of seta 3. epipodites iv–v with processes about 1.5 of epipodite length. exopodite v weakly bilobed. male unknown. differential diagnosis flavalona natalensis comb. nov. has a characteristic sculpture of the valves, composed of thick, closely spaced longitudinal lines. type material holotype parthenogenetic ♀, zoological museum of moscow state university, ml-44. paratypes seven parthenogenetic ♀♀, zoological museum of moscow state university, ml-45. material studied earlier see sinev (2008) for a list of material from the republic of south africa and lesotho. type locality south africa: lakes cave tarn 3, cobham, kwazulu-natal province, 29°39ʹ50″ s, 29°18ʹ47″ e, altitude 2190 m.a.s.l. distribution inhabits small, possibly temporary, water bodies (pools and tarns) in the east escarpment portion of the drakensberg mountains in kwazulu–natal and lesotho (sinev 2008). at the time of sampling, ph was 6 to 7.6, and temperature 16.4 to 22ºc. it belongs to the complex of endemic species of the drakensberg mountains, which includes five other benthic cladocerans viz. pleuroxus carolinae (methuen, 1910), dumontiellus africanus smirnov, 2007, alona meridionalis sinev, 2006, a. martensi sinev, 2009 (chydoridae) and ilyocryptus martensi kotov & štifter, 2005 (ilyocryptidae) (see van damme et al. 2013). all are restricted to the eastern escarpment of the drakensberg, which lies within an area with an effective temperature index below 14°c (stuckenberg 1969). the index of temperature expresses the relative duration of the warm period of year. flavalona setigera (brehm, 1931) comb. nov. fig. 3d alona guttata var. setigera brehm, 1931: 499–501, figs 5–7. biapertura setigera smirnov & timms, 1983: 58–59, figs 67–68. biapertura rusticoides smirnov & timms, 1983: 64, fig. 75. alona setigera sinev, 1999a: 138–142, figs 5–8. non alona setigera – chen & peng 1993: 19, figs 1–5. sinev a.y. & dumont h.j., revision of the costata-group of alona (cladocera) 17 diagnosis female length 0.39–0.49 mm. body regular oval, maximum height at mid-body, height/length ratio ca 0.6. valves from previous molts not retained. posteroventral angle without setules protruding beyong valve margins. ventral margin with about 40 setae. valves covered by thin, sparsely spaced longitudinal lines. eye larger than ocellus. rostrum short, broadly rounded. posterior margin of head shield broadly rounded. two major head pores (fig. 3d) with narrow connection between them. pp ca 1.1–1.3 ip. lateral head pores of ca 0.9–1.0 ip length, lateral pore pockets semicircular, with depth about 1 pore length. labral keel without denticles. postabdomen truncated, length about 2.5 heights. distal margin straight, distal angle prominent, acute with rounded tip. dorsal margin weakly convex in preanal part, concave in anal part, straight in postanal part. distal part of dorsal margin 3 times longer than preanal one, with anal and postanal portion of similar length. preanal and postanal angles weakly defined. postanal margin of postabdomen provided with 10–14 marginal denticles. anal margin with 4–5 clusters of marginal setules. 10–12 lateral fascicles of setules, distal setules in fascicle being longest. postabdominal claw of moderate length, 1.5 times longer than preanal portion of postabdomen. basal spine about 0.15 of length of claw. antennule with longest aesthetasc about 2/3 of length of antennule. scrapers of limb ii of similar morphology. exopodite iii with seta 6 of ca 2/3 length of seta 3. exopodite iv with seta 3 being longest, setae 1–2 slightly shorter, setae 4–6 ca 0.4 length of seta 3. epipodite iv–v without process. exopodite v bilobed. epipodite v with processes as long as epipodite itself. male length of adult 0.38–0.4 mm. body low oval, dorsal margin weakly convex. maximum height at the middle of the body, height/length ratio 0.55–0.58. ocellus larger that in female. postabdomen moderately long, narrowing distally in postanal part. ventral margin almost straight. genital process about 1/3 length of postabdominal claw. distal margin short, distal angle obtuse. dorsal margin concave in anal portion, straight in postanal portion. preanal and postanal angles not defined. clusters of short setules in place of marginal denticles, distalmost cluster consisting of 3–5 thick short setules. postabdominal claw as long as preanal portion of postabdomen, weakly curved. basal spine not found. antennule with male seta arising at 1/4 length from tip, about 1/3 length of antennule. thoracic limb i with idl setae 2 and 3 much thinner and two times shorter than in female, of similar length. male seta curved, slightly shorter than seta 2. ventral face of the limb under the copulatory brush with row of 20 long stiff setules. inner seta (1) of endite 3 longer and thinner than in female, with long setules distally. differential diagnosis flavalona setigera comb. nov. differs from all species of flavalona gen. nov. by the presence of only two major head pores. type material non-existent. european journal of taxonomy 223: 1–38 (2016) 18 material studied earlier see sinev (1999) for a list of material from australia. type locality new zealand. distribution australia, new zealand (sinev 1999a) and brazil (santos-wisniewsky et al. 2001). currently the only species with a transpacific range. however, since the limbs of american populations were not studied, these may turn out to represent a separate species. flavalona weltneri (keilhack, 1905) comb. nov. fig. 3e–f alona weltneri keilhack, 1905: 158–159, figs 13–14. alona weltneri – keilhack 1908: 464, fig. 20. — flössner 1972: 306–308, fig. 144. — smirnov 1971: 365–366, figs 425–426. — flössner 2000: 322–323, fig. 119. — van damme, elais-gutierrez & dumont 2011: 51–53, figs 3–4. diagnosis (according to flössner 2000 and van damme et al. 2011). female length of adult up to 0.45–0.53 mm. body regular oval, maximum height after the middle of body, height/length ratio about 0.57 (the holotype, studied by van damme et al. (2011) was deformed, and its reconstruction (van damme et al. 2011: fig. 3a) is uncertain). valves from previous molts not retained. posteroventral angle without setules protruding beyond valve margins. ventral margin with ca 45 setae. valves covered by thin, sparsely spaced longitudinal lines. eye larger than ocellus. rostrum short, broadly rounded. posterior margin of head shield broadly rounded. three major head pores (fig. 3e), middle pore slightly smaller than others, located near posterior pore. connections between pores narrow. pp ca 0.4 ip. lateral head pores of ca 0.5 ip length, lateral pore pockets not studied in details, but probably shallow. labral keel without denticles. postabdomen (fig. 4f) truncated, length 2.3 heights. unlike all other species, no incursion between postabdominal claw base and distal margin. distal margin straight, distal angle obtuse, not prominent, with rounded tip. dorsal margin straight in preanal part, concave in anal part, straight in postanal part. distal part of dorsal margin 3 times longer than preanal one, with preanal portion only slightly times longer than anal one. preanal and postanal angles weakly defined. postanal margin of postabdomen provided with 6–8 marginal denticles. anal margin with 2 clusters spinules in distal portion. 10–12 lateral fascicles of setules, distal setules in fascicle being longest. postabdominal claw of moderate length, 1.5 times longer than preanal portion of postabdomen. basal spine ca 0.25 of length of claw. antennule with longest aesthetasc of ca ½ length of antennule. exopodite iii with seta 6 of about ⅔ length of seta 3. exopodite iv with seta 3 being longest, setae 1–2 slightly shorter, setae 4–6 ca 2/3 length of seta 3. male unknown. sinev a.y. & dumont h.j., revision of the costata-group of alona (cladocera) 19 differential diagnosis flavalona weltneri comb. nov. differs from all members of the costata-clade by its postabdomen with obtuse distal angle and no incursion between base of claws and distal margin, and by a longer basal spine of postabdominal claw, about 0.25 of claw length, 1.5 times the width of the claw base. type material holotype adult parthenogenetic ♀, in museum für naturkunde, berlin, no. 18948. type locality poland: madü-see (=jezioro miedwie), pyrzyce (pyritz), west pomerania. distribution switzerland, germany, poland, norway, sweden and finland (van damme et al. 2011). rustica-clade characteristic features of the rustica-clade clade include: (1) valves with weakly developed linear or tubercular sculpture; (2) anterior group of setae on valves with longest setules in posterior part of the group; (3) short lateral head pores with small pockets, length of pore less than 0.5 ip. (4) postabdomen with prominent rounded distal angle; (5) lateral fascicles of setules on postabdomen with middle setae being longest; (6) male postabdomen with long genital process, its length over 1/2 length of claw. species of rustica-group have a thicker cuticle than the costata-clade, and more thick, spine-like short setules on antenna and limb i. species of the rustica-clade frequently retain valves from the 1–2 previous molts, a situation never recorded in the costata-clade. the rustica-clade is less uniform than the costata-clade. species of the group differ in size, morphology of posteroventral angle of valves, length of rostrum, morphology and position of major head pores, labrum, armament of postabdomen, morphology of scrapers on limb ii, proportions of setae of exopodite iii and shape of exopodite v. male morphology is known for four taxa, viz. f. rustica rustica comb. nov., f. rustica americana comb. nov., f. iheringula comb. nov. and f. bicolor comb. nov., differing in shape of postabdomen, morphology of postabdominal claw and length of its distal setules. flavalona rustica (scott, 1895) comb. nov. flavalona rustica rustica (scott, 1895) comb. nov. figs 4–5 alona rustica scott, 1895: 189–190, pl. 5, figs 5, 17. alona estonica maëmets, 1958: 260–264, figs 1–3, photos 1–2. alona rustica – frey 1965: 162–169, figs 1–7. — flössner 1967: 443–444, fig. 2. — smirnov 1971: 360–363, figs 416–418. — flössner 1972: 301–304, fig. 142. — van de velde & dumont 1978: 306, fig. 1a–c. — dumont 1983: 102–105, pl. 4. — negrea 1983: 228–190, fig. 117. — margaritora 1985: 316–318, fig. 125. — alonso 1996: 323–325, fig. 144. — flössner 2000: 318, fig. 118. — margaritora & vagaggini 2002: 61–59, figs 1–10. — sinev 1999a: 146, fig. 10. — sinev 2001: 118, figs 26–34. — sinev 2002: 396, figs 3д,о, 4д, 5и. — hudeč 2010: 322–324, fig. 78. alona estonica – maëmets 1960: 35–40, fig. 1, photos 1–2. european journal of taxonomy 223: 1–38 (2016) 20 fig. 4. flavalona rustica (scott, 1895) comb. nov. from bol’shoe torfyanoe lake, bolshoi solovetskiy island (white sea), russia. a–b. juvenile ♀♀ of instars i and ii. c–n. parthenogenetic ♀. c. specimen with oblique valves. d. specimen with tuberculated valves. e. ventral margin of valves. f. anterior setae of valves. g. posterior setae of valves. h. posteroventral corner and posterior margin of valves. i. head shield. j–k. head pores. l–m. postabdomen. n. antenna. sinev a.y. & dumont h.j., revision of the costata-group of alona (cladocera) 21 fig. 5. flavalona rustica (scott, 1895) comb. nov. a–n. labrum, antennule and thoraciс limbs of parthenogenetic ♀ from bol’shoe torfyanoe lake, bolshoi solovetskiy island (white sea), russia. a–b. labrum. c. antennule. d. limb i. e. idl and odl of limb i. f. scrapers of limb ii. g. gnathobase of limb ii. h. exopodite of limb iii. i. inner portion of limb iii. j. exopodite of limb iv. k. inner portion of limb iv. l. limb v. m. gnathobase of limb v. n. limb vi. o–p. adult ♂. o. lateral view. p. postabdomen. figure o redrawn from alonso (1996) and p from sinev (1999a). european journal of taxonomy 223: 1–38 (2016) 22 differential diagnosis flavalona rustica rustica comb. nov. differs from f. iheringula comb. nov. by its middle major head pore located halfway between the posterior and anterior pores and by the oval, an unbilobed exopodite of limb v, from f. sphagnophila comb. nov. by its transverse, not dot-like lateral head pores and by an oval, not bilobed exopodite of limb v, from f. bicolor comb. nov. by a smaller size, short rostrum and labral keel without denticles, from f. hudeci comb. nov. by a smaller size, presence of numerous setules on posteroventral corner of valves, and by seta 6 of exopodite iii almost as long as seta 3. male of f. rustica comb. nov. differs from f. iheringula comb. nov. and f. bicolor comb. nov. by marginal setules of postabdomen in distal groups of moderate length, little more than width of postabdominal claw base (in f. iheringula comb. nov. they are short, less than width of postabdominal claw base, and in f. bicolor comb. nov. they are long, about two postabdominal claw bases). we provide a full description of f. rustica comb. nov. as typical of the rustica-clade. type material non-existent. material studied here four parthenogenetic ♀♀ from lake motjønn, west coast of nisser lake, norway, 59º08.62′ n, 8º27.32′ e, 4 oct. 2015, coll. a.y. sinev & b. walseng; 47 parthenogenetic ♀♀ from bol’shoe torfyanoe lake, bolshoi solovetskiy island (white sea), russia, 24 jul. 1997, coll. a. evseev; 12 parthenogenetic ♀♀ from karelia republic, russia, exact location unknown, from collection of n.n. smirnov. type locality scotland. description parthenogenetic female length of adult 0.35–0.52 mm. in lateral view body (fig. 4a–d) regular oval or slightly ovoid , maximum height at mid-body or somewhat behind it, height-length ratio ca 0.6–0.65. dorsal margin uniformly curved. postero-dorsal and postero-ventral angles broadly rounded. posterior margin uniformly curved. ventral margin (fig. 4e) almost straight, with about 50–60 setae, about 12 first setae long, next 15 setae short, other setae of moderate length. setae of anterior group (fig. 4f) longer than of posterior group (fig. 4g), posteriormost seta of group the longest. groups of small setules located between bases of all ventral setae. posteroventral angle (fig. 4h) with ca 50 long setules protruding beyond margin of valves. a row of about 150 setules along posterior margin on inner side of carapace. sculpture of carapace either thin, weakly developed longitudinal lines without striae, or rows of tubercles (fig. 4d). head relatively small, triangular-round in lateral view, rostrum short, pointing downward. eye larger than ocellus. distance from tip of rostrum to ocellus greater than between ocellus and eye. head shield (fig. 4i) with maximum width behind mandibular articulation, without any sculpture in specimens with linear sculpture of valves, or covered by tubercles. rostrum more elongated than in costata-clade, broadly rounded. posterior margin of head shield broadly rounded, wavy in tuberculated specimens. three major head pores (fig. 4j–k) with thin connection between them, middle pore slightly smaller than others, located at the middle pp about 1–1.5 ip. transverse lateral head pores of about 0.3–0.4 ip length, located about 0.5 ip distance from midline, at a level between anterior and middle main head pores. lateral pore pockets semicircular, with depth ca 0.5–0.9 pore lengths. sinev a.y. & dumont h.j., revision of the costata-group of alona (cladocera) 23 labrum of moderate size (fig. 5a–b). distal labral plate without setulation. labral keel of moderate width, with a blunt or rounded apex. anterior margin of keel convex, posterior margin with two clusters of setules. clusters of lateral setules observed in anterior half of the keel in some specimens. thorax and abdomen subequal in length, dorsal surface of abdominal segments not saddle-shaped. second abdominal segment with two transverse rows of short setules. postabdomen (fig. 4l–m) wide, narrowing distally, with a prominent broadly rounded distal angle, length ca 2.0–2.3 height. ventral margin straight. inflated basis of claws bordered from distal margin by clear incision. distal margin strongly convex. dorsal margin with distal part about 2.5 times longer than preanal part, with postanal portion about 1.3–1.5 times longer than anal one. postanal and anal portions of distal margin weakly concave. preanal angle well-defined, postanal angle weakly defined. preanal margin almost straight. postanal portion of postabdomen with 7–8 well developed composite marginal denticles, each with several spinules on anterior margin, anal portion with 2–3 marginal denticles, similar to postanal ones, and with 2–3 groups of thick setules. length of denticles decrease basally, length of longest denticles about 1.5 widths of postabdominal claw base. ten–twelve lateral fascicles of thin setules; in postanal fascicles middle seta being longest, 1.5 times shorter than marginal denticles. postabdominal claw of moderate length, somewhat longer than preanal portion of postabdomen. basal spines short, their length less than width of postabdominal claw base. antennule (fig. 5c) of moderate length, not reaching the tip of rostrum, with four transverse rows of short thick setules at anterior face. antennular sensory seta slender, 2 times shorter than antennule, arising at ⅔ distance from the base. nine aesthetascs, two of which longer than others, of about 3/4 length of antennule, others of similar length. antenna (fig. 4n) of moderate length. antennal formula, setae 0–0–3/1–1–3, spines 1–0–1/0–0–1. basal segment robust, branches elongated, slender, all segments cylindrical, with short setules around distal margin and groups of thick lateral setules. seta arising from basal segment of endopodite short and thin, not reaching tip of distal segment. seta arising from middle segment of endopodite of similar size with apical setae. spine on basal segment of exopodite shorter then middle segment. apical spines slightly longer than respective apical segments. thoracic limbs six pairs. limb i (fig. 5d–e). epipodite oval, with finger-like process as long as epipodite itself. accessory seta about half the length of odl. idl with three setae and clusters of setules on ventral surface, idl seta 1 small, thin, setae 2 and 3 subequal in length to odl seta, both with thin setules in distal part. endite 3 with inner seta (1) shorter than outer (a–c). endite 2 with three outer setae (d–f) and short inner seta (2) and sensillum near its base, setae e–f short, about half length of limb, seta f slightly longer than seta e. endite 1 with inner seta (3) longer than inner seta of endite 2 and sensillum near its base, two 2-segmented setae (g–h), both setulated in distal part, and a flat seta (i) of characteristic shape, with shortened distal portion, armed with long setules. seven rows of thin long setules on ventral face of limb. two ejector hooks, one of them 1.5 times longer than the other. maxillar process elongated, with single seta. limb ii (fig. 4f–g). exopodite oval, with seta almost as as long as exopodite. eight scraping spines, scrapers 1–5 long, scrapers 6–8 much shorter, armed with small setules. scrapers 3 and 6 of similar length as its neighbours, but thicker and armed with thicker setules. distal armature of gnathobase with four elements. filter plate with seven setae, posteriormost seta considerably shorter and thicker than other, armed with long setules, next seta of same morphology with others, but two times shorter. european journal of taxonomy 223: 1–38 (2016) 24 limb iii (fig. 4h–i). epipodite oval, without process. exopodite (fig. 3i) trapezium-shaped, with seven setae. seta 3 being longest, setae 6 almost as long as seta 3, seta 7 ca 3/4 the length of seta 3, other setae short. setae 6 and 7 unilaterally armed with short setules in distal portion, all other setae flat, plumose, setules on seta 3 considerably longer than on short setae. distal endite with three setae, two distalmost members (1–2) slender, sharp, of similar length, with distal parts unilaterally armed thick sharp denticles; basalmost seta (3) thicker, bilaterally armed with long setules. basal endite and gnathobase as with four plumose outer setae (a–d), increasing in size in basal direction. four inner setae (4–7) increasing in size basally, a small sensillum near the base of distalmost seta (4). distal armature of gnathobase with four elements. the first one elongated, cylindrical sensillum, second thin, bent seta, others two blunt spines with fused bases. filter plate iii with seven setae. limb iv (fig. 5j–k). pre-epipodite setulated; epipodite without process. exopodite quadrangular, with six setae. seta 3 being longest , seta 1–2 slightly about 3/4 length of seta 3, setae 4 ca ⅓ the length of seta 3, setae 5–6 cahalf length of seta 3. setae 1–4 flat, plumose, setae 5–6 slender, unilaterally armed with short setules in distal portion. inner-distal portion of limb iv with four setae and small rounded sensillum. seta 1 slender, sharp, armed with long spinules. distal flaming-torch seta (2) of typical for aloninae shape, broad, with curved distal portion, armed with about 6 long thick setules. two other flaming-torch setae (3–4) thin and straight, with straight distal portion, armed with very thin, hairlike setules, a small sensillum located near their bases. three inner setae (a–c) long, increasing in size basally. gnathobase with a short 2-segmented seta, only little longer than seta 1, and hillock with sensilla distally. filter plate with five setae. limb v (fig. 5l–m). pre-epipodite setulated, epipodite without process. exopodite oval, with 4 plumose setae. seta 1 being longest, setae 2–3 slightly shorter than seta 1, seta 4 two times shorter than seta 1. inner limb portion an elongated oval lobe, with setulated inner margin. at inner face, two setae, one three times longer than other, and a triangular sensillum. filter plate with three setae. limb vi (fig. 5n) an oval lobe, its length about 1.3 times its maximum width, densely setulated along all of its margin. description of ephippial females and males taken from the literature (alonso 1996; hudeč 2010). ephippial female outline similar to that of parthenogenetic female, dorsal margin more highly arched in most specimens. ephippium in preserved specimens yellow-brown. ephippium without sculpture. adult male length up to 0.39 mm. body low oval (fig. 5o), dorsal margin weakly convex. maximum height at midbody, height/length ratio about 0.55. ocellus larger than in female. rostrum short, slightly truncated. postabdomen (fig. 5p) moderately long, narrowing distally in postanal part. ventral margin straight or wavy. basis of claws bordered from distal margin by clear incision. genital process long, about ⅔–¾ length of postabdominal claw. distal margin concave, distal angle brioadly rounded, weakly prominent. dorsal margin weakly concave in anal and in postanal portions. preanal angle not defined, postanal angle weakly defined. clusters of setules in place of marginal denticles, distalmost clusters consisting of numerous long setules, their length exceed width of postabdominal claw. lateral fascicles of setules as in female. postabdominal claw 2 times shorter than preanal portion of postabdomen, weakly curved. basal spine short and thick. sinev a.y. & dumont h.j., revision of the costata-group of alona (cladocera) 25 antennule shorter and broader than in female, with 12 terminal aesthetascs, longest almost as long as antennule, shortest about ½ length of antennule. male seta arising at ¼ length from tip, about ⅓ length of antennule. thoracic limb i (fig. 3) with u-shaped copulatory hook, half as long as limb itself. idl with three setae, idl seta 1 absent. idl setae 2 and 3 much thinner and two times shorter than in female, of similar length. male seta curved, slightly shorter than seta 2. copulatory brush seta about half length of idl seta 2. ventral surface of limb under copulatory brush with row of 20 stiff setules. distribution azores (van de velde & dumont 1978), europe and siberia (smirnov 1971). bryophilic, typical of acid bogs with sphagnum l., 1853, not found in neutral and alkaline waters. in central and southern europe confined to mountainous regions. the only flavalona gen. nov. species found in subterraneous waters (dumont 1983; sabater 1987). flavalona rustica americana (flössner & frey, 1970) comb. nov. alona rustica americana flössner & frey, 1970: 326–328, figs 1–2, 5. differential diagnosis flavalona rustica americana comb. nov. differs from the nominal subspecies in the morphology of the male postabdomen, which lacks the basal spine on the postabdominal claw and has wider clusters of distal setules. type material holotype ♀ in the natural history museum, london, united kingdom, no. 1969.5.27.1. allotype ♂ in the natural history museum, london, united kingdom, same number. type locality united states of america: channel between martin and oliver lakes, la grange co., indiana. distribution canada and u.s.a. flössner & frey (1970) considered all north american populations of flavalona rustica comb. nov. belonging to f. rustica americana comb. nov.. flavalona bicolor (frey, 1965) comb. nov. fig. 6a–f alona bicolor frey, 1965: 169–170, fig. 23–29. alona bicolor – frey 1969: 380–386, fig. 1–6. — smirnov 1971: 363–365, fig. 421 — sinev 2009: 60–65, fig. 1–3. diagnosis female length of adult 0.50–0.62 mm. body (fig. 6a) regular oval, maximum height at the middle of body, height/length ratio about 0.65–0.72. valves from previous molts often retained. posteroventral angle european journal of taxonomy 223: 1–38 (2016) 26 fig. 6. flavalona bicolor (frey, 1965) comb. nov. a–e. parthenogenetic ♀. a. lateral view (specimen retaining valves from previous moult). b. head shield. c. head pores. d. labrum. e. postabdomen. f. postabdomen of adult ♂. — g–j. flavalona hudeci (sinev, 1999) comb. nov., parthenogenetic ♀ g. lateral view. h. head pores. i. posteroventral corner of valves. j. thoracic limb v. — k–o. flavalona iheringula (sinev & kotov, 2004) comb. nov. k–m. parthenogenetic ♀. k. lateral view. l. head pores. m. thoracic limb v. n–o. adult ♂. n. lateral view. o. postabdomen. — p. flavalona sphagnophila (van damme & eggermont, 2011) comb. nov., parthenogenetic ♀, head pores. figures a–f from sinev (2009), g–j redrawn from hudeč (1998), k–m from sinev (2001), n–o from kotov & sinev (2004) and p redrawn from van damme & eggermont (2011). sinev a.y. & dumont h.j., revision of the costata-group of alona (cladocera) 27 without setules protruding beyond valve margins. ventral margin with ca 60 setae. valves without sculpture. eye smaller than ocellus. rostrum elongated, about 1.5–2 length of antennule, with rounded apex. posterior margin of head shield (fig. 6b) triangular with rounded apex. three major head pores (fig. 6c) of similar size, middle pore located closer to posterior pore. connection between pores narrow. pp about 2 ip. lateral head pores of about 0.4 ip length, lateral pore pockets shallow, with depth about ¼ of pore length. labral keel (fig. 6d) with 1–2 denticles on anterior margin. postabdomen (fig. 6e) truncated, length less than 2.5 height. distal margin convex, distal angle prominent, rounded. dorsal margin weakly convex in preanal part, concave in anal part, and almost straight in postanal part. distal part of dorsal margin 3 times longer than preanal one, anal and preanal portions of similar length. preanal and postanal angles weakly defined. postanal margin of postabdomen provided with 6–7 marginal denticles. anal margin with 3–4 more slender denticles and 3–4 clusters of marginal setules. 10–14 lateral fascicles of setules, central setules in fascicle being longest. postabdominal claw 1.5–1.7 times longer than preanal portion of postabdomen. basal spine short, robust, about 0.15 length of claw itself. antennule with longest aesthetasc as long as antennule. limb ii with scrapers 3 and 7 thicker and armed with more robust denticles than other. exopodite iii with seta 6 of about ⅔ length of seta 3. exopodite iv with seta 3 being longest, setae 1–2 slightly shorter, setae 4–6 about half length of seta 3, seta 4 longer than setae 5–6. epipodites iv–v without processes. exopodite v oval. male length 0.51–0.53 mm. body low oval, height/length ratio ca 0.55–0.59. ocellus larger than eye. postabdomen (fig. 6f) evenly truncated. preanal and postanal angles not defined. distal part of postabdomen 3.5 times longer than preanal. distal process long, ca ⅔–¾ length of claw. distal clusters of marginal setules long, about two widths of postabdominal claw base. postabdominal claw short, basal spine short, about 0.1 length of claw. differential diagnosis flavalona bicolor comb. nov. differs from all other species of the rustica-clade by an elongated rostrum, an ocellus larger than the eye, labral keel with blunt denticles and large ip/pp ratio (about 2). males differ by their large size and long distalmost marginal setules at the end of the postabdomen. type material holotype parthenogenetic female, british museum (nat. hist.), london, 1963.6.24.1. material studied earlier see sinev (2009) for list of material from the u.s.a. type locality united states of america: goodwill pond, barnstable county, massachusetts. distribution and ecology flavalona bicolor comb. nov. inhabits east canada and north-east u.s.a. (sinev 2009). according to chengalath (1982, 1987), it is not found west of the ontariomanitoba border. it is found only in areas with rocky or sandy bottom, and is more rare than f. cf. rustica and f. cf. costata. european journal of taxonomy 223: 1–38 (2016) 28 flavalona hudeci (sinev, 1999) comb. nov. fig. 6g–j alona hudeci sinev, 1999: 147. alona rusticoides – hudeč, 1998: 205–207, figs 1–8, 23–31. diagnosis (from hudeč 1998) female length of adult up to 0.7 mm. body regular oval (fig. 6g), maximum height at mid-body, height/ length ratio about 0.62.valves from previous molts not retained. posteroventral angle without setules protruding beyond valve margins (fig. 6i). valves oblique or with weak polygonal reticulation. eye and ocellus of similar size. rostrum short, broadly rounded. posterior margin of head shield broadly rounded. three major head pores (fig. 6h) of same size, middle pore located slightly closer to posterior pore. connection between pores narrow. pp about 0.8 ip. lateral head pores of about 0.25 ip length, lateral pore pockets shallow, with depth about ⅓ of pore length. labral keel without denticles on anterior margin. postabdomen truncated, length about 2.3 heights. distal margin convex, distal angle prominent, broadly rounded. dorsal margin weakly convex in preanal part, concave in anal part, straight in postanal part. distal part of dorsal margin 3 times longer than preanal one. preanal angle well-defined, postanal angle not defined. margin of postabdomen provided with 9–11 marginal denticles and 3–4 clusters of marginal setules. about 10 lateral fascicles of setules, middle setules in fascicle being longest. postabdominal claw 1.3 times longer than preanal portion of postabdomen. basal spine about 0.15 of length of claw. antennule with longest aesthetasc of about ⅔ length of antennule. scrapers of limb ii of similar morphology. exopodite iii (fig. 6j) with seta 6 of about ⅔ length of seta 3. exopodite iv with seta 1 being longest, setae 2–2 slightly shorter, setae 4–5 about half length of seta 1, setae 6 longer than setae 4–5, about 4/5 length of seta 1. epipodites iv–v with processes about 1.5 of epipodite length. exopodite v bilobed. male unknown. differential diagnosis flavalona hudeci comb. nov. differs from f. bicolor comb. nov. by a short rostrum, ocellus smaller than eye, and labral keel without denticles; from all other species of the rustica-branch by its large size, absence of setules on posteroventral corner of valves, and by seta 6 of exopodite iii only ca ⅔ as long as seta 3. type material holotype parthenogenetic ♀, british museum (natural history), no. 1998.784. paratypes parthenogenetic 40 ♀♀ from the type locality at the british museum (natural history), nos 1998.785– 795, 1998.796–797, 1998.798–810, 1998.811–825. sinev a.y. & dumont h.j., revision of the costata-group of alona (cladocera) 29 type locality west venezuela-brazil: small rock pool on roraima tepui (= table mountain). distribution recorded from a humic-water rock-pool on roraima tepui in venezuela (hudeč 1998). flavalona iheringula (kotov & sinev, 2004) comb. nov. fig. 6k–m alona iheringi sars, 1901: 49–51, plate ix, fig. 2a–c. alona iheringula kotov & sinev, 2004: 96–98, figs 1–8. alona iheringi – sinev 2001: 114–118, figs 1–25. alona iheringula – van damme & dumont 2010: 764, fig. 6. non alona iheringula – silva et al. 2014: 2–5, fig. 1. diagnosis female length 0.38–0.45 mm. body oval (fig. 6k), maximum height at mid-body, height/length ratio ca 0.62– 0.65. valves from previous molts retained by some specimens. posteroventral angle with about 50 setules protruding beyond valve margins. ventral margin with about 60 setae. valves without sculpture or tubercles. eye smaller than ocellus. rostrum short. posterior margin of head shield broadly rounded. three major head pores (fig. 6l) of similar size, middle pore located closer to posterior pore. connection between pores narrow. pp ca 0.5–0.7 ip. lateral head pores of ca 0.3 ip length, lateral pore pockets shallow, semicircular, with depth about half length of pore. labral keel without denticles on anterior margin. postabdomen truncated, length about 2.2–2.4 height. distal margin convex, distal angle prominent, rounded. dorsal margin weakly convex in preanal part, concave in anal part, and almost straight in postanal part. distal part of dorsal margin 2.5 times longer than preanal one, with preanal portion 1.5 times longer than anal. preanal angle well-defined, postanal angle weakly defined. postanal margin of postabdomen with 7–8 marginal denticles. anal margin with 2–3 more slender denticles and 2–3 clusters of marginal setules. 10–12 lateral fascicles of setules, central setules in fascicle the longest. postabdominal claw 1.3 times longer than preanal portion of postabdomen. basal spine short, robust, ca 0.2 lengths of claw. antennule with longest aesthetasc ca ⅔ length of antennule itself. limb ii with scrapers 3 and 7 only slightly thicker than other. exopodite iii with seta 6 almost as long as seta 3. exopodite iv with seta 1–3 of similar length, setae 4 about ⅓ length of seta 3, setae 5–6 about ⅔ length of seta 3. epipodites iv–v with long processes. exopodite v strongly bilobed (fig. 6m). male length 0.34 mm. body low ovoid (fig. 6n), height/length ratio ca 0.58. ocellus larger than eye. postabdomen (fig. 6o) evenly truncated. preand postanal angles not defined. distal part of postabdomen 2.5 times longer than preanal. distal process long, about ⅔ length of claw. distal clusters of marginal setules short. postabdominal claw short, basal spine short and thin, slightly shorter than claw. european journal of taxonomy 223: 1–38 (2016) 30 differential diagnosis flavalona iheringula comb. nov. differs from f. rustica rustica comb. nov. and f. rustica americana comb. nov. by the middle major head pore located closest to the posterior pore and by the strongly bilobed exopodite of limb v; from f. bicolor comb. nov. by a smaller size, short rostrum and labral keel without denticles; from f. hudeci comb. nov. by a smaller size, presence of numerous setules on posteroventral corner of valves, and by seta 6 of exopodite iii almost as long as seta 3, and from f. sphagnophila comb. nov. by having transverse lateral head pores. male f. iheringula comb. nov. differ from f. bicolor comb. nov. and f. rustica comb. nov. by shorter distal marginal setules on the postabdomen. type material lectotype adult parthenogenetic female, zoological museum of oslo university, f12336a. paralectotypes see kotov & sinev (2004) for a full list. material studied earlier see sinev (2001) and kotov & sinev (2004) for a list of material from brazil. type locality brazil: são paulo. distribution widely distributed in brazil (elmoor-loureiro 2015), but not yet recorded outside of the country. flavalona sphagnophila (van damme & eggermont, 2011) comb. nov. fig. 6p alona sphagnophila van damme & eggermont, 2011: 71–75, figs 4–5. diagnosis (from van damme & eggermont 2011) female length of adult 0.42–0.48 mm. body regular oval, maximum height at mid-body, height/length ratio ca 0.62–0.65. valves from previous molts not retained. posteroventral angle with about 50 setules protruding beyond valve margins. ventral margin with about 70 setae. valves without sculpture or tubercles. eye smaller than ocellus. rostrum short. posterior margin of head shield broadly rounded. three major head pores (fig. 6p) of similar size, middle pore located closest to posterior pore. connection between pores narrow. pp ca 0.6–0.8 ip. lateral head pores short, rounded, about to 0.1 ip length, lateral pore pockets round, their diameter over 3 lengths of pore. labral keel without denticles. postabdomen truncated, length ca 2.0–2.5 height. distal margin convex, distal angle prominent, rounded. dorsal margin weakly convex in preanal part, concave in anal part, and almost straight in postanal part. distal part of dorsal margin 2.5 times longer than preanal one, with preanal portion 1.5 times longer than anal. preanal angle well-defined, postanal angle weakly defined. postanal margin of postabdomen provided with 7–8 marginal denticles, length of denticle strongly decrease basally. anal margin with 2–3 more slender denticles and 2–3 clusters of marginal setules. 10–12 lateral fascicles of setules, sinev a.y. & dumont h.j., revision of the costata-group of alona (cladocera) 31 central setules in fascicle being longest. postabdominal claw 1.3 times longer than preanal portion of postabdomen. basal spine short, robust, ca 0.2 length of claw. antennule with longest aesthetasc as ca ⅔ length of antennule itself. limb ii with scrapers 3 and 7 only slightly thicker than other. exopodite iii with seta 6 as long as seta 3. exopodite iv with seta 1–3 of similar length, setae 4 ca ⅓ length of seta 3, setae 5–6 about ½ length of seta 3. epipodites iv–v with long processes. exopodite v strongly bilobed. male unknown. differential diagnosis flavalona sphagnophila comb. nov. differs from all species of the group by short, rounded lateral pores with pocket diameter over 3 pore lengths. it differs from f. rustica rustica comb. nov. and f. rustica americana comb. nov. by the middle head pore located closer to the posterior pore and by the strongly bilobed exopodite of limb v; from f. bicolor comb. nov. by its smaller size, short rostrum and labral keel without denticles; from f. hudeci comb. nov. by a smaller size, presence of numerous setules on the posteroventral corner of the valves, and by seta 6 of exopodite iii almost as long as seta 3. type material holotype a parthenogenetic ♀, royal belgian institute of natural sciences, ig.31.685. paratypes seven parthenogenetic ♀♀, royal belgian institute of natural sciences, ig. 31.685. type locality uganda: lake kopello, carex bogs, rwenzori mountains, 00º18.612′ n, 29º53.504′ s, 4054 m.a.s.l.. distribution flavalona sphagnophila comb. nov. appears to be endemic to the rwenzori mountains. it inhabits the carex-sphagnum bogs surrounding lakes kopello (4017 m.a.s.l.) and nsuranja (3834 m.a.s.l), but not the littoral of these lakes (van damme & eggermont 2011). discussion the generic status of flavalona gen. nov. is justified by at least two autapomorphies, viz. elongated lateral head pores with pocket-like cavities, and a male postabdomen with a genital process. within aloninae, pores with cavities are present in ovalona nigra (smirnov, 1996) and camptocercus australis baird, 1843 (see sinev 2015a, 2015b), but in both these species they are small and spherical. other species of ovalona and camptocercus sars, 1896 have normal, alonine dot-like lateral pores. genital process of male postabdomen is present in genera leydigia and armatalona, both quite remote from alona s. lato (see kotov 2009; sinev 2004). like other recently established genera of alona s.l., flavalona gen. nov. is uniform in the morphology of the thoracic limbs, which is a criterion for generic status within the group. it differs from the affinis and intermedia-groups of hexalona by a weakly developed idl seta 1 of limb i, seta i of limb i with reduced distal part, and by differentiated flaming-torch setae of limb iv. in limb morphology, flavalona gen. nov. is similar to the guttata-group, and both might well be sister-groups. synapomorphies of flavalona gen. nov. and the guttata-group include reduced idl seta 1 of limb i, differentiated flamingeuropean journal of taxonomy 223: 1–38 (2016) 32 torch setae of limb iv and weakly-developed lateral fascicles of postabdominal setules, with distalmost setae not much thicker than others, as common for all other alona s.l. the guttata-group, which should probably be given generic status as well, have minute lateral head pores and lack genital process от male postabdomen. species of the guttata-group also clearly differ from flavalona gen. nov. by the morphology of the female postabdomen, which is shorter and wider, and has a prominent preanal angle. the guttata-group also has one distinctive autapomorphy in limb morphology, viz. the fully reduced seta i of limb i (see sinev 1999b; sinev & silva-briano 2012), which is present in flavalona gen. nov. the sister status of flavalona gen. nov. and guttata-group is also supported by dna data. in trees produced by sacherova & hebert (2003), flavalona costata comb. nov. and alona guttata sars, 1862 form a clade, which does not group with alona affinis (leydig, 1860), the only other hexalona species analyzed in their work. while flavalona gen. nov. and the guttata-group thus form a monophyletic clade, their affinities with other groups are unclear. their unique synapomorphy, the differentiated flaming-torch setae of limb iv is not shared by any other alona s.l., and their similarities with other groups of hexalona, such as the affinis and intermedia groups, include only plesiomorphic characters. the genus is clearly separated into two lineages, the costataand the rustica-clades. these groups differ in sculpture and ventral setae of the valves, morphology of lateral head pores, shape and armament of female postabdomen, and morphology of male postabdomen. in contrast, there are no significant differences between them in morphology of appendages. the species-richness of the group appears still to be underestimated. flössner & frey (1970) did not find differences between eurasian and american f. costata comb. nov., and assigned nearctic f. rustica s.l. comb. nov. to subspecies status. but they did not study the morphology of the appendages and of the head pores. the north american f. costata s.l. comb. nov. may belong to a separate species, and north american f. rustica s.l. comb. nov. may include populations differing in limb morphology from each other and from f. rustica rustica comb. nov.. two taxa of flavalona gen. nov. from greenland, described by røen (1992) as alona fabricii røen, 1992 and a. muelleri røen, 1992 were sunk in the synonymy of f. rustica comb. nov. and f. costata comb. nov. by van damme et al. (2010), but in view of the unsettled situation in north american flavalona gen. nov., these taxa need to be restudied. the taxonomic status of flavalona gen. nov. in sub-sahelian africa remains questionable. moreover, alona virago brehm, 1935 from china, is morphologically similar to species of the costata-clade. its postabdomen is similar to that of f. weltneri comb. nov. (see chiang & du 1979), but its head pores and appendages are unknown. flavalona gen. nov. is distributed worldwide, but is more successful in temperate regions of the northern hemisphere. flavalona costata s.l. comb. nov. is a one of the common alonines in the palearctic and nearctic regions. flavalona rustica americana s.l. comb. nov. is common in the nearctic, and f. rustica rustica comb. nov. is widely distributed in europe and siberia. two more taxa with a limited distribution, f. weltneri comb. nov. and f. bicolor comb. nov., are confined to temperate regions as well. in contrast, even widely distributed tropical species, f. cheni comb. nov., f. iheringula comb. nov., f. margipluma comb. nov. and f. setigera comb. nov., as well as african populations of flavalona gen. nov., appear to be rare. the rustica-clade, while more species-rich, is obviously less successful than the costata-clade. most species of the group (f. rustica americana s.l. comb. nov., f. bicolor comb. nov., f. iheringula comb. nov. and f. hudeci comb. nov.) occur in america. flavalona rustica americana s.l. comb. nov. is wide-ranging, but all other american species have limited ranges. in the old world the rustica-clade is represented by only two species, of which f. sphagnophila comb. nov. is a mountain endemic and f. rustica rustica sinev a.y. & dumont h.j., revision of the costata-group of alona (cladocera) 33 comb. nov., while widely distributed in the west palearctic, is confined to dystrophic sphagnum bogs in coniferous forests and mountain belts. species of the rustica-clade have never been recorded in china (chiang & du 1979; ji et al. 2015), south-east asia (korovchinsky 2013), india (chatterjee et al. 2013) or australia (smirnov & timms 1983). in contrast, the costata-clade is distributed worldwide, and includes at least four wide-ranging species: palearctic f. costata s.s. comb. nov., north american f. costata s.l. comb. nov., tropical asian f. cheni comb. nov. and australian f. setigera comb. nov. two species have more limited ranges (f. margipluma comb. nov. and f. weltneri comb. nov.) and two are mountain endemics (f. hudeci comb. nov. and f. natalensis comb. nov.). the distribution of flavalona gen. nov. is similar to that of ovalona. about half of the species in both genera are mountain endemics and species with a limited distribution. the less successful setulosaclade of ovalona and rustica-clade of flavalona mostly occurs in the new world and is absent in tropical asia and australia; species of the setulosa-clade are not present in africa either, while the only african species of the rustica-clade, f. sphagnophila comb. nov., is a mountain endemic. in contrast, the evolutionary advanced ovalona cambouei-clade is distributed on all continents. the most widely distributed species, neotropical o. glabra (sars, 1901), palaeaotropical o. cambouei (guerne & richard, 1893) and australian o. pulchella (king, 1853), are similar morphologically. the costataclade of flavalona gen. nov. is also cosmopolitan, and is much more morphologically uniform than the rustica-clade. acknowledgments we are grateful to yang liang gu (jinan university, guangzhou), thomas jensen and bjorn walseng (norwegian institute for nature research, oslo) for assistance during sampling trips, and to prof. nikolai n. smirnov (a.n. severtsov institute of ecology and evolution, moscow) for material and valuable comments. collecting of material in eurasia and its morphological studies were conducted solely by the first author under support by the russian science foundation grant 14-14-00778. efforts of the second author were supported by the chinese national programs for fundamental research and development (973 program, no. 2012cb956104) and by grant for leading talent scientists of the guangdong province to henri dumont. references alonso m. 1996. crustacea, branchiopoda. fauna iberica 7. museo national de ciencias natural. consejo superior de investigationes cientificas, madrid. behning a.l. 1941. the cladocerans of the caucasus. gruzmedgiz publishing, tbilisi. 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botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands. http://dx.doi.org/10.1111/j.1096-3642.2010.00639.x http://dx.doi.org/10.1111/j.1096-3642.2010.00639.x http://dx.doi.org/10.4081/jlimnol.2013.e36 http://dx.doi.org/10.1007/s10750-008-9607-6 http://dx.doi.org/10.1051/limn/2010034 two new species of the rhyacophila nigrocephala species group from china (insecta, trichoptera, rhyacophilidae) changhai sun department of entomology, nanjing agricultural university, jiangsu 210095, china. email: chsun@njau.edu.cn urn:lsid:zoobank.org:author:dd6e8de5-3e20-4f9e-bc80-5ff92ce798f1 abstract. two new species of the rhyacophila nigrocephala species group, r. voluta sp. nov. and r. linguiformis sp. nov., are described, diagnosed and illustrated. rhyacophila voluta sp. nov. is similar to r. pentagona malicky & sun, 2002 in male genitalia, but can be diagnosed by the inferior appendages of the same length as the complex of preanal appendages and dorsal lobe of segment ix, fused base of the basal segments of inferior appendages, and slightly incised distal margin of apical segment of inferior appendage. rhyacophila linguiformis sp. nov. is similar to r. rima sun & yang, 1995 and r. esorima mey, 1996 in male genitalia, but can be diagnosed by the progressively narrowed complex of preanal appendages and the dorsal lobe of segment x, the short anal sclerites, and the small gap between the upper and lower lobes of apical segment of inferior appendages. an updated checklist of 23 species recorded from china is presented. key words. caddisfly, taxonomy, species group, subgroup. sun c. 2017. two new species of the rhyacophila nigrocephala species group from china (insecta, trichoptera, rhyacophilidae). european journal of taxonomy 300: 1–10. https://doi.org/10.5852/ejt.2017.300 introduction the rhyacophila nigrocephala diagnostic species group was established by ross (1956) to contain 14 species known at that time. schmid (1970) placed it in his rhyacophila divaricata branch, and increased the number of species in the group to 32, with one subspecies (r. putata temba schmid, 1970 from manipur, india). afterwards, several authors reported 29 species from china (sun & yang 1995, 1999; malicky 1995; malicky & sun 2002), vietnam (oláh 1987; mey 1996; armitage & arefina-armitage 2003), nepal (kiss 2011), thailand (malicky 1991; malicky & chantaramongkol 1993; malicky et al. 2004), japan (kobayashi 1987) and korea (botosaneanu 1970; kobayashi 1989). furthermore, additional distribution data have been appended to some species (armitage et al. 2005; chuluunbat & morse 2007; hsu & chen 2003; ito 1999; ivanov 2011; kiss 2013; malicky 2007, 2012, 2014; mey 2005; morse 2016; nozaki & nakamura 2007; nuntakwang et al. 2007; ohkawa & hattori 2007; potikha 2005; satake & kuranishi 2007; wityi et al. 2015; yang et al. 2005). currently, 61 species / subspecies are placed into the r. nigrocephala species group, 53 species / subspecies from the oriental biogeographic region, 7 from the palearctic biogeographic region, and 3 occur in european journal of taxonomy 300: 1–10 issn 2118-9773 https://doi.org/10.5852/ejt.2017.300 www.europeanjournaloftaxonomy.eu 2017 · sun c. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:8bad12e5-a073-42e1-b368-da5319612150 1 mailto:chsun%40njau.edu.cn?subject= http://zoobank.org/urn:lsid:zoobank.org:author:dd6e8de5-3e20-4f9e-bc80-5ff92ce798f1 https://doi.org/10.5852/ejt.2017.300 https://doi.org/10.5852/ejt.2017.300 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:8bad12e5-a073-42e1-b368-da5319612150 both the oriental and palearctic biogeographic region. in addition, 9 species belong to r. manuleata sub-group, 52 species/subspecies fall into the r. nigrocephala sub-group. of 23 chinese species of the rhyacophila nigrocephala species group (table 1), 19 are from oriental china, 1 from palearctic china and 4 occur in both oriental and palearctic china. furthermore, 5 species belong to the r. manuleata sub-group and 18 belong to the r. nigrocephala sub-group. the purpose of this article is to describe two new species, rhyacophila linguiformis sp. nov. and r. voluta sp. nov., both from guangdong, oriental china. as a result, the total number of species of the r. nigrocephala species group now known from china is 25, and the total number of species/subspecies in the species group is 63. material and methods the specimens examined were collected using pan traps with 15-w ultraviolet light tubes set near streams. rhyacophilid adults were then sorted and stored in 100% alcohol. the genitalia preparation procedure follows that used by xu et al. (2014). the structures of the male genitalia were traced in pencil using a drawing tube on a nikon eclipse 80i microscope. original pencil drawings were inked digitally with photoshop® cs4 version to produce illustrations. each male abdomen was transferred to a microvial with 80% ethanol and stored together with the remainder of the specimen in a larger jar. terminology for male genitalia follows that of schmid (1970). type specimens have been deposited in the insect collection, nanjing agricultural university, nanjing, jiangsu province, pr china. abbreviations aed. = aedeagus ab. = apical band aia. = apical segment of inferior appendages as. = anal sclerites bia. = basal segment of inferior appendages com. = complex of the preanal appendages and the apicodorsal lobe of segment ix end. = endotheca ix = segment ix par. = parameres pht. = phallotheca sag. = sagittal appendage tb. = tergal band institutional acronyms: njau = nanjing agricultural university, nanjing, china european journal of taxonomy 300: 1–10 (2017) 2 results family rhyacophilidae stephens, 1836 genus rhyacophila pictet, 1834 rhyacophila nigrocephala species group diagnosis (modified from schmid 1970) the species group can be diagnosed by the combination of following characters. in the male, segment ix is usually long, with the apicodorsal lobe fused with the preanal appendages into a complex, covering the other portion of the male genitalia. the anal sclerites are fused, with the apex slightly incised mesally and curved upwards, its root is large and curved upwards. the tergal band is slender, flattened dorsoventrally, with the apex sagittal, and has been termed as the sagittal appendage by ross (1956). sub-groups and species distribution r. manuleata sub-group altoincisiva hwang, 1957 china (fujian) manuleata martynov, 1934 china (taiwan), japan, korea, oussouri, ussr (far east) melli ulmer, 1926 china (guangdong) shikotsuensis iwata, 1927 china (taiwan), japan, russia linguiformis sp. nov. china (guangdong) rima sun & yang, 1995 china (jiangxi) r. nigrocephala sub-group formosana ulmer, 1927 china (taiwan) kimminsi ross, 1956 china (taiwan) lata martynov, 1918 china (heilongjiang), mongolia, oussouri, russia (southern far east, siberia) nigrocephala iwata, 1927 china (taiwan), japan tecta morton, 1900 bhutan, china (taiwan), india (khasi hills, kameng), myanmar amblyodonta sun & yang, 1999 china (sichuan, yunnan) anakdjeram malicky, 1995 china (sichuan) anaksungai malicky, 1995 china (sichuan) celata sun & yang, 1999 china (guizhou) cuneata sun & yang, 1999 china (zhejiang) euryphylla sun & yang, 1999 china (sichuan) exilis sun & yang, 1999 china (sichuan, yunnan) forcipata malicky & sun, 2002 china (shaanxi) hadestril malicky & sun, 2002 china (zhejiang) hamosa sun, 1995 china (zhejiang) pentagona malicky & sun, 2002 china (yunnan) unisegmentalis malicky & sun, 2002 china (shaanxi) vascula malicky & sun, 2002 china (shaanxi) voluta sp. nov. china (guangdong) table 1. species of the rhyacophila nigrocephala species-group recorded from china. sun c., rhyacophila nigrocephala species group from china 3 the apical band is with two lateral arms articulating, with anal sclerites above and with the tergal band beneath. the phallic apparatus is elongate and cylindrical; the aedeagus with the apex is thin; the parameres, if present, are slender. schmid (1970) recognized two sub-groups, the r. manuleata subgroup and the r. nigrocephala sub-group. rhyacophila manuleata sub-group diagnosis rhyacophila manuleata sub-group is characterized by (1) each apical segment of the inferior appendages deeply incised, (2) phallus apparatus simple, (3) the parameres present, (4) the base of the basal segment of inferior appendages not fused mesally. rhyacophila linguiformis sp. nov. urn:lsid:zoobank.org:act:a2708f26-9f2b-49c3-bca2-d18f180a2ff9 fig. 1 diagnosis this new species is similar to r. rima sun & yang, 1995 from jiangxi, china and r. esorima mey, 1996 from vietnam in the male genitalia, all of them belong to the r. manuleata sub-group, but the new species differs in: (1) the complex of preanal appendages and the dorsal lobe of segment x in dorsal view is progressively narrowed from the base to the apex, but in r. rima the complex is somewhat spindleshaped, and in r. erorima the complex with two parallel lateral margins, which become narrow near the apex; (2) the anal sclerites in lateral view are shorter than the complex, with the apex moderately concave in dorsal view, but in r. rima it extends beyond the apex of the complex, with a deep incision mesally in dorsal view, and in r. esorima it extends to the complex, with the apex slightly incised mesally; (3) the gap between the upper and the lower lobes of each inferior appendage is smaller than the width of the lower lobe, but in both r. rima and in r. esorima the gap at least equals the width of each lower lobe. fig. 1. rhyacophila linguiformis sp. nov., male genitalia. a. lateral view. b. dorsal view. c. ventral view. d. complex of preanal appendages and apicodorsal lobe of segment ix, caudal view. scale bar: 0.5 mm. european journal of taxonomy 300: 1–10 (2017) 4 http://zoobank.org/urn:lsid:zoobank.org:act:a2708f26-9f2b-49c3-bca2-d18f180a2ff9 etymology the species epithet comes from the latin word “linguiformis”, meaning “tongue-shaped”, in reference to the tongue-shaped complex of the preanal appendages and the apicodorsal lobe of segment ix. type material holotype china: ♂, in alcohol, collected in guangdong province, xinyi county, town of dacheng, dawuling nature reserve, stream inside entrance of reserve, 111.19° e, 22.27° n, 1021 m, 26 may 2004, xin zhou and sanbao tang leg. (njau). description male body. length of each forewing: 1.0 mm (n = 1). body (in alcohol) dark brown. head dark brown; eyes black; antennae dark brown, palpi brown. thorax dark brown dorsally, brown laterally and ventrally; legs brown; wings brown. abdomen dark brown dorsally and brown ventrally. male genitalia (fig. 1a–d). segment ix (ix) in lateral view with each anterior margin straight, each posterior margin sinuate, dorsal margins short and ventral margins long (fig. 1a); in dorsal view anterior margin straight, and posterior margin with deep u-shaped incision (fig. 1b). preanal appendages and apicodorsal lobe of segment ix fused tightly into complex (com.), in lateral view complex somewhat trapezoidal with upper margin 3 times as long as the lower margin (fig. 1a), and in dorsal view somewhat tongue-shaped, tapering from base to apex (fig. 1b). anal sclerites (as., fig. 1a) in lateral view barshaped, about ½ times as long as upper margin of complex. apical band (ab.) in lateral view slender, slightly curved posteriad (fig. 1a). tergal band (tb.) membranous, in lateral view elongate. phallotheca (pht.) irregular in lateral view, somewhat rectangular in ventral view; aedeagus (aed.) with a large base, then progressively narrowed to rounded apex; paired parameres (par.) slender and shorter than aedeagus. basal segments of inferior appendages (bia.) setose sparsely, each is subrectangular in lateral view and triangular in ventral view; apical segments (aia.) setose densely, each divided into 2 lobes, each upper lobe slightly shorter than lower one. female unknown. distribution china (guangdong). rhyacophila nigrocephala sub-group diagnosis the rhyacophila nigrocephala sub-group can be diagnosed by the combinations of following characters: (1) none of the apical segments of the inferior appendages is incised; (2) parameres absent; (3) basal segments of inferior appendages fused basomesally. sun c., rhyacophila nigrocephala species group from china 5 rhyacophila voluta sp. nov. urn:lsid:zoobank.org:act:df10323c-1202-4d75-8cb2-9c2c5b87a026 fig. 2 diagnosis the new species belongs to the r. nigrocephala sub-group, and is similar to r. pentagona malicky & sun, 2002 from yunnan, china, in the male genitalia, but differs in: (1) the inferior appendages of about the same length as the complex of the preanal appendages and the apicodorsal lobe of segment ix (obviously shorter than the complex in r. pentagona); (2) the basal segments of the inferior appendages in lateral view are somewhat rectangular and in ventral view with their bases fused (pentagonal in r. pentagona, and their bases not fused obviously); (3) the apical segments of the inferior appendages with each distal margin slightly concave (straight in r. pentagona). etymology the species epithet comes from the latin word “volutus”, meaning “having been curled”, in reference to the subapical margin of the complex of the preanal appendages and the apicodorsal lobe of segment x extending upward into a curled lamella. type material holotype china: ♂, in alcohol, collected in guangdong province, ruyuan county, nanling national nature preserve, lao-peng keng at cascading tributary, route x327, marker 22.5 km, 113.01° e, 24.93° n, 1110 m, 18–19 may 2004, john c. morse, xiaoli tong and xin zhou leg. (njau). paratypes china: 4 ♂♂, in alcohol, collected at the same locality along with the holotype, except marker 17.45 km, 113.03° e, 24.91° n, 935 m, 21–22 may 2004, john c. morse and changhai sun (njau). fig. 2. rhyacophila voluta sp. nov., male genitalia. a. lateral view. b. dorsal view. c. segment ix and inferior appendages, ventral view. d. phallic apparatus, ventral view. scale bar: 0.5 mm. european journal of taxonomy 300: 1–10 (2017) 6 http://zoobank.org/urn:lsid:zoobank.org:act:df10323c-1202-4d75-8cb2-9c2c5b87a026 description male body. length 5.0 mm (n = 3), length of each forewing 6.0–7.0 mm (n = 5). body (in alcohol) brown. head dark brown; antennae brown; palpi light brown, eyes black. thorax brown dorsally, yellow laterally and ventrally; wings dark brown; legs with coxae, trochanters and femora yellow, remainder parts dark brown, spurs dark brown. abdomen dark brown dorsally and yellowish brown ventrally. male genitalia (fig. 2a–c). segment ix (ix) in lateral view with middle portion of each anterior margin (about half the height of segment ix) straight and slightly projecting anteriad, each posterior margin almost straight, with posteroventral angle slightly projecting posteriad, dorsal and ventral margins straight; in dorsal view, anterior margin incised in v-shaped, posterior margin slightly concave; in ventral view anterior margin with broad shallow incision and posterior margin convex slightly. apicodorsal lobe of segment ix and preanal appendages fused into complex (com.), in lateral view it curved downward at middle, with broad base and thin apex, subapex with dorsal margin extended upward to form lamella; in dorsal view complex with the two lateral margins parallel to each other, and apex triangular, the lamella curled. anal sclerites (as.) large, with base fused into complex of apicodorsal lobe and preanal appendages, in lateral view somewhat triangular, in dorsal view its apex incised. apical band (ab.) membranous. sagittal appendage (sag.) in lateral view style-like. basal segment of each inferior appendage (bia.) in lateral view somewhat rectangular, base slightly broader than apex in lateral view, in ventral view bases fused mesally; apical segment of each inferior appendage (aia.) in lateral view trapezoidal, with distal margin shallowly concave, in ventral view somewhat triangular. phallic apparatus simple: phallotheca (pht.) tubular in lateral view and trapezoidal in ventral view, endotheca (end.) membranous, aedeagus (aed.) tubular, with apex narrowed suddenly. female unknown. distribution china (guangdong). discussion the r. nigrocephala species group has the highest diversity of species within the genus rhyacophila, with 63 members up to date. all species are exclusively distributed in both the eastern palearctic and oriental biogeographic regions. twenty-five species of r. nigrocephala species group (including the 2 new species described in this paper) are recorded from china (table 1), which is about 40% of the world fauna of the group. among them, 18 species are endemic to china, distributed in the provinces of guangdong (3 spp.), fujian (1 sp.), jiangxi (1 sp.), guizhou (1 sp.), shaanxi (3 spp.), sichuan (3 spp.), yunnan (1 sp.), both sichuan and yunnan (2 spp.), taiwan (2 spp.), and zhejiang (3 spp.); r. lata martynov, 1918 is the only palearctic species, distributed in heilongjiang, china, mongolia and russia; r. manuleata martynov, 1934, r. nigrocephala iwata, 1927 and r. shikotsuensis iwata, 1927 are widespread in palearctic japan, korea and even russia to oriental taiwan, but are not recognized from mainland china, this situation may be due to poor sampling efforts in mainland china. in recent papers, r. nigrocephala was recorded from the liao river basin in the northeast of china (zhao et al. 2015), and r. manuleata was reported from mt. dabie, hubei province (qiu & yan 2016). so far, r. tecta morton, 1900 is recognized from the oriental region, being widely distributed in bhutan, china (taiwan), india (khasi hills, kameng) and myanmar. in summary, the distribution of members of the r. nigrocephala species group is confined to sun c., rhyacophila nigrocephala species group from china 7 the eastern palearctic and oriental biogeographic regions, but more samples are bound to reveal a more extended distribution of each species within the group. acknowledgements i am grateful to dr. john c. morse of the clemson university, usa, for his long term help and support in the study on trichoptera. thanks are also due to the team members of the collecting trip to the provinces of guang-dong and guang-xi in 2004 and to the province of jiang-xi in 2005, co-organized by dr. morse and prof. yang lian-fang of the nanjing agricultural university, pr china, who are acknowledged for their cooperation and help. i am also grateful to the two anonymous reviewers and the editors for their valuable comments that improved the quality of the manuscript. this research was supported by fundamental research funds for the central university (kyz201106), the national natural science foundation of china (nsfc, no. 412715125), and the united states national science foundation (deb-0316504). references armitage b.j. & arefina t.i. 2003. the genus rhyacophila pictet (trichoptera: rhyacophilidae) in vietman. aquatic insects 25: 95–124. https://doi.org/10.1076/aqin.25.2.95.14037 armitage b.j., mey w., arefina t.i. & schefter p.w. 2005. the caddisfly fauna (insecta: trichoptera) of vietnam. in: tanida k. & rossiter a. 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(eds) proceedings of the 11th international symposium on trichoptera: 441–460. tokai university press, hadano-shi, kanagawa, japan. zhao r., gao x., ding s., zhang y., qu x. & liu s. 2015. tolerance values of macroinvertebrate taxa in liao river basin. acta ecologica sinica 35 (14): 4797–4809. https://doi.org/10.5846/stxb201311082699 manuscript received: 27 april 2016 manuscript accepted: 22 july 2016 published on: 15 march 2017 topic editor: gavin broad desk editor: kristiaan hoedemakers printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands. european journal of taxonomy 300: 1–10 (2017) 10 https://doi.org/10.11646/zootaxa.3785.2.5 https://doi.org/10.11646/zootaxa.3785.2.5 https://doi.org/10.5846/stxb201311082699 european journal of taxonomy 105: 1–6 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2014.105 www.europeanjournaloftaxonomy.eu 2014 · young r.e., vecchione m. & braid h.e. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:fcd51f6f-a5d6-4466-adb6-0adc9f560f66 1 mastigotragus, a new generic name for mastigoteuthis pyrodes young, 1972 (cephalopoda: mastigoteuthidae) richard e. young,1 michael vecchione 2 & heather e. braid 3 1 university of hawaii, honolulu, hawaii, usa. email: dickphyllisyoung@gmail.com 2 nmfs national systematics laboratory, smithsonian institution, washington, dc, usa. email: vecchiom@si.edu 3 institute for applied ecology new zealand, auckland university of technology, auckland, new zealand. email: heather.braid@gmail.com 1 urn:lsid:zoobank.org:author:c5177553-e350-4c6a-9d2e-70581f2ed152 2 urn:lsid:zoobank.org:author:e79ea1da-a7b4-482d-a5e3-ab81e74e4743 3 urn:lsid:zoobank.org:author:a28b5d88-b00c-46a1-9f48-367918caf2f2 abstract. a recent paper on the phylogenetic relationships of species within the cephalopod family mastigoteuthidae meant great progress in stabilizing the classification of the family. the authors, however, left the generic placement of mastigoteuthis pyrodes unresolved. this problem is corrected here by placing this species in a new monotypic genus, mastigotragus, based on unique structures of the photophores and the funnel/mantle locking apparatus. keywords. mastigoteuthidae, morphology, nomenclature, mastigotragus gen. nov., deep-sea squid young r.e., vecchione m. & braid h.e. 2014. mastigotragus, a new generic name for mastigoteuthis pyrodes young, 1972 (cephalopoda: mastigoteuthidae). european journal of taxonomy 105: 1–6. http://dx.doi. org/10.5852/ejt.2014.105 introduction the squid family mastigoteuthidae contains oceanic, mostly bathypelagic, species that are soft bodied and easily damaged in trawls. the inaccessibility of species and their fragile nature are in large degree responsible for problems in the classification of this family. little consistency has existed between classifications, even with the same senior author (e.g., salcedo-vargas & okutani 1994 and salcedovargas 1997). in a revision of the family on the tree of life website, vecchione et al. (2004, with subsequent modifications up to 2014) suggested, because of the uncertainties in classification, “… that modifying the classification is premature and should wait until our knowledge has increased to the point where a full phylogenetic study can be made. we recognize, therefore, only the single genus mastigoteuthis.” vecchione et al. (2004–2014) then placed species (excluding species of doubtful validity and those known only from paralarvae; see below) into the following six species groups: mastigoteuthis agassizii group (three-species group), m. cordiformis (one-species group), m. glaukopis group (threespecies group), m. hjorti (one-species group), m. magna group (two-species group) and m. pyrodes (one-species group). the excluded species from the vecchione et al. species groups are: mastigoteuthis http://dx.doi.org/10.5852/ejt.2014.105 www.europeanjournaloftaxonomy.eu/index.php/ejt/index http://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:fcd51f6f-a5d6-4466-adb6-0adc9f560f66 mailto:dickphyllisyoung%40gmail.com?subject= mailto:vecchiom%40si.edu?subject= mailto:heather.braid%40gmail.com?subject= http://zoobank.org/urn:lsid:zoobank.org:author:c5177553-e350-4c6a-9d2e-70581f2ed152 http://zoobank.org/urn:lsid:zoobank.org:author:e79ea1da-a7b4-482d-a5e3-ab81e74e4743 http://zoobank.org/urn:lsid:zoobank.org:author:a28b5d88-b00c-46a1-9f48-367918caf2f2 http://dx.doi.org/10.5852/ejt.2014.105 http://dx.doi.org/10.5852/ejt.2014.105 european journal of taxonomy 105: 1–6 (2014) 2 latipinna (sasaki, 1916), m. islini macdonald & clench, 1934, m. inermis rancurel, 1972, m. okutani (salcedo-vargas, 1997) and chiroteuthoides hastula berry, 1920. the phylogenetic study that was called for by vecchione et al. (2004-2014) to establish a proper classification for the mastigoteuthidae has now been published by braid et al. (2014). those authors analyzed three mitochondrial genes from 29 mastigoteuthid squids in eight species and re-examined morphological characters. their results agreed with the species groups established by vecchione et al. (2004-2014) and enabled them to establish a formal generic classification for the family that utilized generic and subgeneric names previously present in the literature. the following table compares these two classification systems: braid et al. (2014) left one problem unresolved. they stated, “ unfortunately, no specimens of this species [mastigoteuthis pyrodes] were examined or available for genetic analysis, and therefore, its generic placement remains unclear.” in this paper we examine the generic affinities of mastigoteuthis pyrodes young, 1972 and place it in a new genus. materials and methods we examined all of the specimens of mastigoteuthis pyrodes at the u.s. national museum of natural history (usnm) and at the santa barbara museum of natural history (sbmnh). included were the following type specimens: sbmnh 34983, holotype, ♂, ml 110 mm, and usnm 727462, 3 paratypes (2 ♂♂, ml 85 and 94 mm; 1 sex indet., ml 77 mm). results we strongly support most of the conclusions reached by braid et al. (2014). our examination of mastigoteuthis pyrodes indicates that this species does not belong in mastigoteuthis or in any of the other genera defined by braid et al. (2014). “m.” pyrodes presently remains in mastigoteuthis by default. on superficial examination one could easily conclude that “m.” pyrodes does, indeed, belong in mastigoteuthis. in fin length, fin shape and tentacle structure “m.” pyrodes is most similar to members of mastigoteuthis. the most compelling feature for placing “m.” pyrodes in mastigoteuthis (and the feature that excludes this species from all other genera in the family) is the unique presence of readily visible integumental photophores on the mantle, head and ventral arms in these taxa. a closer comparison of the integument of “m.” pyrodes and the type species of mastigoteuthis, however, shows that they are very different. the photophores of mastigoteuthis agassizii verrill, 1881, the type species of the genus, appear to lie in pockets (fig. 1). the appearance of a pocket is due to the presence of transparent, vacuole-like cells that surround the photocytes and their covering chromatophores. a photophore of “m.” pyrodes appears very different from that of m. agassizii under the dissecting microscope. it is seen as a slight bulge in the skin, comparison of classifications vecchione et al. 2004–2014 braid et al. 2014 1 the mastigoteuthis agassizii group mastigoteuthis spp. 2 mastigoteuthis cordiformis idioteuthis cordiformis 3 the mastigoteuthis glaukopis group echinoteuthis spp. 4 mastigoteuthis hjorti mastigopsis hjorti 5 the mastigoteuthis magna group magnoteuthis spp. 6 mastigoteuthis pyrodes mastigoteuthis pyrodes (no change) young r.e., vecchione m. & braid h.e., a new mastigoteuthid genus 3 not a pocket (i.e., without obvious vacuole-like cells), and is backed by a white reflector and covered by a dark chromatophore (fig. 2b). other features of the skin are also important. mastigoteuthis agassizii has relatively few, scattered chromatophores in the epidermis and each is surrounded by a white ring (the thick edge of a disc that seems to extend beneath the chromatophore); this leaves most of the pigment of the skin in the dermis and not in chromatophore organs of the epidermis (fig. 2a). in contrast, “m.” pyrodes lacks these ringed chromatophores but has more numerous and densely arranged, typical chromatophores which provide most of the skin pigmentation, even though light pigmentation is present in the underlying dermis. mastigoteuthis agassizii also has numerous small, spherical, white structures in the skin whose histology and function are unknown. these structures are lacking in “m.” pyrodes. these large differences in the structure of the integument, by themselves, are enough to conclude that “m.” pyrodes is not a mastigoteuthis. “m.” pyrodes, however, also differs from species of mastigoteuthis in the distribution of the photophores: both taxa have photophores on the ventral surfaces of the mantle, head and ventral arms, but only in “m.” pyrodes are photophores also present on the ventral sides of the fins and the dorsal surfaces of the head and mantle. moreover, the eyelid photophore in “m.” pyrodes is much larger than its integumental photophores, while in mastigoteuthis species the eyelid and integumental photophores are approximately the same size. one other major difference in these taxa is important in the derivation of the new generic name: the funnel component of the funnel/mantle locking apparatus. in “m.” pyrodes this structure has a large tragus but is without an antitragus and without a posterior undercut shelf; these latter two features are always present in mastigoteuthis species, although weakly expressed in some. because of these large morphological differences between “m.” pyrodes and true species of mastigoteuthis, we place “mastigoteuthis” pyrodes as the sole species in a new genus, mastigotragus. fig. 1. longitudinal section through an integumental photophore of mastigoteuthis agassizii verrill, 1881 (modified from chun 1910). european journal of taxonomy 105: 1–6 (2014) 4 fig. 2. photomicrographs of the slightly damaged integument of the head (modified from vecchione et al. 2004–2014). a. mastigoteuthis agassizii. b. “mastigoteuthis” pyrodes [= mastigotragus pyrodes]. – large, white arrows indicate photophores; only two photophores are visible in a but seven are visible in b. small, white arrows indicate two of the many “white spherical structures.” black arrows indicate ringed chromatophores in a and typical chromatophores in b. young r.e., vecchione m. & braid h.e., a new mastigoteuthid genus 5 systematics class cephalopoda cuvier, 1795 subclass coleoidea bather, 1888 order oegopsida d’orbigny, 1845 family mastigoteuthidae verrill, 1881 mastigotragus gen. nov. urn:lsid:zoobank.org:act:5f1ddc20-a391-4b52-ba8a-23ffc9f7888d type species: mastigoteuthis pyrodes young, 1972, by monotypy. diagnosis a mastigoteuthid with (1) numerous small, but easily observed integumental photophores on the ventral surface of arms iv, on the dorsal and ventral surfaces of the head, mantle and fins, and with a relatively large photophore on each eyelid; (2) a broad funnel component of the funnel/mantle locking apparatus, with strong tragus, but without antitragus or undercut posterior margin; (3) comparatively large club suckers (about 0.3 mm in diameter). etymology here “mastigo” refers to the name of the family and “tragus” refers to the prominent tragus of the funnel locking apparatus. mastigo is latin for “whip” which refers to the form of the tentacle; “tragus” is a latin word derived from the greek trágos, indicating an appearance similar to the tragus of the human ear. discussion no additional histological information is available on the structure of the integument of these taxa, but the position of mastigotragus pyrodes as a separate genus is well supported by morphology. we hope that, before long, fresh specimens will be captured that will enable determination of the genetic relationships of this taxon within the family. the species is known only from the eastern north pacific, mostly off southern calfornia. additional information and illustrations of the diagnostic features of mastigotragus pyrodes are presented on the species page of the tree of life website (vecchione et al. 2004–2014). acknowledgements we would like to thank kat bolstad and paul mcbride from the auckland university of technology for their help and the unknown reviewers for their helpful comments. references braid h.e., mcbride p.d. & bolstad k.s.r. 2014. molecular phylogenetic analysis of the squid family mastigoteuthidae (mollusca, cephalopoda) based on three mitochondrial genes. hydrobiologia 725: 145–164. http://dx.doi.org/10.1007/s10750-013-1775-3 chun c. 1910. die cephalopoden. oegopsida. wissenschaftliche ergebnisse der deutschen tiefsee expedition auf dem dampfer „valdivia“ 1898-1899 18 (1): 1–401. salcedo-vargas m.a. 1997. cephalopods from the netherlands indian ocean programme (nio) ii. mastigoteuthid lineage and related forms. beaufortia 47: 91–108. http://zoobank.org/urn:lsid:zoobank.org:act:5f1ddc20-a391-4b52-ba8a-23ffc9f7888d http://dx.doi.org/10.1007/s10750-013-1775-3 european journal of taxonomy 105: 1–6 (2014) 6 salcedo-vargas m.a. & okutani t. 1994. new classification of the squid family mastigoteuthidae (cephalopoda: oegopsida). venus 53: 119–127. vecchione m., young r.e. & lindgren a. 2004–2014. mastigoteuthidae verrill, 1881. mastigoteuthis verrill, 1881. whip-lash squid. the tree of life web project. accessed 21 jan. 2014 (under construction). http://tolweb.org/mastigoteuthis/19453/2014.01.21 young r.e. 1972. the systematics and areal distribution of pelagic cephalopods from the seas off southern california. smithsonian contributions to zoology 97: 1–159. http://dx.doi.org/10.5479/ si.00810282.97 manuscript received: 9 july 2014 manuscript accepted: 13 september 2014 published on: 5 december 2014 topic editor: rudy jocqué desk editor: danny eibye-jacobsen printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; national botanic garden of belgium, meise, belgium; royal museum for central africa, tervuren, belgium; national history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark. http://tolweb.org/mastigoteuthis/19453/2014.01.21 http://dx.doi.org/10.5479/si.00810282.97 http://dx.doi.org/10.5479/si.00810282.97 1 european journal of taxonomy 119: 1–6 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2015.119 www.europeanjournaloftaxonomy.eu 2015 · hughes m. et al. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e begonia yapenensis (sect. symbegonia, begoniaceae), a new species from papua, indonesia mark hughes1, sadie barber1, charlie d. heatubun2 & janet gagul3 1royal botanic garden edinburgh, 20a inverleith row, eh3 5lr, uk. emails: m.hughes@rbge.ac.uk (corresponding author); s.barber@rbge.ac.uk 2fakultas kehutanan dan pusat penelitian lingkungan hidup, universitas papua, jl. gunung salju, amban, manokwari 98314, papua barat, indonesia. email: charlie_deheatboen@yahoo.com 3australian tropical herbarium, james cook university, cairns, qld 4870, australia. email: jgagul@yahoo.com abstract. a new species, begonia yapenensis m.hughes, in begonia section symbegonia (begoniaceae) is described and diagnosed against begonia sympapuana. the new species is endemic to yapen island, papua, indonesia, and is currently known from a single collection. keywords. section symbegonia, new species, endemic, new guinea hughes m., barber s., heatubun c.d. & gagul j. 2015. begonia yapenensis (sect. symbegonia, begoniaceae), a new species from papua, indonesia. european journal of taxonomy 119: 1–6. http://dx.doi.org/10.5852/ ejt.2015.119 introduction begonia section symbegonia (warburg 1894: 149; forrest & hollingsworth 2003: 208) is endemic to the island of new guinea and comprises 13 species (sands 2009). the section was recognised at the genus level prior to a molecular phylogenetic study (forrest & hollingsworth 2003) which found it to be nested within begonia section petermannia (klotzsch 1854: 124; de candolle 1859: 128). the section is well defined morphologically by species having the tepals of the female flowers fused into a tubular corolla, the tepals of the male flowers often fused and usually with columnar androecia and unique endothecial cells in the anthers (tebbitt & maciver 1999). however many of the species within the section are difficult to delimit. it is tempting to speculate that the large amount of highly dissected yet fairly continuous montane habitat on new guinea has promoted rapid yet incomplete diversification in this group, leading to difficult species complexes. the tubular flowers are very different from those in other sections of the genus, and hence the pollination syndrome is likely to differ also. increased population connectivity, possibly mediated by bird pollination, may also contribute to the persistence of widespread species complexes, which would otherwise fragment into different taxa in the absence of gene flow (hughes & hollingsworth 2008). in addition to this biological complexity, the diversity in begonia sect. symbegonia is further difficult to interpret as the species have been described separately over the decades, most without reference to existing taxa and hence without comparative diagnoses. during preparation for a taxonomic revision of the section by two of the authors, it became clear that http://dx.doi.org/10.5852/ejt.2015.119 http://www.europeanjournaloftaxonomy.eu https://creativecommons.org/licenses/by/3.0/ http://dx.doi.org/10.5852/ejt.2015.119 http://dx.doi.org/10.5852/ejt.2015.119 european journal of taxonomy 119: 1–6 (2015) 2 a recent collection from yapen island, papua province, indonesia represents a new species which is described below. materials and methods all available specimens of begonia sect. symbegonia in b, bm, bo, e, fi, k, l, p and sing were examined (272 sheets representing 171 collections), including the types for all names in the section in order to confirm the novelty of the collection from yapen island. the description was based on living material in cultivation at the royal botanic garden edinburgh and later pressed as the holotype. the terminology in the description follows beentje (2010). results class equisetopsida c. agardh (agardh et al. 1825) subclass magnoliidae novák ex takht. (takhtajan 1967) superorder rosanae takht. (takhtajan 1967) order cucurbitales juss. ex bercht. & j.presl (berchtold & presl 1820) family begoniaceae c. agardh (agardh 1824) genus begonia l. (linnaeus 1753) section symbegonia begonia yapenensis m.hughes sp. nov. urn:lsid:ipni.org:names:77145852-1 figs 1–2 type indonesia. cultivated collection 20 aug. 2014, barber sbar86 (holo-: bo; iso-: e, man). cultivated in the royal botanic garden edinburgh from vegetative material collected in the wild (accession 20090830: indonesia, papua province, yapen island, ambaidiru, 1000 m, 18 feb. 2009, argent, barber, ensoll & galloway abeg211). description sprawling much-branched caulescent herb to 20 cm high. stem green, becoming woody at the base, internodes 3–5 cm long, pilose with 3 mm long white hairs. stipules persistent, glabrous, narrowly triangular, caudate, 15–18 × 4–6 mm. leaves: petiole ca. 5 mm, pilose; lamina lanceolate, asymmetric, 8–10 × 2.5–4 cm, midrib 7–9 cm, basifixed, base cordate, lobes not overlapping; upper surface rugose, green with purple-red veins, shortly hispid between veins, hairs ca. 1 mm; underside paler than upper surface, shortly hispid on veins only; venation pinnate palmate; margin biserrate; apex acute-attenuate. inflorescence terminal, total length 3–5 cm, cymose, compressed at first and becoming more elongate at maturity, unisexual or bisexual, protogynous, female flowers solitary or in pairs, basal, male flowers ca. 10–15; primary peduncle 3–13 mm, shortly hispid; secondary peduncles shorter and glabrous; bracts 7–11 × 2–4 mm, linear-lanceolate, apex acute, white, glabrous. male flower: pedicel 4–12 mm, white, shortly hispid; tepals 2, rhombic-ovate, fused just under half way, 10–13 × 8–10 mm, white, shortly hispid at base, base bulbous, apex acute; androecium with 6–9 stamens, basal 5 subsessile, remainder on a thick 1mm long column; filaments 0.5 mm long, white; anthers 1 mm long, burgundy, ellipsoid, pollen white. female flower: pedicel 4–7 mm, pale green, shortly hispid, bracteoles present; ovary whitish green, total size 10 × 19 mm including wings, wings 3, subequal, triangular, up to 10 mm long, margin hispid; capsule ellipsoid, 8 × 5 mm, sparsely hispid, placentae 2 per locule; tepals 5, corolla tubular, ca. 18 × 8 mm, petals fused for ca. ¾ of the length, white, shortly hispid, hairs denser near the base, lobes ca. 5 mm long, apex acute; stigmas 3, on a 1 mm style, length 7 mm, forked for ⅔ of the length, spirally twisted twice, pale yellow. fruit on a stiff ca. 5 mm pedicel, total size including wings 11 × 21 mm. http://www.ipni.org/urn:lsid:ipni.org:names:77145852-1 hughes m. et al., a new symbegonia from papua, indonesia 3 fig. 1. begonia yapenensis m.hughes sp. nov., cultivated specimen at the royal botanic garden edinburgh, accession 20090830. a. whole plant showing spreading habit (scale bar = 5 cm). b. female flower and ovary (left, corolla dissected; right, corolla entire) (scale bar = 1 cm). c. cross section of ovary showing three locules with bilamellate placentae (scale bar = 1 cm). d. stigmas (scale bar = 5 mm). e. male flower (bottom, corolla dissected; upper, corolla entire; scale bar = 10 cm). european journal of taxonomy 119: 1–6 (2015) 4 distribution indonesia. only known from the type locality in central yapen island, papua province (fig. 2). habitat collected in the forest around ambaidiru village, away from disturbed areas on relatively unspoilt steep mountain sides with orange clay soil and large limestone boulders, near the top of the mountain at ca. 1000 m, in an agathis grove showing signs of tapping for resin on the trunks. understory vegetation in the area consists of marattia sw., angiopteris hoffm., laportea gaudich., zingiberaceae martinov, aeschynanthus jack, begonia, cyrtandra j.r.forst. & g.forst., pandanus l.f., davallia sm. and selaginella p.beauv. notes begonia yapenensis sp. nov. appears to be closest to begonia sympapuana (merr. & l.m.perry) l.l.forrest & hollingsw. (merrill & perry 1943: 59; forrest & hollingsworth 2003: 208), which shares the rugose leaves with reddish veins and broadly similar inflorescences and flowers. begonia yapenensis sp. nov. differs in being a smaller, lower growing plant with shorter internodes and a white undumentum (not reddish), the leaves having shorter petioles (ca. 5 mm, not 1–1.5 cm), male flowers with tepals fused halfway (not shortly fused at the base) and fewer stamens (6–9, not ca. 15) which are arranged along a short column (not arising from a short torus), and fruits which have more attenuate wings. plants of b. yapenensis sp. nov. in cultivation in deep shade show a blue iridescence of the upper leaf surface. conservation status data deficient (dd; iucn 2012). the conservation status of b. yapenensis sp. nov. is not known. the distribution and habitat information for this species is based on just one collection from ambaidiru fig. 2. map of tanah papua, indonesia, showing the distribution of begonia yapenensis sp. nov. on yapen island, and of the allied begonia sympapuana in the mountains at the head of the idenburg river. hughes m. et al., a new symbegonia from papua, indonesia 5 village in the highlands of yapen island, papua, indonesia. although the forest around ambaidiru village is part of the central yapen natural reserve, the establishment of road access from south to north across the island within the protected area, as well as the expansion of ambaidiru village, may affect the population of this begonia. more population and distribution data are required to fully assess the conservation status of this species. discussion although known only from a single collection, the authors are confident in the novelty of b. yapenensis sp. nov. as it differs not only in habit, but also in androecium morphology from the most similar member of begonia sect. symbegonia, b. sympapuana. in cultivation, more careful measurement of b. yapenensis sp. nov. shows it to grow to 15(–20) cm in height, with a horizontal spreading habit. the habit of b. sympapuana is difficult to discern as no living material is available, but the original description gives the height as 30–50 cm, and it has much longer internodes consistent with an erect habit. the other species in papua province also differ considerably from b. yapenensis sp. nov.: b. arfakensis (gibbs) l.l.forest & hollingsw. (gibbs 1917: 149; forrest & hollingsworth 2003: 208) and b. symparvifolia (gibbs) l.l.forest & hollingsw. (gibbs 1917: 150; forrest & hollingsworth 2003: 208) have much smaller and more symmetric leaves; b. symgeraniifolia (ridl.) l.l.forest & hollingsw. (ridley 1916: 61; forrest & hollingsworth 2003: 208) has highly dissected leaves; b. pulchra (ridl.) l.l.forest & hollingsw. (ridley 1916: 62; forrest & hollingsworth 2003: 208) has glossy leaves which are glabrous above, and b. symhirta (ridl.) l.l.forest & hollingsw. (ridley 1916: 61; forrest & hollingsworth 2003: 208) has broadly ovate, subentire leaves. acknowledgements the new taxon was collected on yapen island during the papua expedition of the royal botanic garden edinburgh in 2009, which sb and cdh were involved. sb and cdh would like thank kementerian riset dan teknologi republik indonesia, lembaga ilmu pengetahuan indonesia, kebun raya cibodas, balai penelitian kehutanan manokwari, bksda papua i sub seksi pulau yapen, authorities in serui, people and chief of ambaidiru village, george argent, andrew ensoll, louise galloway and herkilaus rumaikewi. financial support for the expedition was provided by the royal horticultural society, the merlin trust, royal botanic garden edinburgh members’ committee, and the james & eve bennett charitable trust. references agardh c.a. 1824. aphorismi botanici. literis berlingianis, lund. agardh c.a., holmberg l.p. & lundstrom p.m. 1825. classes plantarum. literis berlingianis, lund. beentje h.j. 2010. the kew plant glossary: an illustrated dictionary of plant terms. royal botanic gardens, kew, richmond, surrey. berchtold b.v. von & presl j.s. 1820. o přirozenosti rostlin, aneb rostlinár, obsahugjcj: gedánj on žiwobytj rostlinném pro sebe a z ohledu giných žiwotů, podlé stawu nyněgssjho znánj; k rozssjřenj přirodnictwj; w potaženj na užitečnost w rolnictwj, hospodářstwj, řemeslech, uměnj i obchodu a w wztahowánj obzwlásstnjm na lekařstwj. enders, prague. candolle a.l.p.p. de 1859. mémoire sur la famille des bégoniacées. annales des sciences naturelles. botanique, sér 4 11: 93–149. forrest l.l. & hollingsworth p.m. 2003. a recircumscription of begonia based on nuclear ribosomal sequences. plant systematics and evolution 241 (3–4): 193–211. http://dx.doi.org/10.1007/s00606-0020033-y http://dx.doi.org/10.1007/s00606-002-0033-y http://dx.doi.org/10.1007/s00606-002-0033-y european journal of taxonomy 119: 1–6 (2015) 6 gibbs l.s. 1917. a contribution to the phytogeography and flora of the arfak mountains /by l.s. gibbs. taylor & francis, london. http://dx.doi.org/10.5962/bhl.title.894 hughes m. & hollingsworth p.m. 2008. population genetic divergence corresponds with species-level biodiversity patterns in the large genus begonia. molecular ecology 17 (11): 2643–2651. http://dx.doi. org/10.1111/j.1365-294x.2008.03788.x iucn 2012. iucn red list categories and criteria: version 3.1. second edition. iucn, gland, switzerland and cambridge, uk. klotzsch j.f. 1854. b. monatsberichte der königlich preussischen akademie der wissenschaften zu berlin 1854: 119–128. linnaeus c. 1753. caroli linnaei ... species plantarum :exhibentes plantas rite cognitas, ad genera relatas, cum differentiis specificis, nominibus trivialibus, synonymis selectis, locis natalibus, secundum systema sexuale digestas... laurentii salvii, stockholm. merrill e.d. & perry l.m. 1943. plantae papuanae archboldianae xi. journal of the arnold arboretum 24: 34–59. ridley h.n. 1916. report on the botany of the wollaston expedition to dutch new guinea, 1912–1913. transactions of the linnean society of london, 2nd series: botany 9 (1): 1–269. sands m.j.s. 2009. the begonias of new guinea – an overview. blumea biodiversity, evolution and biogeography of plants 54 (1): 272–277. http://dx.doi.org/10.3767/000651909x476274 takhtajan a.l. 1967. sistema i filogeniia tsvetkovykh rastenii (systema et phylogenia magnoliophytorum). soviet science press, moscow. tebbitt m.c. & maciver c.m. 1999. the systematic significance of the endothecium in begoniaceae. botanical journal of the linnean society 131 (3): 203–221. http://dx.doi.org/10.1111/j.1095-8339.1999. tb00765.x warburg o. 1894. begoniaceae. in: a. engler & prantle k. (eds) die natürlichen pflanzenfamilien. teil 3. abt. 6a: 121–150. englemann, leipzig. manuscript received: 19 september 2014 manuscript accepted: 27 february 2015 published on: 10 april 2015 topic editor: thomas janssen desk editor: natacha beau printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark. http://dx.doi.org/10.5962/bhl.title.894 http://dx.doi.org/10.1111/j.1365-294x.2008.03788.x http://dx.doi.org/10.1111/j.1365-294x.2008.03788.x http://dx.doi.org/10.3767/000651909x476274 http://dx.doi.org/10.1111/j.1095-8339.1999.tb00765.x http://dx.doi.org/10.1111/j.1095-8339.1999.tb00765.x two new species of willowsia (collembola: entomobryidae) from yunnan province, china rui chai 1 & yitong ma 2,* 1,2 school of life science, nantong university, nantong 226000, p.r. china. * corresponding author: mayitong@ntu.edu.cn 1 email: chairui2015@126.com 1 urn:lsid:zoobank.org:author:0702f470-4c20-4044-a29f-57a35d38fc29 2 urn:lsid:zoobank.org:author:14f92f90-863a-46eb-b056-41eb3fa341a0 abstract. we describe two new species of willowsia: w. baoshanensis sp. nov. and w. zhaotongensis sp. nov. this is the first report of willowsia from yunnan province, china. w. baoshanensis sp. nov. is mainly characterized by its chaetotaxy, w. zhaotongensis sp. nov. by its color pattern and chaetotaxy. keywords. entomobryinae, taxonomy, chaetotaxy, scales. chai r. & yitong m. 2017. two new species of willowsia (collembola: entomobryidae) from yunnan province, china. european journal of taxonomy 311: 1–12. https://doi.org/10.5852/ejt.2017.311 introduction the genus willowsia was erected by shoebotham (1917) to accomodate the species seira nigromaculata lubbock, 1873 from england, mainly based on its pointed scales. its main generic characters include mucro bidentation and the presence of a basal spine, eyes 8+8, dental spine absent. scale morphology and distribution are important in taxonomy. scales may be absent on the antennae, the legs and the manubrium in some species and are always absent on dens. thirty species have been described from all over the world and their habitats are from arctic regions to tropical areas. thirteen species were described or reported from china. here, two new species from yunnan, southwest china, are described and a key to the chinese species of the genus is given. material and methods specimens were mounted under a coverslip in marc andré ii solution and were subsequently studied with a leica dm2500 microscope. photographs were taken with a mounted leica dfc300 fx digital camera and enhanced with photoshop cs2 (adobe inc.). the nomenclature of the dorsal chaetotaxy of head and interocular chaetae is described following jordana & baquero (2005), szeptycki (1979) and mari-mutt (1986). labial chaetae are designated following gisin (1967). tergal chaetae of the body are designated using the system of szeptycki (1979). morphology of scales is described following zhang, chen & deharveng (2011). european journal of taxonomy 311: 1–12 issn 2118-9773 https://doi.org/10.5852/ejt.2017.311 www.europeanjournaloftaxonomy.eu 2017 · chai r. & ma y. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:bf7a05cc-65a0-41e6-91e9-2c10fd5ec2df 1 mailto:mayitong%40ntu.edu.cn?subject= mailto:chairui2015%40126.com?subject= http://zoobank.org/urn:lsid:zoobank.org:author:0702f470-4c20-4044-a29f-57a35d38fc29 http://zoobank.org/urn:lsid:zoobank.org:author:14f92f90-863a-46eb-b056-41eb3fa341a0 https://doi.org/10.5852/ejt.2017.311 https://doi.org/10.5852/ejt.2017.311 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:bf7a05cc-65a0-41e6-91e9-2c10fd5ec2df abbreviations: abd. = abdominal segment ant. = antennal segment mac = macrochaeta(e) th. = thoracic segment results class collembola lubbock, 1873 order entomobryomorpha börner, 1913 family entomobryidae tömösvary, 1882 subfamily entomobryinae schäffer, 1896 genus willowsia shoebotham, 1917 diagnosis moderate size, usually 1−2 mm; eyes 8+8; four segmented antennae; mucro bidentate and with a basal spine; dentes without spine; various scale types in different species and absent on dens. willowsia baoshanensis sp. nov. urn:lsid:zoobank.org:act:a334fcb0-054e-4527-bc0b-f61a322cd9dc figs 1−3; table 1 diagnosis eyepatches and lateral margin of th. ii and iii with a little blue pigment; abd. i with 4+4 mac; abd. ii usually with 4+4 dorso-central mac; abd. iii with 2+2 dorso-central and 3+3 lateral mac; spinulate type scales present on ant. i, head, terga, legs, ventral tube and ventral side of manubrium. etymology the specific epithet refers to the type locality: baoshan city. fig. 1. willowsia baoshanensis sp. nov. a−c. habitus (a, c = paratypes; b = holotype) scale bars: 500 μm. european journal of taxonomy 311: 1–12 (2017) 2 http://zoobank.org/urn:lsid:zoobank.org:act:a334fcb0-054e-4527-bc0b-f61a322cd9dc type material holotype china: ♀, on slide, collected in guanjia village, xinjie town, longyang district, baoshan city, yunnan province, rotten leaves of bamboo, 10 aug. 2014, xinnan jiang leg. (collection number: 1148, nantong university). paratypes china: 5 ♀♀ on slides, 2 ♀♀ in alcohol, same data as holotype. description colour pattern. body length up to 1.9 mm. ground color pale yellow. eyepatch dark blue. ant. iv and tibiotarsus with blue pigment. lateral margin of th. ii and iii also with a little blue pigment (fig. 1a−c). head. antenna 0.43−0.57 times as long as body. ratios of length of antennal segments i:ii:iii:iv = 1:1.6−2.0:1.6−2.4:2.5−4.1. distal part of ant. iv with many sensory setae, normal ciliate setae and apical bulb bilobed (fig. 2a). dorsal cephalic chaetotaxy with 6−7 antennal (an), 5 (4) median (m1−m4, an additional seta between m2 and m3 rarely absent) and 8 sutural (s0−s5, s4i, s5i) mac. interocular area with p, s, t setae. eyes 8+8, g and h smaller (fig. 2b). labral setae as 4/5, 5, 4, all slender; prelabral setae ciliate, other smooth; distal margin of labrum with 4 papillae, each with 1 denticle (fig. 2c). lateral process of labial palp straight, as thick as normal setae, with tip not reaching apex of labial papilla (fig. 2d). labial triangle setae as in fig. 2e, all finely ciliate. thorax. dorsal macrochaetae shown as in fig. 2f. th. ii with 2 (m1, m2) medio-median, 2 (m4, m4i) medio-lateral and 14−18 posterior mac on each side, 2 specialized setae (s-setae) on antero-lateral margin not clearly seen. th. iii with 13−15 median and 7 (m5, a6, a6i, p5, p6, m6, m6e) lateral mac on each side. trochanteral organ with 26−30 smooth spiny setae (fig. 2g). unguis with 4 inner teeth, one pair located 0.38−0.39 from base of inner edge of unguis, distal unpaired two respectively at 0.68−0.71 and 0.84−0.89 distance from base. unguiculus acuminate and outer edge serrate. tenent hair thick with clavate tip, almost equal length to inner side of unguis (fig. 2h). abdomen. abd. iv 4.2−8.0 times as long as abd. iii along dorsal midline. dorsal mac shown in figs 2i and 3a. abd. i with 4 (m2, m3, m4, m4p) mac. abd. ii with 4 (a2, m3, m3e, m3ep) (rarely 5, a3 present), 1 (m5) lateral mac and 2 s-setae. abd. iii with 2 (a2, m3) dorso-central, 3 (am6, pm6, p6) characters w. baoshanensis sp. nov. w. guangxiensis w. shi labial triangle setae m1m2rel1l2 m1m2m3rrsel1l2 mrel1l2 scales on legs present present absent central macrochaetae on abd. ii 4 (a2, m3, m3e, m3ep, rarely 5, a3 present) 5 (a2, a3, m3ep, m3e, m3) 5 (a2, m3ea, m3ep, m3e, m3) central macrochaetae on abd. iii 2 (a2, m3) 2 (a2, m3) 3 (a2, a3, m3) lateral macrochaetae on abd. iii 3 (am6, pm6, p6) 4 (am6, pm6, p6, m7) 5 (am6, pm6, p6, m7, p7) ungual inner teeth 4 3 4 table 1. comparison of w. baoshanensis sp. nov., w. guangxiensis shi & chen, 2004 and w. shi pan, zhang & chen, 2006. chai r. & ma y., two new species of willowsia from china 3 fig. 2. willowsia baoshanensis sp. nov. a. apex of ant. iv. b. dorsal chaetotaxy of head. c. labrum. d. lateral process of labial palp. e. labium. f. chaetotaxy of th. ii−iii. g. trochanteral organ. h. hind foot complex. i. chaetotaxy of abd. i−iii. scale bars: a, c–e, g–h = 15 μm; b, f, i = 50 μm. european journal of taxonomy 311: 1–12 (2017) 4 fig. 3. willowsia baoshanensis sp. nov. a. chaetotaxy of abd. iv. b. anterior face of ventral tube. c. posterior face of ventral tube. d. lateral flap of ventral tube. e. plaque of manubrium. f. distal part of dens and mucro. g. scale. scale bars: a = 50 μm; b–g = 15 μm. chai r. & ma y., two new species of willowsia from china 5 lateral mac and 3 s-setae (fig. 2i). abd. iv with 11−18 central (rarely 7, a3, a4, a6, b3, b4, b5, b6 always present) and 15−17 lateral mac as in fig. 3a. ventral tube anteriorly with 3 large and about 9 small ciliate setae (fig. 3b), posteriorly with 2 apical smooth setae and about 36 ciliate setae of different sizes (fig. 3c), laterally with 6 smooth and 8 ciliate setae of different sizes (fig. 3d). manubrial plaque with 8−9 ciliate setae and 3 pseudopores on each side (fig. 3e). dens without spines, uncrenulated dens 1.5−1.7 times as long as mucro. mucro bidentate, two teeth subequal and tip of basal spine reaching subapical tooth (fig. 3f). scales. all scales spinulate type and present on ant. i, head, body, legs and ventral tube, ventral side of manubrium (fig. 3g). remarks the new species can be distinguished from other species of the genus in its color pattern. it is very similar to chinese species w. guangxiensis shi & chen, 2004 and w. shi pan, zhang & chen, 2006 in scales and chaetotaxy on abd. i, but their color pattern, labial triangle setae, chaetotaxy on abd. ii and iii and ungual inner teeth are different as shown in table 1. willowsia zhaotongensis sp. nov. urn:lsid:zoobank.org:act:c1d8d5e5-3bf2-4726-b875-2b729f0db073 figs 4−6; table 2 diagnosis dorsal part of abd. ii & iii and median and posterior parts of abd. iv central with blue pigments; abd. i usually with 4+4 mac; abd. ii with 4−5 + 4−5 dorso-central mac; abd. iii with 3+3 dorso-central and 5+5 lateral mac; spinulate type scales present on ant. i, head, body, legs, ventral tube and ventral side of manubrium. etymology the specific epithet refers to the type locality: zhaotong city. fig. 4. willowsia zhaotongensis sp. nov. a−b. habitus (a = paratype; b = holotype). scale bars: 500 μm. european journal of taxonomy 311: 1–12 (2017) 6 http://zoobank.org/urn:lsid:zoobank.org:act:c1d8d5e5-3bf2-4726-b875-2b729f0db073 type material holotype china: ♀, on slide, collected in yuanlong village, yongfeng town, zhaoyang district, zhaotong city, yunnan province, rotten stems of maize, 1 aug. 2014, fudong ding leg. (collection number 1149, nantong university). paratypes china: 7 ♀♀ on slides, 4 ♀♀ in alcohol, same data as holotype. description colour pattern. body length up to 2.3 mm. eyepatch dark blue. dorsal part of abd. ii & iii and median and posterior parts of abd. iv centrally with blue pigment. coxa and distal part of ant. iv weakly pigmented (fig. 4a–b). head. antenna 0.45−0.56 times as long as body. ratios of length of antennal segments i:ii:iii:iv = 1:1.6−2.0:1.3−1.7:2.3−3.1. distal part of ant. iv with many sensory setae, normal ciliate setae and apical bulb biolobed (fig. 5a). dorsal cephalic chaetotaxy with 7 antennal (an), 4 median (m1−m4,) and 9 sutural (s0−s6, s4i, s5i) mac. interocular area with p, s, t setae. eyes 8+8, g and h smaller (fig. 5b). labral setae as 4/5, 5, 4, all slender; prelabral setae ciliate, other smooth; distal margin of labrum with 4 papillae, each with 1 denticle (fig. 5c). lateral process of labial palp straight, as thick as normal setae, with tip not reaching apex of labial papilla (fig. 5d). labial triangle setae as in fig. 5e, all finely ciliate. thorax. dorsal macrochaetae shown as in fig. 5f. th. ii with 2 (m1, m2) (rarely 3, m2i present) mediomedian, 3 (m4, m4i, m4p) medio-lateral and 16−18 posterior mac on each side, 2 s-setae present on medio-lateral margin. th. iii with 16−17 median and 6 (a6, a6i, p5, p6, m6, m6e) lateral mac on each side. trochanteral organ with 42−49 smooth spiny setae (fig. 5g). unguis with 4 inner teeth, one pair located 0.45−0.50 from base of inner edge of unguis, distal unpaired two respectively at 0.70−0.74 and 0.85−0.88 distance from base. unguiculus acuminate and outer edge serrate. tenent hair thick with clavate tip, slightly longer than inner side of unguis (fig. 5h). abdomen. abd. iv 2.8−5.0 times as long as abd. iii along dorsal midline. dorsal mac shown in figs 6a & 6b. abd. i with 4 (m2, m3, m4, m4p) (rarely 3, m4p absent) mac. abd. ii with 4−5 (a2, a3, m3, m3e, m3ep, a3 sometimes absent), 1(m5) lateral mac and 2 sensory chaetae. abd. iii with 3 (a2, a3, m3) dorso-central, 5 (am6, pm6, p6, m7a, p7) lateral mac and 3 sensory chaetae (fig. 6a). abd. iv with 10−12 central (a3, a4, a6, b3, b4, b5, b6 always present) and 17−20 lateral mac as in fig. 6b. ventral tube anteriorly with 3 large and 15−23 small ciliate setae (fig. 6c), posteriorly with 2 apical smooth setae and about 35 ciliate setae of different sizes (fig. 6d); lateral flap not clearly seen. manubrial plaque with characters w. zhaotongensis sp. nov. w. bartkei a1, a2 & a3 on th. iii absent present m3ep on abd. ii present absent a3 on abd. iii present absent ae2, ae3 & ae6 on abd. iv present absent scales on antennae and furcula present absent uncrenulated dens to mucro in length 6−7 4 table 2. comparison of w. zhaotongensis sp. nov. and w. bartkei stach, 1965. chai r. & ma y., two new species of willowsia from china 7 fig. 5. willowsia zhaotongensis sp. nov. a. apex of ant. iv. b. dorsal chaetotaxy of head. c. labrum. d. lateral process of labial palp. e. labium. f. chaetotaxy of th. ii−iii. g. trochanteral organ. h. hind foot complex. scale bars: a, c–e, g–h = 15 μm; b, f = 50 μm. european journal of taxonomy 311: 1–12 (2017) 8 fig. 6. willowsia zhaotongensis sp. nov. a. chaetotaxy of abd. i−iii. b. chaetotaxy of abd. iv. c. anterior face of ventral tube. d. posterior face of ventral tube. e. plaque of manubrium. f. distal part of dens and mucro. g. scale. scale bars: a–b = 50 μm; c–g = 15 μm. chai r. & ma y., two new species of willowsia from china 9 8−11 ciliate setae and 3(4) pseudopores on each side (fig. 6e). dens without spines, uncrenulated dens 1.4−1.7 times as long as mucro. mucro bidentate, two teeth subequal and tip of basal spine reaching subapical tooth (fig. 6f). scales. all scales spinulate type and present on ant. i, head, body, legs and ventral tube, ventral side of manubrium (fig. 6g). remarks the new species can easily be distinguished from the other species of the genus by their color pattern, except for the vietnamese species willowsia bartkei stach, 1965. both new species have blue pigment on the dorsal part of abd. ii and iii and on the median and posterior parts of abd. iv centrally, but there is no blue pigment on abd. i posteriorly and abd. ii and iii laterally in the former, whereas pigment is present in the latter. the other differences, such as chaetotaxy and scales, are listed in table 2. key to the chinese species of the genus willowsia 1. abd. ii with 2+2 mac ………………………………………………………………………………2 – abd. ii with at lest 3+3 central mac ………………………………………………………………3 2. abd. iii with 1+1central mac ……………………………………………w. formosana denis, 1929 – abd. iii with 2+2 central mac …………………………………………w. jacobsoni (börner, 1913) 3. abd. ii with 3+3 mac ………………………………………………………………………………4 – abd. ii with at lest 4+4 mac ………………………………………………………………………11 4. abd. i with 4 +4 mac ……………………………………w. qui zhang, chen & deharveng, 2011 – abd. i at most 3+3 mac ……………………………………………………………………………5 5. abd. iii with 2+2 central mac ……………………………………………………………………6 – abd. iii with 3+3 central mac ……………………………………………………………………9 6. scales present on antennae ……………………w. guangdongensis zhang, xu & chen, 2007 – scales absent on antennae …………………………………………………………………………7 7. body scales of short rib type …………………………………………w. japonica (folsom, 1897) – body scales of long basal rib type …………………………………………………………………8 8. abd. iv with 5+5 central mac ………………………………w. pseudobuskii pan & zhang, 2016 – abd. iv with 7+7 central mac ………………………………w. pseudoplatani zhang & pan, 2016 9. abd. ii without blue transverse band ……………………………w. nigromaculata (lubbock, 1873) – abd. ii with a blue transverse band …………………………………………………………………10 10. metathorax blue pigmented, cephalic s0 absent ……………………w. similis pan & zhang, 2016 – metathorax unpigmented, cephalic s0 present ………………………w. fascia zhang & pan, 2016 11. abd. iii with 2+2 central mac ……………………………………………………………………12 – abd. iii with 3+3 central mac ……………………………………………………………………13 12. abd. iii with 3+3 lateral mac …………………………………………w. baoshanensis sp. nov. – abd. iii with 4+4 lateral mac ………………………………w. guangxiensis shi & chen, 2004 european journal of taxonomy 311: 1–12 (2017) 10 13. scales absent on antennae and manubrium ……w. yiningensis zhang, chen & deharveng, 2011 – scales present on antennae and manubrium ……………………………………………………14 14. abd. ii–iii with blue transverse bands …………………………………w. zhaotongensis sp. nov. – abd. ii–iii with scattered blue pigment and irregular patches ……w. shi pan, zhang & chen, 2006 discussion entomobryidae is the largest family of collembola in china and about 140 species have been reported. willowsia, a genus of the subfamily entomobryinae which includes about 120 chinese species, contains relatively few species with only 13 species reported from china prior to this study. two new species of the genus were added here: w. baoshanensis sp. nov. and w. zhaotongensis sp. nov., and it is the first report of the genus from yunnan province, southwest china. there are different climate zones in yunnan, including northern tropical, subtropical, temperate and highland climate zones suggesting a rich biodiversity might be expected. however, due to the lack of research conducted in the area so far, there are only a few records of collembola from the province. acknowledgements we thank xinnan jiang and fudong ding who collected the specimens. references gisin h. 1967. espèces nouvelles et lignées évolutives de pseudosinella endogés. memórias e estudos do museu zoológico da universidade de coimbra 301: 5–25. jordana r. & baquero e. 2005. a proposal of characters for taxonomic identification of entomobrya species (collembola, entomobryomorpha), with description of a new species. abhandlungen und berichte des naturkundemuseums goerlitz 76 (2): 117−134. mari-mutt j.a. 1986. puerto rican species of lepidocyrtus and pseudosinella (collembola: entomobryidae). caribbean journal of science 22 (1−2): 1−48. pan z.x. & zhang f. 2016. contribution to the willowsia species having body scales of long basal rib type: four new species and a redescription of w. qui (collembola: entomobryidae). european journal of taxonomy 245: 1–25. https://doi.org/10.5852/ejt.2016.245 pan j.l., zhang f. & chen j.x. 2006. a new chinese species of the genus willowsia from tibet (collembola: entomobryidae). journal of the kansas entomological society 79 (3): 261−266. https:// doi.org/10.2317/0511.22.1 schäffer c. 1896. die collembolen der umgebung von hamburg und benachtbaren gebiete. mittheilungen aus dem naturhistorischen museum 13: 147−216. available from http://biodiversitylibrary.org/ page/29521571 [accessed 15 mar. 2017] shi x. & chen j.x. 2004. the genus willowsia (collembola: entomobryidae) and species from china. entomotaxonomia 26 (4): 241−248. shoebotham j.w. 1917. notes on the collembola, part 4. the classification of the collembola; with a list of genera known to occur in the british isles. annals and magazine of natural history series 8, 19: 425−436. available from http://biodiversitylibrary.org/page/22130994 [accessed 15 mar. 1017] szeptycki a. 1979. morpho-systematic studies on collembola. iv. chaetotaxy of the entomobryidae and its phylogenetical significance. polska akademia nauk, kraków. chai r. & ma y., two new species of willowsia from china 11 https://doi.org/10.5852/ejt.2016.245 https://doi.org/10.2317/0511.22.1 https://doi.org/10.2317/0511.22.1 http://biodiversitylibrary.org/page/29521571 http://biodiversitylibrary.org/page/29521571 http://biodiversitylibrary.org/page/22130994 zhang f., chen j.x. & deharveng l. 2011. new insight into the systematics of the willowsia complex (collembola: entomobryidae). annales de la société entomologique de france 47 (1−2): 1−20. https:// doi.org/10.1080/00379271.2011.10697692 manuscript received: 9 may 2016 manuscript accepted: 24 august 2016 published on: 20 april 2017 topic editor: gavin broad desk editor: kristiaan hoedemakers printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands. european journal of taxonomy 311: 1–12 (2017) 12 https://doi.org/10.1080/00379271.2011.10697692 https://doi.org/10.1080/00379271.2011.10697692 1 european journal of taxonomy 657: 1–14 issn 2118-9773 https://doi.org/10.5852/ejt.2020.657 www.europeanjournaloftaxonomy.eu 2020 · mateo-ramírez l. & riina r. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e croton calcareus: a new species of dragon’s blood (euphorbiaceae) from dry forest in the state of chiapas, mexico leticia mateo-ramírez 1 & ricarda riina 2,* 1 department of botany, faculty of biology, university of salamanca, licenciado méndez nieto ave., 37007, salamanca, spain. 2 real jardín botánico, rjb-csic, plaza de murillo 2, 28014, madrid, spain. * corresponding author: rriina@rjb.csic.es 1 email: leticiamr13@usal.es abstract. we describe croton calcareus riina & mateo-ram. sp. nov., a new species in croton section cyclostigma (dragon’s blood trees) from the state of chiapas (mexico). this species is a small tree growing in dry forest on calcareous substrates. both morphological and molecular data support c. calcareus sp. nov. as a new species closely related to c. redolens, another dry forest taxon from northern venezuela. we provide illustrations, a distribution map and suggestions for species conservation status. the new species along with croton draco are the only known representatives of c. section cyclostigma occurring in mexico. keywords. croton section cyclostigma, dry forest, euphorbiaceae, neotropics, phylogeny. mateo-ramírez l. & riina r. 2020. croton calcareus: a new species of dragon’s blood (euphorbiaceae) from dry forest in the state of chiapas, mexico. european journal of taxonomy 657: 1–14. https://doi.org/10.5852/ejt.2020.657 introduction croton l. species popularly known as dragon’s blood (‘sangre de drago’ or ‘sangre de grado’ in spanish) belong to c. sect. cyclostigma griseb. this section is an arborescent clade of 48 species occurring mainly in lowland and montane forests along the andes (riina et al. 2009; van ee et al. 2011) and the atlantic rain forest region in south-eastern brazil (santos et al. 2017; farias et al. 2019). in the northern neotropics, only one species in the section, croton draco schltdl., was previously known to reach mexico. as part of a revision of the common and widespread croton draco, which occurs from mexico to panama, we noticed several specimens from mexico that had obvious morphological differences with c. draco as well as differences in habitat. further examination of these specimens revealed an undescribed taxon, which we assign here to c. sect. cyclostigma based on its morphological affinities with species in this section and support from molecular phylogenetic evidence. members of sect. cyclostigma have laticifers that produce conspicuous reddish latex (wiedenhoeft et al. 2009; feio et al. 2018a) that is widely used for its medicinal properties in several latin american https://doi.org/10.5852/ejt.2020.657 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ mailto:rriina%40rjb.csic.es?subject= mailto:leticiamr13%40usal.es?subject= https://doi.org/10.5852/ejt.2020.657 european journal of taxonomy 657: 1–14 (2020) 2 countries (jones 2003; salatino et al. 2007). species from sect. cyclostigma occur predominantly in mesic forests (riina et al. 2009; arévalo et al. 2017); however, a small number of species are restricted to seasonally dry forest vegetation in different areas of the neotropics, such as c. churutensis riina & cornejo (riina et al. 2007) in ecuador, c. tumbesinus riina (feio et al. 2018b) in peru, c. redolens pittier in northern venezuela and c. charaguensis standl. in bolivia. in this paper, we describe and illustrate a new species from the state of chiapas (mexico), which adds to the list of dry forest specialists of this diverse and medicinally important neotropical croton clade. material and methods we examined collections of the new taxon and c. draco deposited in bm, dav, ma, mexu, mich and mo. acronyms of herbaria follow index herbariorum (thiers, continuously updated). for morphological descriptions we used a 10−60 × magnification stereo microscope. morphological terminology follows recent taxonomic papers on croton (e.g., santos et al. 2017; feio et al. 2018b; riina et al. 2018), and anatomical studies on trichomes and secretory structures (webster et al. 1996; vitarelli et al. 2015; feio et al. 2016; feio et al. 2018a). for the phylogenetic analysis, we used a subset of the nuclear rdna internal transcribed spacers (its) dataset from riina et al. (2009), including species of sect. cyclostigma sampled there, a selection of other croton sections and one outgroup (brasiliocroton mamoninha p.e.berry & cordeiro). to this dataset, we added two newly generated sequences of the new species, c. calcareus riina & mateoram. sp. nov., and two sequences of c. draco from genbank (https://www.ncbi.nlm.nih.gov/genbank/). genbank accession numbers of all the sequences used in the analysis are shown in table 1. dna extraction, amplification and sequencing of its followed the same laboratory procedures as in previous croton phylogenetic studies (riina et al. 2009; van ee et al. 2011). sequences were aligned manually using mesquite ver. 3.2 (maddison & maddison 2017). the aligned matrix, in fasta format, is provided as supplementary file 1. the phylogenetic analysis was performed in mrbayes 3.2.6 (ronquist et al. 2012) using the gtr+i+g substitution model as estimated with jmodeltest 2.1.6 (darriba et al. 2012) using the akaike information criterion (aic). the analysis included two runs of 10 million generations in mrbayes, sampling trees every 1000 generations. one-fourth (25%) of the mcmc samples from each run was discarded as ̒ burninʼ. tracer ver. 1.7.1 (rambaut et al. 2014) was used to determine run convergence and stationarity as indicated by the effective sample size (ess) values, which should be higher than 200. node posterior probability (pp) values were obtained by computing a 50% majority rule consensus of the post-burn-in trees from both mcmc chains using the sumt command. we used figtree ver. 1.3.1 (rambaut 2006) to visualize and edit the consensus tree. we ran a preliminary analysis with all the cyclostigma accessions from riina et al. (2009) to assess the phylogenetic position of c. calcareus sp. nov. within the section (data not shown). based on this preliminary phylogeny, we reduced the sampling within the cyclostigma clade to exclude taxa distantly related from c. calcareus sp. nov. (i.e., all the brazilian species from the atlantic rain forest and most species from the south-central andes). we ran the final bayesian analysis on the reduced dataset (table 1) following the steps indicated above. the species distribution map was produced using r software (https://www.r-project.org) and the dismo package (hijmans et al. 2010). only two of the seven available collections had geographic coordinates on their labels, so we estimated coordinates for the remainder with the locality information using google earth. the proposed conservation status was based on the criteria of the international union for the conservation of nature guidelines (iucn 2012). extent of occurrence (eoo) was assessed using geocat (bachman et al. 2011). https://www.ncbi.nlm.nih.gov/genbank/ https://europeanjournaloftaxonomy.eu/index.php/ejt/article/downloadsuppfile/967/105 https://www.r-project.org mateo-ramírez l. & riina r., a new dragon’s blood croton from mexican dry forest 3 table 1. list of genbank accession numbers of the its sequences used to provide a phylogenetic context to the new species (shown in fig. 1). brasiliocroton mamoninha p.e.berry & cordeiro eu586944 croton arboreus millsp. eu478029 c. beetlei croizat eu586916 c. calcareus riina & mateo-ram. sp. nov.; hampshire et al. 1161; mexico: chiapas mt106663 c. calcareus riina & mateo-ram. sp. nov.; webster et al. 12966; mexico: chiapas mt106664 c. caracasanus pittier dq227525 c. chilensis müll.arg. eu586905 c. conduplicatus kunth eu477957 c. coriaceus kunth eu586921 c. cuneatus klotzsch eu497735 c. cupreatus croizat eu586919 c. curiosus croizat eu586906 c. discolor willd. eu497736 c. draco schltdl. & cham. 1 eu478006 c. draco 2 ef421776 c. echioides baill. eu586907 c. eichleri müll.arg. eu586949 c. emporiorum croizat eu586908 c. gossypiifolius vahl eu586924 c. goudotii baill. eu586946 c. gratissimus burch. (as c. zambesicus in genbank ) ay971260 c. hibiscifolius kunth ex spreng. eu586925 c. lundellii standl. ef421735 c. niveus jacq. eu478046 c. pachypodus g.l.webster (as c. megistocarpus in genbank) eu586953 c. priscus croizat eu586950 c. pungens jacq. ay971241 c. redolens pittier eu586935 c. rimbachii croizat eu586936 c. saltensis griseb. eu586911 c. sampatik müll.arg. ef421792 c. sapiifolius müll.arg. ef421754 c. schiedeanus schltdl. ay971246 c. speciosus müll.arg. ay971251 c. suberosus kunth eu477979 c. verapazensis donn.sm. ef421750 c. xalapensis kunth eu477987 c. yavitensis croizat eu586918 european journal of taxonomy 657: 1–14 (2020) 4 results class magnoliopsida brongn. order malpighiales mart. family euphorbiaceae juss. genus croton l. croton calcareus riina & mateo-ram. sp. nov. urn:lsid:ipni.org:names:77209678-1 figs 2–4 diagnosis croton calcareus riina & mateo-ram. sp. nov. is most closely related to c. redolens pittier and can be distinguished from the latter mainly by its larger fruits (12–13 × 12–15 mm in c. calcareus sp. nov. vs 5–6 × 7–8 mm in c. redolens) and an indumentum of sessile, stellate, multiradiate, porrect trichomes on the adaxial side of leaves (vs stipitate, stellate, porrect trichomes in c. redolens). the new species fig. 1. bayesian majority rule consensus phylogram of nuclear its data from a selection of croton l. taxa including two accessions of croton calcareus riina & mateo-ram. sp. nov. species marked with an asterisk correspond to those with which c. calcareus sp. nov. has been previously confused. the relevant clades are labelled on the left side. numbers next to nodes indicate clade support (bayesian posterior probabilities). http://www.ipni.org/urn:lsid:ipni.org:names:77209678-1 mateo-ramírez l. & riina r., a new dragon’s blood croton from mexican dry forest 5 has been confused with c. draco schltdl., another dragon’s blood from mexico, but it differs from this species in having shorter inflorescences (8–10 cm vs 15–43 cm in c. draco) and larger fruits (only 4–7 × 4–7 mm in c. draco). etymology the specific epithet refers to the calcareous substrates where the species occurs. type material mexico • chiapas, mpio, la trinitaria, 4 km e of la trinitaria along mex 190; 16°8′ n, 92°2′ w; alt. 1480 m; 8 jul. 1990; r.j. hampshire, a. reyes garcía and l. hernández 1161; holotype: mexu[858873]; isotypes: bm, ma[540108], mo[4323513]. paratypes mexico – chiapas • mpio, ocozocoautla, steep-walled canyon at the head of the río de la venta at the chorreadero near derna; 16 dec. 1972; d.e. breedlove, r.f. thorne 30301; mexu • el aguacero, 13 km al no de ocozocoautla; 29 mar. 1985; e. cabrera and h. de cabrera 7870; mexu • el aguacero, canyon of the río la venta; alt. 540–660 m; 19 nov. 1984; g. davidse, m. sousa, o. tellez, e. martinez and j. davidse 30083; ma, mo • 13 km e of ocozocoautla on rte 190, then n on road to aguacero; alt. 630–700 m; 2 oct. 1984; m.j. huft, e. cabrera and r. torres 2253; mexu, mo • “cascada el aguacero en el río la venta” [el aguacero waterfall on the river la venta]; 16°45′17″ n, 93°31′20″ w; alt. 600 m; 12 dec. 1987; e. martinez and a. reyes 22018; mexu, mich • el aguacero, “a 33 km al e de cintalapa” [33 km e of cintalapa]; alt. 540–660 m; 19 nov. 1984; o. tellez, m. sousa, e. martinez, g. davidse and j. davidse 8120; mo, mexu • 7 mi sw of tuxtla gutierrez; alt. 3000 ft.; 11 aug. 1962; g.l. webster, k. miller and l. miller 12966; dav, mo. description small monoecious trees, up to 6 m high; young branches with a dense indumentum of rosulate (sometimes also stipitate) trichomes; latex from the main trunk and branches reddish to dark red (observation from herbarium specimens). stipules short-triangular when young (ca 1.5 mm) to filiform 8–13.5 mm long, covered with a dense indumentum of rosulate and multiradiate trichomes, colleters (ovoid glands) at base, margin and apex. leaf blade ovate to broadly ovate, 5–13 × 3.5–11 cm; base rounded to cordate; apex acute to obtuse; margin entire or slightly dentate; colleters (ovoid glands) along the margin; venation 3–5-plinerved, brochidodromous; primary and secondary veins raised on both surfaces; petiolar nectary glands 2–6, patelliform, sessile, subsessile or stipitate, attached to the petiole on the adaxial side, but also sometimes visible from the abaxial side; petiole 1.5–8 cm long, with a more or less dense indumentum of stellate adpressed trichomes; adaxial leaf indumentum very dense on young leaves to nearly glabrous on old leaves, sessile, stellate-porrect and stellate-multiradiate trichomes; abaxial leaf surface very densely pubescent on young leaves to sparser with age, with sessile or stipitate stellate or multiradiate trichomes. inflorescences terminal, erect thyrses 8–10 cm long, axis costate, with a dense indumentum of stellatemultiradiate trichomes; bracts triangular, ca 1 × 1 mm; cymules usually bisexual, regularly spaced at the base and more congested toward the apex. staminate flowers shortly pedicellate, pedicels 1–2 mm long; sepals valvate, triangular-ovate, 2–2.8 × 1.1–2 mm, adaxial surface with scattered simple trichomes, abaxial surface with a dense indumentum of stellate-multiradiate trichomes; petals narrow-elliptical, 1.7–1.8 × 0.7–0.8 mm, with scattered simple or stellate hairs on both surfaces, margin lanate; receptacle densely pilose; stamens 16; filaments glabrous, 2–3 mm long, anthers 0.6–0.9 × 0.4–0.5 mm. pistillate flowers with pedicels 1.5–2 mm long, with stellate-multiradiate trichomes; sepals valvate, triangularovate, 2–2.2 × 1.2–1.3 mm, adaxial and abaxial surfaces with dense covering of stellate-multiradiate trichomes; petals absent or reduced to a filament with a colleter at the tip; ovary subglobose, densely covered with stellate-multiradiate trichomes; styles bifid, with six terminal tips, the lower half of the european journal of taxonomy 657: 1–14 (2020) 6 fig. 2. croton calcareus riina & mateo-ram. sp. nov. a. image of an isotype (ma). b. apical portion of a young branch showing the short young stipule (right arrow) and fully developed stipule (left arrow); the arrows point to the base of each stipule. c. petiolar nectary glands visible from the adaxial side on a leaf with a rounded base. d. petiolar nectary glands visible from the abaxial side on a leaf with a cordate base. e. branch showing a young inflorescence. f. section of inflorescence with detail of male flowers. g. mature fruit. scale bars = 1 mm. vouchers: a, c, e, g: hampshire et al. 1161 (ma); b, d, f: davidse et al. 30083 (ma). mateo-ramírez l. & riina r., a new dragon’s blood croton from mexican dry forest 7 fig. 3. image of one of the paratypes of croton calcareus riina & mateo-ram. sp. nov. (cabrera and cabrera 7870, mexu), previously indentified as c. xalapensis kunth, to show the variation in leaf shape and indumentum density. it also shows immature fruits (younger than those in the type specimens). european journal of taxonomy 657: 1–14 (2020) 8 styles with stellate-multiradiate trichomes. capsules depressed-subglobose, slightly trilobate, 12–13 × 12–15 mm; columella 9–10 mm long; seeds ovoid, 7–8 × 5.5–6 mm, brown, surface smooth, shiny, caruncle trapezoid, 0.6 × 1.2 mm. phylogenetic placement and relationships the its alignment consisted of 686 aligned positions. the resulting tree topology (fig. 1) was congruent with previous analyses of croton using the its region for the clades involved (riina et al. 2009; van ee et al. 2011). this phylogeny indicates that croton calcareus sp. nov. is indeed a member of croton sect. cyclostigma and that it is most closely related to c. redolens and c. gossypiifolius vahl (fig. 1). the two sequences of c. calcareus sp. nov. are identical and share two unique positions with respect to the rest of sequences in the matrix. the accessions of mexican species with which c. calcareus sp. nov. has been confused in the past (i.e., c. draco, c. xalapensis kunth and c. verapazensis donn.sm.) are indicated with an asterisk in the phylogeny (fig. 1). of these, c. draco is the most closely related to the new species, coming out in the same clade (sect. cyclostigma), the other two species belong to two different sections/clades of croton (fig. 1). distribution, ecology and phenology croton calcareus sp. nov. appears to be restricted to the state of chiapas (mexico) where it grows on limestone outcrops, in low deciduous forests on rocky slopes, steep river canyons and river floodplains, fig. 4. distribution map showing the localities (black squares) of croton calcareus riina & mateoram. sp. nov., all known from southern mexico. mateo-ramírez l. & riina r., a new dragon’s blood croton from mexican dry forest 9 between 550–1500 m elevation (fig. 4). flowering between august and february and probably fruiting between march and july. conservation status the species could be categorized as vulnerable (vu d2; iucn 2012). the extent of occurrence of croton calcareus sp. nov. is only 2718 km2 and it is only known from four different localities in two notadjacent municipalities. the species appears to be under low human pressure since three of the known localities (municipality of ocozocoautla) are within the buffer zone of the selva el ocote biosphere reserve (ochoa-gaona 1996; unesco 2012). most of the known localities are of limited access due to their vicinity to limestone rocky formations along rivers. discussion our phylogenetic analysis supports croton calcareus sp. nov. as a distinct lineage in c. sect. cyclostigma sensu van ee et al. (2011), which, combined with the observed distinct morphological features, warrants recognition at the species level. based on the its phylogeny and morphology, the new species is most closely related to c. redolens, a dendroid shrub (up to 3 m tall) occurring in dry forest vegetation associated with rocky outcrops in northern venezuela. croton calcareus sp. nov. also belongs to the same clade that includes c. draco, the only other species in sect. cyclostigma known to occur in mexico. characters shared among c. calcareus sp. nov., c. draco, c. redolens and other members of sect. cyclostigma include, but are not limited to, arborescent habit, indumentum of stellate trichomes, red latex, conspicuous stipules, petiolar nectary glands, bisexual cymules at the lower portion of the inflorescence axis, bifid styles and more than 15 stamens. for this reason, it is not surprising that specimens of c. calcareus sp. nov. have been confused with c. draco in the past. among the many morphological differences between c. calcareus sp. nov. and both c. redolens and c. draco (table 2), the most obvious ones are fruit size and inflorescence length; although the latter is only relevant in the case of c. draco. inflorescences are much longer in c. draco and fruits are much larger in c. calcareus sp. nov. (table 2). unfortunately, only one of the examined collections (hampshire et al. 1161, the type) had fully mature fruits and seeds. plant stature and habitat preference are also important features setting the two mexican species ( c. draco and c. calcareus sp. nov.) apart (table 2). there are also differences, although with some overlap, in the leaves among the three species in question. in croton calcareus sp. nov. leaves can have a rounded or cordate base (fig. 1c–d) and acute to obtuse apex, whereas in c. redolens the base is mostly rounded and rarely cordate and the apex is acute, and in c. draco the base of the leaves is usually cordate and the apex acuminate (table 2). in addition, the texture of the adaxial leaf surface is soft to touch in c. draco and c. redolens, while in c. calcareus sp. nov. it has a rough texture. stipules are usually informative distinguishing species within sect. cyclostigma (riina et al. 2009; farias et al. 2019). in croton calcareus sp. nov., there is variation in the shape of stipules depending on their state of development. when young, stipules in c. calcareus sp. nov. are triangular in shape and they become more linear and much longer as they grow (fig. 1b), whereas in c. redolens and c. draco young and old stipules have more or less the same shape (table 2). croton calcareus sp. nov. occurs in a diverse and relatively poorly explored area of mexico in which species from different plant groups continue to be described (e.g., martínez-meléndez et al. 2014; hirai et al. 2014; ruiz-sánchez 2015; ortiz-díaz et al. 2016; steinmann & carrillo-reyes 2018; areces-berazain & vega-lopez 2019). several specimens of c. calcareus sp. nov. have previously been misidentified in herbaria as c. draco, c. xalapensis or c. verapazensis. besides c. draco, which belongs to the same section as the new species, the other two species belong to different sections or clades (fig. 1). croton xalapensis is a shrub in c. sect. adenophylli griseb., the largest section of european journal of taxonomy 657: 1–14 (2020) 10 croton and the sister clade of sect. cyclostigma (van ee et al. 2011). it is also one of the few sections of croton with a distinct synapomorphy, i.e., the presence of three inflated lobes at the apex of the fruit columella (riina et al. 2009; van ee et al. 2011). so, c. xalapensis can be easily distinguished from c. calcareus sp. nov. by its shrubby habit and its trilobed columella apex. croton verapazensis, from central america, is superficially similar to c. calcareus sp. nov. and it also grows in dry forest. however, it is member of a different clade of neotropical croton, namely c. sect. corylocroton g.l.webster, and can be distinguished from any member of sect. cyclostigma, including c. calcareus sp. nov., mainly by its unique petiolar nectary gland morphology (apical part with almost the same diamater as the stipe; in sect. cyclostigma it is much wider than the stipe), presence of stellate-lepidote trichomes, leaf margins serrate to coarsely dentate and presence of unisexual cymules at the base of the inflorescence axis. finally, as in c. draco, both c. xalapensis and c. verapazensis also have much longer mature inflorescences and smaller capsules and seeds than c. calcareus sp. nov. acknowledgements we thank the staff of herbarium ma for their support as well as david s. gernandt and ubaldo melo samper palacios from mexu for providing information and images of specimens. we are grateful to montserrat martínez ortega (universidad de salamanca), alberto herrero (rjb-csic) and mercedes feature croton calcareus sp. nov. croton draco croton redolens stipules short triangular to long filiform subulate to broadly subulate short filiform leaf size (max. cm) 5–13 × 3.5–11 9.9–22.9 × 6.6–17.7 7.8–8.3 × 4.8–6.3 leaf base rounded or cordate cordate, rarely rounded rounded, rarely cordate leaf apex acute to obtuse acuminate acute adaxial leaf texture rough soft soft adaxial leaf trichomes sessile, stellate porrect and stellate multiradiate stipitate, stellate multiradiate stipitate, stellate porrect adaxial leaf indument sparse to glabrescent dense to very dense dense inflorescence length (cm) 8–10 14.6–42.5 5.7–15.6 fruit size (mm) 12–13 ×2–15 4–7 × 4–7 5–6 × 7–8 seed surface smooth rugose slightly rugose plant stature (max. height, m) 6 25 3 habitat dry forest moist forest dry forest distribution restricted to chiapas widespread (mexico to panama) northern venezuela table 2. main differences between croton calcareus riina & mateo-ram. sp. nov., c. redolens pittier and c. draco l. 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received: 13 january 2020 manuscript accepted: 6 march 2020 published on: 2 june 2020 topic editor: frederik leliaert desk editor: radka rosenbaumová printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. https://doi.org/10.3732/ajb.1500017 https://doi.org/10.1111/j.1095-8339.1996.tb00744.x https://doi.org/10.1163/22941932-90000209 european journal of taxonomy 657: 1–14 (2020) 14 supplementary material supplementary file 1 its alignment (fasta file) used in phylogenetic analysis including the newly generated sequences of croton calcareus riina & mateo-ram. sp. nov. https://europeanjournaloftaxonomy.eu/index.php/ejt/article/downloadsuppfile/967/105 european journal of taxonomy 870: 76–86 issn 2118-9773 https://doi.org/10.5852/ejt.2023.870.2119 www.europeanjournaloftaxonomy.eu 2023 · razzaq a. et al. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e 76 inocybe subhimalayanensis (agaricales, inocybaceae), a new smooth spored inocybe species from pakistan annum razzaq1,*, arooj naseer1,2 & abdul nasir khalid1,3 1fungal biology and systematics research laboratory, institute of botany, university of the punjab, lahore, pakistan. *corresponding author: annum.phd.botany@pu.edu.pk 2email: arooj.hons@pu.edu.pk 3email: drankhalid@gmail.com abstract. during mycological explorations, a new smooth spored species, inocybe subhimalayanensis razzaq, naseer & khalid sp. nov. was collected from moist temperate sub-himalayan region, pakistan. phylogeny of its and lsu regions of nrdna, and morphoanatomical data make it distinct from other known species of the genus. the taxon is characterized by: a yellowish orange to brown pileus with prominent fibrillose, prominent umbo; ellipsoid to amygdaliform smooth larger basidiospores (8.4‒)8.6‒12.2(‒12.6) × (4.9‒)5.1‒7.1(‒7.3) µm; and lack of velipellis. molecular phylogenetic analyses further support the recognition of the new species. keywords. inocybaceae, its, lsu, punjab, taxonomy. razzaq a., naseer a. & khalid a.n. 2023. inocybe subhimalayanensis (agaricales, inocybaceae), a new smooth spored inocybe species from pakistan. european journal of taxonomy 870: 76–86. https://doi.org/10.5852/ejt.2023.870.2119 introduction the inocybaceae is one of the most diverse families of agaricales with approximately more than 1050 species worldwide (matheny et al. 2020). matheny et al. (2020) performed a six-loci phylogenetic analysis of the inocybaceae and classified the family into seven genera; auritella matheny & bougher ex matheny & bougher, inocybe (fr.) fr., inosperma (kühner) matheny & esteve-rav., mallocybe (kuyper) matheny, vizzini, esteve-rav., nothocybe matheny & k.p.d. latha, pseudosperma matheny & esteve-rav., and tubariomyces esteve-rav. & matheny. among these, inocybe is the most speciose genus represented by 850 species worldwide (matheny et al. 2020; bandini et al. 2022) and has a cosmopolitan distribution. more than 200 new species of the genus have been reported during the last five years, the number is still growing as a result of new discoveries (crous et al. 2017; matheny & bougher 2017; bandini et al. 2021; dovana et al. 2021; mešić et al. 2021; muñoz et al. 2022). inocybe plays a significant role in ectomycorrhizal communities of temperate and boreal forests (cripps et al. 2010; kokkonen & vauras 2012). this genus is characterized by the presence of cheilocystidia and generally also pleurocystidia with more or less thick walls, whereas this is not the case with the other six genera (matheny et al. 2020). https://doi.org/10.5852/ejt.2023.870.2119 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ mailto:annum.phd.botany%40pu.edu.pk?subject= mailto:arooj.hons%40pu.edu.pk?subject= mailto:drankhalid%40gmail.com?subject= https://doi.org/10.5852/ejt.2023.870.2119 razzaq a. et al., a new species of inocybe from pakistan 77 the himalayan moist temperate forests of pakistan are included in one of the twenty-five hotspots for biodiversity (myers et al. 2000). these forests are floristically rich, covered with evergreen vegetation and home to a number of plant species, therefore, an equally high diversity of fungal species is expected. however, only 33 species of this diverse genus have been reported from pakistan (naseer et al. 2019; jabeen & khalid 2020; saba et al. 2020; khan et al. 2022). during the course of a macrofungal survey from different parts of himalayan forests of pakistan, a novel species of inocybe was discovered. in this paper, we present our collections as a new species with brief diagnosis, an extensive description, and phylogenetic analyses based on its and lsu sequences data. materials and methods study area the basidiomata have been collected from himalayan temperate forests of bhurban during the rainy seasons (september 2020‒2021). bhurban is a small town of the punjab province of pakistan (33°56′26.24″ n to 73°26′59.48″ e). this area is a part of the sub-himalayan tract with an approximate elevation of 1800‒2200 m a.s.l. the area receives precipitation during the entire year with a mean annual rainfall of about 1800 mm (30 year period: world weather information service 2018). the climate is humid and variable due to a broad range of elevation and the vegetation is mainly comprised of pinus wallichiana a.b.jacks., olea ferruginea royle, quercus leucotrichophora a.camus and cedrus deodara (roxb. ex d.don) g.don providing a perfect environment for the growth of mycorrhizal fungi (hameed et al. 2012). sampling and morphoanatomical studies the specimens were photographed in the field using a nikon d70s digital camera and morphological features of fresh basidiomata including size, shape and color of pileus, stipe, lamellae, presence or absence of annulus and volva were noted. color codes were given according to munsell (1975). the specimens were dried by using an electric fan heater (ecostar geh-800) and packed in labeled polythene bags. samples were submitted in the lahore (lah) herbarium, institute of botany, university of the punjab, quaid-e-azam campus, lahore, pakistan. anatomical structures were studied with a trinocular olympus ch30 microscope and measurements were taken with the help of calibrated piximètre software attached to the microscope with a digital camera (hdce-x5) under 400× and oil immersion 1000× magnification. for microscopic analyses, 5% koh was used to rehydrate the tissues (pileus, gills and stipe) of the basidiomata and congo red (1%) was used as stain to clarify microscopic features. the abbreviation n/m/p specifies n basidiospores from m basidiomata of p collections. measurements for basidiospores are taken as length × width (l × w). the average length and average width of all the microscopic characters were measured and given as avl × avw. extreme values are given in parenthesis, q value shows l/w ratio of the spore (bas 1969; yu et al. 2020) and avq means average q of all basidiospores (liang & yang 2011). molecular and phylogenetic analyses for molecular analyses, genomic dna was extracted using modified ctab method (gardes & bruns 1993). for the qualitative analysis of extracted dna, gel electrophoresis was done by using 1% agarose gel (voytas 2001). the extracted dna was amplified using polymerase chain reaction by a thermocycler (bio-rad t100tm, bio-rad laboratories pte. ltd., jurong east, singapore). two genetic markers (its and lsu) were amplified in this study. in order to amplify its region, primer pairs its1f and its4 were used (gardes & bruns 1993) while lr0r and lr5 primer pairs were used for the amplification of lsu region (moncalvo et al. 2000). sequencing of the pcr products were done by tsingke, china. a total of six its sequences and two lsu sequences were generated. european journal of taxonomy 870: 76–86 (2023) 78 consensus sequences were generated using bioedit software ver. 7.0.4.1 (hall 1999). these were blast searched at ncbi (https://www.ncbi.nlm.nih.gov/) and closely related inocybe sequences were retrieved from genbank. these sequences were aligned with the sequences generated in this study using the online muscle tool at embl-ebi (http://www.ebi.ac.uk/). for phylogenetic analyses, maximum likelihood (ml) bootstrap analyses were performed in mega ver. 11 (tamura et al. 2021). results molecular phylogenetic analyses the aligned its dataset, including the results of blast searching against genbank and the published work of bandini et al. 2022, consisted of 43 nucleotide sequences including i. spuria jacobsson & e. larss. as an outgroup (bandini et al. 2022). muscle ver. 3.8 (edgar 2004) was used for the alignment of the sequences and bioedit ver. 7.2.5. (hall 1999) was used for manual adjustment. the its dataset’s ml analysis was done using mega 11. in both its and lsu based phylograms, i. subhimalayanensis razzaq, naseer & khalid sp. nov. get separated from other species with a strong bootstrap. taxonomy phylum basidiomycota r.t.moore class agaricomycetes doweld order agaricales underw. family inocybaceae jülich genus inocybe (fr.) fr. inocybe subhimalayanensis razzaq, naseer & khalid sp. nov. mycobank mb 845094 figs 1–2, table 1 diagnosis the new species differs from inocybe parvicystis f.j.rodr.-campo & esteve-rav. by the combination of these characters: prominent umbo, larger spores up to 9.9 μm (vs 6.5 μm), hymenal cystidia with noncrystalliferous apex, non-crystalliferous caulocystidia and absence of velipellis. etymology the epithet ‘subhimalayanensis’ is referring to the sub-himalayan region where the type specimen was collected. type material holotype pakistan • punjab province, rawalpindi district, bhurban town; 33.9554° n, 73.4519° e; alt. 1828 m a.s.l.; on soil below pinus; 7 sep. 2020; annum razzaq, gb-18; genbank nos on810645 (nrits), on810652 (lsu); lah[37437]. additional material examined pakistan • punjab province, rawalpindi district, bhurban town; 33.9554° n, 73.4519° e; alt. 1828 m a.s.l.; on soil below pinus; 7 sep. 2020; annum razzaq, gb-26; genbank nos on810643 (nrits), on810653 (lsu); lah[37438] • same data as for preceding; 10 sep. 2021; annum razzaq, gc-35; genbank no on810644 (nrits); lah[37439] • kashmiri bazar; 10 sep. 2021; annum razzaq, br88; genbank no on810646 (nrits); lah[37440] • same data as for preceding; 4 sep. 2021; annum https://www.ncbi.nlm.nih.gov/ http://www.ebi.ac.uk/ https://www.mycobank.org/page/name%20details%20page/845094 razzaq a. et al., a new species of inocybe from pakistan 79 fig. 1. macromorphological characters of inocybe subhimalayanensis razzaq, naseer & khalid sp. nov. a. lah37438. b. lah37439. c‒d. lah37437 (holotype). european journal of taxonomy 870: 76–86 (2023) 80 fig. 2. inocybe subhimalayanensis razzaq, naseer & khalid sp. nov. holotype (lah37437). a. basidiospores. b. pleurocystidia. c. cheilocystidia. d. basidia. e. caulocystidia. f. stipitipellis. g. pileipellis. razzaq a. et al., a new species of inocybe from pakistan 81 razzaq, br-22; genbank no on810647 (nrits); lah[37441] • same data as for preceding; 12 sep. 2020; annum razzaq, br-57; genbank no on810648 (nrits); lah[37442] • murree; 33.9070° n, 73.3943° e; alt. 2291 m a.s.l.; 18 aug. 2019; arooj naseer an-36; genbank no on911331 (nrits); lah[151676]. description pileus 15–25 mm wide, (sub)campanulate to subumbonate when immature becoming broadly convex or expanded at maturity, with prominent large umbo; margin decurved at young stage later becoming straight, incised to eroded, concolorous, pileus depressed near the umbo; prominent dark brown radial striations at pileal surface extended up to the central umbonate region; color yellowish orange (10yr 7/8) to orange (7.5yr 8/8) when fresh becoming yellowish brown (7.5yr 6/8) to brown hue (7.5yr 5/8) when old. hymenophore regular, close, adnexed, ventricose, whitish at first becoming brownish with age; edge entire. stipe 21–35 × 2–4 mm, cylindrical or curved, fine whitish tomentum when young, later becoming longitudinally striate or glabrous, yellowish orange (10yr 7/8) to brown (7.5yr 5/8) colored. annulus and volva absent. context thin. habit pluteoid. odor unspecific. taste not recorded. basidiospores [100/3/3] (8.4‒)8.6‒12.2(‒12.6) × (4.9‒)5.1‒7.1(‒7.3) µm, avl × avw = 9.9 × 5.9 µm, q = 1.62‒1.73, avq = 1.69, smooth, (sub)amygdaloid with rounded base and rounded to acute apex in side view, oblong to ovoid with rounded to (sub)acute apex in frontal view, often with explicit suprahilar depression, prominent hilar appendix, moderately thick-walled, guttulated, pale to dark brown in 5% koh and water. basidia (24.8‒)26.2–35.2(‒38.7) × (12.2‒)13.3–19.2(‒19.7) µm, avl × avw = 29.8 × 8.4 µm, thin-walled, hyaline in 5% koh, tetrasterigmate, broadly clavate, content present. pleurocystidia (35.2‒)37.2–49.4(‒49.6) × (7.7‒)8.1–12.6(‒14.2) µm, avl × avw = 42.6 × 16.6 µm, clavate or (sub)clavate to broadly clavate or (sub)cylindrical or fusiform, metuloidal,thick walled, at apex generally wide, apex non crystalliferous. cheilocystidia (28.3‒)32.3–48.3(‒49.4) × (13.1‒)14.2–22.4(‒23.6) µm, avl × avw = 37.2 × 16.9 µm, similar to pleurocystidia in size and shape. pileipellis a cutis of parallel hyphae 3.5–11.5 µm wide, subcutis with wider and paler to colorless hyphae. stipitipellis a cutis of parallel hyphae, 1.73‒16.2 µm thick (av = 7.6 µm), septate, branched. caulocystidia only near the apex of the stipe, (56.2‒)58.4–97.8(‒101.1) × (17.7‒)19.2–35.7(‒37.6) µm, avl × avw = 78.1 × 23.4 µm, long and narrow subcylindrical to (sub)utriform; intermixed. habitat found from nutrient-rich soil, under pinus wallichiana (pinaceae). distribution this species is reported for the first time from sub-himalayan forests of punjab, pakistan. discussion in this study, a new species of inocybe collected from punjab, pakistan, is identified on the bases of its morpho-anatomical features and phylogenetic analyses of its and lsu sequences dataset (fig. 3). the small sized basidiomata, campanulate to convex yellowish orange to brown pileus with radial striations, prominent large umbo, smooth and amygdaloid to oblong basidiospores, presence of pleurocystidia and caulocystidia and presence of clamp connections allocate this species in the genus inocybe (figs 1–2). the its sequences of i. subhimalayanensis sp. nov. showed 99.8% similarity with an unpublished soil ecm sequence from pakistan (genbank accession numbers kf679813) (fig. 4). in phylogenetic analyses of the its region, the sequence i. subhimalayanensis sp. nov. form a clade in strong supported sister relationship to i. parvicystis (ky349121, ky349127, ky349122, ky349120, ky349126), a species described from comunidad de madrid (spain), but separated with strong bootstrap value european journal of taxonomy 870: 76–86 (2023) 82 fig. 3. molecular phylogenetic analysis of lsu sequences of inocybe subhimalayanensis sp. nov. inferred by using the maximum likelihood method. sequences generated from local collection are marked with bullets. table 1 comparison of the diagnostic characters of i. subhimalayanensis razzaq, naseer & khalid sp. nov., with phylogenetically closed species. characters i. subhimalayanensis i. parvicystis i. mystica i. dvaliniana i. cryptocystis pileus size 15–25 15–40 up to 30 up to 20 up to 35 (mm) pileus surface fibrillose surface smooth, finely fibrous smooth to finely smooth and compact radially fibrillose felted and fibrillose with finely scaly at margin umbo prominent large not or hardly present at prominent prominent present early stages velipellis absent present – present present (only in young basidiomata) spores 8.4‒12.6 × 7.5–11.5 × up to about 7.3–10.4 × 7–8 × 5 µm 4.9‒7.3 µm 4.5–6. μm 9(–9.5) µm 4.7–5.9 µm caulocystidia non-crystalliferous, crystalliferous, – – numerous at the top at apex of stipe accompanied by paracystidia razzaq a. et al., a new species of inocybe from pakistan 83 fig. 4. molecular phylogenetic analysis of its sequences of inocybe subhimalayanensis sp. nov. inferred by using the maximum likelihood method. sequences generated from local collection are marked with bullets. (100) from i. parvicystis (fig. 4). inocybe subhimalayanensis sp. nov. differs morphoanatomically in having smaller pileus, 15–25 mm wide (vs 15–40 mm wide) with prominent dark brown fibrillose (vs surface smooth, radially fibrillose at margin), prominent large umbo (vs absent or hardly present), and absence of velipellis (vs present), caulocystidia with non-crystalliferous apex (vs crystalliferous apex caulocystidia) (esteve-raventós et al. 2017). another closely related taxon is i. dvaliniana bandini & b.oertel, which also has the cystidia with noncrystalliferous apex but differs from the new taxon by mottled straw-colored to honey-colored pileus, european journal of taxonomy 870: 76–86 (2023) 84 presence of whitish velipellis and relatively short hymenial cystidia (17–42 × 5–11 µm, av. 28 × 7 µm) (https://www.inocybe.org) whereas our new taxon has prominent dark brown radial striations on the pileus, absence of velipellis and large sized cheilocystidia (28‒49 × 13‒23 µm, av. 37 × 16 µm). the two lineage forming taxa i. mystica stangl & glowinski and i. cryptocystis d.e.stuntz are close to the new taxon. inocybe mystica differs in having relatively large size basidiospores (up to 9–9.5 µm), short cheilocystidia with crystalliferous apex (https://www.inocybe.org) and i. cryptocystis shows clear morphoanatomical differences with i. subhimalayanensis sp. nov. in having smooth and compact pileal surface, and a partial white veil at young stage (murrill et al. 1932). a comparison of diagnostic characters of phylogenetically close species is given in table 1. himalayan forests of pakistan exhibit large altitudinal variation, with climatic conditions and a diverse vegetation that supports a diverse and conspicuous lichen biota. the nature reserves have abundant biological resources, it is expected that more new species of macrofungi may be discovered in the future. acknowledgements we are thankful to ms. rubab khurshid (fungal biology and systematics research laboratory, 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meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; leibniz institute for the analysis of biodiversity change, bonn – hamburg, germany; national museum of the czech republic, prague, czech republic. https://doi.org/10.11646/phytotaxa.450.1.2 european journal of taxonomy 336: 1–14 issn 2118-9773 https://doi.org/10.5852/ejt.2017.336 www.europeanjournaloftaxonomy.eu 2017 · csuzdi c. et al. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:a61daf10-1819-4c90-a64a-1f1dddac220d 1 three new species of kynotus from the central highlands of madagascar (clitellata, megadrili) csaba csuzdi 1, malalatiana razafindrakoto 2 & yong hong 3,* 1 department of zoology, eszterházy károly university, eger, hungary. 2 department of animal biology, university of antananarivo, madagascar and laboratory of radio-isotopes, university of antananarivo, madagascar. 3 department of agricultural biology, college of agriculture & life sciences, chonbuk national university, jeonju, republic of korea. * corresponding author: geoworm@hanmail.net 1 email: csuzdi.csaba@ektf.hu 2 email: malalasraz@yahoo.fr 1 urn:lsid:zoobank.org:author:86f75d9d-740d-4d98-849b-b744c4438d76 2 urn:lsid:zoobank.org:author:f21c5094-de1f-4f7e-bf0a-c59c8aaf7d7a 3 urn:lsid:zoobank.org:author:49f2d6fb-b6ba-438c-a452-ed4c109db544 abstract. the earthworm fauna of madagascar is scarcely known. a recently launched exploration of the soil fauna (“global change and soil macrofauna diversity in madagascar”) resulted in the discovery of six new earthworm species belonging to the malagasy endemic family kynotidae. the success of the collecting campaign carried out between 2008 and 2011 inspired a new exploration of the earthworm fauna across the central highland region of the island in the spring of 2015. during this expedition, two new species of kynotus, k. ankisiranus sp. nov. and k. voimmanus sp. nov., were discovered. barcoding of the recently collected species of kynotus revealed that the unpigmented worms referred previously to k. alaotranus michaelsen, 1897 also represented a new, still undescribed species, k. blancharti sp. nov. keywords. oligochaeta, malagasy region, endemism, kynotidae, barcoding, coi. csuzdi c., razafindrakoto m. & hong y. 2017. three new species of kynotus from the central highlands of madagascar (clitellata, megadrili). european journal of taxonomy 336: 1–14. https://doi.org/10.5852/ejt.2017.336 introduction madagascar, with its territory of 592,800 km2, is the fourth largest island in the world and, due to its tectonic history and environmental variability, is listed among the 25 megadiverse regions of the earth (myres et al. 2000). this is why the island’s fauna has always been the focus of research (goodman & benstead 2003). interestingly though, data on the earthworm fauna of madagascar is quite scarce; since the description of the first earthworm species geophagus darwini (=kynotus darwini) by keller (1887) just a couple of papers dealing with the earthworm fauna of madagascar (e.g., michaelsen 1891, 1897, 1907, 1931; rosa 1892) have been published. after the last paper by michaelsen (1931), no new https://doi.org/10.5852/ejt.2017.336 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:a61daf10-1819-4c90-a64a-1f1dddac220d mailto:geoworm%40hanmail.net?subject= mailto:csuzdi.csaba%40ektf.hu?subject= mailto:malalasraz%40yahoo.fr?subject= http://zoobank.org/urn:lsid:zoobank.org:author:86f75d9d-740d-4d98-849b-b744c4438d76 http://zoobank.org/urn:lsid:zoobank.org:author:f21c5094-de1f-4f7e-bf0a-c59c8aaf7d7a http://zoobank.org/urn:lsid:zoobank.org:author:49f2d6fb-b6ba-438c-a452-ed4c109db544 https://doi.org/10.5852/ejt.2017.336 european journal of taxonomy 336: 1–14 (2017) 2 taxonomic works on the earthworm fauna of the island were published until a new project, entitled “changement global et diversité de la macrofaune du sol à madagascar” (global change and soil macrofauna diversity in madagascar), was launched in 2008. during this project, a comprehensive collecting campaign resulted in the discovery of several rare, as well as new, earthworm species (razafindrakoto et al. 2010, 2011, 2016; csuzdi et al. 2012). these results inspired new collecting activities carried out by the latter authors, which resulted in the discovery of two further species new to science, belonging to the endemic family kynotidae. barcoding of the recently collected material of kynotus revealed that the unpigmented specimens, previously relegated to the red-pigmented k. alaotranus michaelsen, 1907 (csuzdi et al. 2012), also represent a new species. descriptions of the three new species of kynotus, together with their barcodes, are herewith provided. material and methods study area the new collecting sites are situated in the central highlands region of the island (fig. 1). fig. 1. main soil types of madagascar, with indication of the sampling localities. 1 = moramanga district, andasibe, voimma community park; 2 = antsirabe district, sambaina; 3 = ambalavao district, mt ankisira national park. csuzdi c. et al., three new kynotus species from madagascar 3 methods earthworms were collected primarily by using the diluted formaldehyde method (raw 1959) supplemented by digging and hand-sorting. the material collected was killed in 75% ethanol and fixed in 4% formaldehyde solution. parallel material from each morpho-species was conserved in 96% ethanol for dna studies. deposition the material collected is deposited in the hungarian natural history museum (hnhm) and in the collection of chonbuk national university, korea (chbnu). molecular methods a small piece of the postclitellate body wall was cut out from the paratype specimens for barcoding, as indicated in table 1, and processed according to the methodology described in szederjesi & csuzdi (2015). additional coi sequences of kynotus were acquired from the bold database (table 1). coi sequences were aligned with clustalw (thompson et al. 1994), using the default settings. the final dataset was 671 bp long and contained no internal gaps. maximum likelihood analysis was carried out with mega v. 6.06 (tamura et al. 2013) using the best fitting substitution model gtr g + i and 1000 bootstrap replicates. bayesian inference was performed with beast v. 1.8.2 software (drummond et al. 2012) with the gtr g + i substitution model selected by mega using the akaike information criterion. beast was run for 10 million generations, saving trees at every 1000th generation. the first 2000 trees were discarded “burn in” in treannotator v. 1.8.2. the resulting tree was visualized with fig tree v. 1.4.2 (rambaut 2014). interand intraspecific genetic distances were calculated with mega using the k2p substitution model. results class clitellata michaelsen, 1919 subclass oligochaeta grube, 1850 superorder megadrili benham, 1890 order opisthopora michaelsen, 1929 suborder crassiclitellata jamieson, 1988 family kynotidae jamieson, 1971 genus kynotus michaelsen, 1891 kynotus ankisiranus sp. nov. urn:lsid:zoobank.org:act:ddafa680-a9e8-4a8b-ad38-9fb52f7378f2 figs 2, 5 diagnosis length 70–90 mm, diameter 3 mm. colour grey with slight red pigmentation at the head on dorsum. segments 1–3 simple, 4–10 biannulate. setae appear from segment 2 and clearly visible, male pores ventral on 16. clitellum circular on 18–27, ½28. three rows of spermathecae of elongated oval-shape in 13/14, 14/15, 15/16. no septa notably thickened. genital setal glands in 14, 15. etymology the species epithet refers to the type locality, mt ankisira national park. http://zoobank.org/urn:lsid:zoobank.org:act:ddafa680-a9e8-4a8b-ad38-9fb52f7378f2 european journal of taxonomy 336: 1–14 (2017) 4 material examined holotype madagascar: clitellate, ambalavao district, sendreisoa, namoly, mt ankisira national park, aristida grassland at the edge of primary forest, 22°06′45.8″ s, 46°56′33.8″ e, 1500 m a.s.l., 24 feb. 2015,y. hong and m. razafindrakoto leg. (hnhm/af5658). paratypes madagascar: 2 clitellate, 3 aclitellate adults (hnhm/af5659), 2 aclitellate (chbnu md-06al5), locality and date same as for holotype. species voucher no. locality accession no. k. ankisiranus sp. nov. hnhm/af5659 madagascar, ambalavao, namoly, mt ankisira national park kx527886 k. alaotranus 1 hnhm af/5528 madagascar, vohimana reserve, secondary pandanus forest kx372247 k. alaotranus 2 hnhm af/5528 madagascar, vohimana reserve, secondary pandanus forest kx372248 k. blancharti sp. nov. hnhm/af5539 madagascar, antsirabe district, sambaina kx517831 k. blancharti sp. nov. hnhm/af5539 madagascar, antsirabe district, sambaina kx517832 k. michaelseni 1 hnhm af/5530 madagascar, sambaina, open bush-land kx372243 k. michaelseni 2 hnhm af/5530 madagascar, sambaina, open bush-land kx372244 k. minutus 1 hnhm af/5235 madagascar, vohimana reserve, secondary pandanus forest kx372245 k. minutus 2 hnhm af/5235 madagascar, vohimana reserve, secondary pandanus forest kx372246 k. proboscideus ewsjc848 madagascar, mangalaza ambatondrazaka, near villages jn260829 k. proboscideus ewsjc849 madagascar, mangalaza ambatondrazaka, near villages jn260830 k. proboscideus ewsjc850 madagascar, mangalaza ambatondrazaka, near villages jn260831 k. proboscideus ewsjc851 madagascar, mangalaza ambatondrazaka, near villages jn260832 k. proboscideus ewsjc852 madagascar, mangalaza ambatondrazaka, near villages jn260833 k. proboscideus ewsjc855 madagascar, vohitsoa ambatondrazaka, along the road jn260834 k. proboscideus hnhm af/5512 madagascar, vohitsoa ambatondrazaka, tsarahonenana kx527885 k. sihanakus ewsjc924 madagascar, bemanevika bealanana, grassland jn260884 k. sihanakus ewsjc925 madagascar, bemanevika bealanana, grassland jn260885 k. sihanakus ewsjc927 madagascar, bemanevika bealanana, grassland jn260886 k. sihanakus ewsjc928 madagascar, bemanevika bealanana, grassland jn260887 k. sihanakus ewsjc929 madagascar, bemanevika bealanana, grassland jn260888 k. sihanakus ewsjc930 madagascar, ambatosoratra ambatondrazaka, banana plantation jn260889 k. sihanakus ewsjc931 madagascar, ambatosoratra ambatondrazaka, banana plantation jn260890 k. sihanakus ewsjc939 madagascar, ambatosoratra ambatondrazaka, banana plantation jn260895 k. voimmanus sp. nov. hnhm af/5661 madagascar, moramanga, andasibe, voimma community park kx527887 k. voimmanus sp. nov. hnhm af/5661 madagascar, moramanga, andasibe, voimma community park kx517833 k. voimmanus sp. nov. hnhm af/5661 madagascar, moramanga, andasibe, voimma community park kx517834 k. voimmanus sp. nov. hnhm af/5661 madagascar, moramanga, andasibe, voimma community park kx517835 k. voimmanus sp. nov. hnhm af/5662 madagascar, moramanga, andasibe, voimma community park kx517836 k. voimmanus sp. nov. hnhm af/5662 madagascar, moramanga, andasibe, voimma community park kx517837 k. voimmanus sp. nov. hnhm af/5662 madagascar, moramanga, andasibe, voimma community park kx517838 table 1. localities and coi genbank accession numbers of the specimens of kynotus analysed (ewsjc samples were obtained by sam james). csuzdi c. et al., three new kynotus species from madagascar 5 description holotype 50 mm in length, diameter after clitellum 3 mm, segment number 98, tail missing. paratypes 72–86 mm in length, 3 mm in diameter, segment number 170–179. colour grey with red pigmentation at head on dorsum. head zygolobous, segments 1–3 simple, 4–10 biannulate. dorsal pores lacking. setae small, ab and cd clearly observed from segment 2. setal arrangement (holotype) after clitellum aa:ab:bc:cd:dd = 15:1.25:9.5:1:18, dd = 0.32 u (fig. 2a). clitellum annular on segments 18–27, ½ 28. fig. 2. kynotus ankisiranus sp. nov. a. setal arrangement, a, b, c, d representing setal lines. b. spermathecae, schematic representation. c. genital seta. d. tip of the genital seta. scale bars = 0.1 mm. european journal of taxonomy 336: 1–14 (2017) 6 male pores ventral on 16. female pores not seen. spermathecal pores in 13/14 (2), 14/15 (2), 15/16 (2) from bc to cd. genital setal pores in variable position between ab and cd on 14, 15. internal characters large muscular gizzard in 5. septa all membranous. calciferous glands, lamellae and typhlosolis lacking. dorsal blood vessel simple throughout, last pair of hearts in 11. excretory system holoic, vesiculate. two pairs of testes and sperm funnels in 10, 11 enclosed in large peri-oesophageal testis sacs. seminal vesicles lacking. ovaries in 13. a pair of large, oval copulatory chambers occupying the ventral space of segments 16–17. each copulatory chamber bearing an irregular prostate-like gland (pseudoprostate) reaching segment 21. spermathecae elongated, oval-shaped, with short duct attached at 13/14(2), 14/15(2), 15/16(2) in variable position arranged from between bc to cd (fig. 2b). two pairs of genital setal glands present in segments 14, 15 above ab and around cd respectively. the genital setae are slightly curved, with spoon-shaped apex, length 1.6–1.7 mm, diameter at the middle 0.05 mm, ornamentation dense serrations (fig. 2c–d). remarks the new species belongs to the alaotranus group (table 2) characterized by biannulate segments between 4–10. with its smaller size and red pigmentation on the dorsum, k. ankisiranus sp. nov. most closely resembles k. minutus csuzdi, razafindrakoto & blanchart, 2012, but differs from it by the longer clitellum (18–26 vs 18–27, ½ 28) and the number of genital setal glands (single pair in 15 vs two pairs in 14, 15). k. amkisiranus sp. nov. is clearly different from all the examined alaotranus group’s species by its coi barcode as well (table 3, fig. 5). kynotus blancharti sp. nov. urn:lsid:zoobank.org:act:89ca5809-835d-4698-b6ea-2bdf9178f718 figs 3, 5 kynotus alaotranus michaelsen, 1907 – csuzdi et al. 2012: 37. diagnosis length 80–90 mm, diameter 4–5 mm, segment number 180–195. segments 1–3 simple, 4–10 biannulate. colour light grey, pigmentation lacking. setae appear from segment 2, clearly visible. male pores ventral on 16. clitellum circular on ½18–27. three rows of spermathecae of oval shape, with long and thin duct in 13/14, 14/15, 15/16. genital setal glands present in 14 and 15. etymology we dedicate this species to eric blanchart (l’institut de recherche pour le développement (ird), antananarivo), who initiated the recent earthworm research in madagascar. material examined holotype madagascar: clitellate, antsirabe district, sambaina, open bushland, 19°38′12″ s, 47°8′49″ e, 1700 m a.s.l., 21 apr. 2008, cs. csuzdi, e. blanchart and m. razafindrakoto leg. (hnhm/af5652). paratypes madagascar: 3 clitellate ex., locality and date same as holotype (hnhm/af5539). http://zoobank.org/urn:lsid:zoobank.org:act:89ca5809-835d-4698-b6ea-2bdf9178f718 csuzdi c. et al., three new kynotus species from madagascar 7 ta bl e 2. d is tin gu is hi ng c ha ra ct er s in th e k . a la ot ra nu s sp ec ie s gr ou p. sp ec ie s a ve ra ge s iz e (l en gt h / w id th ) (m m ) p ig m en t b ia nn ul at e se gm en ts f ir st se ta l se gm en t c lit el lu m sp er m at he ca e (n um be r) g en it al se ta l gl an ds g en it al s et ae (m m ) a nd a pe x sh ap e k . a nk is ir an us s p. n ov . 90 /3 sl ig ht re d 4– 10 2 18 –½ 2 8 13 /1 4, 1 4/ 15 , 1 5/ 16 (2 ) o va l, sh or t d uc t 14 , 1 5 1. 7 sp oo nsh ap ed k . a la ot ra nu s m ic ha el se n, 1 90 7 12 0/ 6 sl ig ht re d 4– 10 3 ½ 1 8– 27 13 /1 4, 1 4/ 15 , 1 5/ 16 (2 -4 ) o va l, lo ng d uc t 14 , 1 5 1. 5 la nc eo la te k . b la nc ha rt i s p. n ov . 90 /5 la ck in g 4– 10 2 ½ 1 8– 27 13 /1 4, 1 4/ 15 , 1 5/ 16 (2 -4 ) o va l, lo ng d uc t 14 , 1 5 1. 9 sp oo nsh ap ed k . m ic ha el se ni r os a, 1 89 2 13 0/ 8 da rk re d 4– 10 3 19 –2 5 13 /1 4, 1 4/ 15 (1 ) fi ng er -s ha pe d 13 , 1 4 2. 0 la nc eo la te k . m in ut us c su zd i e t a l., 2 01 2 70 /4 da rk re d 4– 10 2 18 –½ 2 7 13 /1 4, 1 4/ 15 , 1 5/ 16 (2 -3 ) o va l, sh or t d uc t 15 1. 2 la nc eo la te k . p ar vu s c su zd i e t a l., 2 01 2 10 0/ 4 sl ig ht re d 4– 10 2 18 –½ 2 7 13 /1 4, 1 4/ 15 , 1 5/ 16 (2 -3 ) o va l, sh or t d uc t 14 , 1 5 0. 9 la nc eo la te k . p itt ar el lii c og ne tti , 1 90 6 23 0/ 10 la ck in g 4– 10 6 18 –2 8 14 /1 5, 1 5/ 16 (2 -4 ) fi ng er -s ha pe d 14 , 1 6 3. 6 la nc eo la te k . r os ae c og ne tti , 1 90 6 10 5/ 5 sl ig ht re d 4– 10 2 18 –2 8 13 /1 4, 1 4/ 15 , 1 5/ 16 , 1 6/ 17 (1 -4 ) o va l, sh or t d uc t 15 2. 0 la nc eo la te k . s ik or ai m ic ha el se n, 1 90 1 20 5/ 10 da rk re d 4– 10 9 – 13 /1 4, 1 4/ 15 , 1 5/ 16 (1 -2 ) fi ng er -s ha pe d 14 , 1 5 3. 5 la nc eo la te k . v oi m m an us s p. n ov . 17 0/ 7 sl ig ht re d 4– 10 2 18 –2 8 13 /1 4, 1 4/ 15 , 1 5/ 16 (2 -3 ) o va l, lo ng d uc t 15 2. 3 la nc eo la te european journal of taxonomy 336: 1–14 (2017) 8 description holotype 90 mm in length, diameter after the clitellum 5 mm, segment number 194, tail missing. paratypes 80–90 mm in length, 4–5 mm in diameter, segment number 180–195. colour greyish, pigmentation lacking. head zygolobous, segments 1–3 simple, 4–10 biannulate. dorsal pores lacking. setae small, ab and cd clearly observed from segment 2. setal arrangement after clitellum aa:ab:bc:cd:dd = 22.5:1.5:13:1:20, dd = 0.27 u (fig. 3a). clitellum circular on ½18–27. male pores ventral on 16. fig. 3. k. blancharti sp. nov. a. setal arrangement, a, b, c, d representing setal lines. b. spermathecae (14/15) schematic representation. c. genital seta. d. tip of the genital seta. scale bars = 0.1 mm. csuzdi c. et al., three new kynotus species from madagascar 9 female pores not seen. spermathecal pores in 13/14(2–3), 14/15(3–4), 15/16(2–3) irregularly arranged from bc to above cd. genital setal pores in variable position between ab and cd on 14, 15. nephridial pores between setal lines ab and cd. internal characters large muscular gizzard in 5. septa 7/8–9/10 moderately thickened, 5/6–6/7, 10/11 slightly strengthened. calciferous glands, lamellae and typhlosolis lacking. dorsal blood vessel simple throughout, last pair of hearts in 11. excretory system holoic, vesiculate. two pairs of testes and sperm funnels in 10, 11 enclosed in large peri-oesophageal testis sacs. seminal vesicles lacking. ovaries in 13. a pair of large, oval copulatory chambers occupying the ventral space of segments 16–18. each copulatory chamber bearing an irregular prostate-like gland (pseudoprostate) reaching segment 26. spermathecae ovalshaped, with long and thin duct attached at 13/14(2–3), 14/15(3–4), 15/16(2–3), arranged from ab to above cd (fig. 3b). two pairs of genital setal glands present in segment 14, 15 above ab and around cd. the genital setae are slightly curved, with elongated spoon-shaped apex, length 1.8–1.9 mm, diameter just under the apex 0.05 mm, ornamentation with dense short serrations (fig. 3c–d). remarks k. blancharti sp. nov. was previously classified as an unpigmented k. alaotranus (csuzdi et al. 2012). besides the lack of red pigments, the penial setae are also somewhat different. instead of being lanceolate as in alaotranus, in the new species the tip is flanked with slightly dentate rims making the seta more spoon-shaped. k. blancharti sp. nov. is clearly different from all the examined alaotranus group species by its coi barcode as well (table 3, fig. 5). kynotus voimmanus sp. nov. urn:lsid:zoobank.org:act:8998c949-547b-454e-86e5-50b2ef4dbcb0 figs 4–5 diagnosis length 145–170 mm, diameter 6–7 mm. colour greyish with slight reddish hints at the head on dorsum. segments 1–3 simple, 4–10 biannulate. setae appear from segment 2 and clearly visible; male pores ventral on 16. clitellum circular on 18–½28, 28.three rows of spermathecae of oval shape with long and thin duct in 13/14, 14/15, 15/16. septa 6/7–8/9 strongly thickened. genital setal glands in 15. table 3. average interspecific (below the diagonal) and intra specific (bold, in the diagonal) k2p genetic distances of the species of kynotus michaelsen, 1891 analysed. k. ank. k. alao. k. blan. k. mich. k. min. k. prob. k. sih. k. voi. k. ankisiranus – k. alaotranus 0.206 0.0 k. blancharti 0.217 0.197 0.003 k. michaelseni 0.264 0.255 0.256 0.003 k. minutus 0.237 0.221 0.224 0.252 0.0 k. proboscideus 0.256 0.262 0.235 0.282 0.242 0.005 k. sihanakus 0.237 0.243 0.233 0.261 0.215 0.241 0.004 k. voimmanus 0.215 0.143 0.219 0.265 0.262 0.264 0.245 0.001 http://zoobank.org/urn:lsid:zoobank.org:act:8998c949-547b-454e-86e5-50b2ef4dbcb0 european journal of taxonomy 336: 1–14 (2017) 10 etymology the species epithet refers to the type locality voimma community park. material examined holotype madagascar: clitellate, ambalavao district, sendreisoa, namoly, mt ankisira national park, in primary forest, 18°55′38″ s, 48°24′50″ e, 900 m a.s.l., 2 mar. 2015. y. hong and m. razafindrakoto leg. (hnhm/af5660). paratypes madagascar: 1 clitellate, 9 aclitellate adults (hnhm/af5661); 2 clitellate, 5 aclitellate adults (hnhm/af5662); 3 aclitellate (chbnu md-10-al4); 7 aclitellate (chbnu md-10-al8); 9 aclitellate (chbnu md-10-c). locality and date same as for holotype description holotype clitellate, 100 mm in length, diameter after clitellum 7 mm, segment number 115, tail missing. paratypes 145–170 mm in length, 6–7 mm in diameter, segment number 234–268. colour greyish, with slight reddish pigmentation at head on dorsum. head zygolobous, segments 1–3 simple, 4–10 biannulate. fig. 4. kynotus voimmanus sp. nov. a. setal arrangement, a, b, c, d representing setal lines. b. spermathecae (13/14) schematic representation. c. genital seta. d. tip of the genital seta. scale bars: c = 0.6 mm; d = 0.1 mm. csuzdi c. et al., three new kynotus species from madagascar 11 dorsal pores lacking. setae small, ab and cd clearly observed from segment 2. setal arrangement after clitellum aa:ab:bc:cd:dd = 23:1.5:13:1:25, dd = 0.32 u (fig. 4a). clitellum annular on segments 18–½ 28, 28. male pores ventral on 16. female pores not seen. spermathecal pores in 13/14(2–3), 14/15(2–3), 15/16(2) irregularly arranged from bc to above cd. genital setal pores in variable position between ab and cd on 15. internal characters large muscular gizzard in 5. septa 6/7–8/9 strongly thickened, 5/6, 9/10 less so. calciferous glands, lamellae and typhlosolis lacking. dorsal blood vessel simple throughout, last pair of hearts in 11. excretory system holoic, vesiculate. two pairs of testes and sperm funnels in 10, 11 enclosed in large peri-oesophageal testis sacs. seminal vesicles lacking. ovaries in 13. a pair of large, oval copulatory chambers occupying ventral space of segments 16–19. each copulatory chamber bearing an irregular prostate-like gland (pseudoprostate) reaching segment 22. spermathecae oval-shaped, with long and thin duct attached at 13/14(2–3), 14/15(2–3), 15/16(2) arranged from above ab to between cd and m (fig. 4b). single pair of genital setal glands present in segment 15 above ab and around cd. genital setae slightly curved, with lanceolate apex, length 2.25–2.3 mm, diameter just under the apex 0.07 mm; ornamentation dense short serrations (fig. 4c–d). remarks this new species, with biannulate segments between 4 and 10 belongs to the alaotranus group (table 2). in the alaotranus group there are only two other species possessing a single pair of genital setal glands: k. rosae cognetti, 1906 and k. minutus csuzdi, razafindrakoto & blanchart, 2012. k. voimmanus sp. nov. differs from k. rosae by its smaller number of spermathecal segments (13/14, 14/15, 15/16, 16/17 vs 13/14, 14/15, 15/16) and also in the structure of spermathecae (k. rosae possesses oval spermathecae with very short ducts, k. voimmanus sp. nov. possesses oval spermathecae with thin and very long ducts, 2–3 times longer that the ampoule). the new species differs from k. minutus in its size and fig. 5. bayesian inference tree of some species of kynotus michaelsen, 1891 using coi sequences. numbers above branches indicate bayesian posterior probabilities, below branches ml bootstrap supports. values lower than 80% are not shown. european journal of taxonomy 336: 1–14 (2017) 12 coloration, and furthermore in the position of the clitellum (18–26 vs 18–½ 28), as well as the shape of the spermathecae (short vs long spermathecal duct). k. voimmanus sp. nov. is clearly different from all the examined alaotranus group species by its coi barcode as well (table 3, fig. 5). discussion before launching the project “changement global et diversité de la macrofaune du sol à madagascar” (global change and soil macrofauna diversity in madagascar) in 2008, only 13 species were known from the malagasy endemic earthworm family kynotidae. our activity in the recent years has resulted in the description of a further 9 kynotus species new to science (razafindrakito et al. 2011; csuzdi et al. 2012; razafindrakoto et al. 2017), increasing the number of valid species in the family to 22. it is worth noting that, due to difficulties accessing the rural sites in the rainy season (which is the most suitable for earthworm collections), larger regions of the country, especially in the western and southern part of the island, still remain unexplored. it is also important to highlight that several new species were discovered in highly disturbed habitats such as around sambaina (k. blancharti sp. nov. in secondary bushland, with scattered eucalyptus l’hér.) or antsirabe (k. parvus csuzdi, razafindrakoto & blanchart, 2012 in secondary aristida l. grassland). this indicates that not only the very few pristine regions should be sampled, but also disturbed areas, where native, still undescribed, species occur. our barcoding results clearly separated all the newly described species (fig. 5), supporting the conclusion that this method is quite powerful in recognizing earthworm species, as demonstrated recently by decaëns et al. (2016), chang & james (2011) and chang et al. (2009). the coi tree revealed that the alaotranus group’s species (except k. michaelseni) form the second largest clade and show a clear south-north geographical distribution pattern. the most basal species of this clade, k. ankisiranus sp. nov., possesses the southernmost distribution and the terminal species pair k. voimmanus sp. nov. – k. alaotranus is distributed in the north (figs 1, 5). however, it seems that coi alone is not suitable for inferring the phylogenetic relationships inside the genus. all deeper branches revealed showed no statistical support. kimura 2-parameter genetic distances (kimura 1980) showed intraspecific and interspecific separation values ranging between 0%–0.5 % and 14.3%–28.2% respectively (table 3). this data fits well with that observed by decaëns et al. (2016) reporting 0%–5.25% (mean 1.27%) intra-motu divergence and (13.68%–31.01% (mean 23.33%) inter-motu divergence for earthworms collected in french guiana. acknowledgements our thanks are due to s.w. james (iowa, usa) who provided his unpublished barcodes for several species of kynotus. we are grateful to j. philbert, j. randriatsarafara, e.j. razainjara, i.m. mamoude, j.w. maminiaina and m. rakotonandrasana for their generous help during the fieldwork. emma sherlock (nhm, london) is thanked for polishing the english of the text. this work was supported by the faune-m project of the institut français de la biodiversité (ifb) and by the national research foundation of korea (nrf) funded by the ministry of education (nrf-2015r1d1a2a01057305). references chang ch.-h. & james s.w. 2011. a critique of earthworm molecular phylogenetics. pedobiologia 54s: s3–s9. https://doi.org/10.1016/j.pedobi.2011.07.015 chang ch.-h., rougerie r. & chen j.-h. 2009. identifying earthworms through dna 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336: 1–14 (2017) 14 tamura k., stecher g., peterson d., filipski a. & kumar s. 2013. mega6: molecular evolutionary genetics analysis version 6.0. molecular biology and evolution 30: 2725–2729. https://doi.org/10.1093/ molbev/mst197 thompson j.d., higgins d.g. & gibson t.j. 1994. clustal w: improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position-specific gap penalties and weight matrix choice. nucleic acids research 22: 4673–4680. https://doi.org/10.1093/nar/22.22.4673 manuscript received: 7 july 2016 manuscript accepted: 26 september 2016 published on: 13 july 2017 topic editor: rudy jocqué desk editor: kristiaan hoedemakers printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain. https://doi.org/10.1093/molbev/mst197 https://doi.org/10.1093/molbev/mst197 https://doi.org/10.1093/nar/22.22.4673 european journal of taxonomy 112: 1–10 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2015.112 www.europeanjournaloftaxonomy.eu 2015 · grootaert p. et al. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:a92b8545-3b96-4596-ad29-b36d33e69770 1 two new coastal species of elaphropeza macquart (diptera: hybotidae) from bali, indonesia patrick grootaert 1,3, isabella van de velde 1,4 & igor v. shamshev 1,2,5 1entomology, royal belgian institute of natural sciences, vautierstraat 29, b-1000 brussels, belgium. email: patrick.grootaert@naturalsciences.be and isa.vandevelde@naturalsciences.be 2zoological institute, russian academy of sciences, universitetskaja nab. 1, st. petersburg 199034, russia. email: shamshev@mail.ru 3 urn:lsid:zoobank.org:author:b80bc556-9087-4d0d-9d69-7fa9be5779c4 4 urn:lsid:zoobank.org:author:cb1b2335-553c-4a40-bb10-19c62e3875c6 5 urn:lsid:zoobank.org:author:569f41cc-ec2b-4cf0-802a-8d7056c72c93 abstract. two new species of elaphropeza macquart, 1827 are described from the northeast coast of bali (indonesia): elaphropeza triseta sp. nov. belonging to the ephippiata-group and e. balinensis sp. nov. belonging to the biuncinata-group. a coi neighbour-joining tree is given for the southeast asian elaphropeza species showing large genetic distances between the species. the morphological characterisation of the ephippiata-group and the biuncinata-group sensu shamshev & grootaert 2007 is confirmed. key words. elaphropeza, new species, hybotidae, southeast asia. grootaert p., van de velde i. & shamshev i. 2015. two new coastal species of elaphropeza macquart (diptera: hybotidae) from bali, indonesia. european journal of taxonomy 112: 1–10. http://dx.doi.org/10.5852/ejt.2015.112 introduction shamshev & grootaert (2007) revised the genus elaphropeza in the oriental realm, reporting 79 species. apart from the redescription of the holotypes of previously known species, 51 new species were described mainly from singapore. these new species were the result of a one-year collecting campaign with malaise traps in various biotopes in singapore, e.g., swamp forest, secondary rain forest and mangrove. later, three species were described from vietnam by grootaert & shamshev (2009). these species were remarkable in having brush-like clusters of long hairs in the intertergal spaces between tergites 3 and 4 and sometimes 4 and 5 as well. these brushes might help in the evaporation of pheromones secreted by the glands below the tergites. subsequently, grootaert & shamshev (2012) published a monograph on the tachydromiinae of singapore which was partly based on a one-month site assessment study performed in 2009, when insects were collected with malaise traps in thirteen mangrove fragments all along the coast of singapore. in that monograph another eight elaphropeza species were described, all being coastal. finally, wang et al. (2012) described two new elaphropeza species from taiwan (wang et al. 2012). thus, 137 species of elaphropeza are now known for the oriental realm, including the two new species described in the http://dx.doi.org/10.5852/ejt.2015.112 www.europeanjournaloftaxonomy.eu/index.php/ejt/index http://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:a92b8545-3b96-4596-ad29-b36d33e69770 mailto:patrick.grootaert%40naturalsciences.be?subject= mailto:isa.vandevelde%40naturalsciences.be?subject= mailto:shamshev%40mail.ru?subject= http://zoobank.org/urn:lsid:zoobank.org:author:b80bc556-9087-4d0d-9d69-7fa9be5779c4 http://zoobank.org/urn:lsid:zoobank.org:author:cb1b2335-553c-4a40-bb10-19c62e3875c6 http://zoobank.org/urn:lsid:zoobank.org:author:569f41cc-ec2b-4cf0-802a-8d7056c72c93 http://dx.doi.org/10.5852/ejt.2015.112 european journal of taxonomy 112: 1–10 (2015) 2 present paper. this number is considered only a fraction of the number of species that might eventually be found to occur in the oriental realm, since most species have only been recorded from the limited territory of singapore with an emphasis on fauna restricted to mangroves. at the end of 2013 the first two authors collected a small number of specimens of elaphropeza from shrubs in the supralittoral zone of a shingle beach with black volcanic sands on the northeast coast of bali. it is the first record of the genus in bali and the specimens belong to two species that are new to science. they are described, illustrated, coi-barcoded and compared with the other species known from the oriental realm in order to confirm the separation of the genus elaphropeza in two species groups (shamshev & grootaert 2007). material and methods specimens were collected by sweep netting and preserved in 70% ethanol. holotypes are conserved in the museum zoologici bogor, indonesia (mzb). the specimens from which dna was extracted are conserved in the tissue collection at the royal belgian institute of natural sciences, brussels (rbins), with the cuticular parts still being recognisable. for the barcoding technique we refer to nagy et al. (2013). the evolutionary history was inferred using the neighbor-joining method mega6 (tamura et al. 2013). the optimal tree with the sum of branch lengths = 1.87955927 is shown. the percentages of replicate trees in which the associated taxa clustered together in the bootstrap test (1000 replicates) are shown next to the branches. the tree is drawn to scale, with branch lengths in the same units as those of the evolutionary distances used to infer the phylogenetic tree. the evolutionary distances were computed using the maximum composite likelihood method and are in the units of the number of base substitutions per site. the analysis involved 39 nucleotide sequences. codon positions included were 1st + 2nd + 3rd + noncoding. all positions containing gaps and missing data were eliminated. there was a total of 640 positions in the final dataset. evolutionary analyses were conducted in mega6 (tamura et al. 2013). results class insecta linnaeus,1758 order diptera linnaeus,1758 superfamily empidoidea latreille, 1804 family hybotidae macquart, 1823 subfamily tachydromiinae meigen, 1822 elaphropeza macquart, 1827 elaphropeza macquart, 1827: 86 (as genus). type-species: tachydromia ephippiata fallén, 1815: 11, by monotypy. ctenodrapetis bezzi, 1904: 351 (as subgenus of drapetis). type-species: ctenodrapetis ciliatocosta bezzi, 1904 by subsequent designation of melander (1928: 309). elaphropeza triseta sp. nov. urn:lsid:zoobank.org:act:cd141962-6947-4418-ab0b-25a25a2f41e7 figs 1–4 diagnosis a species of the e. ephippiata group; recognised by almost entirely yellow thorax, only postnotum brownish along posterior margin; antenna with scape and pedicel yellow; postpedicel short, about 1.5 times as long as wide, brown; legs yellow, with only tarsomere 5 dark brown, hind tibia with 3 brownish black anterodorsal bristles; halter yellow. http://zoobank.org/urn:lsid:zoobank.org:act:cd141962-6947-4418-ab0b-25a25a2f41e7 grootaert p. et al., new coastal elaphropeza (hybotidae) from bali 3 etymology the specific epithet refers to the presence of three anterodorsal bristles on the hind tibia. type material holotype indonesia: ♂, bali, amed, bunutan, beach, 8°20’19.44” s, 115°39’53.39” e, 2 jan. 2014, leg. p. grootaert & i. van de velde (mzb). paratypes indonesia: 2 ♂♂, 2 ♀♀, bali, amed, same provenance as holotype (rbins); 2 ♂♂, 2 ♀♀, bali, arya amed, bunutan, 8°20’38.89” s, 115°40’21.81” e, 25 dec. 2013, leg. p. grootaert & i. van de velde (rbins). description male (fig. 1) length. body 2.3 mm, wing 2.1 mm. head. occiput entirely black, largely shining, with yellowish setae; inner verticals long inclinate, outer verticals hardly prominent. anterior ocellars long proclinate, posterior ocellars minute. frons subshining, very narrow, above antennae about as broad as anterior ocellus. antenna with scape and pedicel yellow, postpedicel and stylus brownish; pedicel with circlet of subequally short setulae; postpedicel short, fig. 1. elaphropeza triseta sp. nov. male habitus (photo j. brecko). european journal of taxonomy 112: 1–10 (2015) 4 nearly 1.5 times as long as wide; stylus very long, nearly 4.0 times as long as postpedicel, with short pubescence. proboscis brownish yellow. palpus yellow, small, rounded, silvery pubescent, bearing scattered setulae, subapical seta short. thorax. almost entirely yellow, only antepronotum with brownish spot dorsally, postnotum brownish along posterior margin, hypopleuron brownish on extreme apex and metapleuron yellowish brown on upper part anteriorly; shining, with yellowish setation; anterior spiracle brownish yellow, posterior spiracle brown. prothoracic episterna lacking long upturned seta just above fore coxa, with minute setula on upper part. postpronotal seta hardly prominent. mesonotum with 2 long notopleurals, 1 shorter postsutural supra-alar, 1 long postalar and 4 scutellars (apical pair very long, cruciate; lateral pair short); acrostichals short, arranged in 4 irregular rows anteriorly but becoming sparser and biserial posteriorly, distinctly separated from dorsocentrals, lacking on prescutellar depression; dorsocentrals arranged in 3–4 irregular rows, as long as acrostichals, 1 prescutellar pair very long. legs. long, slender, almost entirely yellow, only tarsomere 5 of all legs dark brownish. coxae and trochanters with unmodified setation. fore femur slightly thickened, with rows of minute anteroventral and posteroventral setae, long seta near base and 1 subapical bristle anteriorly. fore tibia lacking prominent bristles (except subapicals). mid femur with rows of minute anteroventral and posteroventral spinule-like setae (more numerous near base), 1 long seta near base and 1 long subapical bristle anteriorly. mid tibia only with 1 black, short, subapical spine, lacking prominent setae (except subapicals). hind femur with short anteroventrals and 3 erect dorsal setae near base. hind tibia with 3 brownish black anterodorsal bristles; apical projection small, rounded, brownish. tarsi of all legs unmodified, with unmodified setation. figs 2–4. elaphropeza triseta sp. nov. male terminalia. 2. right epandrial lamella. 3. epandrium with cerci. 4. left surstylus. c = cerci; ls = left surstylus; rs = right surstylus. scale bar = 0.1 mm. grootaert p. et al., new coastal elaphropeza (hybotidae) from bali 5 wings. normally developed, finely infuscate, covered with uniform microtrichia; veins mostly yellowish brown; basal section of m1+2, crossveins bm-cu and r-m pale. costal vein with moderately long, uniform setulae along anterior margin. costal bristle long, brown. costal index: 41/31/31. vein rs long, slightly longer than crossvein bm-cu. vein r2+3 evenly bowed. veins r4+5 and m1+2 somewhat divergent near wing apex, r4+5 slightly bowed, m1+2 straight. vein cua1 reaching wing margin. vein a1 lacking. crossvein bm-cu slightly oblique. crossvein r-m before middle of cell bm. squama yellowish, with long, brownish yellow setae. halter yellow. abdomen. tergite 1 pale yellow; tergites 2–3 brownish yellow, subtriangular viewed laterally, divided medially, with scattered unmodified setae; tergite 4 brown, broadest, with squamiform setae; tergite 5 narrow, undivided, with squamiform setae; an ampoule-like structure below tergite 5; tergites 6–7 brownish yellow, tergite 6 undivided, with minute unmodified setae; tergite 7 undivided, with long posteromarginal setae; tergite 8 unmodified. sternites yellowish, with scattered minute setae, sternite 8 with long, posteromarginal setae. gland-like structure present between tergites 4 and 5, ampoule-like. terminalia (figs 2–4) rather small, with cerci, apical part of right epandrial lamella and left surstylus brownish, otherwise yellow. right epandrial lamella bifid (fig. 2); dorsal lobe with setae on dorsal margin. tip with a brush of densely set, short setae. ventral lobe with strong brown setae on apex. right and left cerci equally long. right cercus with a single long subapical seta, other setae short. left cercus (fig. 3) bifid; outward lobe with 3 strong brown setae on apex. tip of left epandrial lamella with 4 setae: 1 weak and 3 stronger. left surstylus also bifid, with short setae (fig. 4). female similar to male; abdomen without ampoule-like gland-like structure; intersegmental area between tergites 4 and 5 set with short hairs, anteriorly with 2 openings; segment 8 moderately long, tergite and sternite 8 separated, yellowish brown; tergite 8 entire, with several short to moderately long setae posteriorly; sternite 8 entire, covered with moderately long setae; cercus small, yellowish. distribution bali, indonesia. remarks elaphropeza triseta sp. nov. is actually the only yellow oriental species with three anterodorsal setae on the hind tibiae. in the key of grootaert & shamshev (2012), e. triseta sp. nov. resolves closest to e. acrodactyla (melander, 1928), described from a female from the philippines, but the latter has only two anterodorsal bristles on the hind tibiae and has the postpedicel about three times as long as wide. in e. triseta sp. nov. the postpedicel is nearly 1.5 times as long as wide. elaphropeza balinensis sp. nov. urn:lsid:zoobank.org:act:d744e850-5f65-4c29-a671-9f9d0872a6a9 figs 5–10 diagnosis a species of the e. biuncinata group; thorax yellow, scutum with narrow, short, brownish patch above wing; antenna with short postpedicel, about 2.0 times as long as wide; legs yellow, but all tibiae and tarsi faintly brownish tinged, including apical tarsomere of all legs; hind tibia with 1 short, black anterodorsal bristle at about middle and 1 similar subapical bristle anteriorly; halter brownish. scutum entirely covered with setae (acrostichals and dorsocentrals not differentiated), reaching base of scutellum. http://zoobank.org/urn:lsid:zoobank.org:act:d744e850-5f65-4c29-a671-9f9d0872a6a9 european journal of taxonomy 112: 1–10 (2015) 6 etymology the specific epithet refers to the provenance from bali. type material holotype indonesia: ♂, bali, amed, bunutan, beach, arya amed, 8°20’38.89” s, 115°40’21.81” e, 2 jan. 2014, leg. p. grootaert & i. van de velde (mzb). paratypes indonesia: 3 ♂♂ (1 ♂ dissected), 2 ♀♀, bali, amed, bunutan, beach, 8°20’19.44” s, 115°39’53.39” e, 26 dec. 2013, leg. p. grootaert & i. van de velde (rbins). description male (fig. 5) length. body 1.8 mm, wing 1.6 mm. head. occiput entirely black, largely shining, with yellowish setae; inner verticals long inclinate, outer verticals short. anterior ocellars long proclinate, posterior ocellars minute. frons subshining, very narrow, above antennae about as broad as anterior ocellus. antenna with scape and pedicel yellow, postpedicel and stylus brownish; pedicel with circlet of subequally short setulae; postpedicel short, nearly 2 times as long as wide; stylus with short pubescence, very long, 2.5–3.0 times as long as postpedicel. proboscis brownish yellow. palpus yellow, small, rounded, bearing scattered setulae, subapical seta short. thorax. almost entirely yellow, scutum with narrow, short, brownish yellow patch above wing on each side (indistinct in paler specimens), metanotum slightly shadowed on middle part (sometimes only posteriorly or entirely yellow), mesopleuron posteriorly and sternopleuron on lower part usually with small, brownish yellow patch; shining, with yellowish setation; spiracles brownish yellow. prothoracic episterna with long, upturned seta just above fore coxa. postpronotal seta not prominent. mesonotum with 2 long notopleurals, 1 short postsutural supra-alar, 1 moderately long postalar and 4 scutellars (apical pair long, cruciate; lateral pair short); scutum covered with uniform, minute setae, extending to base of scutellum. acrostichals and dorsocentrals not differentiated, except for 1 pair of long, prescutellar dorsocentrals. legs. robust, yellow, but all tibiae and tarsi yellowish-brownish tinged. coxae and trochanters with unmodified setation. fore femur slightly thickened, with rows of minute anteroventral and posteroventral setae, long seta near base and 1 subapical bristle anteriorly. fore tibia lacking prominent bristles (except subapicals). mid femur with rows of minute anteroventral and somewhat longer posteroventral setae, 1 long seta near base and 1 moderately long subapical bristle anteriorly. mid tibia with row of black, short, anteroventral spines on apical 3/4, lacking prominent setae (except subapicals). hind femur with minute anteroventrals and 3 erect, dorsal setae near base. hind tibia with 1 short, black anterodorsal bristle on about middle and 1 similar subapical bristle anteriorly. tarsi of all legs unmodified, with unmodified setation. wings. normally developed, finely infuscate, covered with uniform microtrichia; veins mostly yellowish brown; basal section of m1+2, crossveins bm-cu and r-m pale. costal vein with moderately long, uniform setulae along anterior margin. costal bristle moderately long, brown. costal index: 29/22/26. vein rs long, nearly as long as crossvein bm-cu. vein r2+3 evenly bowed. veins r4+5 and m1+2 somewhat divergent near wing apex, both slightly bowed. vein cua1 reaching wing margin. vein a1 lacking. crossvein bm-cu oblique. crossvein r-m near middle of cell bm. squama yellowish brown, with long, brownish setae. halter brownish. grootaert p. et al., new coastal elaphropeza (hybotidae) from bali 7 abdomen. tergite 1 largely pale yellow, very narrow, divided medially; tergite 2 broadly concave medially, undivided; tergite 3 broadest, subrectangular, with squamiform setae; tergites 4–5 narrow, both with squamiform setae; tergites 6–7 broader than tergite 5, the latter with moderately long, posteromarginal setae; sternites 3–4 broadly divided, sternite 5 entire, concave anteriorly, sternites 6–7 entire, subrectangular, with scattered minute setae. gland-like structure present between tergites 3–4 and 4–5, plate-like. terminalia (figs 6-10) rather small; epandrium brown, otherwise yellow. right epandrial lamella (fig. 6) with base brown, apex (surstylus) yellowish; basal half of dorsal margin with 4 long, strong setae. left epandrial lamella without the usual set of setae dorsally. left surstylus composed fig. 5. elaphropeza balinensis sp. nov. male habitus (photo j. brecko). european journal of taxonomy 112: 1–10 (2015) 8 of 3 pieces (fig. 10), hidden by the large left cercus: ls1 with apex indented (fig. 9), on inner side with at least 6 setae; ls2 with apex saw-toothed; ls3 with short setae (fig. 10). phallus long, with partly coiled tip (fig. 8). right cercus short, with rather long setae. left cercus very wide, covering the left surstylus; with only minute setae. female similar to male but mid tibia without anteroventral spines; abdominal tergites 3–7 of subequal width; segment 8 short, tergite 8 separated from sternite 8, deeply cleft posteriorly; cercus short, yellowish. distribution bali, indonesia. remarks in the key of grootaert & shamshev (2012), e. balinensis sp. nov. resolves closest to e. monospina shamshev & grootaert, 2007, known only from singapore. the latter also has a somewhat maculated scutum, but it has the hind tarsomere 1 with a spine-like ventral seta near the base. the structure of the male genitalia is also very different. figs 6–10. elaphropeza balinensis sp. nov. male terminalia. 6. right epandrial lamella. 7. detail of right surstylus. 8. epandrium with cerci. 9. left surstylus, ls1. 10. left surstylus. lc = left cercus; ls1, ls2, ls3 = 3 sclerites of the left surstylus; ph = phallus; rc = right cercus. scale bar = 0.1 mm. grootaert p. et al., new coastal elaphropeza (hybotidae) from bali 9 discussion shamshev & grootaert (2007) proposed two species groups in the oriental elaphropeza based on morphological characters. in species of the e. biuncinata group a long upturned bristle is present just above the fore coxa and the abdominal tergites 3, 4 and 5 bear squamiform setae (fig. 5). in species of the e. ephippiata group the proepisternum is bare just above the fore coxa and the abdominal tergite 3 is covered with ordinary setulae, lacking squamiform setae, which are present on tergites 4 and/or 5 only. in addition, there are several characters in the male genitalia, such as fusion or not of the right surstylus with the right epandrial lamella, fusion or not of both cerci, short or very long phallus and more (shamshev & grootaert 2007). this preliminary and informal classification is confirmed here by the coi gene (fig. 11), with a bootstrap value of 86 for the biuncinata-group. however, it would not be practical to elevate the two species groups to genus level since the morphological characters are not always easy to see. as can be seen further on fig. 11, the genetic distance between all species is large and always more than 5%, showing that at least for the selected species from the oriental realm, this separation is already of considerable age. fig. 11. neighbour-joining tree of the coi barcodes of a number of selected southeast asian elaphropeza species showing the position of e. triseta sp. nov. and e. balinensis sp. nov. (both underlined) in the ephippiata-group and biuncinata-group, respectively. the bootstrap value is indicated at the nodes. the scale of the genetic distance is 2%. european journal of taxonomy 112: 1–10 (2015) 10 acknowledgements the authors thank mr johan de gruyter for the barcoding of the species. mr jonathan brecko helped with the photography. references bezzi m. 1904. empididae indo-australiani raccolti dal signor l. biro. annales historico-naturales musei nationalis hungarici 2: 320–361. fallén c.f. 1815. empididae sveciae. berlingianus, lund. grootaert p. & shamshev i.v. 2009. new species of the genus elaphropeza macquart (diptera: hybotidae) with remarkable abdominal structures from viet nam. studia dipterologica 15: 165–174. grootaert p. & shamshev i.v. 2012. the fast-running flies (diptera, hybotidae, tachydromiinae) of singapore and adjacent regions. european journal of taxonomy 5: 1–162. http://dx.doi.org/10.5852/ ejt.2012.5 macquart j. 1827. insectes diptères du nord de la france. platypezines, dolichopodes, empides, hybotides. l. danel, lille. melander a.l. 1928. diptera, fam. empididae. in: wytsman p. (ed.) genera insectorum 185: 1–434. http://www.biodiversitylibrary.org/item/60119#page/315/mode/1up nagy z., sonet g., mortelmans j., vandewynkel c. & grootaert p. 2013. using dna barcodes for diversity assessment in hybotidae (diptera, empidoidea). zookeys 365: 263–278. http://dx.doi. org/10.3897/zookeys.365.6070 shamshev i.v. & grootaert p. 2007. revision of the genus elaphropeza macquart (diptera: hybotidae) from the oriental region, with a special attention to the fauna of singapore. zootaxa 1488: 1–164. tamura k., stecher g., peterson d., filipski a. & kumar s. 2013. mega6: molecular evolutionary genetics analysis version 6.0. molecular biology and evolution 30: 2725–2729. http://dx.doi. org/10.1093/molbev/mst197 wang j., zhang l. & yang d. 2012. two new species and one newly recorded species of elaphropeza macquart from taiwan (diptera, empididae, tachydromiinae). zookeys 203: 15–25. http://dx.doi. org/10.3897/zookeys.203.3284 manuscript received: 22 october 2014 manuscript accepted: 10 december 2014 published on: 13 february 2015 topic editor: koen martens desk editor: danny eibye-jacobsen printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; national botanic garden of belgium, meise, belgium; royal museum for central africa, tervuren, belgium; national history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark. http://dx.doi.org/10.5852/ejt.2012.5 http://dx.doi.org/10.5852/ejt.2012.5 http://www.biodiversitylibrary.org/item/60119#page/315/mode/1up http://dx.doi.org/10.3897/zookeys.365.6070 http://dx.doi.org/10.3897/zookeys.365.6070 http://dx.doi.org/10.1093/molbev/mst197 http://dx.doi.org/10.1093/molbev/mst197 http://dx.doi.org/10.3897/zookeys.203.3284 http://dx.doi.org/10.3897/zookeys.203.3284 european journal of taxonomy 87: 1–19 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2014.87 www.europeanjournaloftaxonomy.eu 2014 · boström s. & holovachov o. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:691ba51c-6882-4437-a037-0e8861dcbf17 1 descriptions of species of stegelleta thorne, 1938 (nematoda, rhabditida, cephalobidae) from california, new zealand and senegal, and a revision of the genus sven boström 1 & oleksandr holovachov 2 1, 2 department of zoology, swedish museum of natural history, box 50007, se-104 05 stockholm, sweden 1 email: sven.bostrom@nrm.se (corresponding author) 2 email: oleksandr.holovachov@nrm.se 1 urn:lsid:zoobank.org:author:528300cc-d0f0-4097-9631-6c5f75922799 2 urn:lsid:zoobank.org:author:89d30ed8-cfd2-42ef-b962-30a13f97d203 abstract. populations of stegelleta are described from california, new zealand and senegal. an amphimictic population from california is identified as belonging to s. incisa and compared with type specimens from utah and an amphimictic population from italy. one population from new zealand is close to s. incisa but considered to represent a new species, stegelleta laterocornuta sp. nov. it is particularly characterised by a 379–512 µm long body in females and 365–476 µm in males; cuticle divided into 16 rows of blocks at midbody (excluding lateral field); lateral field with four incisures; three pairs of asymmetrical lips, u-shaped primary axils without guarding processes, each lip asymmetrically rectangular with a smooth margin, only lateral lips have slender acute tines; three labial probolae, bifurcated at half of their length; vulva without flap; spermatheca 17–31 µm long; postuterine sac 7–24 µm long; spicules 21.5–23.5 µm long. other specimens from new zealand are identified as belonging to s. tuarua. a parthenogenetic population from senegal is identified as belonging to s. ophioglossa and compared with type specimens from mongolia and records of several other populations of s. ophioglossa. the generic diagnosis is emended and a key to the species of stegelleta is provided. keywords. morphology, new species, sem, stegelleta, taxonomy. boström s. & holovachov o. 2014. descriptions of species of stegelleta thorne, 1938 (nematoda, rhabditida, cephalobidae) from california, new zealand and senegal, and a revision of the genus. european journal of taxonomy 87: 1–19. http://dx.doi.org/10.5852/ejt.2014.87 introduction the genus stegella was erected by thorne (1937) with stegella incisa thorne, 1937 as type species, for cephalobids having: cuticle divided into plate-like segments over entire body; labial probolae furcate, prongs simple or with few branches; cephalic probolae variable, rounded axil borders or flap-like with membranes; and wing area unusually broad with a medial line and crenate borders. http://dx.doi.org/10.5852/ejt.2014.87 https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:691ba51c-6882-4437-a037-0e8861dcbf17 http://www.europeanjournaloftaxonomy.eu http://zoobank.org/urn:lsid:zoobank.org:author:528300cc-d0f0-4097-9631-6c5f75922799 http://zoobank.org/urn:lsid:zoobank.org:author:89d30ed8-cfd2-42ef-b962-30a13f97d203 http://dx.doi.org/10.5852/ejt.2014.87 european journal of taxonomy 87: 1–19 (2014) 2 due to a homonomy with a hydroid, stegella stechow, 1919, it was later renamed and became stegelleta thorne, 1938. stegelleta currently includes six valid species of which the most recent one, s. arenaria boström & holovachov, 2012, was described from kelso dunes, mojave national preserve, california. specimens of s. incisa were also found in the samples from kelso dunes. this population is here described from studies by light (lm) and scanning electron microscope (sem) and compared with type specimens from utah and an amphimictic population from italy described under the name s. ophioglossa andrássy, 1967 (thorne 1937; orselli & vinciguerra 2002). another population from new zealand, described here from studies by light and scanning electron microscope, is close to s. incisa but considered to represent a new species. some specimens of s. tuarua yeates, 1967 from new zealand are also described and compared with the original description (yeates 1967). a recent parthenogenetic population of s. ophioglossa from senegal is described and compared with the original description of s. ophioglossa from mongolia and records of populations from uzbekistan, senegal, iran and spain (andrássy 1967; mavljanov 1978; de ley et al. 1990; shokoohi et al. 2008; abolafia et al. 2011). the generic diagnosis is emended and a key to the species of stegelleta is provided. material and methods during a tour in the kelso dunes area, mojave national preserve, southern california, the junior author collected several samples of sand underneath different species of desert plants. nematodes were recovered from sand samples using a modified baermann funnel extraction. for lm, specimens were relaxed by gentle heat, fixed in cold 4% formaldehyde solution, transferred to pure glycerine by a slow evaporation method and mounted on permanent slides in glycerine with paraffin wax as support for the coverslip. after measuring and observations, some specimens were removed from slides and rehydrated by first gradually adding drops of s1 (1% glycerine, 20% ethanol and 79% distilled water) to glycerine in an embryo-dish until the volume tripled and then gradually adding distilled water until the volume tripled again. the specimens were then washed in distilled water before resuspension in formaldehyde. for sem, specimens were post-fixed in 1% osmium tetroxide (oso4) and transferred to pure acetone through an acetone/distilled water series. specimens were critical point dried in liquid co2, mounted on stubs, gold-plated under vacuum to a thickness of 200 å in an agar high resolution sputter coater model 20, and examined in a hitachi s-4300 sem at an accelerating voltage of 5 kv. specimens from new zealand were collected by christine (gamble) powell in the abel tasman national park and subsequently cultured as the strain stegelleta sp. jb-75, which was sequenced and included in a phylogenetic study on cephalobina (nadler et al. 2006). additional material from the castlecliff beach in new zealand was supplied by dieter sturhan and gregor yeates. this is one of the localities where the original populations of s. iketaia yeates, 1967 were collected. specimens from senegal were collected in the kaolack region, thyssé-kaymor by cécile villenave. for lm, the specimens from new zealand and senegal were processed using different modifications of a slow evaporation method. specimens from the strain stegelleta sp. jb-75 were processed to sem by dan bumbarger and photographs were put at our disposal. morphometric characters applied herein and their abbreviations are as defined for cephalobidae in de ley et al. (1999); terminology of the labial and cephalic region and stoma terminology follows holovachov et al. (2009). type and other specimens are deposited in the invertebrate collections of the department of zoology, swedish museum of natural history, stockholm, sweden (smnh). boström s. & holovachov o., species of stegelleta and revision of the genus 3 results class chromadorea inglis, 1983 order rhabditida chitwood, 1933 family cephalobidae filipjev, 1934 genus stegelleta thorne, 1938 stegella thorne, 1937: 4 [junior homonym of stegella stechow, 1919]. stegelleta thorne, 1938: 64–65. type species stegella incisa thorne, 1937. diagnosis (emended after holovachov et al. 2009) cuticle annulated, without distinctly annulated internal layer; annuli with longitudinal incisures (tessellated). lateral field with two or four wings (appearing as three, four or five incisures); ending at tail terminus in females and in males. lip region weakly offset, consisting of six globular lips arranged in three pairs: one dorsal and two subventral. pairs of lips separated by primary axils; guarding processes absent; secondary axils undeveloped. cephalic probolae absent or in the shape of one acute tine extending along the primary axil occurring only on lateral lips or on all six lips. labial probolae elongate-conoid with broad basis, bifurcated about halfway forming two slender prongs, in some species further bifurcated apically. six labial and four cephalic papilliform sensilla arranged in a cephaloboid manner. amphidial aperture slit-shaped, located on lateral lips. stoma divided into cheilo-, gymnoand stegostom: cheilostom barrel-shaped with strongly sclerotized bacilliform cheilorhabdia; gymnostom weakly developed, as wide as stegostom, with weak gymnorhabdia; stegostom consists of a funnel-shaped prostegostom and variably shaped mesostego-, metastegoand telostegostom parts. metastegostom denticle present. pharynx cephaloboid: pharyngeal procorpus and metacorpus cylindrical, lining of procorpus and metacorpus are of same thickness; isthmus narrower than metacorpus; basal pharyngeal bulb oval, with strongly developed valves. nerve ring encircling posterior part of metacorpus or anterior part of isthmus. excretory pore opens at level of nerve ring, at posterior part of metacorpus or anterior part of isthmus. deirids present. female reproductive system cephaloboid; posterior part of ovary straight; spermatheca present; postvulval uterine sac present; vagina straight; vulva flat, vulval flap absent or present. male reproductive system cephaloboid; spicules cephaloboid with manubrium and corpus of approximately equal width; gubernaculum plate-like or wedge-shaped; cornua crurum present. male genital papillae: at least one ventrosublateral pair located anterior to cloaca; one ventrosublateral pair located at the level of cloacal opening; two pairs located at middle of tail length; and three pairs (lateral, subventral and subdorsal) near tail terminus; there is a midventral papilla on anterior cloacal lip. rectum short (1–2 times longer than anal body diameter). phasmids located at about one-third to half of tail length in both sexes. female tail conoid or subcylindrical, straight, tail terminus bluntly rounded or truncate; male tail conoid, arcuate ventrad, tail terminus bluntly rounded. valid species stegelleta arenaria boström & holovachov, 2012 stegelleta georgica bagaturija, 1973 stegelleta iketaia yeates, 1967 stegelleta incisa (thorne, 1937) stegelleta ophioglossa andrássy, 1967 = stegelleta cylindrica mavljanov, 1978: 1889, figs 1–6 stegelleta tuarua yeates, 1967 european journal of taxonomy 87: 1–19 (2014) 4 stegelleta incisa (thorne, 1937) figs 1a–e, 2, table 1 stegella incisa thorne, 1937: 14–15, fig. 4g–i. stegelleta incisa – thorne 1938: 65. stegelleta ophioglossa – orselli & vinciguerra 2002: 216–219, figs 5, 6a–c. diagnosis stegelleta incisa is characterised by a 465–580 µm long body in females and 428–600 µm in males; cuticle divided by regular longitudinal striations into 16 rows of blocks at midbody (excluding lateral field); lateral field with four incisures extending almost to tail terminus in females and to midtail papillae in males, two outer incisures extend to tail terminus in males; three pairs of asymmetrical lips, pairs of lips separated by u-shaped primary axils without guarding processes, secondary axils demarcated by a shallow incisure, each lip asymmetrically rectangular with a smooth margin and without tines; three labial probolae, bifurcated at half of their length, prongs bent toward one another apically, “snaketongue”-shaped; pharyngeal corpus 2.1–2.9 times isthmus length; nerve ring and excretory pore at level of metacorpus to metacorpus-isthmus junction, deirids at level of metacorpus-isthmus junction to isthmus; vulva with anterior flap; spermatheca 24–43 µm long; postuterine sac 22–36 µm long; spicules 22–36 µm long. material examined usa: 13 ♀♀, 9 ♂♂, smnh 135940–135942, 28 mar. 2010, california, mojave national preserve, kelso dunes, soil around roots of desert plants (34°53.698’ n, 115° 42.155’ w and 34° 53.754’ n, 115° 42.248’ w), legit o. holovachov & p. de ley. description adult body slightly arcuate in different ways when killed by heat, males often strongly arcuate ventrad in tail end. cuticle coarsely annulated, annuli 2.4–3.4 µm wide at midbody and 2.2–3.0 µm wide in pharyngeal region. cuticle tessellated: longitudinal striae giving it a tiled appearance, each block often with a central indentation as seen under sem. anteriorly, 9–10 first annuli, the longitudinal striation is irregular and blocks rounded, followed by a more regular striation creating rectangular blocks, 16 rows of blocks at midbody (excluding lateral field). lateral field consisting of two wings separated by a broad groove, appearing as four incisures under lm, occupying about 20–25% of body diameter, extending almost to tail terminus in females and to midtail papillae in males, two outer incisures extend to tail terminus in males. lip region slightly offset, carrying 6 + 4 papillae and two round amphids. three pairs of asymmetrical lips, one dorsal and two ventrolateral. pairs of lips separated by u-shaped primary axils without guarding processes. each pair with a shallow incisure demarcating a secondary axil. each lip asymmetrically rectangular with a smooth margin, without any tines. three labial probolae, 8.0–9.5 µm high, bifurcated at half of their length, prongs bent toward one another apically (“snake-tongue”-shaped). stoma somewhat longer than lip region diameter. stomatal parts not clearly discernible. cheilorhabdia bacilliform in lateromedian view; metastegostom with a dorsal denticle. pharynx cephaloboid. pharyngeal corpus cylindrical; isthmus narrow, often folded, and not clearly demarcated from corpus; bulb oval, with valves. nerve ring at level of metacorpus to metacorpus-isthmus junction. excretory pore and deirids at level of metacorpus-isthmus junction to isthmus. female reproductive system monodelphic, prodelphic, in dextral position in relation to intestine. ovary reflexed posteriorly at oviduct, ovary straight posterior to vulva. spermatheca well developed. postvulval uterine boström s. & holovachov o., species of stegelleta and revision of the genus 5 fig. 1. a–e. stegelleta incisa (thorne, 1937). a. pharyngeal region. b. female gonad. c. anterior end, surface view. d. female tail. e. male tail. f–j. stegelleta laterocornuta sp. nov. f. pharyngeal region. g. female gonad. h. anterior end, surface view. i. female tail. j. male tail. scale bar = 20 µm. european journal of taxonomy 87: 1–19 (2014) 6 sac one to one-and-a-half times vulval body diameter long. vagina about one-third of vulval body diameter. vulval lips not or slightly protruding, anterior lip with cuticular flap directed posteriad and partly covering vulval opening. vulva covered by genital plug in some specimens. tail conoid with 13–15 ventral annuli, terminus truncate. one specimen has a tail with only 10–11 annuli and broadly rounded terminus. phasmids located at about one-third to half of tail length. male reproductive system monorchic, dextral in position; testis reflexed ventrad anteriorly. spicules paired and symmetrical, curved ventrad; with oval manubrium and subcylindrical, gradually narrowing shaft. gubernaculum plate-like, cornua crurum present. tail strongly arcuate ventrad, conoid with rounded terminus. the two outer lateral lines extend posterior to the phasmid, transforming into a cuticular ridge that reaches tail terminus. genital papillae arranged as follows: two pairs subventral precloacal (at 24–33 µm and at 42–58 µm anterior to cloaca), one pair subventral adcloacal, a single midventral papilla fig. 2. stegelleta incisa (thorne, 1937), sem micrographs. a–b. anterior end, left lateral view. c. anterior part of lateral field. d. deirid (arrow). e. vulval region. f. female tail, subventral view. g. female tail, left sublateral view (arrow points at phasmid). h. female tail, lateral view. scale bars = 5 µm. boström s. & holovachov o., species of stegelleta and revision of the genus 7 table 1. measurements (in µm) of stegelleta incisa (thorne, 1937) from kelso dunes, california compared with the type population of s. incisa from utah and a bisexual population described by orselli & vinciguerra (2002) from italy under the name s. ophioglossa andrássy, 1967 (presented as mean ± s.d. and (range) or only range). * calculated from data in original publication; ** number of annuli from anterior end to nerve ring, excretory pore and deirid, respectively; ¤ estimated from drawings in original publication; indicates that data is not available or not applicable; figures within [ ] indicate the number of specimens measured. population utah kelso dunes, california italy no. & sex 1 ♀ 1 ♂ 10 ♀♀ 11 ♂♂ 5 ♀♀ 5 ♂♂ body length 500 460 490 ± 23 (465–534) 463 ± 28 (428–526) 550 ± 30 (510–580) 550 ± 30 (510–600) body diameter (bd) 24* 20* 23.2 ± 1.0 (22–25) 21.2 ± 1.7 (18–24) 29 ± 1.4 (27.5–30.8) 25.3 ± 2.6 (20.9–27.5) pharynx length 139* 115* 130.6 ± 1.0 (115–141) 123.9 ± 7.9 (107–135) 181.4 ± 6.1 (177–192) 184.8 ± 24.2 (161–225) tail length 34* 34* 38.8 ± 1.7 (32–41) 32.2 ± 2.5 (29–36) 50.1 ± 3.6 (44–53.9) 48.6 ± 3.6 (45.1–53.9) anal or cloacal diam. (abd) 14.4 ± 0.2 (14.0–14.5) 16.1 ± 1.0 (14.5–18.0) 18.4 ± 0.4 (17.6–18.7) 22 ± 1.1 (20.9–23.1) vulva or testis 307 ± 14 (291–335) 269 ± 23 (223–300) v-a/t 3.8 ± 0.2 (3.4–4.1) a 21 23 21.2 ± 0.9 (19.6–22.5) 21.9 ± 1.3 (19.5–24.2) 19 ± 2 (17–21) 22 ± 2.2 (15–25) b 3.6 4 3.8 ± 0.2 (3.5–4.2) 3.7 ± 0.2 (3.4–4.3) 3 ± 0.1 (2.8–3.2) 2.9 ± 0.2 (2.6–3.3) c 15.0 13.5 12.7 ± 0.6 (11.8–13.4) 14.4 ± 1.0 (12.7–15.5) 11 ± 0.5 (10.4–11.5) 11.3 ± 0.6 (10.9–12.2) c’ 2.7 ± 0.1 (2.5–2.8) 2.0 ± 0.2 (1.8–2.3) 2.6 ± 0.2 (2.3–2.9) 2.1 ± 0.1 (1.9–2.3) v or t (%) 66 41 62.7 ± 1.1 (61–65) 58.1 ± 3.4 (52–64) 62.5 ± 0.6 (61.4–63) lip region diameter 8.3 ± 0.3 (7.5–8.5) 8.0 ± 0.6 (7.0–8.5) stoma length 9.8 ± 0.9 (8.5–11.0) 9.7 ± 0.6 (9–11) 13.2 ± 2 (11–15.4) 13.5 ± 0.4 (13.2–14.3) corpus length 83.2 ± 2.8 (78–86) [5] 81.2 ± 3.9 (76–87) [5] 123 ± 5.4 (117–129) 123.6 ± 21.5 (102–159) isthmus length 34.2 ± 2.7 (29–36) [5] 31.8 ± 2.4 (30–36) [5] 30.3 ± 7.5 (17.6–36.3) 32.7 ± 4.6 (25.3–37.4) bulb length 16.6 ± 0.7 (16–18) 15.7 ± 0.9 (14.5–17.0) [6] 20.2 ± 2.1 (17.6–23.1) 18.8 ± 0.6 (18.1–19.8) bulb diameter 12.2 ± 0.7 (11–13) 11.8 ± 0.7 (11–13) [6] corpus/isthmus ratio 3 2.4 ± 0.2 (2.3–2.9) [5] 2.6 ± 0.2 (2.1–2.8) [5] nerve ring from ant. end 88.5 ± 4.5 (78–94) 90.2 ± 7.1 (78–102) 137.5 ± 6.3 (129–144) 141.8 ± 23.1 (118–179) european journal of taxonomy 87: 1–19 (2014) 8 on anterior cloacal lip; two pairs (one ventrosublateral and one lateral) at midtail; three pairs (one lateral, one subventral and one dorsosublateral) closer to tail terminus. phasmids located at about two-fifths to half of tail length. remarks the population collected from sand in the kelso dunes area, mojave desert, california and described here, agrees in many respects with the type specimens of stegelleta incisa described from utah by thorne (1937) and a bisexual population described from italy by orselli & vinciguerra (2002) under the name s. ophioglossa. one major difference is the presence of a cuticular flap on the anterior vulval lip directed posteriad and partly covering the vulval opening, as seen by sem in the present specimens. this feature is not easily seen with light microscopy, which probably explains why it was not mentioned by thorne (1937) in his original description. orselli & vinciguerra (2002) also did not mention or illustrate it in their specimens, presumably for the same reason. population utah kelso dunes, california italy no. & sex 1 ♀ 1 ♂ 10 ♀♀ 11 ♂♂ 5 ♀♀ 5 ♂♂ excretory pore from ant. end 93.8 ± 2.9 (88–99) 95.6 ± 7.0 (84–108) 122–147 deirid from ant. end 101.1 ± 3.4 (94–107) 101.5 ± 8.3 (82–114) rnr** 32 ± 1.5 (30–35) 32 ± 2 (30–36) rep** 34 ± 2 (32–38) 34 ± 2 (31–38) rdei** 37 ± 2 (35–41) 37 ± 2 (34–41) annuli width at midbody 2.4–3.4 2.4–3.0 2.0–2.5 annuli width anteriorly 2.4–2.7 2.2–3.0 vagina or testis flexure length 7.3 ± 0.4 (7–8) 39.5 ± 6.7 (27–48) 7–8 spermatheca or spicule length 30.9 ± 6.7 (24–43) 24.0 ± 1.3 (22–27) 24–26.5 34.5 ± 1.6 (31.9–36.3) pus or gubernaculum length 27.3 ± 2.3 (24–31) 13.2 ± 1.1 (12–16) 30.6 ± 5.9 (22.5–36.3) 20.4 ± 1.4 (19.8–23.1) pus/vbd 1.3 ± 0.1 (1.1–1.4) 0.7–1.3 rectum 16.0 ± 1.6 (13–18) 21.6 ± 0.3 (16.5–24.7) rectum/abd 1.1 ± 0.1 (0.9–1.2) phasmid 14.9 ± 2.2 (12–20) 14.1 ± 1.8 (11–18) phasmid (% of tail) 30 ¤ 47 ¤ 38.3 ± 4.6 (32–49) 43.8 ± 4.7 (37–50) 15–30 25–35 boström s. & holovachov o., species of stegelleta and revision of the genus 9 stegelleta laterocornuta sp. nov. urn:lsid:zoobank.org:act:7669b17a-d71a-46c0-8ba0-eb92f28b7111 figs 1f–j, 3, table 2 stegelleta sp. jb-75 – nadler et al. 2006: 701, table 1. diagnosis stegelleta laterocornuta sp. nov. is characterised by a 379–512 µm long body in females and 365–476 µm in males; cuticle divided by regular longitudinal striations into 16 rows of blocks at midbody (excluding lateral field); lateral field with four incisures extending almost to tail terminus in females and to midtail papillae in males, two outer incisures extend to tail terminus in males; three pairs of asymmetrical lips, pairs of lips separated by u-shaped primary axils without guarding processes, secondary axils demarcated by a shallow incisure, each lip asymmetrically rectangular with a smooth margin, only lateral lips have slender acute tines; three labial probolae, bifurcated at half of their length, prongs bent toward one another apically, “snake-tongue”-shaped; pharyngeal corpus 2.3–3.0 times isthmus length; nerve ring and excretory pore at level of metacorpus to metacorpus-isthmus junction, deirids at level of isthmus; vulva without flap; spermatheca 17–31 µm long; postuterine sac 7–24 µm long; spicules 21.5–23.5 µm long. etymology the species name is derived from the latin lateris (= side) and cornu (= horn), referring to the long acute tine extending along the primary axil on the lateral lips. material examined new zealand: holotype ♀, paratypes 13 ♀♀ and 9 ♂♂, smnh type-8602, 1999, south island, tasman district, abel tasman national park, legit christine (gamble) powell, cultured as the strain stegelleta sp. jb-75. description adult body slightly arcuate in different ways when killed by heat, males often strongly arcuate ventrad in tail end. cuticle coarsely annulated, annuli 1.9–2.7 µm wide at midbody and 2.0–2.4 µm wide in pharyngeal region. cuticle tessellated: longitudinal striae giving it a tiled appearance, 16 rows of blocks at midbody (excluding lateral field). lateral field consisting of two wings separated by a broad groove, appearing as four incisures under lm, occupying about 20–25% of body diameter, extending almost to tail terminus in females and in males. lip region slightly offset, carrying 6 + 4 papillae and two round amphids. three pairs of asymmetrical lips, one dorsal and two ventrolateral. pairs of lips separated by u-shaped primary axils without guarding processes. each pair with a shallow incisure demarcating a secondary axil. each lip asymmetrically rectangular with a smooth margin. lateral lips only with long, slender, acute tine extending along the primary axil. three labial probolae, 7.0–8.5 µm high, bifurcated at half of their length, prongs bent toward one another apically (“snake-tongue”-shaped). stoma somewhat longer than lip region diameter. stomatal parts not clearly discernible. cheilorhabdia bacilliform in lateromedian view; metastegostom with a dorsal denticle. pharynx cephaloboid. pharyngeal corpus cylindrical; isthmus narrow, often wrinkled, and not clearly demarcated from corpus; bulb oval, with valves. nerve ring and excretory pore at level of metacorpus-isthmus junction to isthmus. deirids generally at level of isthmus. female reproductive system monodelphic, prodelphic, in dextral position in relation to intestine. ovary reflexed posteriorly at oviduct, ovary straight posterior to vulva. spermatheca well developed. postvulval uterine http://zoobank.org/urn:lsid:zoobank.org:act:7669b17a-d71a-46c0-8ba0-eb92f28b7111 european journal of taxonomy 87: 1–19 (2014) 10 sac about half to one-and-a-half times vulval body diameter long. vagina about one-third of vulval body diameter. vulval lips not or slightly protruding, without flaps. vulva with genital plug seen in one specimen. in four females one intrauterine egg each was found, measuring 36–57 x 14.5–15.5 µm. tail conoid with 16–22 ventral annuli, terminus broadly rounded or truncate. phasmids located at about onethird to two-fifths of tail length. male reproductive system monorchic, dextral in position; testis reflexed ventrad anteriorly. spicules paired and symmetrical, curved ventrad; with oval manubrium and subcylindrical, gradually narrowing shaft. gubernaculum plate-like, cornua crurum present. tail strongly arcuate ventrad, conoid with rounded terminus. the two outer lateral lines extend posterior to the phasmid, transforming into a cuticular ridge that reaches tail terminus. genital papillae arranged as follows: two pairs subventral precloacal (at 4–6 µm and at 23–28 µm anterior to cloaca), one pair subventral adcloacal, a single midventral papilla on anterior cloacal lip; two pairs (one ventrosublateral and one lateral) at midtail; three pairs (one lateral, one subventral and one dorsosublateral) closer to tail terminus. phasmids located at about two-fifths to half of tail length. remarks the population of stegelleta laterocornuta sp. nov. from new zealand described here agrees in many respects with the descriptions of s. incisa. one major difference is the presence of a long, slender and fig. 3. stegelleta laterocornuta sp. nov., sem micrographs. a. vulval opening. b. anal opening. c–d. anterior end, left subventral view. e. anterior end, left lateral view (arrows in c–e point at the long acute tine extending along the primary axil on the lateral lips). scale bars = 2 µm. boström s. & holovachov o., species of stegelleta and revision of the genus 11 table 2. measurements (in µm) of stegelleta laterocornuta sp. nov. and s. tuarua yeates, 1967 from castlecliff, new zealand, the latter compared to the type population (presented as mean ± s.d. and (range) or only range).* number of annuli from anterior end to nerve ring, excretory pore and deirid, respectively; indicates that data is not available or not applicable. stegelleta laterocornuta sp. nov. stegelleta tuarua population new zealand jb-75 type population castlecliff no. & sex holotype 14 ♀♀ (incl. holotype) 9 ♂♂ {1 aberr.} 1 ♀ 3 ♂♂ 2 ♂♂ body length 447 433 ± 41 (379–512) 428 ± 35 (365–476) 635 563–739 650; 688 body diameter (bd) 20.5 19.1 ± 1.7 (15.5–21.5) 17.4 ± 1.1 (15–19) 45; 46 pharynx length 118 116.7 ± 5.2 (109–126) 117.8 ± 4.6 (109–123) {138} 144; 165 tail length 38 35.9 ± 2.9 (30–42) 24.2 ± 0.9 (22–25) {36} 43; 48 anal or cloacal diam. (abd) 11 11.1 ± 0.8 (9.5–12.5) 11.9 ± 0.4 (11.0–12.5) {13} 30; 30 vulva or testis 273 270 ± 22 (235–315) 223 ± 15 (200–235) 432; 476 v-a/t 3.6 3.6 ± 0.3 (3.0–3.9) a 21.8 22.7 ± 1.7 (18.6–26.0) 24.6 ± 1.3 (21.5–26.5) 21.2 17.2–18.8 14.4; 15.0 b 3.8 3.7 ± 0.3 (3.4–4.3) 3.6 ± 0.2 (3.3–4.0) 3.8 3.9–4.8 4.5; 4.2 c 11.8 12.1 ± 0.7 (10.5–13.0) 17.7 ± 1.4 (15.2–19.8) {13.1} 25.4 14.1–16.0 15.1; 14.3 c’ 3.5 3.2 ± 0.2 (2.7–3.6) 2.0 ± 0.1 (1.8–2.2) {2.8} 1.6 1.6–1.9 1.4; 1.6 v or t (%) 61 62.3 ± 1.2 (60–65) 54.2 ± 3.7 (49–60) 69.2 56.8–62.7 66; 69 lip region diameter 7.0 6.8 ± 0.4 (6.5–8.0) 6.7 ± 0.4 (6–7) 11; 12 stoma length 9.0 8.9 ± 0.3 (8.5–9.5) 8.7 ± 0.5 (8.0–9.5) 14.5; 19 corpus length 70 69.6 ± 4.1 (63–75) 69.6 ± 1.9 (66–72) {81} 92; 110 isthmus length 26 26.7 ± 2.3 (24–30) 27.1 ± 1.7 (24–29) {36} 22; 20.5 bulb length 14.5 14.8 ± 1.0 (13–17) 14.3 ± 0.7 (13.0–15.5) {17} 23; 24 bulb diameter 11 11.5 ± 0.7 (10–13) 11.0 ± 0.7 (10–12) 19; 20.5 corpus/isthmus ratio 2.7 2.6 ± 0.2 (2.3–3.0) 2.6 ± 0.2 (2.4–2.9) {2.3} 4.2; 5.4 nerve ring from ant. end 92 83.0 ± 7.7 (70–96) 86.6 ± 6.4 (77–96) {102} 108; 121 european journal of taxonomy 87: 1–19 (2014) 12 acute tine extending along the primary axil on the lateral lips. this is a somewhat intermediate stage between s. incisa, which has no tines, and s. arenaria, in which each lip has one acute tine extending along the primary axil. another difference is the absence of a cuticular flap on the anterior vulval lip, as seen by sem in the specimens of s. incisa from california. stegelleta ophioglossa andrássy, 1967 fig. 4a–d, table 3 stegelleta ophioglossa andrássy, 1967: 208–210, fig. 3. diagnosis stegelleta ophioglossa is characterised by a 315–490 µm long body in females; cuticle divided by regular longitudinal striations into 12–16 rows of blocks at midbody (excluding lateral field); lateral field with four incisures extending almost to tail terminus in females; three pairs of asymmetrical lips, pairs of lips separated by u-shaped primary axils without guarding processes, secondary axils demarcated by stegelleta laterocornuta sp. nov. stegelleta tuarua population new zealand jb-75 type population castlecliff no. & sex holotype 14 ♀♀ (incl. holotype) 9 ♂♂ {1 aberr.} 1 ♀ 3 ♂♂ 2 ♂♂ excretory pore from ant. end 94 86.3 ± 8.1 (72–97) 92.0 ± 6.6 (81–99) {106} ?; 130 deirid from ant. end 100 92.9 ± 7.2 (81–104) 100.2 ± 5.6 (89–105) {107} 141; 171 rnr* 42 35 ± 4 (28–42) 35 ± 2 (32–39) {47} 35; 38 rep* 43 37 ± 4 (32–43) 37 ± 2 (33–39) {49} ?; 41 rdei* 46 40 ± 4 (35–48) 42 ± 2 (37–44) {50} 46; 55 annuli width at midbody 2.7 2.0–2.7 2.0–2.7 2.4; 2.4 annuli width anteriorly 2.0 2.0–2.4 2.0–2.4 3.0–3.4; 3.0–3.4 vagina or testis flexure length 6.0 6.0 ± 0.6 (5–7) 39.8 ± 3.9 (35–46) 90 (dors); 82 spermatheca or spicule length 18 20.6 ± 4.6 (17–31) 22.5 ± 0.7 (21.5–23.5) 36–40 43; 42 pus or gubernaculum length 17 18.1 ± 5.9 (7–24) 11.4 ± 0.5 (11–12) 21–24 25; 24 pus/vbd 0.9 1.0 ± 0.3 (0.6–1.3) rectum 17 16.6 ± 1.1 (15.5–19.0) rectum/abd 1.5 1.5 ± 0.1 (1.4–1.6) phasmid 12 11.6 ± 0.7 (10–13) 10.6 ± 1.1 (8.5–12.0) {18} 18; 18 phasmid (% of tail) 32 32.2 ± 1.7 (30–37) 44.0 ± 3.6 (38–50) ~40 42; 38 boström s. & holovachov o., species of stegelleta and revision of the genus 13 a shallow incisure, each lip asymmetrically rectangular with a smooth margin and without tines; three labial probolae, bifurcated at half of their length, prongs bent toward one another apically, “snaketongue”-shaped; pharyngeal corpus 3–5 times isthmus length; nerve ring, excretory pore and deirids at level of isthmus; vulva without flap; spermatheca 7–13 µm long; postuterine sac 7–17 µm long. fig. 4. a–d. stegelleta ophioglossa andrássy, 1967. a. pharyngeal region. b. anterior end, surface view. c. female gonad. d. female tail. e–g. stegelleta tuarua yeates, 1967. e. pharyngeal region. f. anterior end, surface view. g. male tail. scale bar = 20 µm. european journal of taxonomy 87: 1–19 (2014) 14 table 3. measurements of females of stegelleta ophioglossa andrássy, 1967 from senegal compared with other populations of the same species (presented as mean ± s.d. and (range) or only range). * calculated from data in original publication. ** number of annuli from anterior end to nerve ring, excretory pore and deirid, respectively; indicates that data is not available or not applicable. population mongolia uzbekistan senegal iran spain senegal reference andrássy 1967 mavljanov 1978 de ley et al. 1990 shokoohi et al. 2008 abolafia et al. 2011 this paper no. & sex ? ♀♀ 19 ♀♀ 8 ♀♀ 4 ♀♀ ? ♀♀ 6 ♀♀ body length 470–490 335–390 347 ± 16 (324–356) 347 ± 17.5 (324–356) 330–420 353 ± 22 (315–382) body diameter (bd) 17.7* 15–18 16.1 ± 1.2 (15–18) 17.9 ± 1.5 (15.5–20.5) pharynx length 114.7* 104 ± 5 (99–111) 93.9 ± 3.2 (90–98) 102.7 ± 3.8 (97–107) tail length 34.8* 31 ± 2 (28–33) 30.0 ± 1.5 (29–32) 27–32 31.6 ± 1.8 (29–34) anal body diam. (abd) 10 9–10 9.6 ± 0.7 (9–10) 10.8 ± 0.6 (9.5–11.5) vulva 253* 219.7 ± 12.6 (209–238) 224 ± 13 (200–243) v-a/t 3.1 ± 0.1 (2.8–3.2) a 20–21 18.1–22.9 21.3 ± 1.2 (19.7–23.1) 21.6 ± 0.6 (20.8–21.9) 19–23 19.8 ± 0.8 (18.6–20.4) b 3.3–3.4 3.1–3.6 3.3 ± 0.1 (3.2–3.5) 3.7 ± 0.1 (3.7–3.8) 3.2–3.6 3.4 ± 0.2 (3.2–3.7) c 11–12 10.4–11.8 11.2 ± 0.6 (10.4–12.9) 11.6 ± 0.2 (11.3–11.8) 12–13 11.2 ± 0.4 (10.5–11.6) c’ 3.0–3.5 3.3 ± 0.2 (3.0–3.6) 3.1 ± 0.3 (2.9–3.4) 2.3–2.7 2.9 ± 0.2 (2.6–3.2) v (%) 60–62 63.0–66.8 61–63 63.2 ± 0.6 (63–64) 62–63 63.5 ± 0.8 (63–65) lip region diameter 4.9 ± 0.7 (4–6) 6.6 ± 0.3 (6–7) stoma length 7–9 7.8 ± 1.2 (6–8) 8.8 ± 0.3 (8.5–9.0) corpus length 67 ± 3 (63–71) 63.5 ± 3.6 (60–68) 65.9 ± 2.9 (61.5–69.0) isthmus length 22 ± 2 (18–25) 14.7 ± 2.8 (12–18) 15.9 ± 1.1 (14.5–17.0) bulb length 13–14 16.0 ± 0.9 (16–17) 14.1 ± 0.9 (12.0–14.5) bulb diameter 11.3 ± 0.9 (9.5–12.0) corpus/isthmus ratio 3 3.8–5.0 3 4.2 ± 0.2 (4.0–4.4) nerve ring from ant. end 68 ± 4 (60–71) 67.7 ± 2.9 (64–71) 76 ± 3.6 (69–81) boström s. & holovachov o., species of stegelleta and revision of the genus 15 material examined senegal: 6 ♀♀, smnh 135943–135944, kaolack region, thyssé-kaymor, legit c. villenave. description adult body slightly arcuate ventrad when killed by heat. cuticle annulated, annuli 1.6–2.2 µm wide at midbody and 1.7–2.0 µm wide in pharyngeal region. cuticle tessellated: longitudinal striae giving it a tiled appearance, 16 rows of blocks at midbody (excluding lateral field). lateral field consisting of two wings separated by a narrow groove, appearing as four incisures under lm, occupying about 20% of body diameter, extending almost to tail terminus in females. lip region slightly offset, carrying 6 + 4 papillae and two round amphids. three pairs of asymmetrical lips, one dorsal and two ventrolateral. pairs of lips separated by u-shaped primary axils without guarding processes. each pair with a shallow incisure demarcating a secondary axil. each lip asymmetrically rectangular with a smooth margin. three labial probolae, 6–7 µm high, bifurcated at half of their length, prongs bent toward one another apically (“snake-tongue”-shaped). stoma somewhat longer than lip region diameter. stomatal parts not clearly discernible. cheilorhabdia bacilliform in lateromedian view; metastegostom with a dorsal denticle. pharynx cephaloboid. pharyngeal corpus cylindrical; isthmus narrow, not clearly demarcated from corpus; bulb oval, with valves. nerve ring, excretory pore and deirids at level of isthmus. population mongolia uzbekistan senegal iran spain senegal excr. pore from ant. end 66 ± 3 (61–71) 69.4 ± 3.6 (70–74) 79.5 ± 3.5 (72–83) deirid from ant. end 75 ± 2 (72–78) 73.6 ± 3.3 (70–78) 82.3 ± 4.0 (75–87) rnr** 36 ± 1 (35–37) rep** 36–39 37 ± 1 (36–38) rdei** 43 39 ± 1 (37–40) annuli width at midbody 1.7–2.0 2.1 1.5–2.1 1.7–2.3 1.6–2.2 annuli width anteriorly 1.5 1.7–2.0 vagina length 3–5 6.2 ± 0.6 (6–7) 5.8 ± 0.3 (5.5–6.0) spermatheca length ≤ 13 8.1 (n=1) 13 8.9 ± 1.7 (7–12) pus length 11–14 13.6 ± 4.9 (8–17) 10 7.0–8.5 pus/vbd ~1 1.1–1.4 0.5–0.9 0.4–0.5 rectum 13 ± 2 (10–16) 12.7 ± 0.8 (12–14) 12.7 ± 1.5 (9.5–14.5) rectum/abd 1.7–2.0 1.3–1.4 > 1 1.2 ± 0.1 (1.0–1.3) phasmid 8.8 ± 0.4 (8.5–9.5) phasmid (% of tail) ~30 21–29 24–31 33–46 27.5 ± 2.4 (25–32) rows of cuticle blocks 14 16 16 12 16 european journal of taxonomy 87: 1–19 (2014) 16 female reproductive system monodelphic, prodelphic, in dextral position in relation to intestine. ovary reflexed posteriorly at oviduct, ovary straight posterior to vulva. spermatheca small, not developed. postvulval uterine sac short, about one-half of vulval body diameter long. vagina about one-third of vulval body diameter. vulval lips not protruding. tail conoid with 12–20 ventral annuli, terminus truncate. phasmids located at about one-fourth to one-third of tail length. male not found. remarks the specimens described here agree in many respects with the population of stegelleta ophioglossa from senegal described by de ley et al. (1990) and several other populations of s. ophioglossa (andrássy 1967; mavljanov 1978; shokoohi et al. 2008; abolafia et al. 2011). stegelleta tuarua yeates, 1967 fig. 4e–g, table 2 stegelleta tuarua yeates, 1967: 536–538, fig. 5. diagnosis stegelleta tuarua is characterised by a 635 µm long body in females and 563–739 µm in males; cuticle divided by regular longitudinal striations into 40–44 rows of blocks at midbody; lateral field with five incisures extending almost to tail terminus in females and in males; three pairs of asymmetrical lips, pairs of lips separated by u-shaped primary axils without guarding processes, secondary axils demarcated by a shallow incisure, each lip asymmetrically rectangular with a smooth margin and one tine; three labial probolae, bifurcated at about 1/5 of their length; pharyngeal corpus 4.2–5.4 times isthmus length; nerve ring and excretory pore at level of isthmus and deirids at level of bulb; vulva and spermatheca undescribed, postuterine sac about 1 vulval body diameter long; spicules 21–25 µm long. material examined new zealand: 2 ♂♂, smnh 135945–135946, 18 may 2008, north island, wanganui, castlecliff beach, partly stabilized coastal sand dunes with ammophila arenaria, 50 cm deep, legit g. yeates. description adult body slightly arcuate ventrad when killed by heat. cuticle annulated, annuli 2.4 µm wide at midbody and 3.0–3.4 µm wide in pharyngeal region. cuticle tessellated: longitudinal striae giving it a tiled appearance, tiles are not equal in size and are not arranged in straight rows, approximately 40–44 rows of blocks at midbody. lateral field with five incisures occupying about 20% of body diameter, extending almost to tail terminus in females. lip region slightly offset, carrying 6 + 4 papillae and two round amphids. three pairs of asymmetrical lips, one dorsal and two ventrolateral. pairs of lips separated by u-shaped primary axils without guarding processes. each pair with a shallow incisure demarcating a secondary axil. each lip asymmetrically rectangular with a smooth margin and one acute tine extending along the primary axil. three labial probolae, 7.0–8.5 µm high, bifurcated at about 1/5 of their length. stoma somewhat longer than lip region diameter. stomatal parts not clearly discernible. cheilorhabdia bacilliform in lateromedian view; metastegostom with a dorsal denticle. pharynx cephaloboid. pharyngeal corpus cylindrical; isthmus narrow, not clearly demarcated from corpus; bulb oval, with valves. nerve ring and excretory pore at level of isthmus and deirids at level of bulb. boström s. & holovachov o., species of stegelleta and revision of the genus 17 male reproductive system monorchic, dextral in position; testis reflexed ventrad anteriorly. spicules paired and symmetrical, curved ventrad; with oval manubrium and subcylindrical, gradually narrowing shaft. gubernaculum wedge-shaped, cornua crurum present. tail strongly arcuate ventrad, conoid with rounded terminus. the two outer lateral lines extend posterior to the subdorsal papilla close to tail terminus, transforming into a cuticular ridge that reaches tail terminus. genital papillae arranged as follows: two pairs subventral precloacal (at 36 µm and at 74 µm anterior to cloaca), one pair subventral adcloacal, a single midventral papilla on anterior cloacal lip; two pairs (one ventrosublateral and one lateral) at midtail; three pairs (one lateral, one subventral and one dorsosublateral) closer to tail terminus. phasmids located at about two-fifths to half of tail length. female not found in our study. remarks the specimens described here agree well in main morphological and morphometric features with the original description by yeates (1967) of the males of s. tuarua from new zealand. key to species 1. lateral field with five incisures; cuticle with over 40 longitudinal rows of blocks; labial probolae biacute apically ...............................................................................................s. tuarua yeates, 1967 – lateral field with three-four incisures; cuticle with 12–26 longitudinal rows of blocks; labial probolae bifurcate half of length ...................................................................................................................... 2 2. labial probolae with minute secondary bifurcations at tips ...........................s. iketaia yeates, 1967 – labial probolae without minute secondary bifurcations at tips ........................................................ 3 3. cephalic probolae in shape of acute tines present on all six lips along the primary axils; cuticle with 24–26 longitudinal rows of blocks ...................................s. arenaria boström & holovachov, 2012 – cephalic probolae absent, or present only on lateral lips along the primary axils; cuticle with 12–22 longitudinal rows of blocks ............................................................................................................... 4 4. cuticle with 22 longitudinal rows of blocks ......................................... s. georgica bagaturija, 1973 – cuticle with 12–16 longitudinal rows of blocks ............................................................................... 5 5. parthenogenetic species; spermatheca 7–13 µm long; postuterine sac about 7–17 µm long ............. ............................................................................................................s. ophioglossa andrássy, 1967 – amphimictic species; spermatheca 24–43 µm long; postuterine sac about 22–36 µm long ............ 6 6. cephalic probolae absent ............................................................................. s. incisa (thorne, 1937) – cephalic probolae setose, present on lateral lips only ................................ s. laterocornuta sp. nov. discussion representatives of the family cephalobidae filipjev, 1934 are mostly terrestrial and bacteria-consuming nematodes with a worldwide distribution, including antarctica. they occur in tropical and temperate regions as well as in hot and cold arid areas globally. they seem to be especially diverse and abundant in deserts and many species have been described from warm and dry habitats like the namib desert in southern africa (see, e.g., rashid & heyns 1990a, b; rashid et al. 1990a, b) and the mojave desert in southern california (see, e.g., de ley et al. 1999; taylor et al. 2004; waceke et al. 2005). sand dunes european journal of taxonomy 87: 1–19 (2014) 18 appear to be another suitable habitat for cephalobids, as evidenced by several studies (see, e.g., boström & holovachov 2012, 2013a, b; bussau 1991; orselli & vinciguerra 2002; yeates 1967). species of the genus stegelleta are rather rare inhabitants of terrestrial habitats, although they occur on all continents except antarctica. in the present study, a new species of stegelleta, s. laterocornuta sp. nov., is described from new zealand, which brings the total number of species in this genus to seven. descriptions of new material of some already known species from dry areas in california and senegal are also included. this leads to an addition of morphological data which broadens the diagnosis of stegelleta and increases the number of character combinations useful for species identification in the genus. acknowledgements the second author was supported in part by an award from the nsf partnerships for enhancing expertise in taxonomy (peet) program grant “peet: training the next generation of nematode taxonomists: applying the tools of modern monography across free-living and parasitic tylenchina”(deb-0731516). sampling in the mojave desert was performed by o. holovachov and p. de ley under the permit # moja2010-sci-0003 from the national park service, united states department of interior. we thank d. sturhan, g. yeates and c. villenave for providing us with samples and slides, and d. bumbarger for sem pictures of stegelleta jb-75 ( = s. laterocornuta sp. nov.). references abolafia j., guerrero-rodríguez p. & peña-santiago r. 2011. nematoda, rhabditida. fauna ibérica 34, museo nacional de ciencias naturales, madrid. andrássy i. 1967. ergebnisse der zoologischen forschungen von dr. z. kaszab in der mongolei. 92. weitere bodennematoden aus den jahren 1964 und 1965. opuscula zoologica budapestinensis 6 (2): 203–233. boström s. & holovachov o. 2012. description of chilodellus eremus gen. n., sp. n. and stegelleta arenaria sp. n. 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(acrobelinae, rhabditida), a new species of nematodes from cotton. zoologicheskii zhurnal 57: 1889–1891 (in russian). http://www.nrm.se/download/18.9ff3752132fdaeccb6800015606/cephaloboidea%5b1%5d.pdf http://www.nrm.se/download/18.9ff3752132fdaeccb6800015606/cephaloboidea%5b1%5d.pdf boström s. & holovachov o., species of stegelleta and revision of the genus 19 nadler s.a., de ley p., mundo-ocampo m., smythe a.b., stock s.p., adams b.j., de ley i.t., holovachov o. & baldwin j.g. 2006. phylogeny of cephalobina (nematoda): molecular evidence for recurrent evolution of probolae and incongruence with traditional classifications. molecular phylogenetics and evolution 40 (3): 696–711. http://dx.doi.org/10.1016/j.ympev.2006.04.005 orselli l. & vinciguerra m.t. 2002. nematodes from italian sand dunes. 6. two new and three rare species of cephalobidae (nematoda). nematologia mediterranea 30: 211–220. rashid f. & heyns j. 1990a. chiloplacus and macrolaimellus species from south west africa/namibia (nematoda: cephalobidae). phytophylactica 22 (2): 189–199. rashid f. & heyns j. 1990b. description of namibinema scaphovulva n. gen., n. sp. and zeldia punctata (thorne, 1925) from namibia (nematoda: cephalobidae). phytophylactica 22 (4): 397–403. rashid f., heyns j. & coomans a.1990a. paracrobeles and acrobeles species from south west africa/ namibia with a description of a new acrobeles species (nematoda: cephalobidae). phytophylactica 22 (1): 41–49. rashid f., heyns j. & coomans a.1990b. species of seleborca from south west africa/namibia (nematoda: cephalobidae). phytophylactica 22 (1): 51–62. shokoohi e., abolafia j., kheiri a. & zad j. 2008. nematodes of the order rhabditida from tehran province (iran). some known species of the family cephalobidae. journal of nematode morphology and systematics 11 (1): 67–85. taylor t.m., baldwin j.g. & mundo-ocampo m. 2004. paracrobeles mojavicus sp. n. (nematoda: cephalobidae) from the mojave desert, california. journal of nematode morphology and systematics 6 (2): 151–160. thorne g. 1937. a revision of the nematode family cephalobidae chitwood and chitwood, 1934. proceedings of the helminthological society of washington 4 (1): 1–16. thorne g. 1938. notes on free-living and plant-parasitic nematodes. iv. proceedings of the helminthological society of washington 5 (2): 64–65. waceke j.w., bumbarger d.j., mundo-ocampo m., subbotin s.a. & baldwin j.g. 2005. zeldia spannata sp. n. (nematoda: cephalobidae) from the mojave desert, california. journal of nematode morphology and systematics 8 (1): 57–67. yeates g.w. 1967. studies on nematodes from dune sands. 5. acrobelinae. new zealand journal of science 10 (2): 527–547. manuscript received: 19 february 2014 manuscript accepted: 23 april 2014 published on: 16 june 2014 topic editor: rudy jocqué desk editor: charlotte thionois printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark. http://dx.doi.org/10.1016/j.ympev.2006.04.005 european journal of taxonomy 159: 1–6 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2015.159 www.europeanjournaloftaxonomy.eu 2015 · ricchiardi e. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:2971b804-8aa2-4713-8c66-756640d0331a 1 description of a new monospecific genus of south african trichiina, with a key to the related genera (coleoptera: scarabaeidae) enrico ricchiardi corso a. tassoni 79/4, 10143 torino, italy. e-mail: alericor@fastwebnet.it urn:lsid:zoobank.org:author:f09bcf76-35de-4821-91c4-eedc5114810a abstract. a new monospecific genus, pseudostegopterus gen. nov., endemic to the northwestern region of south africa, is erected. the type species is described as pseudospegopterus melonthinoides sp. nov. and is currently known only from male specimens. a provisional dichotomic key of the african trichiina genera is also provided, in order to facilitate the identification of male specimens to the genus level. key words. cetoniinae, pseudostegopterus, afrotropical region, new genus, new species. ricchiardi e. 2015. description of a new monospecific genus of south african trichiina, with a key to the related genera (coleoptera: scarabaeidae). european journal of taxonomy 159: 1–6. http://dx.doi.org/10.5852/ejt.2015.159 introduction the number of described trichiina species (sensu smith et al. 2006) endemic to south africa has increased substantially during the last two decades (evans 1987; ricchiardi 1997, 1998, 1999, 2000; ricchiardi et al. 2004, 2008; sipek et al. 2012; ricchiardi & perissinotto 2013, 2014) and currently ranges around 40. these species are grouped into nine genera, with three of them regarded as tropical intruders and represented in south africa by one single species each. the biodiversity richness of this country is well established, but many taxa remain as yet underscribed, among them several trichiina. while undertaking a close analysis of the specimens preserved at the iziko south african museum (isam) in cape town and in my private collection, i have recently identified a series of eight males belonging to an undescribed species, which superficially resembles members of the genus stegopterus burmeister & schaum, 1840. however, on closer scrutiny it is clear that a new genus is involved. both genus and species are hereby described as pseudostegopterus gen. nov. and p. melolonthinoides gen. et sp. nov. the species is currently known only from the two type localities of elands bay and pella mission in the western cape, south africa. material and methods the following abbreviations are used to denote the housing location of the study material: erc = enrico ricchiardi collection, torino, italy isam = iziko south african museum, cape town, south africa http://dx.doi.org/10.5852/ejt.2015.159 https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:2971b804-8aa2-4713-8c66-756640d0331a mailto:alericor%40fastwebnet.it?subject= http://zoobank.org/urn:lsid:zoobank.org:author:f09bcf76-35de-4821-91c4-eedc5114810a http://dx.doi.org/10.5852/ejt.2015.159 european journal of taxonomy 159: 1–6 (2015) 2 the specimen length was measured between the apex of the pygidium and the anterior margin of the pronotum. the specimen width is the maximum elytron width. the clypeus length was measured laterally, between the frontal margin and the antennal basal attachment. photographs were taken with a nikon coolpix p7700, sometimes attached to one of the eyepieces of a wild dissecting microscope. photographs were processed with photo stacking software (zerene stacker version 1.04 build t201411272115, http://www.zerenesystems.com [accessed 13 nov. 2015]). finally, backgrounds were removed from photos using gimp 2.8.14, in order to increase contrast. results class insecta order coleoptera superfamily scarabaeoidea latreille, 1802 family scarabaeidae latreille, 1802 subfamily cetoniinae leach, 1815 tribe trichiini fleming, 1821 subtribe trichiina fleming, 1821 discussion this work is part of a series of studies aimed at identifying and clarifying the composition and morphological relations of the exceptionally diverse south african trichiina. the phylogenetic relationships among the trichiina of south africa and those occurring in other parts of the continent are yet to be investigated and in need of further species descriptions and better definition of the genera. as a contribution to the understanding of the morphological relationship between the south african genera of trichiina an updated dichotomic key is here proposed. preliminary key to south african trichiina genera 1. abdominal tergites not covered by elytra in dorsal view ................... calometopus blanchard, 1850 – abdominal tergites covered by elytra in dorsal view ........................................................................ 2 2. elytral lateral ridge near lateral margin absent .................................................................................. 3 – elytral lateral ridge present on most of lateral margin ...................................................................... 4 3. posterior pronotal corners widely rounded; mesosternal process present, but not visible in side view ...................................................................................myodermum burmeister & schaum, 1840 – posterior pronotal corners rounded or angled; mesosternal process absent .........diploa kolbe, 1892 4. first metatarsal segment longer than second ..................................................................................... 5 – first metatarsal segment approximately as long as second ............................................................... 8 5. pronotum length > than 0.5 times elytral length; body large (length 20.3 mm, width 11.0 mm; glabrous, slightly shiny, black...............................................................camapterus ricchiardi, 2000 – pronotum length < than 0.5 times elytral length; body smaller, orange-red, but female often black or dark brown ......................................................................................................................................... 6 6. clypeus as wide as long ............................................................................... campulipus kirby, 1827 – clypeus wider than long. posterior pronotal margin not emarginate ................................................ 7 ricchiardi e., description of pseudostegopterus melolonthinoides 3 7. pronotum covered with very scattered setae (even if locally dense); female with normal wings ....... ...................................................................................................stripsipher gory & percheron, 1833 – pronotum covered with dense, long setae; male elytra light brown; female smaller, completely black or dark brown, brachypterous .......................................... eriopeltastes burmeister & schaum, 1840 8. metatibial apex modified ................................................................................................................... 9 – metatibia apex unmodified .............................................................................................................. 10 9. metatibial apex of male much longer than single metatibial spur; female metatibial apices truncate with two spurs ..........................................................................................brachagenius kraatz, 1890 – metatibial apex slightly pronouced in males, truncate in females, with two spurs ............................. ......................................................................................................................elpidus péringuey, 1907 10. metafemura enlarged (maximum width larger than 0.5 its length); length of protarsal claws > 0.8 that of last tarsal segment ...................................................................... pseudostegopterus gen. nov. – metafemora slender (width less than 0.25 its length); length of protarsal claws at least 0.7 that of last tarsal segment .................................................................... stegopterus burmeister & schaum, 1840 genus pseudostegopterus gen. nov. urn:lsid:zoobank.org:act:3feb8498-b2a1-4136-904a-7eb7f37ce568 type species pseudostegopterus melolonthinoides gen. et sp. nov. diagnosis this new genus differs from all other trichiini genera occurring in south africa by the following combination of diagnostic characters: 1) the eye-canthus covers about ½ of the ocular hemisphere; 2) the frons and vertex are covered with thick, long, inclined, testaceous setae; 3) the length of the pronotum is < 0.5 times that of the elytra; 4) the posterior pronotal margin does not exhibit any ridge centrally; 5) the pronotal surface is generally covered with thick, long, inclined, testaceous setae; 6) the pronotal posterior corners are obtuse; 7) the elytral apex is rounded; 8) the elytral lateral ridge is visible on the entire lateral border; 9) the elytra cover the entire pygidium, except its very apex; 10) the length of protarsal claws is 0.70–0.90 times that of the last tarsomere; 11) the mesotibia are slightly curved outwardly; 12) the metatibia exhibits two spurs; 13) the first metatarsal segment is as long as the second or shorter; 14) the metatibial apex is strongly enlarged; 15) the internal corner of the metatibial apex is deprived of long spines; 16) the mesosternal process is absent; 17) the metafemur width is 0.4–0.45 times its length; 18) the males are fully winged. etymology the name comes from its superficial resemblance to some representatives of the genus stegopterus burmeister & schaum, 1840. pseudostegopterus melolonthinoides sp. nov. urn:lsid:zoobank.org:act:42397679-9840-4330-aedd-a9ee3cdd955b fig. 1 etymology while very similar to the smaller species of the genus stegopterus, this species also resembles in color and shape a small melolonthinae. http://zoobank.org/urn:lsid:zoobank.org:act:3feb8498-b2a1-4136-904a-7eb7f37ce568 http://zoobank.org/urn:lsid:zoobank.org:act:42397679-9840-4330-aedd-a9ee3cdd955b european journal of taxonomy 159: 1–6 (2015) 4 type material holotype republic of south africa: ♂ (isam), western cape, leipoldtville, eland’s bay (32°18’48” s, 18°21’00” e), oct. 1947, museum expedition (sam-co-a043595). paratypes republic of south africa: 6 ♂♂ (isam), same data as the holotype (but without sam number); 1 ♂ (erc), western cape, pella mission (33°37’30” s, 18°37’30” e), 14 aug. 1947, n.j. duke legit. description holotype measurements. length: 10.1 mm; width: 5.4 mm. head. clypeus black, margins reflexed (fig. 1c), anterior margin not sinuate, covered with thick, large punctures and long, recumbent, testaceous setae; vertex and frons black. antennae. brown, with three lamellae, darkening at apex, slightly curved, about 2.0 times the length of clypeus. fig. 1. pseudostegopterus melolonthinoides gen. et sp. nov. holotype, ♂ (isam). a. dorsal habitus. b. ventral view. c. head. d. parameres, frontal view. e. parameres, lateral view. ricchiardi e., description of pseudostegopterus melolonthinoides 5 pronotum. black, slightly shiny and rugulose, wider than long; anterior angles rounded, posterior acute, with slightly rounded tip; posterior margin without carina; surface covered with thick, long, reclining, testaceous setae. scutellum. black, slightly shiny, triangular, as long as wide, apex slightly rounded; covered with scattered, rounded punctures. elytron. mostly fulvous, transparent, glabrous, with a large black stripe along the suturae, larger at elytral apex; both striae and interstriae effaced; apicosutural angle rounded; sutural half of elytral apex scabrous. abdomen. sternites black, shiny; covered with reclining, long, testaceous setae, becoming glabrous and matt on anal sternite; tergites black, covered with same setae as sternites. propygidium. black, not visible, covered by elytron. pygidium. black, rugulose, longer than wide, triangular, with apex rounded; covered with scattered, long, reclining, light testaceous setae. protibia. brown, glabrous, shiny, with two robust teeth on external margin. mesotibia. black, slightly curved outwardly, with strong tooth at middle of external margin; proximal half covered with scattered, long, reddish, bristles; apex strongly enlarged; mesofemur black, shiny, covered with testaceous, reclined setae. metatibia. black, shiny, with apex strongly enlarged; covered with scattered, long, reddish, bristles; metatarsomeres black, twice as long as metatibia (claws included); first segment about as long as second; spurs long, with apex rounded, especially on ventral one; dorsal spur 1.2 times longer than ventral and arcuate; metafemur enlarged, width 0.5 times its length. remarks the female is unknown. type series variability some paratypes have darker elytra and/or scattered instead of dense setae on head and pronotum, which does not seem to represent simply wear and tear due to age. distribution the single species belonging to this new genus is known from two places in the western cape province. these two localities are especially relevant, as they are part of a unique biome, the fynbos, which is seriously threatened by climate change and land use. acknowledgements i thank dawn larsen and simon van noort of isam (cape town) and renzo perissinotto of the nelson mandela metropolitan university (port elizabeth) for kindly organizing the loan of material for this study. references evans a. 1987. new species of the genus brachagenius kraatz (= xiphoscelidus peringuey syn. nov.) with notes on the synonymy of elpidus hopei (burmeister) (coleoptera: scarabaeidae: trichiinae). journal of the entomological society of southern africa 50 (2): 351–356 european journal of taxonomy 159: 1–6 (2015) 6 ricchiardi e. 1997. notes on south african trichiini. on the genus eriopeltastes burmeister & schaum, 1840, with description of three new species (coleoptera: cetoniidae: trichiinae). elytron 11: 121–132. ricchiardi e. 1998. notes for the revision of the genus stripsipher gory & percheron, 1833, with descriptions of four new species (coleoptera, cetoniidae, trichiinae, trichiini). mitteilungen der münchner entomologischen gesellschaft 88: 45–64. ricchiardi e. 1999. description of three new species and the brachypterous females of the genus eriopeltastes burmeister & schaum, 1840 (=diploeida péringuey, 1907), (coleoptera: cetoniidae: trichiinae). elytron 11: 133–147. ricchiardi e. 2000. camapterus, a new brachypterous trichiini genus from south africa (coleoptera, cetoniidae, trichiinae). elytron 14: 201–206. ricchiardi e., perissinotto r. & clennell l. 2004. parapeltastes, a new subgenus of eriopeltastes burmeister & schaum (1840) from south africa, with description of a new species (coleoptera, cetoniidae, trichiinae). lambillionea 114: 223–229. ricchiardi e., perissinotto r. & clennell l. 2008. taxonomic revision of the south african genus stripsipher gory & percheron, 1833, with descriptions of four new species (coleoptera, cetoniidae, trichiinae). bollettino della società entomologica italiana 140 (3): 155–178. ricchiardi e. & perissinotto r. 2013. description and ecology of a new species of eriopeltastes burmeister & schaum 1840, from kwazulu-natal (south africa). zootaxa 3630 (2): 379–384. http:// dx.doi.org/10.11646/zootaxa.3630.2.12 ricchiardi e. & perissinotto r. 2014. new species of eriopeltastes burmeister & schaum, 1840 (coleoptera: scarabaeidae: cetoniinae: trichiini), from south africa. zookeys 422: 103–113. http:// dx.doi.org/10.3897/zookeys.422.7830 sipek p., ricchiardi e. & perissinotto r. 2012. immature stages and ecology of two species of the south african genus stripsipher gory & percheron, 1833 (coleoptera: scarabaeidae; trichiinae). zookeys 180: 19–40. http://dx.doi.org/10.3897/zookeys.180.2315 smith a.b.t., hawks d.c. & heraty j.m. 2006. an overview of the classification and evolution of the major scarab beetle clades (coleoptera: scarabaeoidea) based on preliminary molecular analyses. papers in entomology 121: 34–46. manuscript received: 2 september 2015 manuscript accepted: 28 october 2015 published on: 7 december 2015 topic editor: koen martens desk editor: charlotte thionois printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark. http://dx.doi.org/10.11646/zootaxa.3630.2.12 http://dx.doi.org/10.11646/zootaxa.3630.2.12 http://dx.doi.org/10.3897/zookeys.422.7830 http://dx.doi.org/10.3897/zookeys.422.7830 http://dx.doi.org/10.3897/zookeys.180.2315 1 european journal of taxonomy 138: 1-8 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2015.138 www.europeanjournaloftaxonomy.eu 2015 · jongkind c.c.h. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e rhaphiostylis minima jongkind (icacinaceae), a new liana species from ivory coast & liberia carel c.h. jongkind botanic garden meise, nieuwelaan 38, 1860 meise, belgium. e-mail: carel.jongkind@kpnmail.nl abstract. a new species of rhaphiostylis, r. minima sp. nov. from liberia and ivory coast, is described and illustrated here. it is characterised by its comparatively small size, an ovate to lanceolate leaf blade with long drip tip and a partly glabrous ovary that is in shape in between those of r. elegans engl. and r. preussii engl. rhaphiostylis elegans is lectotypified and illustrated here. keywords. rhaphiostylis, tropical forest, taxonomy, conservation. jongkind c.c.h. 2015. rhaphiostylis minima jongkind (icacinaceae), a new liana species from ivory coast & liberia. european journal of taxonomy 138: 1–8. http://dx.doi.org/10.5852/ejt.2015.138 introduction rhaphiostylis is an icacinaceae genus of ca. 10 liana species endemic to tropical africa (boutique 1960, villiers 1973a, 1973b). only 4 species were known from the upper guinean flora subregion (the closed forest from senegal to togo) where the new species was discovered, r. beninensis (hook.f. ex planch.) planch. ex benth., r. cordifolia hutch. & dalziel, r. ferruginea engl. and r. preussii engl. (hepper 1958: 638; hawthorne & jongkind 2006: 346). in 2013 a small rhaphiostylis climber with glabrous leaves and branches was found flowering and fruiting in the evergreen forest in the south-east of liberia. with the small and slender leaves with a long drip tip it is already at first sight different from r. preussii. rhaphiostylis preussii is the only other rhaphiostylis with glabrous leaves often found in the undergrowth in that forest area. later it became clear that the shape and indumentum of the ovary is also distinct (fig. 1f). in rhaphiostylis the leaf shape and the indumentum of the ovary are important characters for identification of the species (hepper 1958). the new species, r. minima sp. nov., most resembles r. elegans engl. from cameroon, being also a smaller liana with more or less similar leaves. the new species differs from r. elegans by its less stretched and more hairy ovary (fig. 1f, table 1). moreover, there exists also a large geographical gap between the two. the here illustrated flower of r. elegans shows the stretched ovary with a long narrow middle part, while the ovaries of the two r. preussii flowers, from ivory coast and gabon, have only a narrow constriction. the ovary of r. minima sp. nov. is in shape and indumentum in between r. elegans and r. preussii. the ovary of the only other rhaphiostylis species from upper guinea with glabrous leaves and branches, r. beninensis, is in shape and size more or less equal to the one of r. preussii but it is glabrous or almost so (villiers 1973a: 35). the flower of r. preussii, illustrated by villiers (1973a: 35) after le testu 7491 from gabon, is similar to that of the specimens illustrated here. http://dx.doi.org/10.5852/ejt.2015.138 http://www.europeanjournaloftaxonomy.eu https://creativecommons.org/licenses/by/3.0/ mailto:carel.jongkind@kpnmail.nl http://dx.doi.org/10.5852/ejt.2015.138 european journal of taxonomy 138: 1–8 (2015) 2 the new species is named here r. minima sp. nov., it being the smallest species in the genus. all specimens are small climbers only a few meters high, much smaller than all other rhaphiostylis species from upper guinea. except for r. preussii, the hairy r. cordifolia is the only other rhaphiostylis species found in the same evergreen forests. rhaphiostylis beninensis and r. ferruginea are growing in drier forest types like semi-deciduous forest (hawthorne & jongkind 2006: 3). in a table (table 1) all rhaphiostylis species with glabrous leaves and branches from upper guinea and r. elegans are compared. when preparing the icacinaceae issues of the floras of cameroon and gabon, villiers could not find type material of r. elegans and could not place it (villiers 1973a: 41). the original description (engler 1909: 184) was based on fruiting material only and the flowers were never described, this made it difficult to understand the taxon. in 1986 a duplicate of the destroyed holotype was found in the mo herbarium by al gentry and duncan thomas (noted on the sheet). during the research on the new species it was possible to match this isotype of r. elegans in herbaria with other specimens. in this publication an illustration is included from a newly identified flowering specimen (fig. 1). in the fresh flowers of r. preussii and r. beninensis the flat filaments touch each other along the edge and together they close around the ovary from their base to about half their length (figs 2, 3). it is likely that the stamens of r. minima sp. nov. and those of other species in the genus share this character because they also share the special shape of the filaments. materials and methods the new species was studied and collected in 2013 and 2014 by the author in the forest in the southeast of liberia. the br, k, p and wag herbaria were searched for additional specimens of the new species and r. elegans. the herbarium, where the specimens are located, is indicated by the international code (herbarium acronym) registered in index herbariorum (thiers continuously updated). preliminary assessments of the iucn red list categories of threat were performed using the iucn criteria and the royal botanic gardens kew website http://geocat.kew.org. results order “unplaced” (kårehed 2001) family icacinaceae miers genus rhaphiostylis planch. ex benth. key to the species in upper guinea (forest area west of nigeria) 1. leaves and twigs of plant almost glabrous .......................................................................................... 2 – leaves and twigs of plant densely appressed or erect hairy, at least on the young parts .................... 4 2. ovary for most of its surface, and the fruit around the base of the style, hairy ................................... 3 – ovary glabrous or with a small patch of hairs on top, fruit glabrous .................................................. ....................................................................... r. beninensis (hook.f. ex planch.) planch. ex benth. 3. constricted middle of the ovary glabrous. all leaves with drip tip 1.5–3 cm long. slender liana up to 4 m high .................................................................................................................. r. minima sp. nov. – constricted middle of the ovary hairy. drip tip on leaves, if present, shorter. usually a larger liana ............................................................................................................................. r. preussii engl. http://geocat.kew.org jongkind c.c.h., a new rhaphiostylis species (icacinaceae) from africa 3 4. leafbase rounded to cordate. flower pedicel up to 3 mm long. in evergreen forest ............................ .............................................................................................................. r. cordifolia hutch. & dalziel – leafbase cuneate or obtuse. flower pedicel 6 to 9 mm long. mainly in semideciduous forest .............................................................................................................................. r. ferruginea engl. rhaphiostylis minima jongkind sp. nov. urn:lsid:ipni.org:names:77150009-1 table 1, figs 1d–g, 2 diagnosis slender woody liana up to 4 m high, resembling r. elegans engl. from cameroon in leafshape. differing from r. elegans in the first place by its shorter and more hairy ovary, 2.5–3 mm versus 3.5–4 mm. type ivory coast. ca. 140 km n of tabou, crossing hana river, on the way to taï, fl., 10 oct. 1963, w. de wilde 1061 (holo-: wag; iso-: br, k n.v.) description slender woody liana going up to 3–4 m high. leaves alternate, glabrous; slender petiole 2–3 mm long; blade ovate to lanceolate, 5–10 cm long and 1.5–3.5 cm wide, papery, midrib impressed above, 3 or 4 pairs of main laterals of which the first are often long and ascending, margin entire, base rounded, apex long acuminate, up to 3 cm long with a spathulate tip. inflorescences fasciculate, supra-axillary with 1–3 flowers. flowers bisexual; pedicel 2–4 mm long; calyx 5 lobed, lobes about 1 mm long, puberulous; petals 5, ca. 5 mm long, valvate, free, glabrous. stamens 5, glabrous; filaments subulate, gradually narrowing to filiform above, about 4 mm long, alternipetalous; anthers ca. 1 mm long. ovary 2.5–3 mm high, hairy on the lower ½ and at the top, almost glabrous on the narrow part in between; style 3.5–4 mm long, excentric but erect. fruit 2-lobed, flattened through the longitudinal axis, fleshy with thin exocarp, smooth and glabrous, but conspicuously hairy on and around the base of the persistent style; endocarp woody, reticulate, about 1.5 cm wide and 1 cm high. distribution and habitat south-east liberia and south-west ivory coast below 250 m altitude. in the shade of evergreen forest with a rainfall of more than 2200 mm a year and a weak dry season. additional material examined ivory coast. entre niebe et bihoué, fl., 9 aug. 1962, jangoux 401 (br). liberia. east of greenville-zwedru road, fl., fr., 23 sep. 2013, jongkind 12109 (br, mo, p); west of sapo national park, 15 sep. 2013, jongkind 12321 (br), jongkind 12322 (wag); west of greenville, 18 sep. 2013, jongkind 12323 (br); north of sapo national park, 27 sep. 2013, jongkind 12324 (br); ca. 50 km east of greenville, 10 mar. 2014, jongkind 12361 (br), 11 mar. 2014, jongkind 12381 (wag). conservation status the “extent of occurrence” (eoo) is 6,056 km2 and the “area of occupancy” (aoo) is 32 km2, the eoo counts as “vulnerable” and the eoo counts as “endangered”. the aoo is based on a cell width of 2 km. none of the 9 specimens was collected in a protected area, but it is very likely that the species occurs in sapo national park in liberia, as this protected area is completely included in the species eoo. the plant and its flowers are not conspicuous, so it is probably more common than what the few collections suggest. however, looking at all the economical development planned and in progress in this http://www.ipni.org/urn:lsid:ipni.org:names:77150009-1 european journal of taxonomy 138: 1–8 (2015) 4 fig. 1. a–c. rhaphiostylis elegans engl. a. flower b. flower without petals and stamens. c. leaf. d–g. rhaphiostylis minima sp. nov. d. branch with flowers. e. flower. f. flower without petals and stamens. g. leaf. h–m. rhaphiostylis preussii engl. h. flower. i. flower without petals and stamens. j. leaf. k. flower. l. flower without petals and stamens. m. leaf. all flowers are on the same scale (scale bar with b) and all single leaves too (scale bar with g). a–c from tchouto & elad 3310 (wag), d–g from w.de wilde 1061 (wag), h–j from beentje 1353 b from ivory coast (wag), k–m from breteler 14204 from gabon (wag). drawn by hans de vries. jongkind c.c.h., a new rhaphiostylis species (icacinaceae) from africa 5 r. beninensis r. elegans r. minima sp.nov. r. preussii length when fertile ˃ 15 m 5–10 m 3–4 m ˃ 10 m leaf apex short acuminate long driptip 1.5–3 cm long driptip 1.5–3 cm acuminate ovary shape 1.5–2 mm high, short constriction between lower part and top 3.5–4 mm high, with long constricted part 2.5–3 mm high, shape in between r. preussii and r. elegans 1.7–2.5 mm high, short constriction between lower part and top ovary indumentum glabrous or only hairy on top only lower ca. ⅓ hairy hairy on the lower half and at the top, almost glabrous on the narrow part in between completely hairy or only glabrous around the base table 1. main differences between rhaphiostylis beninensis, r. elegans, r. minima sp. nov. and r. preussii. fig. 2. rhaphiostylis preussii engl. showing flowers with the flattened filaments closing around the ovary while the petals are bending down. photograph by ehoarn bidault (missouri botanical garden) from bidault 786 (mo) from gabon. european journal of taxonomy 138: 1–8 (2015) 6 part of liberia, and while it is not sure that the species grows in a protected area, “vulnerable” should be the correct status for the moment (b1 & b2 ab(iii), iucn 2015). remarks all specimens cited under the new species were collected after the publication of the icacinaceae in the flora of west tropical africa (hepper 1958). the leaves of seedlings of rhaphiostylis beninensis and r. preussii may resemble those of mature r. minima sp. nov. rhaphiostylis elegans engl. (fig. 1a–c) botanische jahrbücher für systematik, pflanzengeschichte und pflanzengeographie 43: 184 (engler 1909). – type: cameroon, bipindi, fr., jan. 1908, zenker 3615 (holo-: b destroyed; lecto-: mo [mo260675] designated here). additional material examined cameroon. between kienke river and km 7 on kribi-ebolowa road, fr., 17 oct. 1969, bos 5535 (br, k, p, wag); 7.5 km from kribi, few km n. of ebolowa road, fl. buds, 24 aug. 1970, bos & fig. 3. rhaphiostylis beninensis (hook.f. ex planch.) planch. ex benth., like fig. 2, but also showing ovary and style in older flowers. photograph by bart wursten from boyekoli ebale botany 785 (br) from congo kinshasa (copyright botanic garden meise). jongkind c.c.h., a new rhaphiostylis species (icacinaceae) from africa 7 breteler 7257 (br, k, p, wag); bidou, nkol mbouda on “mont d’elephant”, fl., 16 oct. 2001, tchouto mbatchou & elad 3310 (wag). remarks specimens of rhaphiostylis elegans key out as r. preussii with the flore du gabon (villiers 1973a) and flore du cameroun (villiers 1973b). according to notes in the wag herbarium there should be a duplicate specimen of the type in kew but i could not find it. acknowledgements i am grateful to hans de vries for preparing the drawing, and i also want to thank ehoarn bidault and bart wursten for the use of their photographs. references boutique r. 1960. icacinaceae. in: robyns w., staner p., demaret f., germain r., gilbert g., hauman l., homès m., jurion f., lebrun j., vanden abeele m. & boutique r. (eds) flore du congo belge et du ruanda-urundi: vol. 9. institut national pour l’étude agronomique du congo belge, brussels. fig. 4. distribution map of rhaphiosylis minima sp. nov. european journal of taxonomy 138: 1–8 (2015) 8 engler e. 1909. icacinaceae africanae. beiträge zur flora von afrika xxxiv. botanische jahrbücher für systematik, pflanzengeschichte und pflanzengeographie 43: 179–188. hawthorne w.d. & jongkind c.c.h. 2006. woody plants of western african forests, a guide to the forest trees, shrubs and lianes from senegal to ghana. royal botanic gardens, kew. hepper f.n. 1958. icacinaceae. in: keay r.w.j. (ed.) flora of west tropical africa, 2nd ed. vol. 1 (2): 636–644. crown agents for oversea governments and administrations, london. iucn 2015. iucn red list categories and criteria: version 3.1. prepared by the iucn species survival commission. iucn, gland, switzerland/cambridge, u.k. kårehed j. 2001. multiple origin of the tropical forest tree family icacinaceae. american journal of botany 88 (12): 2259–2274. http://dx.doi.org/10.2307/3558388 thiers b. continuously updated. index herbariorum: a global directory of public herbaria and associated staff. new york botanical garden’s virtual herbarium [online]. new york botanical garden’s virtual herbarium, new york. available from http://sweetgum.nybg.org/ih/ [accessed 9 jan. 2015]. villiers j.-f. 1973a. icacinacées. in: aubréville a. & leroy j.-f. (eds) flore du gabon 20. muséum national d’histoire naturelle, paris. villiers j.-f. 1973b. icacinacées. in: aubréville a. & leroy j.-f. (eds) flore du cameroun 15. muséum national d’histoire naturelle, paris. manuscript received: 10 february 2015 manuscript accepted: 14 july 2015 published on: 22 september 2015 topic editor: thomas janssen desk editor: natacha beau printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; national botanic garden of belgium, meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark http://dx.doi.org/10.2307/3558388 http://sweetgum.nybg.org/ih/ 45 european journal of taxonomy 828: 45–60 issn 2118-9773 https://doi.org/10.5852/ejt.2022.828.1851 www.europeanjournaloftaxonomy.eu 2022 · liu l.-y. & sittichaya w. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:57be020f-f904-498e-a4a1-a7585ad0a8b6 the oriental genera of xyloperthini (coleoptera: bostrichidae: bostrichinae), with a new genus and species from thailand, and a key to the genera lan-yu liu 1,*& wisut sittichaya 2 1 department of science communication, national pingtung university, no.4-18, minsheng rd, pingtung city, pingtung county 90049, taiwan. 2 agricultural innovation and management division, faculty of natural resources, prince of songkla university, 15 karnjanavanich rd, hat yai, songkhla 90110 thailand. * corresponding author: liulysky@gmail.com 2 email: wanakorn62@hotmail.com 1 urn:lsid:zoobank.org:author:8a4ece7c-2607-440d-b1bc-6e3b05ef02bb 2 urn:lsid:zoobank.org:author:12c0c726-fff1-4e84-95ae-58f22ce095b3 abstract. we describe a new genus, infrantenna gen. nov. and a new species infrantenna fissilis gen. et sp. nov., from thailand and briefly review the oriental genera currently placed in the tribe xyloperthini  lesne, 1921. a key to the oriental genera of xyloperthini is provided. keywords. xyloperthini, oriental fauna, new genus, new species, key. liu l.-y. & sittichaya w. 2022.the oriental genera of xyloperthini (coleoptera: bostrichidae: bostrichinae),  with a new genus and species from thailand, and a key to the genera. european journal of taxonomy 828: 45–60. https://doi.org/10.5852/ejt.2022.828.1851 introduction in the fourth part of his monograph of the family bostrichidae latreille, 1802, lesne (1901) included sixteen genera under the heading “les xylopertha” in the subfamily bostrichinae latreille, 1802. lesne (1921) formally erected the tribe xyloperthini lesne, 1921 to include those genera with a lamelliform  intercoxal process of the first abdominal ventrite. the tribe xyloperthini is the most species-rich in the  bostrichidae, and currently includes 35 genera (borowski & węgrzynowicz 2007; ivie 2010; park et al. 2015; liu et al. 2016; liu & beaver 2017; liu 2021a; zhang et al. 2022). the  tribe  has  a  worldwide  distribution,  but  the  individual  genera  are  mostly  confined  to  a  single  zoogeographical region or subregion (morrone 2002; borowski & węgrzynowicz 2007; beaver et al. 2011; holt et al. 2013; sittichaya et al. 2013; park et al. 2015; liu et al. 2016, 2021; liu & beaver  2017; borowski 2018; liu 2021a, 2021b; zhang et al. 2022), with the oriental region having the most  genera-rich fauna, and the african region the most species-rich fauna. https://doi.org/10.5852/ejt.2022.828.1851 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:57be020f-f904-498e-a4a1-a7585ad0a8b6 https://orcid.org/0000-0002-6874-2671 https://orcid.org/0000-0001-6200-1285 mailto:wanakorn62%40hotmail.com?subject= mailto:wanakorn62@hotmail.com http://zoobank.org/urn:lsid:zoobank.org:author:8a4ece7c-2607-440d-b1bc-6e3b05ef02bb http://zoobank.org/urn:lsid:zoobank.org:author:12c0c726-fff1-4e84-95ae-58f22ce095b3 https://doi.org/10.5852/ejt.2022.828.1851 european journal of taxonomy 828: 45–60 (2022) 46 in this paper, we describe a new genus for an unusual species collected in northern thailand. we provide an initial profile of oriental xyloperthini and a key to all oriental genera of the tribe xyloperthini. material and methods in the course of this study, the senior author has examined all available types, and other specimens of xyloperthini  in the paris museum, numerous additional european museums, private collections,  the online database of insect types of mcz and the results of the examination of types in the ifri by  borowski & singh (2017). institutional abbreviations ifri = indian forest research institute, uttar pradesh, dehra dun, india lyl = private collection of dr liu lan-yu, yilan, taiwan mcz  =  insect type database, museum of comparative zoology, harvard university, usa mnhn = muséum national d’histoire naturelle, paris, france nhmu = natural history museum, london, uk pmcsnhm = princess maha chakri sirindhorn natural history museum, prince of songkla university, songkla, thailand wst = private collection of dr wisut sittichaya, songkla, thailand the biogeographic regions used are based on morrone (2002) and holt et al. (2013). photographs were taken with an olympus e-m5mii digital camera and a canon 6d digital camera with  a canon mp-e 65 mm macro photo lens (canon, tokyo, japan) and stackshot-macrorail (cognisys inc, mi, usa). the photographs were then combined using the program helicon focus ver. 6.8.0. and ver. 7.0 (helicon soft, ukraine), and optimized with adobe photoshop ver. cs2 and ver. cs6 (adobe systems, california, usa) the discussion of the characters of the oriental xyloperthini genera are mainly based on lesne (1901,  1921), with additional characters obtained by the examination of specimens from the museums and collections listed above. results taxonomy class insecta linnaeus, 1758 superfamily bostrichoidea latreille, 1802 family bostrichidae latreille, 1802 subfamily bostrichinae latreille, 1802 tribe xyloperthini lesne, 1921 infrantenna gen. nov. urn:lsid:zoobank.org:act:5187d839-b85d-48b4-8321-738f3e5cfd6d type species infrantenna fissilis gen. et sp. nov., here designated. diagnosis a member of  the  tribe xyloperthini as characterized by the antennal club segments elongated,  the  mandibles crossed at the tips, and the lamelliform intercoxal process of the first abdominal ventrite  (lesne 1901; fisher 1950; liu & schönitzer 2011). this taxon is distinguished from other genera of  http://zoobank.org/urn:lsid:zoobank.org:act:5187d839-b85d-48b4-8321-738f3e5cfd6d liu l.-y. & sittichaya w., the oriental genera of xyloperthini 47 xyloperthini by the following combination of characters: frons weakly convex, without tuft of long  upwardly directed hairs on the head in either sex. antennal fossa inserted next to lower intero-lateral margin of eyes below the fronto-clypeal suture; eyes strongly detached from cheek. posterior part of  elytral disc without costae or teeth, infero-lateral margin of elytral declivity strongly raised to form a false epipleuron; female elytra declivity with deep clefts extending a little above the middle of the  declivity, dividing the outer part of the declivity from a pair of elongate, raised and grooved sutural lobes; posterior margin of 5th abdominal ventrite strongly emarginate in female; 5th abdominal ventrite of male with pleural pieces. etymology the genus name is feminine, and refers to the unusually low position of the antennal fossa. description body. elongate, cylindrical. head deeply inserted in prothorax, not visible from above. head. frons weakly convex, without tuft of long upwardly directed hairs on the head in either sex, apart from a pair next to the inner margin of the eyes; fronto-clypeal suture dark, impressed in middle; clypeus  strongly transverse. mandibles subequal, sharply pointed. eyes large, globose, strongly detached from cheeks posteriorly. antennal fossa inserted next to lower intero-lateral margin of eyes below frontoclypeal suture, antenna with ten antennomeres, first and second antennomeres elongate, antennomeres  3–7 forming funicle, each antennomere short, fifth widest, together subequal to first club antennomere in  length; antennomeres 8–10 forming elongate, loose club, each antennomere with short, recumbent hairs,  two distinct c-shaped sensory impressions near anterior margin of antennomeres 8 and 9, indistinct sensory patches ⅓ of length from apex on last antennomere; antennomeres 8 and 9 subequal in length,  last antennomere elongate, oval, longer than penultimate antennomere, but subequal in width. pronotum. slightly wider than long, widest at base, antero-lateral angle with small, strongly upcurved tooth on each side, anterior margin between upcurved teeth slightly concave; sides broadly rounded,  converging more strongly anteriorly, posterior angles broadly rounded, without lateral carina; anterior  slope coarse with 4 large upcurved teeth antero-laterally on each side; discal surface shining, sparsely  punctate. scutellum. small, tongue-shaped, with very sparse punctures. elytra. subequal to pronotum in width, parallel-sided, shining, disc and sides rugulose with small punctures, bearing minute hairs. elytral declivity sexually dimorphic. male with declivity concave, puncturation stronger and becoming smaller and shallower towards apex, declivital margins forming distinct infero-lateral epipleuron (false epipleuron of lesne 1901) gradually narrowing to sutural apex to form carina which runs into the apical margin. female with declivity convex, with deeper and larger punctures, infero-lateral epipleuron distinct and abruptly cut into semi-circular emargination from middle to apex; elytral declivity with deep clefts extending a little above middle of declivity, dividing  outer part of declivity from a pair of elongate, raised and grooved sutural lobes. abdomen. in female, 5th abdominal ventrite strongly emarginated in middle on posterior margin. last visible abdominal ventrite of male with pleural pieces. leg. subequal in length, procoxae separated by narrow intercoxal process of first thoracic ventrite,  mesocoxae  very  narrowly  separated;  tibiae  expanded  toward  apices,  protibiae  broadly,  shallowly  grooved on external face, protarsi longer than the length of protibiae, with series of long hairs beneath, mesoand meta-tarsi longer than their respective tibiae. european journal of taxonomy 828: 45–60 (2022) 48 remarks infrantenna gen. nov. can be distinguished from all other xyloperthine genera by the combination of the frons with only one long hair on each side next to eyes, the antennal fossa placed very low on the head below fronto-clypeal suture, 10-segmented antennae with two c-shaped sensory areas near anterior margin of 1st and 2nd club segments, without lateral carina of pronotum, the female with deep clefts on the elytral declivity and strongly emarginated 5th abdominal ventrite and male with pleural pieces of 5th abdominal ventrite. five genera which appear to be related to infrantenna gen. nov. in their form and morphology have been selected, and characters considered to be potentially informative in the determination of phylogenetic relationships are compared in table 1. psicula lesne, 1941 is the most similar genus morphologically to infrantenna gen. nov. it has similar modifications of the apex of the elytral declivity and last abdominal  ventrite of the female, and has a similar distribution. however, psicula has nine-segmented antennae and, like all the other genera compared, a more dorsal location of the antennal fossa. the last ventrite of the male also lacks pleural pieces. the african genus xylion differs in the modifications of the female  abdominal ventrites with variously modified from 3rd to 5th ventrites. the australian genus, xylobosca lesne, 1901, differs in the sexually dimorphic frons, the first abdominal ventrite of female enlarged, and  the widest of the protibia toward to middle. the north american genus, xyloblaptus lesne, 1901, which is the only selected genus with entire declivital apex in both sexes, lacks male pleural pieces, and has the widest of the protibia toward to middle. the protibia of the mediterranean genus, xylopertha guérinméneville, 1845, has the same form as infrantenna gen. nov., but it differs in the nine-segmented antenna, and the higher position of the antennal insertions. based on either geographical distribution or morphological characters, psicula is the most similar genus to infrantenna gen. nov. distribution northern thailand. infrantenna fissilis gen. et sp. nov. urn:lsid:zoobank.org:act:4beb3bb9-d678-4c4a-b25c-334872f069b9 figs 1–4 etymology the name ‘fissilis’ is from the latin word for ‘cleft’ which refers to the deep clefts of the elytral declivity of the female. type material holotype thailand • ♂; mueang district, maehongson province; 19°08.01′ n, 98°12.30′ e; 7 mar. 2019; ex.  semi-dry twig of unknown fagaceae l.; col. w. sittichaya; nhmuk014433955. allotype thailand • ♀; same collection data as for holotype; nhmuk014433956. paratype thailand • 1 ♀; same collection data as for holotype; pmcsnhm. http://zoobank.org/urn:lsid:zoobank.org:act:4beb3bb9-d678-4c4a-b25c-334872f069b9 liu l.-y. & sittichaya w., the oriental genera of xyloperthini 49 g en us c ha ra ct er in fr an te nn a ge n. n ov . p si cu la l es ne , 1 94 1 x yl io n l es ne , 1 94 1 x yl ob os ca l es ne , 1 90 1 x yl ob la pt us l es ne , 1 90 1 x yl op er th a g ué ri nm én ev ill e, 1 84 5 n o. o f a nt en na l s eg m en ts 10 9 10 10 10 10 a nt en na l f os sa in se rt io n b el ow fr on to -c ly pe al su tu re . a bo ve fr on to -c ly pe al su tu re . a bo ve fr on to -c ly pe al su tu re . o n  fr on to -c ly pe al  s ut ur e. a bo ve fr on to -c ly pe al su tu re . a bo ve fr on to -c ly pe al s ut ur e. l on g ha ir s on fr on s x d ir ec te d up w ar dl y x d ir ec te d up w ar dl y, lo ng to v er y lo ng in fe m al e, s ho rt o r a bs en t in m al e. l on g up w ar dl y at s id es , sh or t u pw ar d in m id dl e. d ir ec te d up w ar dl y p os te ri or a ng le s of pr on ot um w ith ou t l at er al c ar in a w ith ou t l at er al c ar in a w ith la te ra l c ar in a w ith ou t l at er al c ar in a w ith ou t l at er al c ar in a w ith ou t l at er al c ar in a p le ur al p ie ce s v x v v x v sh ap e of d is c of e ly tr al de cl iv it y m al e: c on ca ve fe m al e: c on ve x m al e: c on ca ve fe m al e: c on ve x sl ig ht ly c on ve x m al e: c on ca ve fe m al e: c on ve x sl ig ht ly , e ve nl y co nc av e m al e: w ea kl y to s tr on gl y co nc av e fe m al e: c on ve x sc ul pt ur e of a pi ca l m ar gi n of d ec liv it y m al e: w ith a s m al l v -s ha pe d em ar gi na tio n at a pe x. fe m al e: a pe x of e ac h el yt ro n st ro ng ly em ar gi na te , em ar gi na tio n  fil le d  by  a   pa ir o f v en tr al ly di re ct ed p ro ce ss es n ex t to s ut ur e. m al e: w ith ou t a n ap ic al em ar gi na tio n. fe m al e: a pe x of e ac h el yt ro n st ro ng ly em ar gi na te , em ar gi na tio n  fil le d  by  a   pa ir o f v en tr al ly di re ct ed p ro ce ss es n ex t to s ut ur e. b ot h se xe s w ith d ee p em ar gi na tio n at a pe x an d ri se d tip . m al e: n ot e m ar gi na te a t ap ex fe m al e: e m ar gi na te . e nt ir e, ra is ed s lig ht ly a t ap ex . m al e: w ith a s m al l v -s ha pe d em ar gi na tio n at a pe x. fe m al e: a pe x of e ac h el yt ro n st ro ng ly em ar gi na te . m od ifi ed v en tr it es o f f em al e ab do m en 5t h v en tr ite s tr on g em ar gi na te in m id dl e po st er io rl y. 5t h v en tr ite s tr on g em ar gi na te in m id dl e po st er io rl y. v ar io us ly  m od ifi ed   fr om 3 rd to 5 th ve nt ri te s, s tr on g em ar gi na tio n at po st er io r m ar gi n of 5t h v en tr ite 1s t  v en tr ite  e nl ar ge d;  4 th an d 5t h v en tr ite s em ar gi na te in m id dl e po st er io rl y;  4 th v en tr ite w ith a p ai r o f s pi ne s in so m e sp ec ie s. 5t h v en tr ite s tr on g em ar gi na te in m id dl e po st er io rl y. 5t h  v en tr ite  v ar io us ly  m od ifi ed . p ro ti bi a w id er a t a pe x th an in m id dl e, e xt er na l f ac e br oa dl y gr oo ve d. w id es t c lo se to a pe x, ex te rn al  fa ce  fl at te ne d. w id es t a t a pe x, ex te rn al fa ce m or e or le ss  fl  a tte ne d w id es t t ow ar ds m id dl e, w ith ou t a  fl at  e xt er na l  fa ce w id es t t ow ar ds m id dl e, w ith ou t a  fl at  e xt er na l  fa ce w id es t a t a pe x, g ro ov ed o n ex te rn al fa ce g eo gr ap hi ca l d is tr ib ut io n n or th er n t ha ila nd in di a an d t ha ila nd a fr ic a a us tr al ia u sa a nd m ex ic o m ed ite rr an ea n ta bl e 1. c om pa ri so n of c ha ra ct er s of in fr an te nn a  ge n.  n ov . w ith  s el ec te d  ge ne ra  o f x yl op er th in i l es ne , 1 92 1. european journal of taxonomy 828: 45–60 (2022) 50 description male (figs 1–2) measurements. 3.3–3.6 mm long, about 2.6–3.0 × as long as wide. coloration. head, pronotum, proand mesotibiae and tarsi, ventral side reddish brown, labrum, antennae, elytral disc, proand mesofemora and whole hind legs brown, pronotum and elytra gradually becoming darker brown anteriorly and posteriorly respectively, elytral declivity dark brown. head. mandibles dark, stout, apex strongly pointed, inner margin sharp. labrum trapezoid, narrowed towards base, fringe of dense, short, golden hairs along anterior margin, and fringe of long, yellowish hairs along anterior and lateral margins (fig. 1a). clypeus transverse, finely punctured, anterior margin  emarginated in middle to cover base of labrum and pair of small, broadly rounded teeth on either side. fronto-clypeal suture dark, attenuated laterally, with median fovea (fig. 1a). frons rugulose with sparse, fine punctures, each puncture with short, white hair; one long upwardly-directed hair close  to intero-lateral margin of eye, triangular shining area on lower middle part, and triangular area with reticulations on upper part above shining area (fig. 1a), upper part with row of small punctures between each two rugosities. vertex with sparse small punctures, longitudinal rugosities slightly angled toward midline arranged along whole vertex (fig. 1a). eyes large, strongly detached from cheeks posteriorly (fig. 1a). antennal fossa inserted next to lower intero-lateral margin of eyes below fronto-clypeal suture (fig. 1b). antennae 10-segmented, 1st and 2nd antennomeres subequal in length, 3–7 antennomeres together subequal to first club antennomere and shorter than last club antennomere in length, first and  second club antennomeres subtriangular and subquadrate, wider than long, with small, visible c-shaped areas of dense sensillae close to middle part of anterior margin, last club antennomere more elongate, and longer than penultimate antennomeres, sensory areas indistinct patches at one-third of length from apex (fig. 1c). pronotum. 1.1–1.2 × as wide as long, widest at base, antero-lateral angles with small, strongly upcurved tooth, semicircular area above anterior margin with transverse rugosities; sides of pronotum broadly  rounded, converging more strongly anteriorly, posterior angles broadly rounded (fig. 2a), without lateral carina (fig. 2b), postero-lateral area with fine rugosities (fig. 2c); anterior slope coarse with  4 large upcurved teeth antero-laterally on each side, 2–3 teeth next to antero-lateral upcurved teeth small, remaining teeth gradually reduced in size towards summit, and arranged approximately in arcs (fig. 2a–c); discal surface shining with sparse punctures, sparser toward posterior angles, posterolateral areas with small, slightly elongate rugulosities and series of granuloistes along posterior angles (fig. 2b–c). anterior slope with short, semi-erect hairs between teeth, recumbent hairs on sides. scutellum. small, tongue-shaped, with very sparse punctuation. elytra. 1.8–2 × as long as wide, parallel-sided, shining, disc and sides rugulose with small punctures, bearing minute hairs; epipleuron narrow, elongated triangular; declivity concave, puncturation larger  than on disc, but becoming smaller and shallower toward apex with triangular, nearly glabrous area at apex, sutural margin raised, more strongly in apical three-quarters of declivity (fig. 2d), declivital margins forming distinct rim at sides and apex, infero-lateral epipleuron distinct gradually narrowing to sutural apex to form carina which runs into apical margin (fig. 2c, e); margin slightly upwardly raised  at sutural apex and with tiny v-shaped emargination at apex of declivity (fig. 2d). legs. protibiae expanded from base to apex, broadly, shallowly grooved on external face (fig. 2a–b). protarsi with long hairs underneath. second and third segments of tarsi subequal in length and shorter than last segment. liu l.-y. & sittichaya w., the oriental genera of xyloperthini 51 abdomen. abdominal ventrites finely, moderately densely punctured, punctures with moderately long,  whitish hairs; last ventrite with pleural pieces, posterior margin slightly concave with fringe of dense,  short, yellowish hairs and fringe of very long, golden hairs along posterior margin (fig. 2c). female (figs 3–4) measurements. 4–4.5 mm long, about 3 × as long as wide. generally similar to male, but pronotum 1–1.1 × as wide as long and elytra 2–2.2 × as long as wide. elytra. discal punctures become larger posteriorly, declivity convex, with deeper and larger punctures (fig. 3a, d), piceous and darker on raised sutural margin and latero-apical margin (fig. 3a, c–d);  infero-lateral epipleuron distinct and abruptly cut into semi-circular emargination from middle to apex with only posterior margin continuing to apex (fig. 3b–c); elytra with two narrow fissure-like  emarginations extending from close to apex about two-thirds of height of declivity, separating a narrow inner, leglike, grooved process from the outer part of declivity (fig. 3d), each process minutely and sparsely punctured, with sinuate lateral margin, constricted before apex and then slightly raised to form spoon-like apex; inner margin of outer part of declivity thickened, inner area abruptly, strongly raised,  gradually sloping downward towards apex, forming an arc (fig. 3a, c–d). abdomen. last abdominal ventrite strongly curved upwards at sides to form triangular profile (fig. 4b),  ventrally with deep, broad emargination in one-third of middle part forming trapezoidal arc with pair of strongly sclerotised protrusions on sides, margin of trapezoidal arc and protrusion sclerotised, thickened (fig. 4a–c), and sinuate with a pair of tubercles in middle, and pointed, slightly upwardly curved hook on inner side of protrusion, and postero-dorsal upwardly curved hook on postero-lateral angle on both sides (fig. 4a, c), fringe of long, golden hairs along inner margin of trapezoidal arc and external side of sclerotised, thickened margin of protrusion (fig. 4a–c). abdominal 5th tergite convex with transverse prominence in middle third, two oval pads of very dense, short, white hairs on apical half behind prominence ventrally (fig. 4a, c). fig. 1. head of infrantenna fissilis gen. et sp. nov., ♂ , holotype (nhmuk014433955). a. frontal view. b. fronto-lateral view. c. antennae. not to scale. european journal of taxonomy 828: 45–60 (2022) 52 biology as a member of the family bostrichidae, the species is likely to be polyphagous. the junior author extracted the specimens from a semi-dry twig of an unknown species of fagaceae l. we suggest that the hind legs could move vertically in the clefts to clean wood frass. the strongly developed last abdominal ventrite and tergite of the female may help to hold the male genitalia during mating and support the ovipositor  during  oviposition.  observations  of  mating  and  oviposition  behaviours  are  necessary  to  confirm this hypothesis. distribution northern thailand. key to the oriental genera of xyloperthini lesne, 1921 (see also table 2) 1. antennae with 10 segments .............................................................................................................. 6 – antennae with less than 10 segments ............................................................................................... 2 2. antennae with 8 segments ................................................................................................................ 3 – antennae with 9 segments ................................................................................................................ 4 3.  antennae distinctly shorter than pronotum. funicle subequal in length to first antennomere of club.  last antennomere with two distinct circular, sensory impressions. elytral declivity with a large spine on each elytron. female with unmodified third ventrite. male without pleural pieces of last  abdominal ventrite ......................................................................................octodesmus lesne, 1901 –  antennae distinctly longer than pronotum. funicle much shorter in length than first antennomere of  club. last antennomere without sensory impressions. elytra with more than one costa, lacking spines on declivity. female with third ventrite thickened and modified, overlapping and concealing fourth  ventrite. male with pleural pieces of last abdominal ventrite ....... octomeristes liu & beaver, 2016 fig. 2. infrantenna fissilis gen. et sp. nov., ♂ , holotype (nhmuk014433955). a. dorsal view. b. lateral view. c. ventral view. d. declivity. e. false epipleuron. scale bar = 1 mm. liu l.-y. & sittichaya w., the oriental genera of xyloperthini 53 fig. 3. infrantenna fissilis gen. et sp. nov., ♀, allotype (nhmuk014433956). a. dorsal view. b. ventral view. c. lateral view. d. declivity. scale bar = 1 mm. fig. 4. apical posterior of female of infrantenna gen. nov. and psicula lesne 1941. a–c. infrantenna fissilis gen. et sp. nov., allotype (nhmuk014433956). a. 2nd to 5th visible abdominal ventrites and apical part of last tergite with two very dense, short hairy oval pads on apex. b. ventral view of last two ventrites. c. apical-lateral view of last ventrite and declivity. d. psicula heterogama lesne 1941, ventral view of last two visible abdominal ventrites. european journal of taxonomy 828: 45–60 (2022) 54 4. antennal club segments without clear sensory areas. pronotum with lateral carinae and posterolateral angles pointed .............................. xylopsocus lesne, 1901(capucinus, intermedius, radula) – antennal club segments with clearly visible sensory areas. pronotum without lateral carinae and postero-lateral angles rounded .......................................................................................................... 5 5. dense sensory hairs along anterior margin of 1st and 2nd antennomeres of club. frons without long erect hairs. declivital disc flattened. external face of protibia grooved. male with pleural pieces of  last abdominal ventrite .................................................................................xylophorus lesne, 1906 – antennal club segments without distinct impressions, only areas with denser pores. frons with long erect hairs. declivital disc concave in male and convex in female. external face of protibia flattened.  male without pleural pieces of last abdominal ventrite ......................................psicula lesne, 1941 6. antennal club segments without clear sensory areas. pronotum with lateral carinae and posterolateral angles pointed ........................................................................................................................ 7 – antennal club segments with clearly visible sensory areas. pronotum without lateral carinae and postero-lateral angles rounded .......................................................................................................... 8 7. body 3.5–4 mm in length. frons without long erect hairs. male without pleural pieces of last abdominal ventrite ............................................................................. xylopsocus lesne, 1901 (part) – body 6–7.5 mm in length. frons with long erect hairs. male with pleural pieces of last abdominal ventrite ...........................................................................................................xylothrips lesne, 1901 8. frons with long erect hairs. apical margin of declivity entire. last abdominal ventrite of female without emargination ........................................................................................................................ 9 – frons without long erect hairs. apical margin of declivity emarginated. last abdominal ventrite of female with emargination ............................................................................................................... 10 9.  body 4.5–5 mm in length. declivital disc flattened. declivity obliquely sloping and without spines  on upper margin ........................................................................................... paraxylion lesne, 1941 – body 6–8.5 mm in length. declivital disc slightly convex. declivity steep with three pairs of spines on upper margin ............................................................................................xylodrypta lesne, 1901 10. sensory areas on antennal club segments indistinct. declivital disc not sexually dimorphic. male without pleural pieces of last abdominal ventrite ............................................................................11 – sensory areas on antennal club segments distinct. declivital disc sexually dimorphic. male with pleural pieces of last abdominal ventrite ........................................................................................ 12 11. body 2.2–3 mm in length. antennal club segments without distinct impressions, only areas with denser pores. declivital disc evenly concave. protibia expanded from base to apex, external face convex. in female, 4th abdominal ventrite extended over 5th as a thin, leaflike, plate, convex ventrally;  the middle part of the 5th abdominal ventrite slightly emarginate .................calonistes lesne, 1936 – body 3.5–6 mm in length. antennal club segments with two indistinct areas of sensory hairs at anterior margin of 1st and 2nd antennomeres of club. declivital disc slightly convex. protibia expanded  from  base  to  apex,  external  face  flattened.  abdominal  ventrites  not  sexually  dimorphic ...................................................................................................... xylodectes lesne, 1901 12. antennal fossa inserted next to lower intero-lateral margin of eyes. antennal club with two areas of concentration of sensory pores on 1st and 2nd antennomeres. protibia expanded from base to apex with broadly, shallowly grooved external face. in male, upper margin of declivity without processes, a small v-shaped emargination at apex of declivity. in female, apex of each elytron liu l.-y. & sittichaya w., the oriental genera of xyloperthini 55 strongly emarginate, emargination filled by a pair of ventrally-directed processes next to suture, 5th abdominal ventrite strongly emarginate in middle posteriorly ....................... infrantenna gen. nov. – antennal fossa inserted next to middle intero-lateral margin of eyes. antennal club with dense sensory hairs along anterior margin on 1st and 2nd antennomeres. protibia expanded from base to apex with broadly grooved external face. in male, upper margin of declivity with a pair of tiny teeth, entire apex. in female, declivity with a y-shaped carina on lower half, apex entire with one pair of distinct tubercles at the apex, 5th abdominal ventrite broadly emarginate posteriorly ....xylocis lesne, 1901 discussion table 2 compares the characters of the twelve oriental genera of xyloperthini based on the senior  author’s observations of specimens (except for xylophorus lesne, 1906 for which specimens could not be examined, and the original description has been used). among the twelve oriental genera, there are  two genera with eight antennal segments, two genera and three species of xylopsocus lesne, 1901 with nine antennal segments, and the remainder have ten antennal segments. the sensory impressions on club segments are absent or indistinct in calonistes lesne, 1936, psicula, xylopsocus and xylothrips lesne, 1901. there are no long erect hairs on the frons in calonistes lesne, 1936, octodesmus lesne, 1901, xylocis lesne, 1901, xylodectes lesne, 1901, xylophorus and xylopsocus and only one long hair on each side near the eyes in infrantenna gen. nov. lateral carinae of pronotum are only present in xylopsocus and xylothrips. the pleural pieces of the last abdominal ventrite of male are absent in calonistes, octodesmus, psicula, xylodectes and xylopsocus. these characters and differences in the sculpture of the elytral declivity, and modifications of the abdominal ventrites of the female shown in table 2 permit  us to easily key out the twelve genera. including the new genus, there are 36 genera in the tribe xyloperthini, and the geographical distribution  of the tribe covers all zoogeographic regions except the arctic-siberian (liu 2016). the faunal elements of the tribe xyloperthini (table 3) shows the largest number of genera occur in the oriental region, and  nine genera are endemic to that region, but more species (about 82 species) occur in the african region. twelve genera occur in the oriental region, of which six are confined to india and the indochinese  peninsula, including calonistes, infrantenna gen. nov., octodesmus, octomeristes liu & beaver, 2016, psicula and xylodrypta lesne, 1901. they include two genera (infrantenna gen. nov. and octomeristes) described in the last 6 years, implying that more research in the region may discover further genera, and that the oriental region is likely to be a hotspot for the tribe xyloperthini (liu 2016).  following a series of revisions of genera of the tribe xyloperthini (liu et al. 2016; liu & beaver 2017,  2021; liu 2021a), this study added to the profile of the oriental fauna, and provided baseline information  for a large further study project, which will revise the interesting tribe xyloperthini. acknowledgements we are most grateful to department of national parks, wildlife and plant conservation, thailand, for research permission in all conservation areas. special thank also go to all staff of national parks and wildlife sanctuaries for their facilitation in specimen collection. this research was supported by thailand research fund (trf), project number dbg–6180023. we appreciate the following curators who have allowed us access  to  the collections  in  their charge or who have sent specimens for  identification:  a. taghavian (mnhn), m.v.l. barclay and s. shute (nhmuk). european journal of taxonomy 828: 45–60 (2022) 56 g en us c ha ra ct er c al on is te s l es ne , 1 93 6 in fr an te nn a ge n. n ov . o ct od es m us l es ne , 1 90 1 o ct om er is te s l iu & b ea ve r, 2 01 6 p ar ax yl io n l es ne , 1 94 1 p si cu la l es ne , 1 94 1 n o. o f a nt en na l s eg m en ts 10 10 8 8 10 9 se ns or y im pr es si on s on cl ub w ith ou t d is tin ct im pr es si on s, o nl y ar ea s w ith d en se r p or es . tw o zo ne s of co nc en tr at io n of s en so ry po re s on 1 st a nd 2 nd an te nn om er es o f c lu b. d en se s en so ry h ai rs a lo ng an te ri or m ar gi n on 1 st a nd 2n d a nt en no m er es o f c lu b. d en se s en so ry h ai rs a lo ng a nt er io r m ar gi n on 1 st a nd 2 nd a nt en no m er es of c lu b. d en se s en so ry h ai rs al on g an te ri or m ar gi n on 1s t a nd 2 nd a nt en no m er es of c lu b. w ith ou t d is tin ct im pr es si on s, o nl y ar ea s w ith d en se r po re s. c ly pe us a rm ed o n si de s x x x x x x l on g ha ir s on fr on s x x x v v v l at er al c ar in ae o f pr on ot um x x x x x x p le ur al p ie ce s x v x v v x sh ap e of d is c of e ly tr al de cl iv it y c on ca ve e ve nl y m al e: c on ca ve fe m al e: c on ve x sl ig ht ly c on ca ve w ee kl y co nv ex fl at te n m al e: c on ca ve fe m al e: c on ve x sc ul pt ur e of a pi ca l m ar gi n of d ec liv it y e nt ir e, ra is ed s lig ht ly at a pe x. m al e: w ith a s m al l v -s ha pe d em ar gi na tio n at a pe x. fe m al e: a pe x of e ac h el yt ro n st ro ng ly e m ar gi na te , em ar gi na tio n  fil le d  by a p ai r o f v en tr al ly di re ct ed p ro ce ss es n ex t to s ut ur e. r is ed w ith a ti ny v -s ha pe d em ar gi na tio n. e m ar gi na te d e nt ir e m al e: w ith ou t a n ap ic al e m ar gi na tio n. fe m al e: a pe x of e ac h el yt ro n st ro ng ly e m ar gi na te , em ar gi na tio n  fi  lle d  by a p ai r o f v en tr al ly di re ct ed p ro ce ss es ne xt to s ut ur e. m od ifi ed v en tr it es o f fe m al e ab do m en 4t h v en tr ite e xt en de d ov er 5 th a s a th in , le afl ik e,  p la te ,  co nv ex v en tr al ly . t he m id dl e pa rt o f t he 5t h v en tr ite ,s lig ht ly em ar gi na te 5t h v en tr ite s tr on g em ar gi na te in m id dl e po st er io rl y. x 3r d v en tr ite p ro je ct in g ov er a nd co nc ea lin g fo ur th v en tr ite , i ts po st er io r m ar gi n w ith lo be s or te et h. x 5t h v en tr ite s tr on g em ar gi na te in m id dl e po st er io rl y. p ro ti bi a e xp an de d fr om b as e to a pe x, e xt er na l f ac e co nv ex . w id er a t a pe x th an in m id dl e, e xt er na l f ac e br oa dl y gr oo ve d. w id er a t a pe x th an in m id dl e, ex te rn al fa ce fla tte ne d. w id er a t a pe x th an in m id dl e, ex te rn al  fa ce  fl at te ne d. e xp an de d fr om b as e to a pe x, e xt er na l f ac e gr oo ve d. w id es t c lo se to a pe x, ex te rn al  fa ce  fl at te ne d. d is tr ib ut io n m al ay si a an d t ha ila nd n or th er n t ha ila nd in di a an d b ur m a in di a an d t ha ila nd in di a an d so ut he as t a si a in di a an d t ha ila nd ta bl e 2 (c on tin ue d on n ex t p ag e) . c om pa ri so n  of  c ha ra ct er s  of  o ri en ta l g en er a  of  x yl op er th in i l es ne , 1 92 1. liu l.-y. & sittichaya w., the oriental genera of xyloperthini 57 g en us c ha ra ct er x yl oc is l es ne , 1 90 1 x yl od ec te s l es ne , 1 90 1 (x . o rn at us l es ne , 18 97 )* x yl od ry pt a l es ne , 1 90 1 x yl op ho ro us l es ne , 1 90 6 (b as ed o n de sc ri pt io n by l en se ) x yl op so cu s l es ne , 1 90 1 (1 0/ 17 )* * x yl ot hr ip s l es ne , 1 90 1 (x . fl av ip es (i lli ge r, 18 01 )) * n o. o f a nt en na l s eg m en ts 10 10 10 9 9/ 10 10 se ns or y im pr es si on s on c lu b d en se s en so ry ha ir s al on g an te ri or m ar gi n on 1 st a nd 2 nd an te nn om er es o f c lu b in di st in ct tw o zo ne s of se ns or y ha ir s at a nt er io r m ar gi n on 1 st a nd 2 nd an te nn om er es o f c lu b d en se s en so ry h ai rs al on g an te ri or m ar gi n on 1s t a nd 2 nd a nt en no m er es of c lu b d en se s en so ry h ai rs al on g an te ri or m ar gi n on 1s t a nd 2 nd a nt en no m er es of c lu b in di st in ct x c ly pe us a rm ed o n si de s x x v x v v l on g ha ir s on fr on s x x v x x v l at er al c ar in ae o f p ro no tu m x x x x v v p le ur al p ie ce s v x v v x v sh ap e of d is c of e ly tr al d ec liv it y m al e: c on ca ve ev en ly fe m al e: c on ve x s lig ht ly c on ve x sl ig ht ly c on ve x fl at te n fl at te n / s lig ht ly c on ve x sl ig ht ly c on ve x sc ul pt ur e of a pi ca l m ar gi n of de cl iv it y m al e: e nt ir e w ith o ne pa ir o f t in y tip s at up pe r m ar gi n. fe m al e: e nt ir e w ith on e pa ir o f d is tin ct tu be rc le a t t he a pe x. e nt ir e w ith ti ny v -s ha pe d em ar gi na tio n at a pe x. e nt ir e e nt ir e. e nt ir e w ith s lig ht ly ri se d ap ex . e nt ir e w ith ti ny v -s ha pe d em ar gi na tio n at a pe x. m od ifi ed ve nt ri te s of fe m al e ab do m en 5t h v en tr ite b ro ad ly em ar gi na te po st er io rl y x x (c an ’t e xa m in ed ) x c on ve x in m id dl e of po st er io r m ar gi n p ro ti bi a e xp an de d fr om b as e to a pe x, e xt er na l f ac e fla tte n e xp an de d fr om b as e to a pe x, e xt er na l f ac e fla tte n e xp an de d fr om b as e to a pe x, e xt er na l f ac e w ea kl y co nv ex e xp an de d fr om b as e to a pe x, e xt er na l f ac e gr oo ve d e xp an de d fr om b as e to ap ex , e xt er na l f ac e w ea kl y gr oo ve d e xp an de d fr om b as e to ap ex , e xt er na l f ac e gr oo ve d d is tr ib ut io n in di a, s ri l an ka , l ao s, h on g k on g, ta iw an ta iw an , p hi lip pi ne s, in do ne si a, v ie tn am , l ao s, in di a in di a an d t ha ila nd sr i l an ka so ut he 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https://doi.org/10.1046/j.1365-2699.2002.00662.x https://doi.org/10.1016/j.japb.2015.10.015 https://doi.org/10.11646/zootaxa.5091.4.1 european journal of taxonomy 828: 45–60 (2022) 60 manuscript received: 10 feb. 2022 manuscript accepted: 27 may 2022 published on: 7 july 2022 topic editor: tony robillard section editor: max barclay desk editor: eva-maria levermann printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt  consortium:  muséum  national  d’histoire  naturelle,  paris,  france;  meise  botanic  garden,  belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural  sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis  biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid,  spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig,  bonn, germany; national museum, prague, czech republic. amage imajimai sp. nov., a new species of ampharetidae (annelida: polychaeta) from japanese waters michael g. reuscher harte research institute for gulf of mexico studies, texas a&m university – corpus christi, 6300 ocean drive, unit 5869, corpus christi, texas 78412-5869, usa. e-mail: michael.reuscher@tamucc.edu urn:lsid:zoobank.org:author:cee857c6-db17-4c9b-be81-d04ab815e909 abstract. a new polychaete species of the family ampharetidae, amage imajimai sp. nov., is described from deep waters of sagami bay, japan. it is characterized by the possession of four pairs of branchiae, twelve thoracic uncinigers, eleven abdominal uncinigers, and the lack of thoracic notopodial cirri. the new species is named in honor of the renowned japanese polychaetologist minoru imajima. an identification key for all amage species from japanese waters is provided. keywords. ampharetidae, amage, new species, sagami bay, japan. reuscher m.g. 2015. amage imajimai sp. nov., a new species of ampharetidae (annelida: polychaeta) from japanese waters. european journal of taxonomy 154: 1–7. http://dx.doi.org/10.5852/ejt.2015.154 introduction in a recent study on ampharetidae from japan many new species and new records were discovered (imajima et al. 2012, 2013; reuscher et al. 2015a, 2015b). in the last publication of the series (reuscher et al. 2015b), a list of all 58 ampharetid species recorded from japan was provided. six of these species known to occur in japanese waters belong to the genus amage malmgren, 1866: a. cf. adspersa (grube, 1863), a. auricula malmgren, 1866, a. delus (chamberlin, 1919), a. ehlersi reuscher, fiege & imajima, 2015, a. longitorus reuscher, fiege & imajima, 2015 and a. scutata moore, 1923. during the examination of material from the national museum of nature and science in tsukuba (japan), i was able to identify another new species of amage, which is described here. the new species was collected at a depth of about 1000 m in sagami bay off the southeastern honshu coast. material and methods the specimens examined in this study were collected in sagami bay during a research cruise in july 1966. they were fixed in 7% formaldehyde seawater solution and preserved in 70% ethanol. preserved specimens were examined with an olympus szx7 stereo microscope and compound microscopes of the models leica dmlb and olympus cx41. pencil drawings were made using a camera lucida, attached to the leica dmlb. european journal of taxonomy 154: 1–7 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2015.154 www.europeanjournaloftaxonomy.eu 2015 · reuscher m.g. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:f9a2f05e-0f3a-4459-bb5f-fc8563dd683c 1 mailto:michael.reuscher%40tamucc.edu?subject=michael.reuscher%40tamucc.edu http://zoobank.org/urn:lsid:zoobank.org:author:cee857c6-db17-4c9b-be81-d04ab815e909 http://dx.doi.org/10.5852/ejt.2015.154 http://dx.doi.org/10.5852/ejt.2015.154 http://www.europeanjournaloftaxonomy.eu/index.php/ejt https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:f9a2f05e-0f3a-4459-bb5f-fc8563dd683c the drawings were digitized with a wacom intuos drawing tablet and adobe illustrator, according to the methods of coleman (2003). shadings were added in adobe photoshop. the “id card” (imajima et al. 2012) was prepared in adobe illustrator. abbreviations cs = complete specimen af = anterior fragment types and other specimens are deposited in the following institutions: nsmt = national museum of nature and science, japan smf = senckenberg museum frankfurt, germany full details for the material deposited at senckenberg can be found at http://sesam.senckenberg.de/. results phylum annelida lamarck, 1809 class polychaeta grube, 1850 order terebellomorpha hatschek, 1893 family ampharetidae malmgren, 1866 subfamily ampharetinae malmgren, 1866 genus amage malmgren, 1866 amage malmgren, 1866: 370. paramage caullery, 1944: 94. egamella fauchald, 1972: 295. mexamage fauchald, 1972: 309. type species amage auricula malmgren, 1866. diagnosis (emended) prostomium with middle lobe surrounded by inflated lobe, lacking glandular ridges. buccal tentacles smooth. two to four pairs of cirriform branchiae. segment ii usually without chaetae, or exceptionally with minute chaetae. thorax with 9–14 uncinigers. modified or intermediate segments absent. abdomen with rudimentary notopodia. remarks the diagnosis was emended to accommodate the synonymy of the monotypic genus egamella fauchald, 1972 by jirkov (2011). egamella has only two pairs of branchiae and nine thoracic uncinigers. this synonymy needs to be confirmed by the examination of the type specimen of egamella quadribranchiata fauchald, 1972. european journal of taxonomy 154: 1–7 (2015) 2 http://sesam.senckenberg.de/ amage imajimai sp. nov. urn:lsid:zoobank.org:act:cca76c94-c953-418d-afc9-ddbbd7d99c6d fig. 1a–g diagnosis four pairs of branchiae. twelve thoracic uncinigers. notopodia without ventral cirri. eleven abdominal uncinigers. etymology the species is dedicated to the distinguished japanese polychaete taxonomist minoru imajima. specimens examined holotype japan: smf 24087, sagami bay, 35°00.9’ n, 139°35.7’ e – 35°00.7’ n, 139°36.0’ e, 990–1060 m, kt-66-12, st. 7, jul. 1966 (1 cs). paratypes japan: smf 24086, same locality as holotype (3 cs); nsmt-pol. p-600, same locality as holotype (3 cs, 1 af). description length of holotype 3.2 mm, width 0.4 mm. prostomium with middle lobe bearing anterolateral frontal horns, delimited by incision from inflated surrounding lobe (fig. 1a); prostomium without glandular ridges or eyes. single tip of smooth buccal tentacle visible in buccal cavity. four pairs of branchiae in l-shaped arrangement in segments ii–iv (fig. 1b), separated by wide median gap; all branchiae detached from specimen, cirriform, without conspicuous ciliation or annulations; innermost branchiae of anterior transverse row (1) originating from segment ii, outermost branchiae of anterior transverse row (2) originating from segment iii, median branchiae of longitudinal row (3) originating from segment iv, posterior branchiae of longitudinal row (4) originating from segment v (fig. 1b). segment ii without chaetae. notopodia with capillary chaetae from segment iii, present in 15 chaetigers; first three notopodia in close succession due to shortness of segments and slightly elevated above following notopodia (fig. 1c); first notopodia small, increasing in size from first to third pair; notopodial cirri absent. neuropodial tori with uncini from segment vi, present in 12 thoracic uncinigers; tori without cirri. continuous ventral shields conspicuous from anterior thorax to thoracic unciniger 9. modified notopodia or segments absent. intermediate uncinigers absent. eleven abdominal uncinigers with small tuberculate rudimentary notopodia. pinnules with minute tuberculate dorsal cirrus. rudimentary notopodia and pinnules connected by glandular fold. pygidium with one pair of digitiform, ventrolateral anal cirri. left anal cirrus broken off. thoracic uncini with 7 teeth in 2 staggered row over basal prow and rostral tooth (fig. 1d–e). abdominal uncini with numerous teeth in several rows over basal prow and rostral tooth. tube parchment like with needle like spicules embedded. remarks in four of the paratypes the buccal tentacles are better visible and clearly smooth. the tuberculate dorsal cirri of the abdominal pinnules are much better developed in the larger paratype specimens (fig. 1f). the anal cirri are longer and cirriform in the larger paratypes (fig. 1g). however, they also seem to break off easily as three of the six complete paratypes lack both anal cirri. the two other amage species with twelve thoracic uncinigers are a. benhami reuscher, fiege & wehe, 2009 from the northeast pacific and the ross sea and a. longitorus reuscher, fiege & imajima, 2015 reuscher m.g., new amage species from japan 3 http://zoobank.org/urn:lsid:zoobank.org:act:cca76c94-c953-418d-afc9-ddbbd7d99c6d fig. 1. amage imajimai sp. nov. a. anterior end of holotype, dorsal view. b. “id card”. c. anterior end of holotype, lateral view. d. thoracic uncinus, lateral view. e. thoracic uncinus, frontal view. f. abdominal uncinigers, lateral view (from paratype smf 24086). g. posteriormost abdominal uncinigers and pygidium, ventral view (from paratype smf 24086). european journal of taxonomy 154: 1–7 (2015) 4 from japan. the latter species differs from a. imajimai sp. nov. by the possession of only three pairs of branchiae, the very long tori in the first two thoracic uncinigers and the larger number of abdominal uncinigers (13). a. benhami differs from the new species by the presence of club shaped notopodial cirri and the higher number of abdominal uncinigers (15–16). among the other japanese amage species a. auricula, a. delus, a. ehlersi and a. scutata have 11 thoracic uncinigers, a. cf. adspersa has 14 thoracic uncinigers. a. cf. adspersa, a. auricula and a. delus differ from a. imajimai sp. nov. by the presence of notopodial cirri. a. scutata is unusual for the presence of rudimentary notopodia in the anterior segments. a. imajimai sp. nov. has a higher count of abdominal uncinigers (11) than a. auricula (8) and a. ehlersi (10) and a lower count than a. delus (12) and a. longitorus (13). distribution sagami bay on the southeastern pacific coast of honshu, in 990–1060 m. identification key for amage species from japanese waters 1. 11 or 12 thoracic uncinigers ……………………………………………………………………………2 – 14 thoracic uncinigers ………………………………………………amage cf. adspersa (grube, 1863) 2. 11 thoracic uncinigers …………………………………………………………………………………3 – 12 thoracic uncinigers …………………………………………………………………………………6 3. anterior notopodia with notochaetae …………………………………………………………………4 – anterior notopodia lacking notochaetae ………………………………amage scutata moore, 1923 4. thoracic notopodia with ventral cirri …………………………………………………………………5 – thoracic notopodia lacking ventral cirri …………amage ehlersi reuscher, fiege & imajima, 2015 5. 8 abdominal uncinigers ……………………………………………amage auricula malmgren, 1866 – 12 abdominal uncinigers ……………………………………………amage delus (chamberlin, 1919) 6. 3 pairs of branchiae; anterior neuropodia conspicuously elongated; 13 abdominal uncinigers ……………………………………………………amage longitorus reuscher, fiege & imajima, 2015 – 4 pairs of branchiae; anterior neuropodia not conspicuously elongated; 11 abdominal uncinigers ………………………………………………………………………………amage imajimai sp. nov. discussion the variety of habitat types and complex interactions of different environmental gradients in the oceans surrounding the japanese islands attract a variety of species with different physiological and ecological adaptations and thus form the basis of a diverse polychaete fauna. japan has a wide variety of habitats that are colonized by polychaetes, including bays, deep-sea trenches, hydrothermal vents, and cold seeps, among others (e.g., juniper & sibuet 1987; horikoshi et al. 1990). the northern part of japan receives cold water from the oyashio current, whereas southern japan is under the influence of the warm kuroshio current (imajima et al. 2012). therefore, japan’s polychaete fauna includes arctic and sub-arctic species as well as tropical and subtropical species. in the recent series on ampharetidae from japan (imajima et al. 2012, 2013; reuscher et al. 2015a, 2015b), 60% of the examined species were new to science and the number of species known from japan more than doubled. this shows that the current knowledge of species diversity of ampharetid polychaetes from japan is far from exhaustive and more sampling effort is needed to complete the picture. amage imajimai sp. nov. is the 59th species of the family ampharetidae and the seventh species of the genus amage recorded from japan. reuscher m.g., new amage species from japan 5 of the other amage species from japan, a. delus and a. scutata are known only from northern honshu, a. cf. adspersa and a. imajimai sp. nov. have only been recorded from sagami bay, whereas a. auricula, a. ehlersi and a. longitorus have a wider distribution within japanese waters. within ampharetidae amage is probably the most heterogeneous genus as it contains species with two (if the genus egamella is considered a junior synonym), three and four pairs of branchiae, with nine (egamella), eleven, twelve, and fourteen thoracic uncinigers, with and without notopodial cirri. a revision is needed to determine if amage can be upheld as a single genus, or if it should be split into multiple genera. acknowledgements minoru imajima (nsmt) is thanked for the loan of the specimens. i am grateful to richard d. kalke and fabio moretzsohn (harte research institute, texas a&m university–corpus christi) for their permissions to use their camera lucida. paul a. montagna (harte research institute, texas a&m university–corpus christi) is thanked for his financial support through the texas research development post-doctoral support grant. references caullery m. 1944. polychètes sédentaires de l’expedition du siboga. ariciidae, spionidae, chaetopteridae, chlorhaemidae, opheliidae, oweniidae, sabellariidae, sternaspidae, amphictenidae, ampharetidae, terebellidae. siboga-expeditie. uitkomsten op zoölogisch, botanisch, oceanographisch en geologisch gebied verzameld in nederlandsch oost-indie 1899 – 1900 aan boord h.m. siboga onder commando van luitenant ter zee 1. kl. g.f. tydeman 24 (2), brill, leiden. coleman c.o. 2003. “digital inking”: how to make perfect line drawings on computers. organisms, diversity & evolution 3, electronic supplement 14: 1–14. http://dx.doi.org/10.1078/1439-6092-00081 fauchald k. 1972. benthic polychaetous annelids from deep water off western mexico and adjacent areas in the eastern pacific ocean. monographs in marine biology 7, allan hancock foundation, university of 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(2011) discussion of taxonomic characters and classification of ampharetidae (polychaeta). italian journal of zoology 78 (s1): 78–94. juniper s.k. & sibuet m. 1987. cold seep benthic communities in japan subduction zones: spatial organization, trophic strategies and evidence for temporal evolution. marine ecology progress series 40: 115–126. malmgren a.j. 1866. nordiska hafs-annulater. öfversigt af kongliga vetenskaps-akademiens förhandlingar 5: 355-410. european journal of taxonomy 154: 1–7 (2015) 6 http://dx.doi.org/10.1078/1439-6092-00081 http://dx.doi.org/10.11646/zootaxa.3647.1.7 http://dx.doi.org/10.11646/zootaxa.3647.1.7 reuscher m.g., fiege d. & imajima m. 2015a. ampharetidae (annelida: polychaeta) from japan. part iii: the genus amphicteis grube, 1850 and closely related genera. journal of the marine biological association of the united kingdom 95 (5): 929–940. http://dx.doi.org/10.1017/s0025315414001623 reuscher m., fiege d. & imajima m. 2015b. ampharetidae (annelida: polychaeta) from japan. part iv. miscellaneous genera. journal of the marine biological association of the united kingdom 95 (6): 1105–1125. http://dx.doi.org/10.1017/s0025315415000545 manuscript received: 14 august 2015 manuscript accepted: 8 october 2015 published on: 19 november 2015 topic editor: rudy jocqué desk editor: kristiaan hoedemakers printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark. reuscher m.g., new amage species from japan 7 http://dx.doi.org/10.1017/s0025315414001623 http://dx.doi.org/10.1017/s0025315415000545 97 european journal of taxonomy 888: 97–110 issn 2118-9773 https://doi.org/10.5852/ejt.2023.888.2231 www.europeanjournaloftaxonomy.eu 2023 · ahrens d. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:854eacc8-f325-4558-aee4-f9c9d519a018 updates on the neoserica vulpes group (coleoptera, scarabaeidae, melolonthinae, sericini): new species and records dirk ahrens zoologisches forschungsmuseum a. koenig, leibniz institute for the analysis of biodiversity change (lib), adenauerallee 127, 53113 bonn, germany. ahrens.dirk_col@gmx.de; d.ahrens@leibniz-lib.de urn:lsid:zoobank.org:author:d01cbf11-54fa-4974-9cb9-b0e6f930ace2 abstract. here, i describe four new species of the neoserica vulpes species group: neoserica daxue sp. nov., n. mianningana sp. nov., n. myanmarensis sp. nov., and n. yanyuan sp. nov. genitalia and habitus of the new species are illustrated. additional records of species from the group are given and the key to species of the neoserica vulpes species group is updated. keywords. beetles, chafers, neoserica, china, myanmar, new species. ahrens d. 2023. updates on the neoserica vulpes group (coleoptera, scarabaeidae, melolonthinae, sericini): new species and records. european journal of taxonomy 888: 97–110. https://doi.org/10.5852/ejt.2023.888.2231 introduction the genus neoserica brenske, 1894 was recently revised in a series of publications in which many new taxa were established for the fauna of china and adjacent areas (ahrens et al. 2014a, 2014b, 2014c; liu et al. 2014a, 2014b, 2014c, 2015a, 2016, 2019; bohacz & ahrens 2020; ahrens 2021, 2022; ahrens & pham 2021; ahrens & lukic 2022). neoserica (sensu lato) comprises a polyphyletic mix of larger species with multilamellate antenna (ahrens 2003, 2004; liu et al. 2015b; ahrens & fabrizi 2016; eberle et al. 2017). as previously mentioned, they need a revision of their generic systematics and nomenclature, but this is only possible when their taxonomy, morphology, and phylogeny are better understood. the treatment of species of uncertain systematic assignment remains particularly complicated as the relationships to other groups of species remain obscure and would benefit from future investigations, especially once the neighbouring faunas of asia are better known. in the current paper we describe four further new species of the neoserica vulpes group (ahrens et al. 2014c) that were received from diverse collectors and institutions. material and methods the terminology and methods used for measurements, specimen dissection and genital preparation follow ahrens (2004). data from specimens examined are cited in the text with original label contents given in quotation marks, multiple labels are separated by a ‘/’. descriptions and illustrations of new taxa are based on the holotype or lectotype specimen if not otherwise stated, while the variation of specimens is given separately under ‘variation’. male genitalia were glued to a small, pointed card and https://doi.org/10.5852/ejt.2023.888.2231 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ https//zoobank.org/urn:lsid:zoobank.org:pub:854eacc8-f325-4558-aee4-f9c9d519a018 https://orcid.org/0000-0003-3524-7153 mailto:ahrens.dirk_col%40gmx.de?subject= mailto:d.ahrens%40leibniz-lib.de?subject= https://zoobank.org/urn:lsid:zoobank.org:author:d01cbf11-54fa-4974-9cb9-b0e6f930ace2 https://doi.org/10.5852/ejt.2023.888.2231 european journal of taxonomy 888: 97–110 (2023) 98 photographed in both lateral and dorsal view using a leica m125 with a leica dc420c digital camera. in the automontage software as implemented in leica application suite (ver. 3.3.0) single focused images were combined to an entirely focused image. the resulting images were subsequently digitally edited. abbreviations used in the text for collection depositories are as follows: cp = collection p. pacholátko, brno, czech republic zfmk = zoologisches forschungsmuseum a. koenig, bonn, germany results description of new species class insecta linnaeus, 1758 subclass pterygota lang, 1888 superfamily scarabaeoidea latreille, 1802 family scarabaeidae latreille, 1802 subfamily melolonthinae leach, 1819 tribe sericini kirby, 1837 genus neoserica brenske, 1894 neoserica mianningana sp. nov. urn:lsid:zoobank.org:act:999264ad-0fab-473b-9d75-2eb977e3ed9d fig. 1a–e diagnosis neoserica mianningana sp. nov. differs from n. nykli ahrens, liu & fabrizi 2014 by the shape of the ventral phallobasal process, which is well curved and has a less pronounced ventral distal tooth (compared to that of n. nykli) as well as by the shape of both parameres which are longer and more strongly curved. etymology the new species is named (name: adjective in the nominative case singular) for its occurrence close to the city of mianning, china. type material examined holotype china • ♂; “china, daxue shan mts., sichuan, 40 km w mianning, 7-8.vi.1999, 2750m, 28°34’n, 102°00’e, v. siniaev & a. plutenko lgt. / 1152 sericini: asia spec.”; cp. paratype china • 1 ♂; “china, daxue shan mts., sichuan, 40 km w mianning, 7-8.vi.1999, 2750m, 28°34’n, 102°00’e, v. siniaev & a. plutenko lgt.”; zfmk. description measurements. length: 8.8 mm, length of elytra: 6.2 mm, width: 4.6 mm. habitus and coloration (fig. 1e). body oblong, reddish brown, ventral surface and antenna yellow, dorsal surface dull and almost glabrous. https://zoobank.org/urn:lsid:zoobank.org:act:999264ad-0fab-473b-9d75-2eb977e3ed9d ahrens d., updates on the neoserica vulpes group 99 head. labroclypeus subtrapezoidal, widest at base, lateral margins weakly convex and convergent to moderately rounded anterior angles, lateral border and ocular canthus producing a distinct, blunt angle, margins weakly reflexed, anterior margin distinctly sinuate medially; surface flat and shiny, finely and densely punctate, with a few long, erect setae in coarser punctures; frontoclypeal suture feebly incised and medially curved; smooth area in front of eye approximately 1.5 times as wide as long; ocular canthus moderately long and slender, very finely and sparsely punctate, with a fine terminal seta. frons dull, with fine and moderately dense punctures and with a few long setae beside eyes and behind frontoclypeal suture. eyes small, ratio of diameter / interocular width: 0.55. antenna with ten antennomeres; club with four antennomeres, nearly 1.2 times as long as remaining antennomeres combined. mentum convexly elevated anteriorly. pronotum. short and wide, widest shortly before base, lateral margins evenly curved and narrowed anteriorly and posteriorly, anterior angles moderately produced and blunt, posterior angles blunt and slightly rounded at tip; anterior margin convexly produced medially, broad marginal line widely missing; basal margin without marginal line; surface with moderately dense and fine punctures, with minute setae, otherwise glabrous; anterior and lateral borders sparsely setose; hypomeron distinctly carinate at base. scutellum narrow and long, sharp at apex, with fine and moderately dense punctures and minute setae. elytra. oblong, widest in posterior third, striae finely impressed, finely and densely punctate, intervals flat, finely and sparsely punctate, on odd intervals with a few long erect setae, otherwise with minute setae; epipleural edge robust, ending at strongly curved external apical angle of elytra, epipleura densely setose, apical border membranous, with short microtrichomes. ventral surface. dull, with large and dense punctures, sparsely and shortly setose, setae partly adpressed; metacoxa glabrous, with fine setae laterally, apical margin convex and external apical angle rounded; each abdominal sternite with a distinct transversal row of coarse punctures each bearing a short seta between fine and moderately dense punctation. mesosternum between mesocoxae nearly half as wide as mesofemur, with irregularly scattered, fine setae. ratio of length of metepisternum / metacoxa: 1 / 1.5. pygidium strongly convex, finely and densely punctate, without smooth midline, with short setae along apical margin. legs. slender; femora with two longitudinal rows of setae, finely and moderately densely punctate; metafemur ventrally dull, anterior margin sharply carinate, without a submarginal serrated line, posterior margin moderately convex, with a few strong setae medially, only weakly widened externally in apical half and not serrated ventrally in distal half, finely serrated dorsally, with dense, short setae. metatibia slender and long, widest at apex, ratio width / length: 1 / 3.6, dorsal margin sharply carinate, with two groups of spines, basal group shortly before middle, apical one at about three quarters of metatibial length, basally with a few single, fine spines; external face longitudinally convex, with coarse, sparse punctures, glabrous; ventral margin finely serrated, with four fine, equidistant spines; medial face impunctate, apex concavely truncate interiorly near tarsal articulation. tarsomeres dorsally impunctate, with sparse, short setae ventrally; metatarsomeres glabrous dorsally, with a strongly serrated ridge ventrally, and with a fine longitudinal carina immediately beside it; first metatarsomere little longer than following two tarsomeres combined and distinctly longer than dorsal tibial spur. protibia long, bidentate, protarsal claws symmetrical, basal tooth of inner protarsal claw bluntly truncate apically. aedeagus. fig. 1a–d. variation length: 8.8–9.1 mm, length of elytra: 6.2–6.4 mm, width: 4.6–4.8 mm. european journal of taxonomy 888: 97–110 (2023) 100 female unknown. neoserica myanmarensis sp. nov. urn:lsid:zoobank.org:act:921f8fc7-dd6d-45b8-a7ce-692cfbc021bb fig. 1f–i diagnosis neoserica myanmarensis sp. nov. differs from n. baoshana ahrens, liu & fabrizi, 2014 by the shape of the ventral phallobasal process: its distal hook is shorter and only weakly bent towards the right side (dorsal view). fig. 1. a–e. neoserica mianningana sp. nov., holotype, ♂ (cp). f–i. n. myanmarensis sp. nov., holotype, ♂ (zfmk). a, f. aedeagus, left side lateral view. b. distal phallobase, dorsal view. c, g. parameres, dorsal view. d, h. aedeagus, right side lateral view. e, i. habitus (not to scale). scale bars = 0.5 mm. https://zoobank.org/urn:lsid:zoobank.org:act:921f8fc7-dd6d-45b8-a7ce-692cfbc021bb ahrens d., updates on the neoserica vulpes group 101 etymology the new species is named (name: adjective in the nominative case singular) for its occurrence in myanmar. type material examined holotype myanmar • ♂; “myanmar: danasho 2800 m, minshia vi-2004 leg. yin et al. / phil #11-2004:4. / 1137 sericini: asia spec.”; zfmk. paratypes china • 1 ♂; “x-da6902/ x-da6902 china yunnan ne kunming 25°08’35’’n, 102°53’49’’e 2320m leg. m. schülke & v. assing (ch14-6) 13.08.2021 neoserica sptigerthai215”; zfmk • 1 ♂; “x-da6905/ x-da6905 china yunnan mountain wnw wuding 25°38’45’’n, 102°06’55’’e 2390m leg. m. schülke & v. assing (ch14-12a) 01.09.2014 neoserica sptigerthai215”; zfmk. description measurements. length: 8.5 mm, length of elytra: 6.2 mm, width: 5.0 mm. habitus and coloration (fig. 1i). body oblong, dark brown, ventral surface and antenna yellow, dorsal surface dull and with numerous shorter and long, erect setae. head. labroclypeus subtrapezoidal, widest at base, lateral margins weakly convex and convergent to moderately rounded anterior angles, lateral border and ocular canthus producing a blunt angle, margins weakly reflexed, anterior margin distinctly sinuate medially; surface convex and moderately shiny, finely and densely, irregularly punctate, with numerous long erect setae in coarser punctures; frontoclypeal suture feebly incised and medially curved; smooth area in front of eye approximately 1.5 times as wide as long; ocular canthus moderately long and slender, very finely and sparsely punctate, with a fine terminal seta. frons dull, with fine and moderately dense punctures and with long, erect setae. eyes small, ratio of diameter/ interocular width: 0.55. antenna with ten antennomeres; club with four antennomeres, nearly 1.2 times as long as remaining antennomeres combined. mentum convexly elevated anteriorly. pronotum. moderately wide, widest at base, lateral margins in basal half subparallel and almost straight, in anterior half evenly curved and narrowed anteriorly, anterior angles distinctly produced and sharp, posterior angles blunt and slightly rounded at tip; anterior margin convexly produced medially, broad marginal line widely missing; basal margin without marginal line; surface with moderately dense and fine punctures, with minute setae and dense, long, erect setae; anterior and lateral borders sparsely setose; hypomeron distinctly carinate at base. scutellum narrow and long, sharp at apex, with fine and moderately dense punctures and minute setae, a few longer ones. elytra. oblong, widest in posterior third, striae distinctly impressed, finely and densely punctate, intervals weakly convex and not densely punctate, with small and larger punctures, with numerous fine adpressed and long erect setae, small punctures with minute setae; epipleural edge robust, ending at strongly curved external apical angle of elytra, epipleura densely setose, apical border membranous, with short microtrichomes. ventral surface. dull, with large and dense punctures, sparsely and shortly setose, setae partly adpressed; metacoxa glabrous, with fine setae laterally, apical margin convex and external apical angle rounded; each abdominal sternite with a distinct transversal row of coarse punctures each bearing a short seta between fine and moderately dense punctation. mesosternum between mesocoxae nearly half as wide as mesofemur, with irregularly scattered, fine setae. ratio of length of metepisternum / metacoxa: european journal of taxonomy 888: 97–110 (2023) 102 1 / 1.5. pygidium strongly convex, finely and densely punctate, with narrow smooth midline, with moderately dense, long setae on apical half. legs. slender; femora with two longitudinal rows of setae, finely and moderately densely punctate; metafemur ventrally dull, anterior margin sharply carinate, without a submarginal serrated line, posterior margin moderately convex, with a few strong setae medially, only weakly widened externally in apical half and not serrated ventrally in distal half, finely serrated dorsally, with dense, short setae. metatibia slender and long, widest at apex, ratio width / length: 1 / 3.9, dorsal margin sharply carinate, with two groups of spines, basal group of spines at half of metatibial length, apical one at about three quarters of metatibial length, basally with a few single, fine spines; external face longitudinally convex, with fine, sparse punctures, sparsely minutely setose; ventral margin finely serrated, with four fine, equidistant spines; medial face impunctate, apex concavely truncate interiorly near tarsal articulation. tarsomeres dorsally impunctate, with sparse, short setae ventrally; metatarsomeres glabrous dorsally, with a strongly serrated ridge ventrally, and with a fine longitudinal carina immediately beside it; first metatarsomere little longer than following two tarsomeres combined and distinctly longer than dorsal tibial spur. protibia long, bidentate, protarsal claws symmetrical, basal tooth of inner protarsal claw bluntly truncate apically. aedeagus. fig. 1f–h. variation length: 8.4–8.6 mm, length of elytra: 6.1–6.2 mm, width: 4.9–5.0 mm. female unknown. neoserica daxue sp. nov. urn:lsid:zoobank.org:act:5141d981-8c74-4916-ac62-ce039c5ebf60 fig. 2a–e diagnosis neoserica daxue sp. nov. differs from n. xiaguanensis ahrens, liu & fabrizi, 2014 by the shape of the ventral phallobasal process, which is much wider, by the dorsal phallobasal process (dorsal view), which is not widened at apex, as well as by the shape of both parameres being slightly more robust but similar in shape. etymology the new species is named (name: noun in apposition) for its occurrence in the daxue shan, china. type material examined holotype china • ♂; “china, daxue shan mts., sichuan, mianning, 1850 m, 6.vi.1999, 28°33’n, 102°10’e, v. siniaev & a. plutenko lgt. / 1153 sericini: asia spec.”; cp. description measurements. length: 8.4 mm, length of elytra: 5.6 mm, width: 4.6 mm. habitus and coloration (fig. 2e). body oblong, reddish brown, ventral surface and antenna yellow, dorsal surface dull and almost glabrous. https://zoobank.org/urn:lsid:zoobank.org:act:5141d981-8c74-4916-ac62-ce039c5ebf60 ahrens d., updates on the neoserica vulpes group 103 head. labroclypeus subtrapezoidal, widest at base, lateral margins weakly convex and convergent to moderately rounded anterior angles, lateral border and ocular canthus producing a distinct, blunt angle, margins weakly reflexed, anterior margin distinctly sinuate medially; surface flat and shiny, coarsely and densely punctate, with a few long, erect setae; frontoclypeal suture feebly incised and medially curved; smooth area in front of eye approximately 1.2 times as wide as long; ocular canthus moderately long and slender, very finely and sparsely punctate, with a fine terminal seta. frons dull, with fine and moderately dense punctures and with a few long setae beside eyes and behind frontoclypeal suture. eyes moderately large, ratio of diameter / interocular width: 0.65. antenna with ten antennomeres; club with four antennomeres, nearly 1.3 times as long as remaining antennomeres combined. mentum convexly elevated anteriorly. pronotum. moderately wide, widest shortly before base, lateral margins evenly curved and narrowed anteriorly and posteriorly, anterior angles well produced and sharp, posterior angles blunt; anterior margin convexly produced medially, broad marginal line widely missing; basal margin without marginal line; surface with moderately dense and fine punctures, with minute setae, otherwise glabrous; anterior and lateral borders sparsely setose; hypomeron distinctly carinate at base. scutellum narrow and long, sharp at apex, smooth on basal midline, with minute setae in punctures only. elytra. oblong, widest in posterior third, striae distinctly impressed, finely and densely punctate, intervals convex, coarsely and moderately densely punctate, on odd intervals with a few long erect setae, otherwise with minute setae; epipleural edge robust, ending at strongly curved external apical angle of elytra, epipleura densely setose, apical border membranous, with short microtrichomes. ventral surface. dull, with large and dense punctures, sparsely and shortly setose, setae partly adpressed; metacoxa glabrous, with fine setae laterally, apical margin convex and external apical angle rounded; each abdominal sternite with a distinct transversal row of coarse punctures each bearing a short seta between fine and moderately dense punctation. mesosternum between mesocoxae nearly half as wide as mesofemur, with irregularly scattered, fine setae. ratio of length of metepisternum / metacoxa: 1 / 1.6. pygidium moderately convex, finely and densely punctate, without smooth midline, with short setae beside apical margin. legs. slender; femora with two longitudinal rows of setae, finely and moderately densely punctate; metafemur ventrally dull, anterior margin sharply carinate, without a submarginal serrated line, posterior margin moderately convex, with a few strong setae medially, only weakly widened externally in apical half and not serrated ventrally in distal half, finely serrated dorsally, with dense, short setae. metatibia slender and moderately long, widest at apex, ratio width / length: 1 / 3.1, dorsal margin sharply carinate, with two groups of spines, basal group shortly before middle, apical one at about three quarters of metatibial length, basally with a few single, fine spines; external face longitudinally convex, with coarse, sparse punctures, glabrous; ventral margin finely serrated, with four fine, equidistant spines; medial face impunctate, apex concavely truncate interiorly near tarsal articulation. tarsomeres dorsally impunctate, with sparse, short setae ventrally; metatarsomeres glabrous dorsally, with a strongly serrated ridge ventrally, and with a fine longitudinal carina immediately beside it; first metatarsomere little longer than following two tarsomeres combined and distinctly longer than dorsal tibial spur. protibia long, bidentate, protarsal claws symmetrical, basal tooth of inner protarsal claw bluntly truncate apically. aedeagus. fig. 2a–d. female unknown. european journal of taxonomy 888: 97–110 (2023) 104 neoserica yanyuan sp. nov. urn:lsid:zoobank.org:act:91b0585f-fcd7-4582-bdd0-a58a0b56196a fig. 2f–i diagnosis neoserica yanyuan sp. nov. is in external and genital shape very similar to n. rubellula ahrens, liu & fabrizi, 2014. neoserica yanyuan differs from n. rubellula by the shape of the ventral phallobasal process, which has a slightly more extended apical convexity and being more sharply pointed, as well as by the shape of both parameres: the right paramere has a shorter basal piece, with the lateral branch being longer and more curved; the left paramere is more strongly curved (lateral view). fig. 2. a–e. neoserica daxue sp. nov., holotype, ♂ (cp). f–i. n. yanyuan sp. nov., holotype, ♂ (zfmk). a, f. aedeagus, left side lateral view. b. distal phallobase, dorsal view. c, g. parameres, dorsal view. d, h. aedeagus, right side lateral view. e, i. habitus (not to scale). scale bars = 0.5 mm. https://zoobank.org/urn:lsid:zoobank.org:act:91b0585f-fcd7-4582-bdd0-a58a0b56196a ahrens d., updates on the neoserica vulpes group 105 etymology the new species is named (name: noun in apposition) after its type locality, yanyuan, china. type material examined holotype china • ♂; “china s sichuan 2017 s of yanyuan, cguandongzi massiv, ca. 27:21:04-34 / 101:30:2533, 3200m 8-19.6. leg. c. reuter/ 1155 sericini: asia spec.”; zfmk. description measurements. length: 9.2 mm, length of elytra: 7.1 mm, width: 5.2 mm. habitus and coloration (fig. 2i). body oblong, reddish brown, ventral surface light reddish brown, antenna yellow, dorsal surface dull and glabrous. head. labroclypeus subtrapezoidal, widest at base, lateral margins weakly convex and convergent to moderately rounded anterior angles, lateral border and ocular canthus producing a distinct, blunt angle, margins weakly reflexed, anterior margin moderately sinuate medially; surface convex and moderately shiny, coarsely and densely punctate, with a few long setae; frontoclypeal suture feebly incised and medially bluntly angled; smooth area in front of eye approximately 1.5 times as wide as long; ocular canthus moderately long and slender, very finely and sparsely punctate, terminal seta absent. frons dull, with fine and sparse punctures, glabrous. eyes large, ratio of diameter / interocular width: 0.8. antenna with ten antennomeres; club with four antennomeres, nearly 1.3 times as long as remaining antennomeres combined. mentum convexly elevated anteriorly. pronotum. moderately wide, widest at base, lateral margins evenly curved and narrowed anteriorly, anterior angles well produced and sharp, posterior angles blunt; anterior margin convexly produced medially, broad marginal line widely missing; basal margin without marginal line; surface with moderately dense and fine punctures, with minute setae, otherwise glabrous; anterior and lateral borders sparsely setose; hypomeron distinctly carinate at base. scutellum narrow and long, sharp at apex, punctures sparse on basal midline, with minute setae in punctures only. elytra. oblong, widest in posterior third, striae finely impressed, finely and densely punctate, intervals weakly convex, finely and moderately densely punctate, on odd intervals with a few short adpressed setae, otherwise with minute setae; epipleural edge robust, ending at strongly curved external apical angle of elytra, epipleura densely setose, apical border membranous, with short microtrichomes. ventral surface. dull, with large and dense punctures, sparsely and shortly setose, setae partly adpressed; metacoxa glabrous, with fine setae laterally, apical margin convex and external apical angle rounded; each abdominal sternite with a distinct transversal row of coarse punctures each bearing a short seta between fine and moderately dense punctation. mesosternum between mesocoxae nearly half as wide as mesofemur, with irregularly scattered, fine setae. ratio of length of metepisternum / metacoxa: 1 / 1.33. pygidium moderately convex, finely and densely punctate, without smooth midline, with short and sparse setae on apical half. legs. slender; femora with two longitudinal rows of setae, finely and moderately densely punctate; metafemur ventrally dull, anterior margin sharply carinate, without a submarginal serrated line, posterior margin moderately convex, with a few strong setae medially, only weakly widened externally in apical half and not serrated ventrally in distal half, finely serrated dorsally, with dense, short setae. metatibia slender and long, widest at apex, ratio width / length: 1 / 4.1, dorsal margin sharply carinate, with two groups of spines, basal group shortly before middle, apical one at about four fifths of metatibial length, european journal of taxonomy 888: 97–110 (2023) 106 between them with another or two robust spines, basally with a few single, fine spines; external face longitudinally convex, with coarse, sparse punctures, glabrous; ventral margin finely serrated, with four fine, equidistant spines; medial face impunctate, apex concavely truncate interiorly near tarsal articulation. tarsomeres dorsally impunctate, with sparse, short setae ventrally; metatarsomeres glabrous dorsally, with a strongly serrated ridge ventrally, and with a fine longitudinal carina immediately beside it; first metatarsomere little longer than following two tarsomeres combined and distinctly longer than dorsal tibial spur. protibia long, bidentate, protarsal claws symmetrical, basal tooth of inner protarsal claw bluntly truncate apically. aedeagus. fig. 2f–h. female unknown. further records of species of the neoserica vulpes group neoserica biuncinata ahrens, liu & fabrizi, 2014 additional material examined china • 1 ♂; “china, s. sichuan, kiangshan mts., s. xichang h=3000m, 1.vii.2002 leg. s. murzin, i. shokhin”; cp. neoserica nykli ahrens, liu & fabrizi, 2014 additional material examined china • 1 ♂; “china, yunnan 1993 50 km n lijiang, 24.-29.6. yulongshan nat. res. leg. e. jendek & o. sausa”; cp. updated key to species of neoserica vulpes group (males) 1. phallobase with a ventral process ..................................................................................................... 4 – phallobase without a ventral process, but with a lateral one ............................................................ 2 2. right paramere large and spherical, longitudinal extension half of phallobasal length ..................... ......................................................................................n. baishuiensis ahrens, liu & fabrizi, 2014 – right paramere small, spherical, longitudinal extension one third of phallobasal length ................ 3 3. lateral process sharply pointed (dorsal view) .......... n. lateriuncinata ahrens, liu & fabrizi, 2014 – lateral process convexly rounded at apex (dorsal view) .................................................................... .......................................................................................... n. leiboensis ahrens, liu & fabrizi, 2014 4. phallobase with a dorsal process ...................................................................................................... 5 – phallobase without a dorsal process ................................................................................................. 6 5. ventral phallobasal process narrow. dorsal phallobasal lobe widened at apex. both parameres slenderer ..................................................................... n. xiaguanensis ahrens, liu & fabrizi, 2014 – ventral phallobasal process much wider. dorsal phallobasal lobe not widened at apex. both parameres more robust ............................................................................................................. n. daxue sp. nov. 6. ventral process of phallobase at apex widened, with sharp hooks or bent processes ....................... 7 – ventral process of phallobase at apex convexly rounded or pointed .............................................. 18 ahrens d., updates on the neoserica vulpes group 107 7. ventral process of phallobase at apex convexly widened, without sharp hooks ................................ ........................................................................................ n. luzhouana ahrens, liu & fabrizi, 2014 – ventral process of phallobase at apex with sharp hooks or bent processes ...................................... 8 8. right paramere deeply bifurcate. hooks small ................................................................................. 9 – right paramere simple, not bifurcate. hooks large .........................................................................11 9. ventral process in cross section circular ......................................................................................... 10 – ventral process dorsoventrally flattened ............................n. parausta ahrens, liu & fabrizi, 2014 10. ventral phallobasal process with less extended apical convexity and more less pointed. right paramere with longer basal piece, its lateral branch is short and straight; left paramere straight (lateral view) ..................................................................... n. rubellula ahrens, liu & fabrizi, 2014 – ventral phallobasal process with a slightly more extended apical convexity and more sharply pointed. right paramere with shorter basal piece, its lateral branch is longer and more curved; left paramere strongly curved (lateral view) ............................................................................. n. yanyuan sp. nov. 11. left paramere at apex strongly curved ventrally ............................................................................ 12 – left paramere at apex straight ....................................... n. biuncinata ahrens, liu & fabrizi, 2014 12. eyes smaller, ratio diameter / interocular with: 0.58. ventral process of phallobase exceeding parameres .................................................................................. n. usta ahrens, liu & fabrizi, 2014 – eyes moderate to large, ratio diameter / interocular with > 0.7 ....................................................... 13 13. ventral process of phallobase exceeding parameres ....................................................................... 14 – ventral process of phallobase ending distally at same point as parameres ..................................... 15 14. ventral phallobasal process straight (lateral view), its ventral distal tooth well pronounced. parameres shorter and less strongly curved ..............................................n. nykli ahrens, liu & fabrizi, 2014 – ventral phallobasal process well curved and its ventral distal tooth little pronounced. parameres longer and more strongly curved ................................................................ n. mianningana sp. nov. 15. ventral process of phallobase basally very enlarged and strongly dorsoventrally produced on right side, its width at base subequal to half of phallobase length .......................................................... 16 – ventral process of phallobase basally only weakly enlarged and not dorsoventrally produced on right side, its width at base subequal to less than third of phallobase length .......................................... 17 16. left paramere one third as wide as long ........................ n. sichuanica ahrens, liu & fabrizi, 2014 – left paramere half as wide as long .......................................n. dundai ahrens, liu & fabrizi, 2014 17. ventral process of phallobase at middle strongly bent dorsally ......................................................... .................................................................................... n. pseudovulpes ahrens, liu & fabrizi, 2014 – ventral process of phallobase nearly straight or only weakly bent at middle dorsally ....................... ......................................................................................n. ganhaiziana ahrens, liu & fabrizi, 2014 18. ventral process of phallobase bent or strongly curved ................................................................... 19 – ventral process of phallobase straight ............................................................................................ 24 19. left paramere bifurcate, divided in two long lobes ........................................................................ 20 – left paramere simple, not divided in two long lobes ..................................................................... 21 european journal of taxonomy 888: 97–110 (2023) 108 20. ventral lobe of left paramere distinctly wider than dorsal one (lateral view) ..................................... .................................................................................. n. ningyuanensis ahrens, liu & fabrizi, 2014 – ventral lobe of left paramere as wide as dorsal one (lateral view) ..................................................... ........................................................................................n. heishuiana ahrens, liu & fabrizi, 2014 21. ventral process of phallobase sharply pointed at apex, curved slightly to the right ........................... ................................................................................... n. kunmingensis ahrens, liu & fabrizi, 2014 – ventral process of phallobase convex at apex, curved slightly to the left ...................................... 22 22. curvation of ventral process moderate, not exceeding width of process at middle ........................... ................................................................................................n. kereni ahrens, liu & fabrizi, 2014 – curvation of ventral process strong, exceeding width of process at middle .................................. 23 23. distal hook of ventral phallobasal process longer (dorsal view) . ....................................................... .......................................................................................... n. baoshana ahrens, liu & fabrizi, 2014 – distal portion of ventral phallobasal process short and little hooked (dorsal view) .......................... ................................................................................................................... n. myanmarensis sp. nov. 24. ventral process of phallobase in basal cross section circular ......................................................... 25 – ventral process of phallobase in basal cross section flattened ............................................................ .......................................................................................... n. laocaiana ahrens, liu & fabrizi, 2014 25. ventral process of phallobase evenly widened towards apex ............................................................. .................................................................................... n. weishanensis ahrens, liu & fabrizi, 2014 – ventral process of phallobase not distinctly widened towards apex ............................................... 26 26. right paramere spherical, at apex with a single process ................................................................ 27 – right paramere spherical, but at apex with two separate processes ............................................... 28 27. left paramere wider (dorsal view), strongly blown up and spherical (lateral view); ventral process of phallobase distinctly widened apically .................................................... n. vulpes (arrow, 1946) – left paramere narrower (dorsal view), less blown up and less spherical (lateral view); ventral process of phallobase not widened apically ...................... n. yangjiapingensis ahrens, liu & fabrizi, 2014 28. left paramere at apex slightly less than total width of paramere (dorsal view). parameres nearly elongate in lateral view ................................................................................................................... 29 – left paramere at apex distinctly less than one third of total width of paramere. parameres subsphaerical in lateral view ..................................................................... n. ruzickai ahrens, liu & fabrizi, 2014 29. parameres narrow, strongly bent to the left side (dorsal view) .................. n. alaoshanensis sp. nov. – parameres wider, nearly straight (dorsal view) ......... n. shinkaisiensis ahrens, liu & fabrizi, 2014 discussion this is the third contribution on the knowledge of the species of the neoserica vulpes group, and it shows that the extensive and comprehensive revisionary works done so far are still far from being conclusive, which makes updates with descriptions of individual species technically and scientifically necessary. we expect further new discoveries with more rigorous light trapping programs all over asia, although that is technically not always easy for logistical reasons. this study once more reveals a large amount of endemism in asian sericini and demonstrates that night active chafers are still rather poorly represented in material from occasional, non-targeted field surveys. ahrens d., updates on the neoserica vulpes group 109 acknowledgements i am grateful to petr pacholátko (brno) for the loan of his interesting specimens and to michal langer and christoph reuter (hamburg) for the donation of their specimens. furthermore, i thank max barclay and the anonymous referees for the helpful comments on the manuscript. references ahrens d. 2003. zur identität der gattung neoserica brenske, 1894, nebst beschreibung neuer arten (coleoptera, melolonthidae, sericini). koleopterologische rundschau 73: 169–226. ahrens d. 2004. monographie der sericini des himalaya (coleoptera, scarabaeidae). dissertation.de – verlag im internet gmbh, berlin. ahrens d. 2021. further two new species of the neoserica (sensu stricto) group from china (coleoptera: scarabaeidae: sericini). journal of natural history 54 (45–46): 2927–2936. https://doi.org/10.1080/00222933.2021.1884761 ahrens d. 2022. a new gynaecoserica brenske, 1897 species and further new bicolored species of the neoserica calva group (coleoptera: scarabaeidae: melolonthinae: sericini) (coleoptera: scarabaeidae: sericini). zootaxa 5165 (2): 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scarabaeidae, sericini). zookeys 448: 83–121. https://doi.org/10.3897/zookeys.448.8429 liu w.g., bai m., yang x.k. & ahrens d. 2015a. new species and records of the neoserica (sensu stricto) group (coleoptera: scarabaeidae: sericini). journal of natural history 49 (39–40): 2379–2395. https://doi.org/10.1080/00222933.2015.1034208 liu w.g., bai m., eberle j., yang x.k. & ahrens d. 2015b. a phylogeny of sericini with particular reference to chinese species using mitochondrial and ribosomal dna (coleoptera: scarabaeidae). organisms, diversity and evolution 15: 343–350. https://doi.org/10.1007/s13127-015-0204-z liu w.g., fabrizi s., bai m., yang x.k. & ahrens d. 2016. a taxonomic revision of neoserica (sensu lato): the species groups n. lubrica, n. obscura, and n. silvestris (coleoptera, scarabaeidae, sericini). zookeys 635: 123–160. https://doi.org/10.3897/zookeys.635.9915 liu w.g., fabrizi s., bai m., yang x.k. & ahrens d. 2019. a taxonomic revision of chinese neoserica (sensu lato): final part (coleoptera: scarabaeidae: sericini). bonn zoological bulletin supplementum 64: 1–71. manuscript received: 22 october 2022 manuscript accepted: 15 march 2023 published on: 11 august 2023 topic editor: tony robillard section editor: max barclay desk editor: pepe fernández printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; leibniz institute for the analysis of biodiversity change, bonn – hamburg, germany; national museum of the czech republic, prague, czech republic. https://doi.org/10.3897/zookeys.448.8368 https://doi.org/10.3897/zookeys.448.8429 https://doi.org/10.1080/00222933.2015.1034208 https://doi.org/10.1007/s13127-015-0204-z https://doi.org/10.3897/zookeys.635.9915 1 european journal of taxonomy 670: 1–14 issn 2118-9773 https://doi.org/10.5852/ejt.2020.670 www.europeanjournaloftaxonomy.eu 2020 · zaova d. et al. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e tertiarius minutulus sp. nov. (stephanodiscaceae, bacillariophyta) – a new fossil diatom species from lake ohrid dušica zaova 1,*, aleksandra cvetkoska 2, danijela mitic-kopanja 3, elena jovanovska 4, nadja ognjanova-rumenova 5 & zlatko levkov 6 1,3,6 institute of biology, faculty of natural sciences, arhimedova 3, 1000 skopje, r. north macedonia. 2 department of aquatic ecology, netherlands institute of ecology (nioo-knaw), wageningen, netherlands. 4 institute for ecology, evolution and diversity, goethe university, frankfurt, germany. 4 department of animal ecology and systematics, justus liebig university giessen, giessen, germany. 5 department of palaeontology, stratigraphy and sedimentology, geological institute, bulgarian academy of sciences, sofia, bulgaria. * corresponding author: zaovadusica@gmail.com 2 email: acvetkoska@yahoo.com 3 email: mitickopanja@yahoo.com 4 email: jovanovska.eci@gmail.com 5 email: ognjanovanadja@gmail.com 6 email: zlevkov@pmf.ukim.mk abstract. a new fossil diatom species, tertiarius minutulus sp. nov., is described from a sediment sequence deep-5045-1 of lake ohrid. the species is characterized by small valves (3.0–8.0 µm) with a round shape, a marginal area with radially arranged costae and a central uneven area with scattered areolae. externally, the alveoli are occluded by cribra perforated by irregularly arranged pores. internally, the alveoli are simple and areolae are occluded with domed cribra. the marginal fultoportulae are situated on costae close to the valve margin, located on every 5th or 7th thick internal costa. one to three fultoportulae are present on the valve face, each surrounded by two to three satellite pores. one rimoportula is present, positioned on a costa at the valve face / mantle junction. the species is compared with morphologically similar taxa and a detailed differential diagnosis is provided. tertiarius minutulus sp. nov. is known only as a fossil taxon from lake ohrid, with a first occurrence during the early stages of lake basin development. keywords. ancient lake, diatom fosil record, new species, taxonomy. zaova d., cvetkoska a., mitic-kopanja d., jovanovska e., ognjanova-rumenova n. & levkov z. 2020. tertiarius minutulus sp. nov. (stephanodiscaceae, bacillariophyta) – a new fossil diatom species from lake ohrid. european journal of taxonomy 670: 1–14. https://doi.org/10.5852/ejt.2020.670 https://doi.org/10.5852/ejt.2020.670 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ mailto:zaovadusica%40gmail.com?subject= mailto:acvetkoska%40yahoo.com?subject= mailto:mitickopanja%40yahoo.com?subject= mailto:jovanovska.eci%40gmail.com?subject= mailto:ognjanovanadja%40gmail.com?subject= mailto:zlevkov%40pmf.ukim.mk?subject= https://doi.org/10.5852/ejt.2020.670 european journal of taxonomy 670: 1–14 (2020) 2 introduction lake ohrid, in the balkan peninsula, was formed between 1.9 and 1.3 ma ago (wagner et al. 2017). it is the oldest existing european lake and one of the few long-lived lakes around the world (stanković 1960; radoman 1985; dumurdžanov et al. 2004; albrecht & wilke 2008). the lake has accumulated a wealth of sediments throughout its limnological history, which are well constrained in age and hold well-preserved fossils, of which diatoms (bacillariophyta karsten) are one of the most diverse and abundant groups of organisms (e.g., cvetkoska et al. 2016). in order to investigate in greater details the age and origin of lake ohrid, and link its geological and biological evolution, a deep drilling project was performed in 2013 within the frames of the icdp project “scientific collaboration on past speciation conditions in lake ohrid (scopsco)” (wagner et al. 2014). a total of ~ 2100 m of sediment was recovered from four different sites – deep, cerava, gradište and peštani. the drilling in the central part of the lake resulted in a recovery of a 569 m long sediment sequence (deep-5045-1). observations of the diatom communities along the entire length of the core revealed the dominance of planktonic diatoms, mostly belonging to genera of the family stephanodiscaceae gleser & markarova (wagner et al. 2014; cvetkoska et al. 2016; jovanovska et al. 2016a). the majority of them belongs to cyclotella (kützing) brébisson s. l., while high abundances of species belonging to stephanodiscus ehrenberg and cribrionella jovanovska, cvetkoska, tofilovska, ognjanova-rumenova & levkov were also observed in the older parts of the sediment sequence (wagner et al. 2014, cvetkoska et al. 2016). in previous studies on this core, several new diatom species such as cribrionella ohridana jovanovska, cvetkoska, tofilovska, ognjanova-rumenova & levkov (jovanovska et al. 2016b), cyclotella cavitata tofilovska, cvetkoska, jovanovska, ognjanova-rumenova & levkov and cyclotella sollevata tofilovska, cvetkoska, jovanovska, ognjanova-rumenova & levkov (tofilovska et al. 2016) were described. further studies on this core resulted in a record of a new species bearing unique morphological features that is described here as tertiarius minutulus sp. nov. additional analyses of core samples from lake ohrid are of great importance for improving the understanding of the palaeoecology and biochronology of freshwater diatoms, their evolution and the main environmental factors that shaped the diatom community structure over geological time. material and methods lake ohrid (north macedonia / albania) is located in a tectonically active graben system in western macedonia. it is situated in a karstic graben within the southern balkan extensional regime (lindhorst et al. 2010). earlier studies hypothesized that the age of the lake is between 2 and 10 ma, thus considering it the oldest european lake (stanković 1960; radoman 1985). most recent results from the analyses of the deep site sediment sequence suggested that the lake established between 1.9 and 1.3 ma (wagner et al. 2017). the lake is ~ 30 km long and 15 km wide, located at an altitude of 693 m above sea level. the maximum water depth is 289 m, and the total volume of the lake is 55.4 km3 (lindhorst et al. 2014). the total water inflow can be estimated to 37.9 m3s-1, with ca 25% originating from direct precipitation and 25% from riverine inflow (wagner et al. 2010). about 50% of the total inflow derives from karst aquifers, of which ca 8 m3s-1 are supposed to come from lake prespa (wagner et al. 2010). within the scopsco project in 2013, a total of 2100 m of sediments was recovered from four different drilling locations (deep, cerava, gradište and peštani). in this study, only core samples from the deep site were used. this site is located in the central part of the lake at a water depth of 245 m (franche et al. 2016), where drilling resulted in a maximum penetration of 569 m below the lake floor (wagner et al.2014). the new species, tertiarius minutulus sp. nov., was observed at 452 m of the composite sequence that corresponds to an age older than 1.3 ma (wagner et al. 2019). the sediment in this part is a mixture of sand, silt and clay (wagner et al. 2014) for diatom analyses, ca 0.1 g of wet sediment was sampled and stored in sterilin tubes at 4°c. samples were prepared using a modification of renberg’s technique (1990). diatom samples were acid cleaned zaova d. et al., tertiarius minutulus, a new fossil diatom from lake ohrid 3 by adding a few drops of cold 10% hcl and 35% h2o2, and left overnight to remove carbonates. the samples were then boiled ca 30 min in 37% hcl to oxidize the organic matter. samples were rinsed several times with distilled water and subsequently centrifuged. diatom slides were prepared using naphrax®. slides were observed under oil immersion at 1500 × magnification with a nikon eclipse 80i microscope, and diatom images were produced using a nikon coolpix p6000 camera. for scanning electron microscopy (sem), the material was prepared by drying clean diatom suspension onto cover slips that were carbon tape attached to the sem stubs and coated with gold-palladium (polaron sc7640 sputter coater, quorum technologies, ashford, uk). sem observations were performed using a cambridge s4 stereoscan at 10 kv (cambridge instruments ltd, cambridge, uk) at the friedrich hustedt study centre for diatoms (bm), bremerhaven, germany. the holotype slide is designated and deposited at the macedonian national diatom collection (mkndc), faculty of natural sciences, skopje, republic of north macedonia. an isotype slide is hosted at the friedrich hustedt study centre for diatoms (bm), bremerhaven, germany. terminology followes håkansson & khursevich (1997), houk et al. (2010) and ognjanova-rumenova et al. (2015). results phylum bacillariophyta karsten class mediophyceae medlin & kaczmarska order stephanodiscales nikolaev & harwood family stephanodiscaceae gleser & markarova genus tertiarius håkansson & khursevich tertiarius minutulus sp. nov. figs 1–3 etymology the specific epithet ʻminutulusʼ refers to its small size. type material holotype republic of north macedonia • lake ohrid, core deep-5045-1; 41.049167° n, 20.715000° e; sediment sample from 451.92 m depth of the composite core; mar.–jun. 2013; mkndc 010093. isotype republic of north macedonia • 1 spec.; same collection data as for holotype; bm 81918 (slide). type locality lake ohrid core 5045-1, site deep, at a depth of 451.92 m of the composite sequence. description light microscope (lm) valves circular, 3.5–8.0 µm in diameter, central area 3.0–6.5 µm in diameter (fig. 1). valve face uneven with two parts of distinctly different morphology. marginal area with short, radiating striae exceeding 1/8–¼ of valve diameter, with ca 24–30 striae in 10 µm. central area uneven and colliculate with scattered central areolae that are not radially arranged. frustules rectangular in connective view. european journal of taxonomy 670: 1–14 (2020) 4 sem external view (fig. 2) valve face uneven and colliculate (fig. 2a–c), with small granules in central area (fig. 2d–e). areolae present in central area with simple round openings. marginal striae short, occluded by cribra, perforated with pores of variable size. line of bigger pores bordering each costa (fig. 2d–e). marginal fultoportulae situated on costae, close to margin edge (fig. 2a–b), situated on every 6th–7th costa. externally marginal fultoportulae with simple, slightly elevated and round opening (fig. 2c, e). one to three valve face fultoportulae present, with external small openings, round (fig. 2a–b). single solid ligulate girdle band present (fig. 2a). sem internal view (fig. 3) internal valve face flat with several central areolae occluded with domed cribra (arrow in fig. 3b). marginal areolae loculate, crossed with inwardly raised circumferential silica trabeculae (arrowhead in fig. 3b), while central areolae occluded with domed cribra (fig. 3b–c). costae strongly thickened, ca 30 in 10 µm (fig. 3a, e–f). marginal fultoportulae with short tube located on short and narrow costae (fig. 3a). each marginal fultoportula with two satellite pores in lateral position (arrow in fig. 3a). one to three valve face fultoportulae present. valve face fultoportulae with short central tube surrounded with 2–3 satellite pores (arrow in fig. 3e). one rimoportula present, located at valve face / mantle junction in the middle of costa (arrow in fig. 3f) or on its side (arrow in fig 3d). rimoportula consists of short and narrow labium with oblique slit (fig. 3d–f). distribution freshwater fossil species observed only at its type locality, lake ohrid. differential diagnosis the main differential features of t. minutulus sp. nov. that can be observed in lm are its small valve size (diameter = 3.5–8.0 µm) and the presence of central scattered areolae that are not radially arranged. tertiarius pygmaeus (pantocsek) håkansson & khursevich (1997: 22) is characterized by a valve diameter ranging from 8.0–16.0 µm, and occasionally the areolae are not arranged in radial striae (e.g., pantocsek 1892: figs 2: 22, 4: 59; houk et al. 2010: fig. 296: 1, 4). important differences between t. minutulus sp. nov. and t. pygmaeus can be observed with sem. the marginal striae in t. pygmaeus do not have a complex alveolar structure and are composed of regular rows of fine pori. externally, distinct fig. 1. tertiarius minutulus sp. nov., lm valve views, lake ohrid, north macedonia. the asterisk indicates the holotype. scale bar = 10 µm. zaova d. et al., tertiarius minutulus, a new fossil diatom from lake ohrid 5 fig. 2. tertiarius minutulus sp. nov., sem external valve views. a. valve view showing the marginal openings of the fultoportulae (see white arrow). b–d. view of the whole frustule. e. close view of marginal openings of the fultoportula (see white arrow). f. close view of the external openings of the valve face fultoportulae (see white arrows). scale bars: a–d = 2.0 µm; e–f = 1.0 µm. european journal of taxonomy 670: 1–14 (2020) 6 fig. 3. tertiarius minutulus sp. nov., sem internal valve views. a. view of the whole frustule and the marginal fultoportulae surrounded by two satellite pores, cowlings absent (see white arrow). b. close view of marginal areolae with circumferential silica trabeculae and central areolae occluded by domed cribra (see white arrow). c. view of the whole frustule. d. close view of rimoportula (see white arrow) located on costa at junction between valve face / mantle. e. internal valve view showing the central fultoportulae with short central tube surrounded with 2–3 satellite pores (see white arrows). f. valve view. scale bars: a, c, e–f = 2.0 µm; b, d = 1.0 µm. zaova d. et al., tertiarius minutulus, a new fossil diatom from lake ohrid 7 spines are present on every costa. internally, the alveolus is round and small in t. pygmaeus whereas it is larger and more elongated in t. minutulus sp. nov. in t. pygmaeus, the marginal fultoportulae are present on each costa, the number of valve face fultoportulae is one to seven scattered in the central area, and the rimoportula is situated inside of the alvelolar opening on a side of costa. additionally, the costae are much thicker and with a lower density (8–10 in 10 µm) in t. pygmaeus than in t. minutulus sp. nov. another species with small-sized valves is t. mariovensis ognjanova-rumenova, jovanovska, cvetkoska & levkov (diameter 4.0–19.0 µm), recently described from a diatomite near the village manastir, mariovo neogene basin, republic of north macedonia (ognjanova-rumenova et al. 2015: 56, figs 36–105). this species was also observed in several sediment samples from lake ohrid deep5045-1 core. however, both species can easily be differentiated by the structure of the central area. in t. mariovensis, the areolae in the central area form radially arranged striae. the number of valve face fultoportulae in t. mariovensis is higher (2–9) and the tube of the central fultoportula is always surrounded by three satellite pores. additionally, the marginal fultoportulae are positioned on every costa, while the rimoportula is located inside and in the middle of the alveolus, but not connected to a costa as in t. minutulus sp. nov. another small-sized species, tertiarius distinctus khursevich & kociolek (2002: 333, figs 1–5, 12–22), has a comparable valve size as t. minutulus sp. nov. (diameter 4.5–14.0 µm), but coarser areolae arranged in short or long radial striae composed of large areolae, while the marginal striae are composed of 4–6 rows of small pori. marginal fultoportulae are present on every costa, while in the central area there are 1–5 valve face fultoportulae. the rimoportula is located near the base of a costa within the alveolus. differences between t. distinctus and t. minutulus sp. nov. can be noticed in the structure of the marginal and central striae, the number of valve face and marginal fultoportulae, and the position of the rimoportula. smaller specimens of t. indigenus khursevich & kociolek (2002: 336, figs 6–11, 23–32) have a valve size comparable to t. minutulus sp. nov. (diameter 5.0–27.5 µm). however, differences between these two species can be noticed in the striae structure (longer marginal striae composed of 4–5 regular rows or pori), central area (colliculate central area with 3–5 large areolae and several ʻbumpsʼ), presence of fultoportulae on each costa and one to several rimportulae located on thinner costae within the alveolar chamber. tertiarius baicalensis khursevich & fedenya (in khursevich et al. 2003: 306, figs 1: 1, 2, 4, 5, 14) is characterized by circular valves with flat face and diameter of 6.4–16.6 µm, three to 17 valve face fultoportulae arranged in radial rows and one to four sessile rimoportulae located in the submarginal zone of the valve face. differences between t. minutulus sp. nov. and t. baicalensis can be noticed in the ornamentation of the central area (areolae organized in radial striae in t. baicalensis), number and position of valve face fultoportulae, number of satellite pores on marginal fultoportulae (three satellite pores in t. baicalensis), and position of rimoportula(e) that are located in the submarginal zone of the valve face (not connected with costa or within the alveolar chamber). another small-celled species, cribrionella ohridana, was recently described from the same core from lake ohrid (jovanovska et al. 2016b). both species share several characters such as small size (2.0– 7.5 µm in diameter), the presence of areolae in the central area and a single rimoportula situated on a costa, marginal fultoportulae located on each 4th–5th costa surrounded by two satellite pores and the presence of inwardly raised circumferential silica trabeculae. however, differences between these two species can be noticed in the absence of valve face fultoportula in c. ohridana and areolae internally not occluded with domed cribra as in t. minutulus sp. nov. one of the dominant and very frequent species in fossil diatom assemblages of lake ohrid is cyclotella minuscula (jurilj) cvetkoska. cyclotella minuscula was recently transferred to lindavia (schütt) de toni & forti as l. minuscula (jurilj) nakov, guillory, julius, theriot & alverson (nakov et al. 2015) and later to the genus pantocsekiella kiss & ács as p. minuscula (jurilj) kiss & ács (ács et al. 2016). european journal of taxonomy 670: 1–14 (2020) 8 sp ec ie s v al ve s iz e (µ m ) c en tr al a re a si ze (µ m ) n um be r of s tr ia e in 1 0 µm st ru ct ur e an d po si ti on o f m an tl e fu lt op or tu la e st ru ct ur e an d po si ti on o f v al ve fa ce fu lt op or tu la e (f p ) st ru ct ur e an d po si ti on o f ri m op or tu la e (r p ) so ur ce te rt ia ri us m in ut ul us sp . n ov . 3. 5– 8. 0 3. 0– 6. 5 25 –3 0 on e ve ry 6 th –7 th co st a, w ith tw o sa te lli te p or es 1– 3 fp w ith 2 –3 sa te lli te p or es on e r p at th e va lv e fa ce / m an tle ju nc tio n on a co st a th is s tu dy te rt ia ri us b ai ca le ns is k hu rs ev ic h & f ed en ya 4– 16 1. 5– 8 10 –2 0 on e ve ry c os ta , n ea r th e va lv e ed ge w ith th re e sa te lli te p or es 3– 17 f p w ith tw o sa te lli te p or es 1– 4 r p in th e su bm ar gi na l zo ne o f t he v al ve fa ce k hu rs ev ic h et a l. 20 03 te rt ia ri us c he rn om or ic us k hu rs ev ic h & k oc io le k 10 –1 8 n a d 6– 8 on e ve ry c os ta w ith tw o sa te lli te p or es n a d on e r p at th e en d of th e su bm ar gi na l z on e of th e va lv e fa ce k hu rs ev ic h & k oc io le k 20 12 te rt ia ri us d is tin ct us k hu rs ev ic h & k oc io le k 4. 5– 14 4– 8. 5 10 –2 0 on e ve ry c os ta w ith tw o sa te lli te p or es 1– 5 fp w ith tw o sa te lli te p or es ; on e r p ne ar th e ba se o f co st ae o r w ith in a c ha m be r of th e al ve ol us k hu rs ev ic h & k oc io le k 20 02 te rt ia ri us e lg er i ( h us te dt ) v. h ou k, r .k le e & h .t an ak a 20 –1 10 10 .5 –2 4. 5 13 –1 5 on e ve ry c os ta w ith tw o sa te lli te p or es ab se nt on e r p be tw ee n st ri ae , co nn ec te d w ith a s ho rt ri b at th e va lv e m ar gi n an d on e of th e ad ja ce nt c os ta h ou k et a l. 20 10 te rt ia ri us in di ge nu s k hu rs ev ic h & k oc io le k 5– 27 .5 2. 5– 11 .5 6– 8 on e ve ry c os ta w ith tw o sa te lli te p or es ab se nt 1– 4 r p on th e th in ne r c os ta e w ith in th e al ve ol ar c ha m be rs k hu rs ev ic h & k oc io le k 20 02 te rt ia ri us h id al ge ns is c ab al le ro , k hu rs ev ic h & v el as co d e l eó n 8– 32 11 –2 0 10 –1 5 on e ve ry c os ta w ith tw o sa te lli te p or es 4– 10 f p, fo rm in g a ri ng n ea r t he v al ve ce nt re w ith 2 –3 sa te lli te p or es on e r p be tw ee n tw o co st ae in si de a lv eo la r o pe ni ng c ab al le ro e t a l. 20 09 te rt ia ri us ju ri jli i o gn ja no va -r um en ov a, jo va no vs ka , c ve tk os ka & z .l ev ko v 5. 0– 38 .0 2. 0– 24 .0 5– 8 on e ve ry c os ta w ith tw o sa te lli te p or es 5– 12 f p, w ith th re e sa te lli te p or es on e r p at th e ba se b et w ee n tw o co st ae in fr on t o f al ve ol ar o pe ni ng o gn ja no va -r um en ov a et a l. 20 15 ta bl e 1 (c on tin ue d on th e ne xt p ag e) . m or ph ol og ic al a nd u ltr as tr uc tu ra l c ha ra ct er is tic s of k no w n sp ec ie s of t er tia ri us (g ui ry & g ui ry 2 01 9) . n a d = no a va ila bl e da ta . zaova d. et al., tertiarius minutulus, a new fossil diatom from lake ohrid 9 sp ec ie s v al ve s iz e (µ m ) c en tr al a re a si ze (µ m ) n um be r of s tr ia e in 1 0 µm st ru ct ur e an d po si ti on o f m an tl e fu lt op or tu la e st ru ct ur e an d po si ti on o f v al ve fa ce fu lt op or tu la e (f p ) st ru ct ur e an d po si ti on o f ri m op or tu la e (r p ) so ur ce te rt ia ri us m ar io ve ns is o gn ja no va -r um en ov a, jo va no vs ka , c ve tk os ka & l ev ko v 4. 0– 19 .0 1. 5– 13 .5 6– 12 on e ve ry c os ta w ith th re e sa te lli te p or es 2– 9 fp , a rr an ge d in a ri ng a ro un d va lv e ce nt re w ith th re e sa te lli te p or es on e r p at th e en d of th e su bm ar gi na l z on e of th e va lv e fa ce o gn ja no va -r um en ov a et a l. 20 15 te rt ia ri us p an to cs ek ii (f ri ck e) k hu rs ev ic h & k oc io le k 18 –5 5 25 –3 2 3– 5 on e ve ry c os ta w ith th re e sa te lli te p or es n a d on e r p at th e en d of th e su bm ar gi na l z on e of th e va lv e fa ce k hu rs ev ic h & k oc io le k 20 12 te rt ia ri us p or os us k hu rs ev ic h & k oc io le k 10 –6 0 6. 5– 32 5– 7 on e ve ry c os ta w ith tw o sa te lli te p or es 1– 4 fp , w ith tw o sa te lli te p or es 1– 4 r p at th e ba se o f a n el on ga te d ch am be r k hu rs ev ic h & k oc io le k 20 02 te rt ia ri us p yg m ae us (p an to cs ek ) h åk an ss on & k hu rs ev ic h 8– 14 (1 6) 6. 5– 11 6 on e ve ry c os ta w ith tw o sa te lli te p or es 6– 8 fp , w ith tw o sa te lli te p or es on e sm al l r p po si tio ne d on a si de o f a c os ta h ou k et a l. 20 10 te rt ia ri us r od da e k oc io le k & k hu rs ev ic h 6– 45 2– 27 3– 8 on e ve ry c os ta w ith tw o sa te lli te p or es 3– 6 fp , w ith tw o sa te lli te p or es on e r p at th e su bm ar gi na l zo ne o f t he v al ve fa ce k hu rs ev ic h & k oc io le k 20 02 te rt ia ri us s at su m ae ns is (h .t an ak a & v .h ou k) n ak ov e t a l. 4. 5– 9. 5 3. 5– 7 16 –2 4 on e ve ry 3 rd –9 th c os ta on th e lo w er p ar t o f th e m an tle w ith tw o sa te lli te p or es 2– 3 w ith tw o sa te lli te po re s as so ci at ed w ith th e de pr es si on s on e r p on th ic ke ne d co st a h ou k et a l. 20 10 te rt ia ri us te m pe re ifo rm ic us (k hu rs ev ic h) n ak ov e t a l. 14 .3 –6 0 21 .5 –5 1. 5 7– 10 on e ve ry c os ta w ith tw o sa te lli te p or es ab se nt up to fo ur r p on th ic ke ne d co st a k hu rs ev ic h et a l. 20 01 te rt ia ri us tr an sy lv an ic us (p an to cs ek ) h åk an ss on & k hu rs ev ic h 22 –3 6 17 –2 9 6– 8 on e ve ry c os ta w ith tw o sa te lli te p or es in v ar ia bl e nu m be r w ith tw o sa te lli te po re s on e r p lo ca te d on a s id e of a co st a h ou k et a l. 20 10 ta bl e 1 (c on tin ue d) . m or ph ol og ic al a nd u ltr as tr uc tu ra l c ha ra ct er is tic s of k no w n sp ec ie s of t er tia ri us (g ui ry & g ui ry 2 01 9) . n a d = n o av ai la bl e da ta . european journal of taxonomy 670: 1–14 (2020) 10 both species, tertiarius minutulus sp. nov. and cyclotella minuscula, have a similar valve size (diameter 3–7 µm in c. minuscula), a colliculate central area, a small marginal zone, a single valve face central fultoportula, a marginal fultoportula with two satellite pores located on each 4th–10th costa and a single rimoportula positioned at a rib, beneath the ring of marginal fultoportulae. difference between these two species can be observed only in a presence / absence of areolae in the central area: in c. minuscula areolae are absent, while in t. minutulus sp. nov. they are present. discussion earlier studies have described 17 taxa (16 species and one variety) which belong to the genus tertiarius (guiry & guiry 2019), table 1. the initial description of the genus tertiarius is based on sediment samples of miocene and pliocene age from köpecz (transylvania, romania). furthermore, new species of tertiarius have been described from deposits in the western usa (khursevich & kociolek 2002), pliocene sediments from mexico (caballero et al. 2009) and middle pliocene sediments from lake baikal in asia (khursevich et al. 2003). in addition, two species from this genus, viz., t. jurijlii ognjanova-rumenova, jovanovska, cvetkoska & z.levkov and t. mariovensis, are described from sediments of pliocene age from mariovo basin, north macedonia (ognjanova-rumenova et al. 2015). the presence of tertiarius mariovensis is also recorded from sediments of lake ohrid, at ca 630 ka (francke et al. 2016). according to khursevich & kociolek (2012), the biostratigraphic range of species from the genus tertiarius in the northern hemisphere extends into the miocene to pliocene periods. interestingly, both t. mariovensis and t. minutulus sp. nov. have been observed in middle pleistocene up to quaternary sediments. these two recent records of tertiarius probably represent the latest occurrence of the genus in freshwater habitats, again supporting the notion of lake ohrid being refugium for many species throughout its existence. sem observations indicate that t. minutulus sp. nov. shares characteristics with the genus lindavia (schütt) de toni & forti (= handmania m.peragallo = puncticulata håkansson). the genus tertiarius is considered as fossil with a biostratigraphic range from miocene to pliocene, while lindavia (= handmannia m.peragallo) has a longer range from middle eocene to present. tertiarius is characterized by laterally positioned rimoportula on a fultoportula-bearing costa, while in lindavia, the rimoportula is located on the valve face (nakov et al. 2015). in the latter study, the authors proposed a broad concept of the genus lindavia, but ács et al. (2016) narrowed the concept with a description of the newly erected genus pantocsekiella kiss & ács (in ács et al. 2016: 62). both lindavia and pantocsekiella have the rimoportula situated on the valve face. however, the position of the rimoportula can be variable (houk et al. 2010, cvetkoska et al. 2014). similarly, in tertiarius, the position of the rimoportula is variable: in some species it is located in the alveolus (e.g., t. mariovensis), on the side of costa (e.g., t. pygmaeus), or in the marginal zone (t. baicalensis). in most species of lindavia, the rimoportula is located on the valve face, but in l. thienemannii (jurilj) nakov, guillory, julius, theriot & alverson, it is associated with a costa and situated near the valve face / mantle junction (levkov unpubl. data). another important feature of t. minutulus sp. nov. is the arrangement of marginal fultoportulae (on every 6th–7th costa). in all species of tertiarius so far observed with sem, marginal fultoportulae are located on every costa whereas it is on every 3rd–7th costa in species of lindavia (houk et al. 2010). in general, species of tertiarius and lindavia can be differentiated mainly by the position of the rimoportula. tertiarius minutulus sp. nov. shares the features of both genera and might be placed in any of these two genera. based on the number and position of fultoportulae, t. minutulus sp. nov. can be related to lindavia, but based on the position of rimoportula it is closer to tertiarius. since the occurrence of this fossil species is in relatively deeper geological time, molecular analyses that would uncover its phylogenetic status are yet rather impossible (see for example ancient dna limitation in wilke et al. 2016). zaova d. et al., tertiarius minutulus, a new fossil diatom from lake ohrid 11 khursevich & stachura-suchoples (2008) consider the genus tertiarius to be closely related to the genus pliocaenicus f.e.round & håkansson (round & håkansson 1992: 116) based on the position of the internal domed cribra on valve face areolae, the presence of alveolae mainly on the valve mantle and the external opening of the rimoportula lacking tubuli. the main features that distinguish these two genera are the valve undulation, absence of externally domed interfascicles, different type structure of the rimoportula (i.e., fan-shaped, sessile or raised) and different location of rimoportula – on the valve surface in pliocaenicus and on the valve mantle in tertiarius. based on these differential features, t. minutulus sp. nov. can be distinguished from p. hercynicus f.e.round & håkansson and p. undulatus f.e.round & håkansson (round & håkansson 1992). tofilovska et al. (2016: 230) discussed in detail the validity of the transfer of cyclotella minuscula to lindavia, pointing out the variability of a main synapomorphic character, the position of the rimoportula, that is quite variable in this species (see also levkov et al. 2007 and cvetkoska et al. 2014). similarly, as in cribrionella ohridana and t. minutulus sp. nov., the rimoportula in cyclotella minuscula is located at the base of costa. having in mind that the position of rimoportula should be considered as a main synapomorphic character, then these three species (cyclotella minuscula, cribrionella ohridana and teriarius minutulus sp. nov.) might represent a natural group and could be placed in the same genus. this raises another important question about the stability of characters considered as synapomorphic for any particular genus. as was pointed out, the position and location of rimoportula, the presence of valve face fultoportulae, the number of satellite pores on the fultoportulae, the presence of central lamina, etc. are variable characters within each genus. in such case, the only possible and obviously accepted solution in diatom systematic is to use a combination of characters that is unique for each genus. however, this approach can result in the future in separation and description of new cyclotelloid genera, based on already noted and differentiated morphological groups by khursevich & kociolek (2012). there is a general consensus that the number of diatom genera is underestimated, but the main task is to find appropriate and stable morphological characters that are synapomorphic for all members of the genus. there are several recent studies that resulted in synonymization of newly described genera (e.g., neofragilaria d.m.williams & f.e.round, synedrella round & n.i.maidana, lacunicula langebertalot, cavacini, tagliaventi & alfinito, etc.), since the characters used for their separation are present in other genera. however, whether this is the case with tertiarius and lindavia and whether both genera represent a single genus, remains an open question. it seems that in the older species of tertiarius, the rimoportula is located near the valve margin (within the alveolus or connected with the costa) while in t. minutulus sp. nov., it is located on the costa. in contemporary species of lindavia, the rimoportula is located on the valve face distantly from the costa. it might be hypothesized that during the evolution, the rimoportula moved from the valve margin to the valve face. a similar pattern can be noticed in cyclotella vs pantocskiella, where in cyclotella s. str. the rimoportula is located on the valve mantle while in pantocsekiella it is on the valve face. in some fossil species of cyclotella (e.g., cyclotella iris brun & héribaud-joseph) the rimoportula is located on the transition of valve the mante to the valve face (e.g., houk et al. 2010: fig.s 183: 5). however, additional sem observations especially on fossil species of lindavia and pantocsekiella are necessary to obtain a precise answer. acknowledgements the samples were obtained thanks to the icdp deep drilling campaign in lake ohrid, financed by the research project scientific collaboration on past speciation conditions in lake ohrid (scopsco), the german ministry of higher education and research, the german research foundation, the university of cologne, the british geological survey, the ingv and cnr (both italy), and the governments of the republic of north macedonia and the republic of albania. this study was supported by the alexander von humboldt foundation by providing financial support for zl for sem analyses at the hustedt diatom study centre, bremerhaven, germany. european journal of taxonomy 670: 1–14 (2020) 12 references ács e., ari e., duleba m., dressler m., genkal s.i., jakó e., rimet f., ector l. & kiss k.t. 2016. pantocsekiella, a new centric diatom genus based on morphological and genetic studies. fottea 16 (1): 56–78. 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hauffe t., holtvoeth j., just j., leng m.j., levkov z., penkman k., sadori l., skinner a., stelbrink b., vogel h., wesselingh f. & wonik t. 2016. scientific drilling projects in ancient lakes: integrating geological and biological histories. global and planetary change 143: 118–151. https://doi.org/10.1016/j.gloplacha.2016.05.005 manuscript received: 8 august 2019 manuscript accepted: 16 april 2020 published on: 19 june 2020 topic editor: christian de muizon desk editor: radka rosenbaumová printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. https://doi.org/10.5194/bg-7-3187-2010 https://doi.org/10.5194/sd-17-19-2014 https://doi.org/10.5194/bg-14-2033-2017 https://doi.org/10.1038/s41586-019-1529-0 https://doi.org/10.1016/j.gloplacha.2016.05.005 european journal of taxonomy 529: 1–26 issn 2118-9773 https://doi.org/10.5852/ejt.2019.529 www.europeanjournaloftaxonomy.eu 2019 · sorokin s.j. et al. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:7c0bab7b-f3cd-40bc-b700-19cf4ed3a761 1 a new deep-water tethya (porifera, tethyida, tethyidae) from the great australian bight and an updated tethyida phylogeny shirley j. sorokin 1,*, merrick g. ekins 2, qi yang 3 & paco cárdenas 4 1 south australian museum, north tce, adelaide, south australia; sardi aquatic sciences, 2 hamra ave, west beach, south australia. 2 queensland museum, southbank, queensland, australia; school of biological sciences, university of queensland, st lucia, queensland, australia. 3 center for marine drugs, state key laboratory of oncogene and related genes, department of pharmacy, renji hospital, school of medicine, shanghai jiao tong university, shanghai, china; centre for marine bioproducts development, college of medicine and public health, flinders university, south australia. 4 pharmacognosy, department of medicinal chemistry, bmc, uppsala university, uppsala, sweden; institut méditerranéen de biodiversité et d’ecologie marine et continentale, cnrs, aix marseille université, ird, avignon univ., station marine d’endoume, chemin de la batterie des lions, 13007 marseille, france. 1 corresponding author: shirley.sorokin@samuseum.sa.gov.au 2 email: merrick.ekins@qm.qld.gov.au 3 email: qi.yang@flinders.edu.au 4 email: paco.cardenas@ilk.uu.se 1 urn:lsid:zoobank.org:author:69444578-00e3-4614-8397-0716950cd3ca 2 urn:lsid:zoobank.org:author:103d616e-376b-4ab0-a314-999dcc68e8aa 3 urn:lsid:zoobank.org:author:5f8438eb-c634-49d0-b2ad-1a6d933b035d 4 urn:lsid:zoobank.org:author:9063c523-49fc-427e-9e84-dbc31c5db6d3 abstract. a new species of tethya lamarck, 1815 is described from a depth of 1000 m on the continental slope of the great australian bight (gab), southern australia. the gab slope was explored as part of systematic benthic surveys to understand unexplored communities in the light of current oil and gas exploration activity in the area. tethya irisae sp. nov. was present at 1000 m in six of eight longitudinal depth surveys. three molecular markers were obtained: coi, 28s (d3–d5) and its1-5.8s-its2. coi and 28s phylogenetic analyses show that the new species fits clearly within the genus tethya. this is the 28th species of tethya reported from australia; it is unusual in that it has a stalk. the presence of a stalk as a morphological character to split genera in this family is questioned. the description of this new species is an opportunity to revisit the molecular phylogeny of the tethyida morrow & cárdenas, 2015 using comprehensive datasets of coi and 28s markers. as in previous analyses, four tethya clades were retrieved; we discuss the possibility of using external colour to support some of these clades. despite unclear phylogenetic relationships amongst tethyidae gray, 1848 from australia, our results suggest that tethyid genera tethytimea laubenfels, 1936, tectitethya sarà, 1994, laxotethya sarà & sarà, 2002, stellitethya sarà, 1994, and xenospongia gray, 1858 derive from species of tethya. we show that asters https://doi.org/10.5852/ejt.2019.529 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:7c0bab7b-f3cd-40bc-b700-19cf4ed3a761 mailto:shirley.sorokin@samuseum.sa.gov.au mailto:merrick.ekins@qm.qld.gov.au mailto:qi.yang@flinders.edu.au mailto:paco.cardenas@ilk.uu.se http://zoobank.org/urn:lsid:zoobank.org:author:69444578-00e3-4614-8397-0716950cd3ca http://zoobank.org/urn:lsid:zoobank.org:author:103d616e-376b-4ab0-a314-999dcc68e8aa http://zoobank.org/urn:lsid:zoobank.org:author:5f8438eb-c634-49d0-b2ad-1a6d933b035d http://zoobank.org/urn:lsid:zoobank.org:author:9063c523-49fc-427e-9e84-dbc31c5db6d3 european journal of taxonomy 529: 1–26 (2019) 2 have been secondarily lost at least twice in the hemiasterellidae lendenfeld, 1889: in liosina thiele, 1899 and a potential new genus from northern australia. we formally propose the reallocation of liosina from dictyonellidae van soest, diaz & pomponi, 1990 to hemiasterellidae lendenfeld, 1889. keywords. porifera, dictyonellidae, hemiasterellidae, liosina, marine benthos. sorokin s.j., ekins m.g., yang q. & cárdenas p. 2019. a new deep-water tethya (porifera, tethyida, tethyidae) from the great australian bight and an updated tethyida phylogeny. european journal of taxonomy 529: 1–26. https://doi.org/10.5852/ejt.2019.529 introduction the family tethyidae gray, 1848, reclassified from the order hadromerida topsent, 1894 (order abandoned) to order tethyida morrow & cárdenas, 2015, contains 14 genera of which the genus tethya lamarck, 1815 is by far the most speciose with 93 accepted species (van soest et al. 2019). tethyidae are defined by ‘stylote megascleres mainly stronglyoxeas, generally in radiate tracts and two categories of euasterose microcleres, micrasters and megasters, sometimes rhadbs’ (sarà 2002). occurrence and length of a stalk are included as key morphological characteristics in distinguishing some genera of tethyidae (sarà 1994, 2002; sarà & burlando 1994); although the development of the stalk in the family has been noted as being an adaptation to deep-water, soft sediment habitat (sarà & burlando 1994). the taxonomy of the tethyidae was rearranged by sarà (1994) into eight genera, with the new genera burtonitethya sarà, 1994 and tethycometes sarà, 1994 joining halicometes topsent, 1898, as stalked genera in the family. cladistic analysis by sarà & burlando (1994) placed tethyidae into three clades (a) the genus tethya, (b) genera with stalks and (c) massive and encrusting genera. the australian faunal directory (afd) (hooper 2012) lists six genera of tethyidae in australia (anthotethya sarà & sarà, 2002; laxotethya sarà & sarà, 2002; oxytethya sarà & sarà, 2002; stellitethya sarà, 1994; tethya lamarck, 1815; xenospongia gray, 1858). a seventh genus tethyastra sarà, 2002 (tethyastra oxyaster (burton, 1934), ‘accepted’ (van soest et al. 2019)) is listed in the codes for australian aquatic biota (rees et al. 1999 onwards); in the afd this is listed as tethya oxyaster. the slope of the great australian bight (gab) was explored in 2010 as part of a preliminary exploration of deep-water communities (currie & sorokin 2011) and again in 2015 as part of systematic benthic surveys to understand unexplored communities in the light of current oil and gas exploration activity in the area (mnf 2015; williams et al. 2018). the surveys resulted in the discovery of several new benthic species including sponges. multiple specimens of a small stalked tethyid were found at 1000 m; here, we describe this new species, using morphological characters and molecular markers (coi, 28s (d3–d5) and its1-5.8s-its2) to consider how it fits into the family. the description of this new species is an opportunity to revisit the molecular phylogeny of this group. heim et al. (2007) and heim & nickel (2010) produced the first phylogenetic analyses of tethya, combining coi and morphology. since then, although large demospongiae trees were produced that led to the creation of the order tethyida by morrow & cárdenas (2015), no phylogenetic studies truly focused on the tethya, tethyidae or tethyida. we therefore felt it was time to present an updated molecular phylogeny for coi and ran the first comprehensive 28s phylogenetic analyses focusing on the tethyida. material and methods field collection tethya irisae sp. nov. was first collected, as only one specimen, in the gab in 2010 (currie & sorokin 2011). more specimens were collected in november and december 2015 as part of systematic epibenthic https://doi.org/10.5852/ejt.2019.529 sorokin s.j. et al., a new deep-water tethya (porifera: demospongiae) 3 surveys of the central gab slope. stations were sampled by beam trawl along five longitudinal transects and over depths ranges from 200 m to 5000 m (fig. 1). benthic specimens were collected under australian commonwealth area permit no. aucom2015-284 and commonwealth marine reserve permit no. cmr-15-000344. tethya irisae sp. nov. was found at six out of eight of the 1000 m depth stations. specimens were photographed on board and fixed in 70% ethanol. specimens for molecular analysis were fixed in ethanol (> 95%). most specimens (including the holotype) are lodged at the south australian museum (accession prefix sama), adelaide, south australia; two specimens were deposited at the queensland museum (accession prefix qm), brisbane, australia; four specimens from lot sama s2039 and a thick section are deposited at the museum of evolution, uppsala, sweden (upszty 178608). collection information of specimens examined in this study is archived in the open access pangaea data repository (https://doi.pangaea.de/10.1594/pangaea.894720). light microscope preparation to examine the skeleton, thick sections in resin were prepared from a specimen from collection lot sama s2039, following the method described by boury-esnault et al. (2002). thin sections were also fig. 1. sites where tethya irisae sp. nov. was collected in the great australian bight, including sites that were sampled where the sponge was not found. all specimens of tethya irisae sp. nov. were collected along the 1000 m contour. light shaded polygons show the australian commonwealth marine reserves. the darker polygon strip is the gab marine park benthic protection zone. the 200 m contour is the edge of the continental shelf. abbreviations: gabdmp = great australian bight deepwater marine program (mnf 2015); gabrp = great australian bight research project (williams et al. 2018). cruise ss2010_t02 (currie & sorokin 2011). https://doi.pangaea.de/10.1594/pangaea.894720 european journal of taxonomy 529: 1–26 (2019) 4 made, with the following protocol: sections were cut perpendicular to the surface with a sharp blade and laid onto a slide, covered with a weighted coverslip and dried on a hot plate. sections were covered with mounting media (durcupan™) and dried overnight in an oven at 50oc. spicule slides were prepared by dissolving a small amount (~ 2 mm2) of sponge tissue in 3.9% sodium hypochlorite. the resultant spicules were rinsed with distilled water three times and with 95% ethanol twice before mounting on a microscope slide with dpx™ mountant. sem preparation scanning electron microscope (sem) tissue preparations were made by dissolving the tissue in 12.5% sodium hypochlorite to remove the soft tissue. they were then rinsed twice in distilled water, rinsed twice in 70% ethanol and then finally twice in 98% ethanol and then air dried. sem preparations were sputter coated in gold to improve resolution. the scanning electron micrograph photos were taken using a hitachi tm-1000 sem and plates assembled in adobe photoshop. morphometric measurements of the spicules were done using the same hitachi tm-1000 sem. spicule terminology spicule terminology follows that suggested by bergquist & kelly-borges (1991) for the genus tethya. molecular studies whole genomic dna was extracted from sponge tissue frozen at -80°c. a conventional hexadecyltrimethylammonium bromide (ctab)-based protocol (taylor et al. 2004) was used for isolating dna. briefly, the sponge tissues were ground under liquid nitrogen. the ctab extraction buffer was applied to lyse tissues and then combined with polyvinylpyrrolidone (pvp) and β-mercaptoethanol to help remove phenolic compounds and tannins in the extract. to separate the proteins and polysaccharides from nucleic acids, phenol: chloroform: isoamyl alcohol (25:24:1) was utilised before dna was precipitated with chilled isopropanol. the mitochondrial cytochrome c oxidase subunit 1 (coi) folmer fragment was amplified using the universal primers lco1490 and hco2198 (folmer et al. 1994). the d3–d5 region of 28s rrna gene was amplified by primers nl4f and nl4r (nichols 2005). to amplify its, we used primers originally designed for a unicellular eukaryote, itsra2 (5′-gtc cct gcc ctt tgt aca ca-3′) and its2.2 (5′-cct ggt tag ttt ctt ttc ctc cgc-3′) (adlard & lester 1995), to amplify a 753 bp long sequence including its15.8s-its2 and small fragments of the surrounding 28s and 18s. the thermocycler was programmed following yang et al. (2017). sequences were assembled and blasted using geneious® ver. 8.1 (created by biomatters, http://www.geneious.com). all tethyidae coi and 28s sequences from genbank were collected and aligned with our new sequences in aliview 1.18 (larsson 2014). no its alignment was made since only one other tethyidae its was found on genbank. the full coi and 28s from tethya wilhelma sarà, sarà, nickel & brümmer, 2001 were assembled from the whole genome, https://bitbucket.org/molpalmuc/tethya_wilhelma-genome/src (courtesy of w.r. francis). we included timeidae topsent, 1928 and hemiasterellidae lendenfeld, 1889 (adreus gray, 1867 and axos gray, 1867, hemiasterella carter, 1879, and liosina thiele, 1899) sequences that seemed to group close to the tethyidae in our preliminary analyses and previous studies (erpenbeck et al. 2012; morrow et al. 2012; redmond et al. 2013; thacker et al. 2013). raspailia australiensis ridley, 1884 and ceratopsion axiferum (hentschel, 1912) 28s sequences were included as possible species of adreus, as suggested by morrow et al. (2019). we discarded the 28s sequence of timea lowchoyi hooper, 1986 (ay561871) and hemiasterella sp. wamz12383 (ay561947), both from nichols (2005) that seemed to be mis-identifications: blast results indicated, respectively, that they were close to cymbaxinella gazave, carteron, chenuil, richelle-maurer, boury-esnault & borchiellini, 2010, and a 93% match with hymeniacidon heliophila (wilson, 1911). hemiasterella sp. 1 (ay561901, qmg315767, otu qm2839) http://www.geneious.com https://bitbucket.org/molpalmuc/tethya_wilhelma-genome/src sorokin s.j. et al., a new deep-water tethya (porifera: demospongiae) 5 from nichols (2005) was re-identified as adreus sp. on spongemaps (http://www.spongemaps.org). hemiasterella sp. (ay626310, qm g304645, otu qm0694) from kober & nichols (2007) was reidentified as a tethyid by morrow et al. (2012). hemiasterella sp. ucmpwc1021 (ay56197) from nichols (2005) was re-examined and re-identified as a jaspis sp. (p. cárdenas, unpublished data), we included this sequence in our outgroup. finally, no trustworthy sequences of hemiasterella were left to include in the alignment. we also included 28s (c1–d2) sequences from c. chombard’s phd thesis (chombard 1998), never published or submitted to genbank. these were copied ‘by hand’ from the ‘annexe b2’ from her thesis, and included the following species: tethya sp., tethya taboga (laubenfels, 1936), tectitethya crypta (laubenfels, 1949), tethya aurantium (pallas, 1766) and timea sp. the three first samples were collected by n. boury-esnault in panama in july 1995. preliminary 28s analyses showed that the chombard sequence of t. aurantium (from chausey islands, normandy, france) had 1 bp difference to t. citrina sarà & melone, 1965 (hq379237) from wales, so we considered that the chombard specimen had been mis-identified (both species can be difficult to discriminate but are clearly different genetically) and we thus renamed it t. cf. citrina. we discarded the 28s (d1–d2) sequence of t. aurantium (ay552024) from borchiellini (2004) because it was 100% identical to that of t. taboga 28s (c1–d2) from chombard (1998) and we suspected a mix-up in the sequence submitted. coi alignments were trimmed to contain only the folmer fragment (658 bp). the 28s alignment included essentially sequences from the c1–d2, d3–d5 and/or d6–d8 regions or from the full 28s (3376 bp). when we had different 28s regions for the same species, we merged those sequences into a single one using the ‘merge’ option in aliview; we did this for timea cf. centrifera, tethya citrina, tethya hibernica heim, nickel, picton & brümmer, 2007, laxotethya dampierensis sarà & sarà, 2002, adreus micraster (burton, 1956), adreus fascicularis (bowerbank, 1866) and adreus sp. alignment was done using mafft (katoh et al. 2002) implemented in aliview, then refined by eye; ambiguous regions were kept. we included in both our datasets sequences of placospongiidae that are phylogenetically close to the tethyida, maybe even their sister-group (morrow et al. 2013; thacker et al. 2013). the final coi alignment contained 39 sequences, including five outgroup sequences from paratimea hallmann, 1917 (stelligeridae lendenfeld, 1898), jaspis gray, 1867 (ancorinidae schmidt, 1870) cliona grant, 1826 (clionaidae d'orbigny, 1851) and placospongia gray, 1867 (placospongiidae gray, 1867). the 28s alignment contained 45 sequences, including three outgroup sequences from placospongia and trachycladus (trachycladidae hallmann, 1917). analyses were conducted with the cipres science gateway (http://www.phylo.org) (miller et al. 2010): raxml 8.2.10 (stamatakis 2014) for maximum likelihood (ml) and mrbayes v. 3.2.6 (ronquist et al. 2012) for bayesian analyses. for raxml, 1000 bootstrap iterations were run; bootstrap bayesian analyses consisted of two runs of four chains, each for 5 000 000 generations and sampled every 1000th tree after a 25% burn-in. abbreviations bp = base pairs diam. = diameter h = height l = length w = width collection acronyms abtc = australian biological tissue collection, south australian museum, adelaide, australia am = australian museum, sydney, australia bmnh = the natural history museum (formerly british museum of natural history), london, uk qm = queensland museum, brisbane, australia http://www.spongemaps.org http://www.phylo.org european journal of taxonomy 529: 1–26 (2019) 6 sama = south australian museum, adelaide, australia upszty = museum of evolution, uppsala, sweden results systematics class demospongiae sollas, 1885 order tethyida morrow & cárdenas, 2015 family tethyidae gray, 1848 genus tethya lamarck, 1815 tethya irisae sp. nov. urn:lsid:zoobank.org:act:26151082-02ac-41f5-9e22-15ef212dbdc3 figs 1–3, 4a etymology named after the golden-winged greek goddess iris, grandchild of tethys, who could reach all parts of the cosmos including the deep sea; and in memory of marine naturalist iris sorokin. material examined holotype australia • size 16.6 mm total height (body 11.9 mm (h) × 11.7 mm (w), stalk 4.7 mm (l) × 1.77 mm diam., raised apical osculum); great australian bight (gab); 34.822° s, 132.69° e; 1006 m depth; great australian bight research project (gabrp) leg.; epibenthic sled; sama s3387. paratypes australia • 4 specs; same collection data as for holotype; sama s2913, sama s3388, qm g305000, qm g305001 • 1 spec.; great australian bight; 33.928° s, 131.06° e; 1027 m depth; gabrp leg.; epibenthic sled; upszty 178608. additional material at south australian museum (sighted only) australia • 1 spec.; great australian bight; 33.928° s, 131.06° e; 1027 m depth; gabrp leg.; epibenthic sled; sama s2039 • 1 spec.; great australian bight; 35.152° s, 134.109° e; 1021 m depth; gabrp leg.; epibenthic sled; sama s2371 • 1 spec.; great australian bight; 33.718° s, 130.66° e; 1005 m depth; gabrp leg.; epibenthic sled; sama s2482 • 1 spec.; great australian bight; 34.629° s, 132.35° e; 1021 m depth; great australian bight deepwater marine program (gabdmp) leg.; epibenthic sled; sama s2095 • 1 spec.; great australian bight; 34.705° s, 132.53° e; 987 m depth; gabdmp leg.; epibenthic sled; sama s2096 • 1 spec.; great australian bight; 33.802° s, 130.70° e; 1000 m depth; d. currie leg.; epibenthic sled; sama s1461. comparative material india • 1 section in slide, holotype of burtonitethya gemmiformis sarà, 1994; andaman islands; depth unknown; bmnh 1957.7.15.1. australia – new south wales • 1 spec., syntype and slides of tethya fissurata lendenfeld, 1888; port jackson; “33°51′ s, 151°16′ e [33.85° s, 151.27° e]; depth unknown; am g.9069 (syntype), z6053, z6893 (slides). new zealand • 1 spec., holotype (specimen and slides) of tethya bullae bergquist & kelly-borges, 1991; alderman island; “36°58´ s, 176°05´ e [36.97° s, 176.08° e]; 100 m depth; am z5074. http://zoobank.org/urn:lsid:zoobank.org:act:26151082-02ac-41f5-9e22-15ef212dbdc3 sorokin s.j. et al., a new deep-water tethya (porifera: demospongiae) 7 dna barcoding coi (mh518072), 28s (d3–d5) (mh511148), its1-5.8s-its2 (mh511149). all sequences came from the same individual from lot sama s2913, although this is a different individual than the type specimens. a tissue sample from this voucher is deposited at the australian biological tissue collection at the south australian museum, adelaide (abtc145318). description a small, spherical to oval, stalked, sponge (fig. 2). the sponge body is 11–14 mm diam., with the stalk approximately the same length as the diameter of the sponge. the surface is covered in polygonal platelike tubercules (2–3 mm diam.) separated by grooves (0.5 mm wide, 0.25–0.5 mm deep). the sponge is firm to hard and spiculose. grey/white in life and in ethanol. there is a single raised apical osculum. no sign of any budding. skeleton. a stalk of dense megascleres supports the sponge. there is a ‘nucleus’ where the stalk meets the centre of the sponge body, and although the stalk may divide and/or flatten and thicken externally it emanates from the same point at the base of the sponge. from the nucleus dense bundles (0.3–0.7 mm in fig. 2. a. freshly collected specimens (lot sama s2096) of tethya irisae sp. nov. b. paratype (qm g305000) showing single apical oscule (arrow), and tessellated plate-like polygonal tubercules. c–d. holotype (sama s3387), entire specimen and sem showing surface tubercules with emerging megascleres. e. section of upszty 178608, showing the well-developed cortex and cortical canals around the tubercules. european journal of taxonomy 529: 1–26 (2019) 8 fig. 3. tethya irisae sp. nov. spicules. a–b. straight style/strongyloxeas. c. subtylostyle. d. long-rayed oxyspheraster. e. short-rayed oxyspheraster with small acanthooxyspheraster. f. acanthooxyspheraster. sorokin s.j. et al., a new deep-water tethya (porifera: demospongiae) 9 diameter) of megascleres radiate through the choanosome to the surface tubercles; the bundles slightly fan out in the tubercles. the cortex is a dense layer of micrasters and oxyspherasters interspersed with megascleres emerging through the tubercules, making the surface microscopically hispid (fig. 2d– e). the cortex is well developed and follows the contours of the tubercules, 1–1.7 mm thick. large cortical canals are visible between tubercles (fig. 2e). a thin fibrous layer is below the cortex, it has micrasters in a much lower density. large oxyspherasters are especially found at the base of the cortex. the megascleres of the stalk are covered in a layer of micrasters and regularly interspersed with shortrayed oxyspherasters. the choanosome is rich in sediment-like particles; there are some micrasters and rare oxyspherasters. foraminifera (globigerina d'orbigny, 1826) and radiolaria are common in the cortical canals and the choanosome. spicules. megascleres are straight style/strongyloxeas (size range 900–3060 × 17–52 µm) (table 1, fig. 3a–b) the proximal end is smooth and rounded, the distal end is tapered (not stepped) and either rounded or pointed. there are auxiliary thinner styles to subtylostyles in the medulla between the main styles (260–900 × 7–22 µm) (fig. 3c). megaster microscleres are two types of oxyspherasters: longrayed oxyspherasters (120–185 µm) (fig. 3d) have ~15 rays that can be bent towards the oxeote tips (ray profile is conical); short-rayed oxyspherasters (53–154 µm) (fig. 3e) have a larger centrum ~17 rays with a conical profile and oxeote tips. micraster microscleres are acanthooxyspherasters (12–20 µm) (fig. 3f), with a centrum and spined tips, and lightly spined on the rays. ecology and distribution found on the continental slope in the great australian bight at a depth of 1000 m, in soft sediment (clay/ silt). remarks the morphology as well as molecular markers confirm that our new sponge is a tethya. table 2 shows morphological comparisons between other species of tethya from australia and new zealand. the external appearance of tethya irisae sp. nov. is similar to t. fissurata from port jackson, new south wales, australia, which is spherical with polygonal tubercules and has a stalk. however, t. fissurata differs from t. irisae sp. nov. in body size (~ 4 cm diam.), tubercule shape, and number of oscula (2–4). tethya fissurata has megascleres with stepped ends unlike t. irisae sp. nov., which are smooth and t. fissurata lacks the short-rayed oxyspherasters seen in t. irisae sp. nov. although we do not know the exact depth at which t. fissurata was collected, port jackson (viz. sydney harbour) is not deeper than 45 m (johnston et al. 2015), so this is presumably a shallow species. tethya bullae is a deep-water (100 m) sponge that is of comparable size to t. irisae sp. nov., although it has prominent raised tubercules rather than the flat plate-like tubercules of t. irisae sp. nov. (fig. 4). the holotype from the australian museum does not include a stalk but the description and photograph in bergquist & kelly-borges (1991) shows “basal flattened branched rooting processes”. the long-rayed oxyspherasters of t. irisae sp. nov. are similar to those of t. bullae. the short-rayed oxyspherasters in t. irisae sp. nov. do not fork as those of t. bullae. tethya irisae sp. nov. has lightly spined acanthooxyspherasters compared to the completely spined acanthooxyspherasters of t. bullae. in addition to t. fissurata and t. bullae, other tethya with rooting processes/stolons are shown in table 2 (descriptions in bold text). it is difficult to tell how similar the rooting processes are to each other but these species differ in spicule forms and dimensions from t. irisae sp. nov. for example: species with megascleres < 2000 µm (t. acuta sarà & sarà, 2004, t. bergquistae hooper in hooper & wiedenmayer, 1994, t. burtoni sarà & sarà, 2004, t. dendyi sarà & sarà, 2004, t. robusta (bowerbank, 1873), t. seychellensis (wright, 1881), t. stolonifera bergquist & kelly-borges, 1991); species with megasters not of a ‘spheraster’ form (t. amplexa bergquist & kelly-borges, 1991, and t. fastigata bergquist & kelly-borges, 1991); species with very different micrasters (t. ingalli bowerbank, 1858, t. flexuosa sarà & sarà, 2004 and t. monstrosa (burton, 1924)). european journal of taxonomy 529: 1–26 (2019) 10 in addition, t. irisae sp. nov. is collected at the start of the bathyal zone (~1000 m). the deepest of the tethya is t. compactus bergquist, 1961 (402 m), which has very different external morphology. it occurred to us that when using the key to genera of tethyidae (sarà 2002), tethya irisae sp. nov. appears closest to the monospecific genus burtonitethya, a tethyid with a stalk of equal length to the diameter of the sponge. the type of burtonitethya (b. gemmiformis), was collected from the andaman sea at an unknown depth (sarà 1994). burtonitethya gemmiformis was originally assigned to tethya (labelled as tethya gemmiformis burton & rao, 1957 on the nhm microscope slide) but was re-assigned to a new genus burtonitethya by sarà (1994) on account of the stalk, the conspicuous nucleus with strongyles, the reduced lacunar cortex, the specialised surface tubercules and the giant oxyaster megasters. our new species clearly differs from this species in having different microscleres and does not have the giant megasters present in b. gemmiformis. as there is no specimen of the type species of burtonitethya and thus no potential to sequence the sponge, we cannot test if burtonitethya is a junior synonym of tethya. as seen above, the genus tethya shows many different modes of attachment including basal stolons, basal roots, curved peduncles, flattened rooting processes as well as attachment discs and narrow skirts of tissue. our results suggest that the stalk may not be a good genus-defining character within the family. heim et al. (2007) in their analysis of tethya species, for which they used morphological characters and molecular markers, suggest that characters pertaining to ecological influences may have developed several times. similarly, we suggest that some of the external morphological characters used to separate genera of tethyidae are homoplasious, probably appearing several times in different clades of tethya and we question whether they should be grouped as definitive characters in morphological identifications. in the same way the genus amphitethya lendenfeld, 1907 (family tetillidae sollas, 1886) was created based on its stalk, but phylogenies show it is a cinachyrella wilson, 1925 (szitenberg et al. 2013; schuster et al. 2017). table 1. spicule dimensions of tethya irisae sp. nov. holotype (sama s3387) and paratypes (qm g305000, qm g305001). spicule type specimen # n size range (µm) mean value underlined strongyloxeas (l × w) s3387 34 897–1361–3060 × 17–27–52 g305000 26 1110–2310–3130 × 19–42–72 g305001 36 1270–2495–3160 × 20–39–64 styles (l × w) s3387 13 262–675–1090 × 7–16–23 g305000 16 830–1067–1730 × 11–20–33 g305001 8 1250–1525–1970 × 19–27–34 short-rayed oxyspherasters (diam.) s3387 75 53–81–154 g305000 41 44–99–177 g305001 59 49–98–133 long-rayed oxyspherasters (diam.) s3387 21 120–153–185 g305000 17 154–196–253 g305001 2 131–134–136 acanthooxyspherasters (diam.) s3387 55 12–15–20 g305000 43 13–15–19 g305001 32 13–16–19 sorokin s.j. et al., a new deep-water tethya (porifera: demospongiae) 11 fig. 4. comparative sizes and external morphology of species of tethya lamarck, 1815. a. tethya irisae sp. nov., holotype (sama s3387). b. tethya bullae bergquist & kelly-borges, 1991, part of the holotype (am z5074). c. tethya fissurata lendenfeld, 1888, syntype (am g.9069) note: the original photo of t. bullae (bergquist & kelly-borges 1991) shows rooting processes. european journal of taxonomy 529: 1–26 (2019) 12 sp ec ie s e xt er na l a pp ea ra nc e si gn ifi ca nt a tta ch m en t in b ol d b od y di am . c ol ou r sk el et al a rr an ge m en t m eg as cl er e tr ac ts (d ia m . in µ m ) & in te rs tit ia l br us he s sp ic ul es m s = m eg as cl er es , m e = m eg as te rs , m i = m ic ra st er s (s iz e in µ m ) ty pe lo ca lit y d ep th (m ) t. ir is ae s p. n ov . sp he ri ca l; pl at elik e po ly go na l t ub er cu le s se pa ra te d by g ro ov es ; ap ic al o sc ul um ; h ar d. st al k, s am e le ng th a s bo dy d ia m . 11 m m g re y/ w hi te tr ac ts o f m eg as cl er e bu nd le s (3 00 –7 00 ) a re un if or m fr om c ho an os om e to c or te x an d fa n sl ig ht ly in tu be rc ul es . m s: s tr on gy lo xe as (t o 30 00 × 7 0) , s ty le s (t o 19 70 × 3 4) . m e: lo ng & s ho rt -r ay ed o xy sp he ra st er s (4 4– 17 7) m i: a ca nt ho ox ys ph er as te rs (1 2– 20 ) g re at a us tr al ia n b ig ht 10 00 t. a cu ta s ar à & s ar à, 20 04 1 sp he ri ca l w ith s m al l, di st in ct tu be rc ul es ; fir m . b ud s an d rh iz oi d pr ot ub er an ce s. 13 m m u nk no w n* t hi ck tr ac ts o f m eg as cl er e bu nd le s (2 50 ) f an ni ng o ut in th e co rt ex . m s: s tr on gy lo xe as , a ni so st ro ng yl es , s ty le s (t o 16 00 × 2 0) m e: o xy sp he ra st er s (1 5– 50 ) m i: o xy as te rs (8 –2 0) pt p hi lli p h ea ds , v ic to ri a, a us tr al ia u nk no w n* a t. a m pl ex a b er gq ui st & k el ly -b or ge s, 1 99 12 sp he ri ca l t o su bs ph er ic al ; su rf ac e en cr us te d w ith an im al s. b ro ad fl at tu be rc ul es . a pi ca l o sc ul e. l on g cu rv ed fl ex ib le pe du nc le . 30 –7 0 m m y el lo w r ob us t b un dl es (4 40 –8 00 ) di ve rg in g in to b ra nc he s in c or te x. b ru sh es o f in te rs tit ia l m eg as cl er es . m s an is os tr on gy lo xe as (t o 23 00 × 3 2) m e: o xy as te rs (1 6– 35 ) m i: v ar io us , i nc l. ac an th ot yl as te rs ac an th oc hi as te rs , a ca nt ho ox ys ph er as te rs (1 0– 20 ) m im iw ha ng at a, n ew z ea la nd 7 t. b er gq ui st ae h oo pe r in h oo pe r & w ie de nm ay er 1 99 41 ,2 sp he ri ca l t o ov at e; su rf ac e m am m ill at e to te ss el la te d w ith b ud s ar is in g on fi la m en ts ; fi rm co m pr es si bl e. o sc ul es in gr ou ps o f 3 –4 ). r oo tlik e pr oc es se s. 20 –2 5 m m r os e pi nk w id el y se pa ra te d th in fl ex uo us tr ac ts o f m eg as cl er e bu nd le s (1 71 – 61 2) ra di at in g in to te rt ia ry br an ch es a t s ur fa ce . in te rs tit ia l m eg as cl er e br us he s. m s: a ni so st ro ng yl ox ea s (t o 16 17 × 1 8) m e: o xy sp he ra st er s (1 8– 52 ), ox ya st er s (2 2– 31 ) m i: a ca nt ho ty la st er s/ ac an th ost ro ng yl as te rs (1 0– 17 ), m ic ro -o xy as te rs (7 –1 0) . w hi tia ng a, n ew z ea la nd 27 t. b ul la e b er gq ui st & k el ly -b or ge s, 1 99 12 sp he ri ca l t o ov at e; la rg e bl un t w el l s ep ar at ed tu be rc ul es . b as al fl at te ne d br an ch ed r oo ti ng pr oc es se s. 10 –1 4 m m y el lo w g re y tr ac ts o f m eg as cl er e bu nd le s (5 00 ) u ni fo rm fr om c ho an os om e to co rt ex , w he re th ey ra di at e sl ig ht ly to s up po rt th e ra is ed tu be rc ul es . m s: a ni so st ro ng yl ox ea s (t o 22 25 × 3 0) m e: lo ng a nd s ho rt -r ay ed o xy sp he ra st er s m i: c om pl et el y m ic ro sp in ed ac an th oo xy sp he ra st er s & ac an th os tr on gy la st er s (1 1– 15 ), m ic ro ox ys ph er as te rs (9 –1 0) a ld er m an is la nd s, n ew z ea la nd 10 0 t. b ur to ni s ar à & s ar à, 20 04 1 sp he ri ca l t o he m is ph er ic al ; tu be rc ul es c on ul at ed , fl at or in co ns pi cu ou s. s po ng es cl um p to ge th er . s m al l m ar gi na l s to lo ns . 10 –1 3 m m pa le o ra ng e br ow n tr ac ts o f m eg as cl er e bu nd le s (5 00 –7 50 ) un br an ch ed . b ru sh es of a ux ili ar y in te rs tit ia l m eg as cl er es . m s: s tr on gy lo xe as /a ni so st ro ng yl es to (1 80 0 × 40 ) + in te rs tit ia l s ty le s m e: o xy -/ sp he ra st er s (4 0– 80 ) m i: ty la st er s (8 –1 4) , c hi as te rs o r t yl ot e ox ya st er s (1 2– 15 ) ta ur an ga , n ew z ea la nd 1 ta bl e 2 (c on tin ue d on fi ve n ex t p ag es ). c om pa ri so n of m or ph ol og y be tw ee n sp ec ie s of t et hy a l am ar ck , 1 81 5 fr om a us tr al ia a nd n ew z ea la nd . sorokin s.j. et al., a new deep-water tethya (porifera: demospongiae) 13 sp ec ie s e xt er na l a pp ea ra nc e b od y di am . c ol ou r sk el et al a rr an ge m en t sp ic ul es ty pe lo ca lit y d ep th (m ) t. c om pa ct a b er gq ui st , 19 61 3 r ho m bo id , w ith la te ra l f an lik e pr oj ec tio ns . s to ny . 8 m m m id b ro w n (i n et ha no l) tr ac ts (3 56 –7 22 ) ( in ec to so m e) . e nd os om e w ith ra di at e st ru ct ur e. m s: s tr on gy lo xe as (t o 15 74 × 2 1) m e: s ph er as te rs (2 5– 43 ) m i: ty la st er s (7 –1 2) c ha th am r is e, n ew z ea la nd 40 2 t. c om m un is b er gq ui st & k el ly -b or ge s, 1 99 12 sp he ri ca l t o su bs ph er ic al ; gr ow in g in c lu m ps . 7– 12 m m d ul l g re y pi nk (i n et ha no l) fl ex uo us tr ac ts o f m eg as cl er e bu nd le s (1 25 –3 00 ) h av e el ab or at e te rt ia ry b ra nc hi ng . m s: a ni so st ro ng yl ox ea s (t o 12 25 × 1 2) m e: o xy sp he ra st er s (3 8– 55 ), ox ya st er s (7 –1 2) m i: in cl ud es p ol yr ha bd s c ap e b an ks , n ew so ut h w al es , a us tr al ia in te rt id al t. d en dy i s ar à & s ar à, 20 04 1 b od y co ve re d in s m al l, fla tte ne d tu be rc ul es . r hi zo id -l ik e st ol on s. 20 m m pa le b uf f ( in et ha no l) tr ac ts o f m eg as cl er e bu nd le s (2 50 –5 00 ) b ra nc h in c ho an os om e an d fo rm fa ns in c or te x. m s: s tr on gy lo xe as a nd s ty le s (t o 98 0 × 15 ) m e: s ph er as te rs 2 0– 30 m i: ty la st er s/ ch ia st er s/ ox ya st er s (1 0– 16 ) pt p hi lli p h ea ds , v ic to ri a, a us tr al ia u nk no w na t. e xp an sa s ar à & sa rà , 2 00 41 ir re gu la r s ha pe , v er y ha rd , ir re gu la r l ab yr in th in e tu be rc ul es . 20 –3 0 m m u nk no w n tr ac ts o f m eg as cl er e bu nd le s (2 50 –5 00 ) f or m se co nd ar y br an ch in g in c or te x w ith fa ns a t su rf ac e. m s: s tr on gy lo xe as a nd a ni so st ro ng yl es (1 40 0 × 28 ) m e: s ph er as te rs /o xy as te rs (3 0– 50 ) m i: ty la st er s/ ch ia st er s/ ox ya st er s (7 –1 3) a hi pa ra b ay , n ew z ea la nd u nk no w n t. fa st ig at a b er gq ui st & k el ly -b or ge s, 1 99 12 o vo id ; 1 –3 la rg e ra is ed ap ic al o sc ul es ; s ur fa ce co nu lo se . s ho rt , t hi ck st ol on s. 40 –6 0 m m b ri gh t or an ge fi ne , w id el y se pa ra te d tr ac ts o f m eg as cl er e bu nd le s (2 75 –5 39 ) r ad ia te sl ig ht ly to w ar ds s ur fa ce . m s: a ni so st ro ng yl ox ea s (t o 24 25 × 3 5) m e: o xy as te rs (1 8– 39 ) m i: v ar io us in cl . a ca nt ho ty la st er s/ ch ia st er s (9 –2 0) po or k ni gh ts is l. n ew z ea la nd 25 –4 0 t. fi ss ur at a l en de nf el d, 1 88 84 ,5 sp he ri ca l; tu be rc ul es se pa ra te d by d ee p gr oo ve s. t hi ck s ta lk w hi ch d iv id es in to r oo ts . 40 m m b ei ge (i n et ha no l) tr ac ts o f m eg as cl er e bu nd le s br an ch in th e tu be rc ul es , s up po rt in g th e ed ge o f t he tu be rc ul es a nd sp ic ul es e xt en d th ro ug h th e su rf ac e. m s: s tr on gy lo xe as (t o 40 00 × 8 0) m e: s ho rt -r ay ed s ph er as te rs (1 60 ), lo ng ra ye d sp he ra st er s (2 40 ) m i: o xy as te rs (5 5) , t yl as te rs (1 9) pt j ac ks on , n ew so ut h w al es , a us tr al ia u nk no w nb t. fl ex uo sa s ar à & sa rà , 2 00 41 ir re gu la r s ph er ic al s ha pe . tu be rc ul es ro un de d an d sl ig ht ly c on ic al a nd c ar ry th in fi la m en ts . e la st ic co ns is te nc y. b as al r hi zo id st ol on s. 18 –3 5 m m b ro w ni sh w hi te tr ac ts o f m eg as cl er e bu nd le s (2 50 –7 50 ) b ra nc h in to s ec on da ry a nd te rt ia ry br an ch es in c or te x. m s st ro ng yl ox ea s an d an is os tr on gy le s (t o 26 00 × 5 0) m e: s ph er as te rs & o xy sp he ra st er s (3 0– 80 ) m i: ty la st er s (1 0– 15 ), ox ya st er s (1 0– 50 ) n w a us tr al ia 30 –6 2 t. g ig an te a (l en de nf el d, 1 88 8) 4 sp he ri ca l; su rf ac e w ith 7 m m h ig h m ou nd s; tu be rc ul os e. 60 –9 0 m m o ra ng ere d w id el y sp ac ed tr ac ts of m eg as cl er e bu nd le s (5 00 –7 00 ), se pa ra tin g on ly s lig ht ly in s ur fa ce tu ft s. m s: s tr on gy le s (2 00 0 × 35 ) m e: o xy sp he ra st er s (3 0– 60 ) m i: c hi as te rs /o xy as te rs (1 0– 20 ) & po ly rh ab ds pt j ac ks on , n ew so ut h w al es , a us tr al ia u nk no w nb ta bl e 2 (c on tin ue d) . c om pa ri so n of m or ph ol og y be tw ee n sp ec ie s of t et hy a l am ar ck , 1 81 5 fr om a us tr al ia a nd n ew z ea la nd . european journal of taxonomy 529: 1–26 (2019) 14 sp ec ie s e xt er na l a pp ea ra nc e b od y di am . c ol ou r sk el et al a rr an ge m en t sp ic ul es ty pe lo ca lit y d ep th (m ) t. g un ni s ar à & s ar à, 20 04 1 sm oo th w ith s m al l co nu la te d tu be rc ul es . 25 m m fa de d pu rp le (i n et ha no l) tr ac ts o f m eg as cl er e bu nd le s (2 50 –5 00 ) br an ch in th e ch oa no so m e en di ng in c om pa ct fa ns in tu be rc ul es . m s st ro ng yl ox ea s (t o 18 50 × 3 5) , s ty le s (t o 80 0 × 10 ) m e: s ph er as te rs (2 0– 55 ) m i: ty la st er s/ ch ia st er s/ ox ya st er s ta sm an ia , a us tr al ia u nk no w n t. h oo pe ri s ar à & sa rà , 2 00 41 b od y el lip so id ; s m oo th su rf ac e, n o tu be rc ul es . 20 –3 0 m m o ch ra ce ou s (i n et ha no l) tr ac ts o f m eg as cl er e bu nd le s (2 50 –5 00 ) r ad ia te in c or te x w ith s ec on da ry an d te rt ia ry b ra nc hi ng . m s: s tr on gy lo xe as (t o 15 30 × 2 0) m e: s ph er as te rs (3 0– 60 ) m i: ty la st er s (8 –1 4) o xy as te rs (1 5– 25 ) h er on , i s. g re at b ar ri er r ee f, a us tr al ia 12 t. in ga lli b ow er ba nk , 18 58 1 sp he ri ca l/s ub -s ph er ic al ; fla tte ne d of te n in co ns pi cu ou s tu be rc ul es . b as al s to lo ns . 30 –5 0 m m pi nk is h tr ac ts o f m eg as cl er e bu nd le s sp lit in to se co nd ar y an d te rt ia ry br an ch es in c or te x. m s: s tr on gy lo xe as /a ni so st ro ng yl es (t o 21 30 × 3 0) m e: o xy sp he ra st er s (7 0– 90 ) m i: ty la st er s (1 0– 15 ), ch ia st er s, o xy as te rs (2 0– 30 ) sw a us tr al ia 30 –5 0 t. ja po ni ca s ol la s, 18 88 6 sp he ri ca l; co nu le s ro un de d; ge m m if er ou s. 13 –2 2 m m g re yi sh w hi te [n o de ta ils o n tr ac ts .] m s: s tr on gy lo xe as (t o 15 00 × 2 6) m e: s ph er as te rs (6 7) m i: c hi as te rs (1 2) e as te rn p hi lip pi ne s 32 t. la ev is (l en de nf el d, 18 88 ) 4 sp he ri ca l/o va te ; s ur fa ce sm oo th to tu be rc ul at e. 30 –4 5 m m l ig ht b ro w n (i n et ha no l) tr ac ts o f m eg as cl er e bu nd le s (5 00 –8 00 ) e xp an d to s ur fa ce tu ft s. m s: s tr on gy lo xe as (t o 20 00 × 3 6) m e: s ph er as te rs (3 0– 60 ) m i: c hi as te rs /o xy as te rs (1 0– 20 ), ty la st er s (1 0– 15 ), po ly rh ab ds pt j ac ks on , n ew so ut h w al es , a us tr al ia u nk no w n2 t. m ag na k ir kp at ri ck , 19 03 1, 7 o va l o r s ph er ic al ; co nu la te d su rf ac e (i n yo un g sp ec im en s) , e xp an di ng to po ly go na l p la te s. t re elik e ro ot le ts . 70 m m pu rp le br ow n (i n et ha no l) tr ac ts o f s to ut m eg as cl er e bu nd le s (5 00 –1 00 0) , un br an ch ed b ut w ith te rm in al fa ns . m s: s tr on gy lo xe as (t o 48 05 × 7 5) m e: s ph er as te rs (1 20 ) m i: a st er s (3 5– 45 ), ch ia st er s (1 2– 17 ) n at al , s ou th a fr ic a 62 t. m ic ro st el la s ar à, 19 90 8 h em is ph er ic al /c us hi on lik e; tu be rc ul es v ar y fr om co nu le s to fl at p ap ill ae . 10 m m b ro w ni sh / lig ht y el lo w tr ac ts o f m eg as cl er e bu nd le s in c ho an os om e ar e ac co m pa ni ed b y st yl es in c or te x. m s: s tr on gy lo xe as (t o 16 00 × 1 5) m e: s ph er as te rs (3 0– 80 ) m i: ty la st er s (4 –6 ) & la rg er ‘o th er m ic ra st er s’ pi on ee r b ay , o rp he us is , q ue en sl an d, a us tr al ia 0– 1 t. m on st ro sa (b ur to n, 19 24 )9 sp he ri ca l; su bgl ab ro us , a fe w s m al l b ud s at ta ch ed . r oo ti ng p ro ce ss es w el l de ve lo pe d. 14 –2 5 m m n ut b ro w n (i n et ha no l) r ad ia l t ra ct s of m eg as cl er e bu nd le s sp re ad ou t i nt o fa ns a t s ur fa ce . m s: s tr on gy lo xe as m e: s ph er as te rs (4 8) m i: ty la st er s (2 0) & o xy as te rs (3 2) . m an y m ic ro sc le re s ha ve a bn or m al iti es w he re th e ty le s & s pi ne s ar e re du ce d to k no bs o r ab se nt le av in g ju st a s ph er e. ta sm an ia u nk no w n ta bl e 2 (c on tin ue d) . c om pa ri so n of m or ph ol og y be tw ee n sp ec ie s of t et hy a l am ar ck , 1 81 5 fr om a us tr al ia a nd n ew z ea la nd . sorokin s.j. et al., a new deep-water tethya (porifera: demospongiae) 15 sp ec ie s e xt er na l a pp ea ra nc e b od y di am . c ol ou r sk el et al a rr an ge m en t sp ic ul es ty pe lo ca lit y d ep th (m ) t. m or to ni b er gq ui st & k el ly -b or ge s, 1 99 12 o vo id ; l ow tu be rc ul es . b ud s on e re ct fi la m en ts . 6– 8 m m m ai ze ye llo w fi ne tr ac ts o f m eg as cl er e bu nd le s (1 22 –4 90 ) un if or m fr om c ho an os om e to c or te x, e xp an d sl ig ht ly in c or te x. in te rs tit ia l sp ic ul es ra di al . m s: a ni so st ro ng yl ox ea s (t o1 09 2 × 21 ) m e: lo ng -r ay ed o xy sp he ra st er s (2 6– 62 ), sh or t r ay ed o xy sp he ra st er s (2 8– 52 ) m i: in cl . a ca nt ho st ro ng yl as te rs & ac an th ot yl as te rs (8 –1 3) m an uk au h ar bo ur , a uc kl an d, n ew z ea la nd 0. 5 t. m ul tis te lla l en de nf el d, 1 88 84 ,1 sp he ri ca l t o su bs ph er ic al ; re gu la rl y tu be rc ul os e. 20 –4 0 m m r ed , fl es h or ro se r ad ia l t ra ct s of m eg as cl er e bu nd le s (3 00 –5 00 ) a re tr um pe tlik e in s ha pe a nd s pr ea d to s ec on da ry a nd te rt ia ry br an ch es in tu be rc ul es . m s: s tr on gy lo xe as (t o 35 00 × 1 8) m e: s ph er as te rs (2 0– 50 ) m i: ty la st er s (1 2– 16 ), ox ya st er s (5 –1 0) pt . j ac ks on , n ew so ut h w al es a us tr al ia u nk no w n t. o rp he i s ar à, 1 99 08 su bs ph er ic al ; l en gt he ne d bu t a pi ca lly fl at te ne d tu be rc ul es a t t op o f s po ng e, sm oo th e ls ew he re . 5 m m b ro w n ye llo w is h [t ra ct s/ bu nd le s no t de sc ri be d. ] m s: s tr on gy lo xe as , ( to 1 60 0 × 15 ) s ty le s (t o 12 00 × 1 0) m e: s ph er as te rs (1 0– 50 ) m i: c hi as te rs & ty la st er s (8 –1 2) , o xy as te rs (1 0– 30 ) pi on ee r b ay , o rp he us is , q ue en sl an d, a us tr al ia 0– 1 t. p el lis b er gq ui st & k el ly -b or ge s, 1 99 12 fl at te ne d su bs ph er ic al , tu be rc ul es d iv id ed b y po re g ro ov es ; fi rm b ut co m pr es si bl e. 40 m m fl es h ro se tr ac ts o f m eg as cl er e bu nd le s w id el y sp ac ed – br an ch in to p th ir d of co rt ex in to fa ns . m s: a ni so st ro ng yl ox ea s (t o 20 75 × 3 0) m e: o xy sp he ra st er s (2 5– 12 0) , o xy as te rs (2 0– 25 ) m i: a ca nt ho -c hi as te rs /ty la st er s/ ox ys ph er as te rs (1 3– 18 ), m ic ro ox ya st er s (5 –1 1) fa ir lig ht , s yd ne y h ar bo ur , n ew so ut h w al es , a us tr al ia in te rt id al t. p op ae b er gq ui st & k el ly -b or ge s, 1 99 12 ir re gu la r s ub sp he ri ca l; ill -d efi ne d tu be rc ul es or s m oo th ; 2 –8 a pi ca l os cu le s; s po ng es c an c lu st er co nn ec te d by s to lo ns . 8– 16 m m b ri gh t d ee p or an ge fl ex uo us tr ac ts o f m eg as cl er e bu nd le s ha ve e la bo ra te te rt ia ry br an ch in g. m s: a ni so st ro ng yl ox ea s (t o 91 0 × 10 ) m e: o xy sp he ra st er s (2 0– 55 ) m i: a ca nt ho ch ia st er s & ac an th os tr on gy la st er s (7 –1 3) , m ic ro ox ya st er s (2 –1 0) c ap e b an ks , n ew so ut h w al es , a us tr al ia in te rt id al t. p ul itz er i s ar à & sa rà , 2 00 41 [n o de sc ri pt io n, s pe ci es de si gn at io n fr om s lid es on ly .] u nk no w n u nk no w n u nk no w n. m s: s tr on gy lo xe as , a ni so st ro ng yl es (t o 11 00 × 1 5) m e: o xy sp he ra st er s (3 0– 50 ) m i: ty la st er s & c hi as te rs (8 –1 0) , o xy as te rs (1 0– 15 ) h er on is la nd , g re at b ar ri er r ee f, a us tr al ia u nk no w n t. r ob us ta (b ow er ba nk , 1 87 3) 1 h em is ph er ic al ; p ol yg on al ar ea s se pa ra te d by g ro ov es a tt ac he d by s ho rt b ro ad ro bu st b as al s to lo ns . 30 –5 0 m m y el lo w -g re y sl im tr ac ts o f m eg as cl er e bu nd le s ex pa nd s lig ht ly in co rt ex . m s: a ni so st ro ng yl ox ea s (t o 18 50 × 3 2) m e: o xy sp he ra st er (3 0– 90 ), ox ya st er s (1 1– 22 ) m i: a ca nt ho -t yl as te rs /s tr on gy la st er s (7 –1 2) , m ic ro ox ys ph er as te rs (4 –5 ) a us tr al ia 0. 2– 30 ta bl e 2 (c on tin ue d) . c om pa ri so n of m or ph ol og y be tw ee n sp ec ie s of t et hy a l am ar ck , 1 81 5 fr om a us tr al ia a nd n ew z ea la nd . european journal of taxonomy 529: 1–26 (2019) 16 sp ec ie s e xt er na l a pp ea ra nc e b od y di am . c ol ou r sk el et al a rr an ge m en t sp ic ul es ty pe lo ca lit y d ep th (m ) t. s ey ch el le ns is (w ri gh t, 18 81 ) 1 ,2 sp he ri ca l o r h em is ph er ic al ; sm oo th w ith fl at te ne d po ly go na l t ub er cu le s se pa ra te d by g ro ov es ; fi rm co m pr es si bl e. a tt ac he d by fin e st ol on s. 15 –2 5 m m b ro w n re d tr ac ts o f m eg as cl er e bu nd le s (2 50 –5 00 ) h av e se co nd ar y br an ch in g in co rt ex – s pi cu le s pr ot ru de w el l b ey on d su rf ac e. m s: a ni so st ro ng yl ox ea s (t o 15 00 × 3 0) m e: o xy sp he ra st er s (1 7– 76 ), ox ya st er s (3 2– 50 ) m i: a ca nt ho ch ia st er s & ac an th os tr on gy la st er s (7 –1 3) , m ic ro as te rs (7 –1 7) d am pi er , w es te rn a us tr al ia in te rt id al t. s te llo de rm is s ar à & sa rà , 2 00 41 [s ha pe n ot d es cr ib ed ]; su rf ac e of v er y fla tte ne d tu be rc ul es ; v er y ha rd . 25 m m pa le p ur pl e br ow n (i n et ha no l) t hi ck tr ac ts o f m eg as cl er e bu nd le s (3 00 –5 00 ) b ra nc h in to s ec on da ry a nd te rt ia ry tr ac ts in c or te x. m s: s tr on gy lo xe as (t o 14 00 × 2 8) m e: s ph er as te rs /o xy as te rs (1 0– 40 ) m i: v ar io us in cl . t yl as te rs (2 0– 25 ), ox ya st er s (5 –1 0) pt p hi lli p h ea ds , v ic to ri a u nk no w n t. s to lo ni fe ra b er gq ui st & k el ly b or ge s, 1 99 12 sp he ri ca l t o su bs ph er ic al ; ci rc ul ar o r p ol yg on al tu be rc ul es ; s of t co m pr es si bl e. e xt en si ve st ol on s. 3– 11 m m r ed di sh ye llo w n ar ro w w el l s ep ar at ed tr ac ts o f m eg as cl er e bu nd le s (2 00 –4 25 ) r ad ia te in c or te x fo rm in g co ni ca l br us he s. m s: a ni so st ro ng yl ox ea s (t o 12 25 × 2 6) m e: o xy sp he ra st er s (1 3– 76 ); m i: a ca nt ho st ro ng yl as te rs , ac an th ot yl as te rs (8 –1 4) ra re m ic ro ox ys ph er as te rs (3 –1 0) w ai te m at a h ar bo ur , a uc kl an d, n ew z ea la nd 0. 5 –1 .5 t. ta sm an ia e sa rà & sa rà , 2 00 41 sp he ri ca l w ith s m al l tu be rc ul es . 20 m m u nk no w n tr ac ts o f m eg as cl er e bu nd le s (5 00 –1 00 0) br an ch in c ho an os om e an d en d in c or te x as w id e co ne s. m s: s tr on gy lo xe as /a ni so st ro ng yl es (t o 12 00 × 2 0) m e: s ph er as te rs 2 0– 50 m i: v ar io us e .g ., ty la st er s, o xy as te rs (3 ty pe s) , c hi as te rs b la ck m an s b ay , ta sm an ia , a us tr al ia u nk no w n so ur ce s. 1 sa rà & s ar à (2 00 4) ; 2 b er gq ui st & k el ly -b or ge s (1 99 1) ; 3 b er gq ui st ( 19 61 ); 4 l en de nf el d (1 88 8) ; 5 h al lm an n (1 91 4) ; 6 s ol la s (1 88 8) ; 7 k ir kp at ri ck ( 19 03 ); 8 sa rà (1 99 0) ; 9 b ur to n (1 92 4) . n ot es . * u nk no w n, o r n ot n ot ed in s ou rc es . a m ax im um d ep th o f p t p hi lli p h ea ds is 5 0 m (b ar to n et a l. 20 12 ); b m ax im um d ep th o f p t j ac ks on is 4 5 m (j oh ns to n et a l. 20 15 ). te th ya d ip lo de rm a sc hm id t, 18 70 is li st ed a s oc cu rr in g in n ew z ea la nd b y k el ly e t a l. (2 00 9) , b ut is n ot in cl ud ed a s th is is c ur re nt ly s ho w n as in ac cu ra te b y va n so es t ( 20 19 ). te th ya c om pa ct a b er gq ui st , 1 96 1 is n ot in cl ud ed in th e lis t o f a us tr al ia a nd n ew z ea la nd s pe ci es b y sa rà & s ar à (2 00 4) d ue to it s sy no ny m is at io n by b er gq ui st w ith t . a ur an tiu m ( pa lla s, 1 76 6) , h ow ev er th ey n ot e th at th e ta xo no m ic s ta tu s of t . c om pa ct a “r em ai ns u nc er ta in ”; it is in cl ud ed h er e as it is s ho w n as ac ce pt ed b y va n so es t ( 20 19 ). t he o cc ur an ce o f t . a ur an tiu m in n ew z ea la nd is s ho w n as in ac cu ra te (v an s oe st 2 01 9) a nd is n ot in cl ud ed h er e. ta bl e 2 (c on tin ue d) . c om pa ri so n of m or ph ol og y be tw ee n sp ec ie s of t et hy a l am ar ck , 1 81 5 fr om a us tr al ia a nd n ew z ea la nd . sorokin s.j. et al., a new deep-water tethya (porifera: demospongiae) 17 results of the phylogenetic analyses 28s and coi trees had similar topologies (fig. 5). the monophyly of the tethyida was not supported (28s, bootstrap of 12; coi, bootstrap of 33) with timeidae sister to a moderately (coi) to poorly supported (28s) tethyidae + hemiasterellidae clade. timea sp. from the ‘3pp cave’, la ciotat, france, (chombard 1998) did not group with the rest of the timeidae, but its paraphyletic position was poorly supported. the hemiasterellidae (adreus, axos, liosina) seemed to group in a moderately to wellsupported clade (28s, 69; coi, 98). the 28s tree suggested that the australian laxotethya dampieriensis (tethyidae) and a tethyida sp. from ireland with no obvious genus assignment (c. morrow, pers. comm.) had more ambiguous positions: in raxml analyses they grouped together (poorly supported), while in mrbayes analyses tethyida sp. branched between the timeidae and the rest of the tethyida. liosina paradoxa thiele, 1899 and liosina blastifera vacelet, bitar, carteron, zibrowius & perez, 2007 did not group together in the 28s trees, while adreus was polyphyletic with two clades (one from australia, the other from the north atlantic). with coi, where we only have tethya sequences, we could identify four moderately to well-supported clades: 1) the t. aurantium clade, 2) the t. wilhelma clade, 3) the t. citrina clade and 4) the t. actinia laubenfels, 1950 clade. our new species t. irisae sp. nov. branched as the sister taxa to clade 4 (in raxml and mrbayes) but this is poorly supported. in our 28s dataset where we found the same four clades, all other tethyidae genera (tethytimea laubenfels, 1936, stellitethya, xenospongia and tectitethya sarà, 1994) were mixed with tethya species, especially in clade 4. t. irisae sp. nov. grouped with tectitethya in clade 4 (in raxml and mrbayes analyses) but this is poorly supported. t. wilhelma was originally described from a tropical aquarium in germany but its original geographical location is unknown. in tethya clade 2, we noticed several species were very close genetically to t. wilhelma: with coi, tethya sp. from israel (mediterranean sea) had only 1 bp difference with t. wilhelma; with 28s, t. taboga from panama had 0–2 bp difference with t. wilhelma (uncertainty due to two ambiguous base pairs) while tethya sp. 2 from saudi arabia had 1 bp difference with t. wilhelma. discussion tethya irisae sp. nov. is a new and distinctive sponge from the slope of the great australian bight (gab). this is the 28th tethya species reported from australia, and the deepest, being found in the bathyal zone at around 1000 m deep. with uncertainties in the validity of genera divisions based on morphological characters e.g., possession of a stalk, a reappraisal of the genera within tethyidae based on molecular sequencing is needed. phylogeny of tethyidae the first phylogenetic analyses of tethya, using coi and morphology (heim et al. 2007; heim & nickel 2010) revealed four main clades: 1) the seychellensis-wilhelma complex, 2 + 3) the citrinia-actinia complex divided in two subclades (european species and western atlantic species + eastern pacific) and 4) the aurantium clade. our coi and 28s analyses with extended datasets retrieve these four clades, but with a higher biogeographical diversity. the seychellensis-wilhelma complex now includes specimens from israel, vietnam, panama, china and queensland; the aurantium clade now includes species from the mediterranean sea, the red sea and panama. all clades are well-supported in the coi tree except for clade 3, the western atlantic/pacific clade. this is precisely the group joined by the coi sequence of t. irisae sp. nov.; its position within this group, however, remains unclear. these same four clades are not as clear in our 28s tree, their inter-relationships are also different, and not supported at all. this may be due to the fact that our 28s alignment is a mix of different 28s domains and different sampling than coi, both of which may influence some of the groupings. the seychellensis-wilhelma and aurantium clades are well supported with 28s as well. on the other hand, the citrinia and actinia subclades are unclear, and this is probably due to the addition in this dataset of many different genera of tethyidae (tethytimea, tectitethya, stellitethya, xenospongia gray, 1858, laxotethya). as suggested by sarà et al. european journal of taxonomy 529: 1–26 (2019) 18 sorokin s.j. et al., a new deep-water tethya (porifera: demospongiae) 19 (2001) and heim et al. (2007), tethya wilhelma and t. gracilis sarà, sarà, nickel & brümmer, 2001, both described from aquaria in germany belong to the seychellensis-wilhelma complex. there is only 1 bp difference between the coi of t. wilhelma and tethya sp. (mediterranean sea, israel) so this specimen should be re-examined to see if it could be conspecific with t. wilhelma. heim et al. (2007) showed that the most reliable characters for tethya taxonomy were morphometric spicule data, but none could actually make good morphological synapomorphies for the two tethya clades supported with coi and 28s. new characters (e.g., chemical compounds, specialized cells, associated microbes) must be explored in order to find independent support for these groups. external colour may be a reliable character to discriminate those clades, as shown previously in some calcareous sponges (rossi et al. 2011). indeed, most shallow water tethya species have a yellow, orange to red surface colour, probably due to different carotenoids (tanaka et al. 1982) some of which they can synthesise themselves (liaaen-jensen et al. 1982) and therefore have a genetic basis. all species currently in the citrina subclade (t. norvegica bowerbank, 1872, t. citrina and t. hibernica) are light-yellow coloured. species from the actinia subclade and aurantium clade are usually bright yellow to orange to bright orange, except for the ‘aquarium’ species t. minuta sarà, sarà, nickel & brümmer, 2001 (white, in artificial conditions at least). finally, the seychellensis-wilhelma clade seems to include especially bright red/carmine surface-coloured species (t. seychellensis, tethya sp. from bocas, t. coccinea bergquist & kelly-borges, 1991, tethya sp. 3 from saudi arabia, t. taboga, t. samaaii ribeiro & muricy, 2011), except for the tethya sp. from israel which was more light orange, and except again for the ‘aquarium’ species (t. wilhelma and t. gracilis). in red surface-coloured species, the choanosome is usually orange. however, more colours exist: some tethya can be green (e.g., tethya brasiliana ribeiro & muricy, 2004), dark blue (e.g., tethya cyanea ribeiro & muricy, 2004), or pink (e.g., tethya bergquistae) but none of these species have been sequenced yet. we can probably dismiss the green colour. it is found in species that can also be orange; laubenfels (1950) suggested the green colour of t. actinia in bermuda was due to symbiotic algae (a specimen may “turn orange” when fixed in alcohol, as the chlorophyll is extracted). more problematic are species with varying colours, from yellow to orange and red (e.g., tethya fastigata). as for the few deep-sea species of tethya, some have lost their colours (e.g., tethya irisae sp. nov.) while others have retained them: e.g., tethya levii sarà, 1988 from new caledonia is light orange, and groups in the actinia clade, in accordance with our hypothesis (p. cárdenas, unpublished data). this groupingby-colour hypothesis should be further tested with the sequencing of new species of tethya. other genera of tethyidae included in our dataset have usually irregular massive forms or are disc-shaped (instead of subspherical forms), and all have dark colours: black-brownish for tectitethya spp., beige-gray for xenospongia, and whitish-brown in ethanol for laxotethya and stellitethya (the live colour is unknown). since all except laxotethya are sister group to a bright orange tethya sp. from south australia (possibly in the actinia clade) (fig. 5, 28s tree), we suppose the common ancestor of these other genera lost its yellow-orange colours, and so its capacity to produce carotenoids. our coi and 28s dataset include type species of four tethyidae genera (of the 14 valid genera): tethya (t. aurantium, coi), tectitethya (t. crypta, 28s), xenospongia (x. patelliformis, 28s) and laxotethya (l. dampierensis, 28s). in addition to that, two other tethyidae genera are represented in our 28s tree: stellitethya and tethytimea. all these genera are essentially defined by different skeletal structures and therefore body shape; all these genera have an indistinct or ill-defined cortex (vs a distinct thick cortex for tethya) and an irregular massive or encrusting shape (vs (sub)spherical shape in tethya). our fig. 5 (opposite page). tethyida coi and 28s maximum-likelihood (raxml) trees. ml bootstrap supports (1000 bootstrap replicates) > 70 are indicated. after the species name, locality of the specimen is given (when known), followed by the genbank accession number(s). for 28s, we also indicated the 28s region that was sequenced as well as the first author + date of the publication where the sequence first appeared. type species of genera are in red boxes while tethya irisae sp. nov. appears in red. european journal of taxonomy 529: 1–26 (2019) 20 28s tree suggests that xenospongia, stellitethya, tectitethya and tethytimea are grouping with tethya (fig. 5), while laxotethya groups with hemiasterellidae, albeit with no support. tethytimea carmelita, tectitethya and stellitethya/xenospongia evolved independently within tethya thus suggesting that the loss of a distinct cortex and of the subspherical shape happened several times. more sequences from australian tethya are needed to understand the origin and relationships of these other tethyidae genera. one clade that is moderately supported (bootstrap of 69) is the sister-group relationship of xenospongia and stellitethya, with a 5–6 bp difference in 28s (d3–d5). both genera have a poorly defined cortex but different shapes: discoid for xenospongia, massive irregular for stellitethya. these two genera also share megasters reaching large sizes (> 150 µm), as in t. irisae sp. nov., grouping nearby (bootstrap of 62) with tectitethya (which does not have very large megasters). phylogeny of tethyida the tethyida also include hemiasterellidae and timeidae. since the ex-hemiasterellid genera stelligera gray, 1867 and paratimea have been reallocated to the stelligeridae lendenfeld, 1898, order axinellida (morrow et al. 2012), the hemiasterellidae include four genera: adreus, axos, hemiasterella and the monospecific leptosastra topsent, 1904. the sister position of hemiasterellidae adreus and axos with the tethyidae has been repeatedly shown by previous coi (morrow et al. 2012, 2013), 28s (thacker et al. 2013; cruz-barraza et al. 2017) and 18s analyses (redmond et al. 2013). liosina, a genus with only four species and a loose arrangement of oxeas/styles, had been tentatively assigned to the family dictyonellidae, order bubarida (van soest et al. 2002). the grouping of liosina paradoxa, type species, with the hemiasterellidae was revealed for the first time by morrow et al. (2012) with 28s. our study confirms this grouping for the first time with coi sequences (l. paradoxa and liosina blastifera), as well as a 28s sequence of l. blastifera. its unambiguous grouping with the hemiasterellidae suggests that species of liosina are in fact hemiasterellidae that have secondarily lost their euasters. furthermore, liosina often have a polygonal surface pattern, and/or pores in shallow grooves, a character present in most tethyidae and timeidae, which often gives rise to the characteristic surface tubercles. since we have sequenced the type species of liosina, we formally propose the reallocation of liosina from dictyonellidae to hemiasterellidae. however, it is unclear in our trees whether liosina is polyphyletic. the small polygons from liosina, also called tubercles or microconules, are also present in the three adreus species from australia (a. australiensis (ridley, 1884), a. axiferum (hentschel, 1912) and adreus sp.) thereby confirming their reallocation. these two branching species also share with the tethyida the typical radial bundles of megascleres fanning out closer to the surface. adreus australiensis and a. axiferum were previously grouped in raspailidae (hooper 1991) although they lacked echinating megascleres, before erpenbeck et al. (2007) showed with 28s that they clustered instead with hemiasterellidae and tethyidae. these two species also secondarily lost their asters, while the vase-shaped adreus sp. from queensland (qm g315767) still has tylasters. this second case of aster loss in the tethyida suggests that, similarly as in the astrophorina (cárdenas et al. 2011), more genera or species without asters, can be expected to be reallocated to the tethyida once they are sequenced. since this adreus clade does not cluster with the clade of adreus fascicularis (type species of the genus), they potentially represent a new genus in the hemiasterellidae. to conclude, the hemiasterellidae now include adreus, axos, hemiasterella, leptosastra, liosina, and a potential new genus. so far, all genbank hemiasterella sp. sequences are doubtful and failed to cluster with the hemiasterellidae (cf. material and methods). the type specimen of hemiasterella typus carter, 1879, has not been revised and sequenced so that the phylogenetic position of hemiasterella remains to be tested. we note, however, that h. typus does not share with most of the tethyida 1) a surface with pores in grooves around tubercles/plates or 2) bundles of megascleres fanning out at the surface. to sum up the main findings of the phylogenetic analysis. sorokin s.j. et al., a new deep-water tethya (porifera: demospongiae) 21 1. four tethya clades were retrieved (as in previous analyses) for which no synapomorphies are currently known; we, however, discuss the possibility of using external colour to support some of these clades. 2. despite unclear phylogenetic relationships amongst tethyidae from australia, our results suggest that tethyidae genera tethytimea, tectitethya, laxotethya, stellitethya, and xenospongia derive from species of tethya, which may challenge their validity in the future. 3. our results suggest that hemiasterellidae is the sister-group of tethyidae while the position of timeidae is still ambiguous (not supported). 4. we show that asters have been secondarily lost at least twice in the hemiasterellidae: in liosina and a potential new genus from northern australia. we formally propose the reallocation of liosina from dictyonellidae to hemiasterellidae. acknowledgements special thanks to franzis althaus (csiro) for providing the collection map, lisa goudie (consultant, victoria) for preparation of slides, and andrea crowther and moon lee (south australian museum) for the loans of sponges to authors. karen gowlett-holmes took the photograph of freshly collected sponges. thanks to steven keable (australian museum) for sourcing the type specimens of tethya bullae and tethya fissurata. ingrid van streepen (volunteer, south australian museum) took the photograph of t. fissurata. thanks to lauren hughes (natural history museum, london) for providing access to the type slides of burtonitethya gemmiformis. thanks to scott nichols (university of denver) for sending a piece of hemiasterella sp. ucmpwc1021. thanks to the riesgo lab (nhm, london) for sharing preliminary coi and 28s alignments of tethya species. thanks to warren r. francis (university of southern denmark) for sharing the full coi and 28s from the t. wilhelmi genome. thanks to christian marschal (imbe, station marine d’endoume, marseille, france) for making the thick sections. s. sorokin wrote this paper while an honorary research associate at the south australian museum and acknowledges her supervisor dr rachael king. qi yang acknowledges the support by the national natural science foundation of china (grant nos 31801954 and 41729002). p. cárdenas received support from the european union’s horizon 2020 research and innovation program through the sponges project (grant agreement no. 679849). this document reflects only the authors’ view and the executive agency for small and medium-sized enterprises (easme) is not responsible for any use that may be made of the information it contains. the authors thank the 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(finland, canada), b. falciceps sp. nov. (finland, canada), b. oelandica sp. nov. (sweden) and b. plusiospina sp. nov. (finland). a few bradysia species, described previously and now found in northern europe, are also redefined and illustrated. key words. sciaroidea, holarctic region, biodiversity, systematics, new taxa heller k., hippa h. & vilkamaa p. 2015. taxonomy of bradysia winnertz (diptera, sciaridae) in the northern holarctic, with the description of four new species. european journal of taxonomy 122: 1–15. http://dx.doi. org/10.5852/ejt.2015.122 introduction within the framework of the research project ‘diversity of the fennoscandian black-winged fungus gnats (diptera, sciaridae),’ financed by the swedish taxonomy initiative, and the research programme ‘deficiently known and threatened forest species’ (putte, see juslén et al. 2008), financed by the ministry of the environment, finland, a number of new species in various genera (vilkamaa et al. 2013a, b, c), including species of the genus bradysia winnertz, 1867, have been found in northern europe, while others are new geographical records. furthermore, conspecific specimens originating from saproxylic studies ongoing in canada have indicated that some of these species are holarctic in distribution. the main aim of this paper is to further expand our knowledge of northern sciaridae by describing species of bradysia that are new to science and redescribing others that are new records. material and methods all specimens of the new species were detected and picked out from unsorted sciarid or general insect material stored in ethanol. most specimens originated from malaise trap samples, but some had been http://dx.doi.org/10.5852/ejt.2015.122 www.europeanjournaloftaxonomy.eu/index.php/ejt/index http://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:26d7aa3f-7d72-45b2-bf6d-69bafdc7b0e7 mailto:kaiheller%40gmx.de?subject= mailto:heikki.hippa%40gmail.com?subject= mailto:pekka.vilkamaa%40helsinki.fi?subject= http://zoobank.org/urn:lsid:zoobank.org:author:8f9a2911-a7ec-4ca7-acac-e9339d258de3 http://zoobank.org/urn:lsid:zoobank.org:author:546524ad-3bd1-4830-842f-eeee7df4799b http://zoobank.org/urn:lsid:zoobank.org:author:a4ead3b5-da5e-4c09-8e3a-9289ca7358ed http://dx.doi.org/10.5852/ejt.2015.122 http://dx.doi.org/10.5852/ejt.2015.122 european journal of taxonomy 122: 1–15 (2015) 2 caught by sweeping or with an aspirator, a photoeclector or pan traps. the specimens were mounted on microscope slides in euparal, after dehydrating them in absolute ethanol. the terminology and methods of measuring and illustrating morphological structures follow hippa & vilkamaa (1991, 1994) and hippa et al. (2010). the material is deposited in the following collections: mzh = zoological museum, finnish museum of natural history, helsinki, finland pkhh = private collection of kai heller, heikendorf, germany prdm = private collection of rob deady, montréal, canada pwmp = private collection of werner mohrig, poseritz, germany smnh = swedish museum of natural history, stockholm, sweden zfmk = zoological museum bonn, germany results class insecta linnaeus, 1758 order diptera linnaeus, 1804 superfamily sciaroidea billberg, 1820 family sciaridae billberg, 1820 genus bradysia winnertz, 1867 bradysia winnertz, 1867: 180. type-species: bradysia angustipennis winnertz, 1867, by subsequent designation of enderlein (1911: 127, 183). bradysia ascenda rudzinski, 1994 fig. 1a–d material examined sweden: torne lappmark, kiruna, abisko, 68°20’ n, 18°52’ e, north slope, 440 m, edge of birch forest, white pan trap, 13–17 sep. 1991, v. tschirnhaus, 3 ♂♂, pwmp. redescription (male) head. dark brown, antenna unicolorous pale brown; maxillary palpus very pale brown. eye bridge 2–3 facets wide. face with 10–12 scattered longer and shorter setae. clypeus with 1–4 setae. maxillary palpus with 3 segments; segment 3 longer than segment 1, segment 2 shortest; segment 1 with 3–4 setae, with dorsal pit of sensilla; antennal flagellomeres with slightly rough surface, body of flagellomere 4 2.15–2.4× as long as wide, neck shorter than broad, longest setae longer than width of flagellomere. thorax. dark brown, setae pale. anterior pronotum with 6–12 setae. episternum 1 with 13–15 setae. scutum with long dorsocentrals, with some longer and shorter laterals, scutellum with 2 longer and some short setae. wing. fumose. length 1.8–1.9 mm. width/length 0.40. veins distinct. r1/r 0.65. c/w 0.70–0.75. r-m shorter than bm, r-m with 1 seta, bm non-setose. legs. pale brown. coxal setae pale. front tibial organ with pale vestiture forming a row. front tibial spur longer than tibial width. claws without teeth. heller k., hippa h. & vilkamaa p., taxonomy of bradysia 3 abdomen. pale brown, setae pale. hypopygium (fig. 1a–d). brown, concolorous with abdomen. gonocoxa. narrow, longer than gonostylus, mesial margin with normal setosity. gonostylus. narrow, rather straight, slightly thickened at middle, slightly narrowed towards apex, with mesial side not impressed; with dense apical setosity, without apical tooth, with 8–9 apical-subapical megasetae; megasetae subequal in size, slightly curved and sharp. tegmen conical, apically truncate, with straight lateral sides, with indistinct basolateral corners, laterally sclerotized, with area of aedeagal teeth. aedeagal apodeme long. remarks bradysia ascenda was described on the basis of three males from the austrian alps (rudzinski 1994). in having its setae pale and in having a distinct sensory pit in the maxillary palpus, it belongs to the bradysia tilicola group of menzel & mohrig (2000). fig. 1. bradysia ascenda rudzinski, 1994 (from sweden). a. part of hypopygium, ventral view. b. gonostylus, ventral view. c. tegmen and cerci, ventral view. d. apical part of gonostylus, ventral view. scales = 0.1 mm. european journal of taxonomy 122: 1–15 (2015) 4 bradysia bigeminata sp. nov. urn:lsid:zoobank.org:act:2b3eb8ac-0d40-48b8-8453-ac9ca78b6316 fig. 2a–b diagnosis small-sized bradysia. dark brown. maxillary palpus with 2 segments. wing length 1.5 mm, anal lobe strong. gonostylus narrow, without apical tooth, with 4 subequal megasetae. etymology the name is derived from the latin words bis(two) and geminus (twin), referring to the two pairs of gonostylar megasetae of the species. material examined holotype finland: ♂, ostrobothnia borealis, pudasjärvi, murto-oja (grid 7267183:3525642), near a brook, malaise trap, 1 may–3 jul. 2006, j. ilmonen & j. salmela, mzh. paratypes finland: karelia borealis, kuhmo, elimyssalo, bred from fomitopsis rosea collected 23–28 may 1998, a. komonen, 1 ♂, mzh. canada, ontario, sudbury co., chapleau, nimitz forest, from wind thrown pinus banksiana trees, photoeclector, 10–24 jul. 2012, r. deady & t. work, 14 ♂♂, 1 [no. 8622] in pkhh, 3 [zmfktis-2527999, zmfk-tis-2527999, zmfk-tis-2528275] in zmfk, 1 in mzh, 9 in prdm; same data but 16–29 may 2012, 4 ♂♂, prdm, 1 ♂, mzh; same data but 28 may–12 jun. 2012, 2 ♂♂, prdm, 1 ♂, mzh; same data but 25 apr.–11 jul. 2012, 2 ♂♂, prdm; same data but 23 jul.–8 aug. 2012, 5 ♂♂, prdm; same data but 7–20 aug. 2012, 1 ♂, prdm; ontario, sudbury co., chapleau, superior forest, mature (99 years old) stand composed of 90% pinus banksiana and 10% p. mariana with an understory shrub layer of vaccinium, very wet with considerable bryophyte coverage, pine log, photoeclector, 21 jul.–6 aug. 2013, r. deady & t. work, 18 ♂♂, prdm; same data but 5–19 aug. 2013, 15 ♂♂, prdm; same data but 7–22 jul. 2013, 2 ♂♂, prdm, 1 ♂, mzh. description (male) head. dark brown, antennal flagellomeres unicolorous brown; maxillary palpus very pale brown. eye bridge 2–3 facets wide. face with 23 scattered longer and shorter setae. clypeus with 1 seta. maxillary palpus with 2 segments; segment 1 longer than segment 2, segment 1 with 1 seta, with dorsal patch of sensilla; surface of antennal flagellomeres smooth, body of flagellomere 4 2.65× as long as wide, neck shorter than broad, longest setae longer than width of flagellomere. thorax. dark brown, setae pale. anterior pronotum with 3 setae. episternum 1 with 7 setae. scutum with long dorsocentrals, with some longer and shorter laterals, scutellum with 4 longer and some short setae. wing. hyalinous. length 1.5 mm. width/length 0.50. anal lobe strong. veins distinct. r1/r 0.55. c/w 0.65. r-m and bm subequal in length, both non-setose. legs. yellow. coxal setae pale. front tibial organ with pale vestiture forming a short row. front tibial spur as long as tibial width. tibial spurs of middle and hind tibia subequal. tarsal setosity normal. claws without teeth. http://zoobank.org/urn:lsid:zoobank.org:act:2b3eb8ac-0d40-48b8-8453-ac9ca78b6316 heller k., hippa h. & vilkamaa p., taxonomy of bradysia 5 abdomen. brown, setae pale. hypopygium (fig. 2a–b). brown, concolorous with abdomen. gonocoxa. basally broad, longer than gonostylus, mesial margin with sparse setosity. gonostylus. narrow, rather evenly broad and straight, tumid, with mesial side impressed at apical half; with dense apical setosity, without apical tooth, with two pairs of sharp subapical megasetae. tegmen truncate, with straight lateral sides, laterally and apically sclerotized, with small area of solitary aedeagal teeth. aedeagal apodeme rather long. remarks bradysia bigeminata sp. nov. is here formally assigned to the genus bradysia because of the presence of a comb-like row of elements in its tibial organ and because of its subequal middle and hind tibial spurs, but it cannot be placed in one of the currently defined species groups. in the structure of its gonostylus it also resembles the species of the corynoptera vagula group (see vilkamaa & hippa 2006). by its elongated gonostylus with four apical megasetae, and by its short front tibial comb placed in a non-setose area at the apical part of the tibia, bradysia bigeminata sp. nov. resembles scatopsciara (xenopygina) vinea rudzinski & baumann, 2013, but differs in having a simpler tegmen and a different placement of the gonostylar megasetae. in a molecular phylogenetic analysis, the corynoptera vagula group of species appeared close to species of scatopsciara edwards (shin et al. 2013). in this analysis, no species of scatopsciara (xenopygina) frey, 1942 was included. it is obvious that a more inclusive analysis is needed to clarify the relationships of b. bigeminata sp. nov., the corynoptera vagula group and the species currently placed in scatopsciara (xenopygina). fig. 2. bradysia bigeminata sp. nov. (holotype). a. part of hypopygium, ventral view. b. gonostylus, ventral view. scales = 0.1 mm. european journal of taxonomy 122: 1–15 (2015) 6 ecology and distribution bradysia bigeminata sp. nov. was found to be very numerous in deadwood sampled in canada. this may suggest deadwood to be the natural habitat of the species. the coi sequence of the collected specimens was allocated to the bin aca4746 on bold (www.boldsystems.org), and, interestingly, the results showed that the species is widely distributed in eastern canada. bradysia chlorocornea mohrig & menzel, 1992 fig. 3a–b material examined finland: regio kuusamoensis, kuusamo, kuohusuo-kalliovaara 30 km s of kuusamo, 65.43° n, 29.05° e, 270 m, spruce/birch swamp forest, malaise trap, 2–15 aug. 2004, m. & c. jaschhof, 4 ♂♂, mzh. redescription (male) head. brown, antennal flagellomeres 1–2 yellow, scapus, pedicellus and flagellomere 3–14 pale brown; maxillary palpus very pale brown. eye bridge 3 facets wide. face with 10–15 scattered longer and shorter setae. clypeus with 1–3 setae. maxillary palpus with 3 segments; segment 1 longer than segment 3, segment 2 shortest; segment 1 with 3–4 setae, with dorsal patch of sensilla; surface of antennal flagellomeres smooth, body of flagellomere 4 2.3–2.7× as long as wide, neck shorter than broad, longest setae as long as width of flagellomere. thorax. brown, setae pale. anterior pronotum with 3–8 setae. episternum 1 with 7–15 setae. scutum with long dorsocentrals, with some longer and shorter laterals, scutellum with 2 longer and some short setae. fig. 3. bradysia chlorocornea mohrig & menzel, 1992 (from finland). a. gonostylus, ventral view. b. part of hypopygium, ventral view. scales = 0.1 mm. www.boldsystems.org heller k., hippa h. & vilkamaa p., taxonomy of bradysia 7 wing. hyalinous. length 1.7–1.9 mm. width/length 0.40. veins distinct. r1/r 0.70–0.85. c/w 0.70– 0.80. r-m and bm subequal in length. bm non-setose, r-m non-setose or with 1 seta. legs. yellow. coxal setae pale. front tibial organ with pale vestiture forming a short row. front tibial spur slightly longer than tibial width. claws with teeth. abdomen. pale brown, setae pale. hypopygium (fig. 3a–b). brown, concolorous with abdomen. gonocoxa. narrow, longer than gonostylus, mesial margin with rather dense setosity. gonostylus. narrow, broadest at middle, narrowed towards apex, with mesial side weakly impressed at apical half; with dense apical setosity, without apical tooth, with 5–6 apical megasetae whip-lash in form. tegmen conical, apically roundish, with straight lateral sides, weakly sclerotized, with large area of aedeagal teeth, laterally straight. aedeagal apodeme short. remarks bradysia chlorocornea was described from three males from japan (mohrig et al. 1992). by having bi-coloured antennal flagellomeres and the tarsal claws with teeth, it belongs to the bradysia fungicola group of menzel & mohrig (2000). by its slender gonostylus with an apical group of slender, subequal megasetae it resembles b. affinis (zetterstedt, 1838), but it can be distinguished by having the megasetae more slender and longer. bradysia falciceps sp. nov. urn:lsid:zoobank.org:act:60fa7b62-2ffa-43fe-a5a2-5ac02cf29b13 fig. 4a–b diagnosis small-sized bradysia. brown. maxillary palpus with 3 segments, segment 1 with shallow sensory area. wing length 1.5–1.7 mm, veins indistinct. gonostylus narrow, mesially impressed, with apical tooth and 5–6 megasetae in one group. etymology the name is derived from the latin words falx (scythe) and -ceps (of the head), referring to the curved apical tooth of the gonostylus of the species. material examined holotype finland: ♂, lapponia inariensis, utsjoki, tundra, malaise trap, 14 aug. 2000, fatouros, mzh. paratypes finland: same data as holotype, 1 ♂, pkhh 3368; regio aboensis, turku, pomponrahka (grid 6719377:3240217), open bog, malaise trap, 1 aug.–28 sep. 2011, j. salmela, 1 ♂, mzh. canada: ontario, sudbury co., chapleau, superior forest, mature (99 years old) stand composed of 90% pinus banksiana and 10% p. mariana with an understory shrub layer of vaccinium, very wet, with considerable bryophyte coverage, photoeclector on pine log, 23 jun.–8 jul. 2013, r. deady & t. work, 3 ♂♂, prdm; same data but 7–22 jul. 2013, 1 ♂, mzh. http://zoobank.org/urn:lsid:zoobank.org:act:60fa7b62-2ffa-43fe-a5a2-5ac02cf29b13 european journal of taxonomy 122: 1–15 (2015) 8 description (male) head. brown, antennal flagellomeres 1–2 yellow, scapus, pedicellus and flagellomeres 3–14 unicolorous pale brown; maxillary palpus very pale brown. eye bridge 3 facets wide. face with 15–21 scattered longer and shorter setae. clypeus with 1–2 setae. maxillary palpus with 3 segments; segment 3 longer than segment 1, segment 2 shortest; segment 1 with 1–2 setae, with small shallow dorsal sensory area; surface of antennal flagellomeres smooth, body of flagellomere 4 1.85–2.3× as long as wide, neck shorter than broad, longest setae as long as width of flagellomere. thorax. brown, setae pale. anterior pronotum with 4–7 setae. episternum 1 with 8–9 setae. scutum with long dorsocentrals, with some longer and shorter laterals, scutellum with 4 longer and some short setae. wing. hyalinous. length 1.5–1.7 mm. width/length 0.45–0.50. veins indistinct. r1/r 0.60–0.65. c/w 0.75–0.85. r-m and bm subequal in length. r-m with 1–2 setae, bm non-setose. legs. yellow. coxal setae pale. front tibial organ with pale vestiture forming short row. front tibial spur longer than tibial width. claws without teeth. abdomen. pale brown, setae pale and fine. hypopygium (fig. 4a–b). brown, concolorous with abdomen. gonocoxa. narrow, longer than gonostylus, mesial margin with normal setosity. gonostylus. narrow, evenly straight, with shallow lateral notch at apical fourth, with mesial side impressed at all its length; with dense apical setosity, with strong curved apical tooth-like megaseta, fig. 4. bradysia falciceps sp. nov. (holotype). a. part of hypopygium, ventral view. b. gonostylus, ventral view. scales = 0.1 mm. heller k., hippa h. & vilkamaa p., taxonomy of bradysia 9 with 5–6 subapical megasetae, megasetae sharp, in single group. tegmen conical, weakly sclerotized, with area of aedeagal teeth. aedeagal apodeme long. remarks bradysia falciceps sp. nov. belongs to the bradysia praecox group and is most similar to b. nitidicollis (meigen, 1818), b. iridipennis (zetterstedt, 1938) and b. quercina menzel & köhler, 2014 by sharing a strong apical tooth at the dorsoapical part of the gonostylus (see köhler et al. 2014). bradysia falciceps sp. nov. can be distinguished from all other species of the b. praecox group in having pale basal antennal flagellomeres with dark necks. bradysia oelandica sp. nov. urn:lsid:zoobank.org:act:d1390d14-f74e-4595-a927-a287b8a331df fig. 5a–c diagnosis large-sized bradysia. brown. maxillary palpus with 3 segments, segment 1 with a small shallow sensory patch. wing length 2.3 mm, anal lobe small. setose intercoxal lobe of hypopopygium present. gonocoxa and gonostylus broad; gonocoxa with elongated ventromesial setae; gonostylus bulged mesiad, with a short apical tooth. etymology the name is derived from the latinized name of the type locality (the island of öland, sweden) of the species. material examined holotype sweden: ♂, öland, mörbylonga, gamla skogsby (kalkstad), 56°37.002’ n, 16°30.457’ e, meadow with bushes, malaise trap (trap id 22), 20 may–28 jun. 2006, swedish malaise trap project, smnh 2652. description (male) head. dark brown, antennal flagellomeres unicolorous, paler brown; maxillary palpus very pale brown. eye bridge 3 facets wide. face with 15 scattered longer and shorter setae. clypeus with 3 setae. maxillary palpus with 3 segments; segment 1 longer than segment 3, segment 2 shortest; segment 1 with 3 setae, with small dorsal patch of sensilla in shallow depression; surface of antennal flagellomeres rough, body of flagellomere 4 2.9× as long as wide, neck shorter than broad, longest setae as long as width of flagellomere. thorax. brown, setae dark. anterior pronotum with 7 setae. episternum 1 with 6 setae. scutum with long dorsocentrals, with some longer and shorter laterals, scutellum with short setae. wing. fumose. length 2.3 mm. width/length 0.45. anal lobe small. veins distinct. r1/r 0.75. c/w 0.70. r-m shorter than bm, both non-setose. legs. yellowish brown. coxal setae pale. front tibial organ with pale vestiture forming a row. front tibial spur longer than tibial width. claws without teeth. abdomen. setae dark and long. http://zoobank.org/urn:lsid:zoobank.org:act:d1390d14-f74e-4595-a927-a287b8a331df european journal of taxonomy 122: 1–15 (2015) 10 hypopygium (fig. 5a–c). brown, concolorous with abdomen. intercoxal area with lobe and possessing ca. 10 long setae. gonocoxa. broad, longer than gonostylus, ventromesial margin with elongated setae, dorsomesial margin with distinct lobes. gonostylus. globular, medially bulged mesiad, with dense apical setosity, with short narrow apical tooth, with apical (lateral) megaseta, with specialized subapical seta. tegmen conical, weakly sclerotized, with area of small aedeagal teeth. aedeagal apodeme short but strong. remarks bradysia oelandica sp. nov. belongs to the bradysia polonica group of menzel & mohrig (2000) in having a broad intercoxal lobe of the hypopygium and an indistinct apical tooth at the end of the gonostylus. the general structure of the gonostylus is fairly similar to the other species of that group, but b. oelandica sp. nov. is easy to distinguish by the elongated setae at the apical part of the gonocoxa. fig. 5. bradysia oelandica sp. nov. (holotype). a. gonostylus, ventral view. b. part of hypopygium, ventral view. c. medial parts of hypopygium, ventral view. scale bars: a = 0.1 mm, b–c = 0.3 mm. 1 = lobe at dorsal mesial margin of gonocoxa, 2 = tegmen, 3 = gonocoxal apodeme, 4 = aedeagal apodeme. heller k., hippa h. & vilkamaa p., taxonomy of bradysia 11 bradysia plusiospina sp. nov. urn:lsid:zoobank.org:act:9efeeac1-6a5a-49fa-881b-1aa96ca76c28 fig. 6a–b diagnosis medium-sized bradysia. brown. maxillary palpus with 3 segments, segment 1 with a dorsal patch of sensilla. wing length 1.8–1.9 mm, anal lobe small. gonocoxa and gonostylus narrow, gonostylus straight, mesially weakly impressed, without apical tooth, with 7 sharp megasetae in two groups. etymology the name is derived from the latinized greek word plousious (wealthy), and the latin word spina (spine), referring to the many gonostylar megasetae of the species. material examined holotype finland: ♂, ostrobothnia borealis, tervola, yrttijänkä (grid 7346827:340814), near a spring, malaise trap, 1–28 jun. 2004, j. salmela, mzh. paratype finland: same data as holotype, ♂, mzh. fig. 6. bradysia plusiospina sp. nov. (holotype). a. part of hypopygium, ventral view. b. gonostylus, ventral view. scales = 0.1 mm. http://zoobank.org/urn:lsid:zoobank.org:act:9efeeac1-6a5a-49fa-881b-1aa96ca76c28 european journal of taxonomy 122: 1–15 (2015) 12 description (male) head. brown, antennal flagellomeres unicolorous, paler brown, maxillary palpus very pale brown. eye bridge 3 facets wide. face with 31–40 scattered longer and shorter setae. clypeus with 1 seta. maxillary palpus with 3 segments; segment 1 longer than segment 3, segment 2 shortest; segment 1 with 4–6 setae, with dorsal patch of sensilla; surface of antennal flagellomeres smooth, body of flagellomere 4 3.1× as long as wide, neck shorter than broad, longest setae slightly longer than the width of flagellomere. thorax. brown, setae pale. anterior pronotum with 6–7 setae. episternum 1 with 8–9 setae. scutum with long dorsocentrals, with some longer and shorter laterals, scutellum with 2 longer and some short setae. wing. fumose. length 1.8–1.9 mm. width/length 0.40. anal lobe small. veins distinct. r1/r 0.80. c/w 0.60–0.70. r-m and bm subequal in length. r-m with 1 seta, bm non-setose. legs. yellow. coxal setae pale. front tibial organ with pale vestiture forming short row. front tibial spur longer than tibial width. claws without teeth. abdomen. pale brown, setae pale and short. hypopygium (fig. 6 a–b). brown, concolorous with abdomen. gonocoxa. narrow, longer than gonostylus, mesial margin with short setosity. gonostylus. narrow, rather straight, slightly narrowed and curved towards apex, with mesial side weakly impressed; with dense apical setosity, without apical tooth, with 7 megasetae, megasetae in apical and subapical groups, subequal in size, sharp. tegmen truncate, with straight lateral sides, weakly sclerotized, with area of aedeagal teeth. aedeagal apodeme short. remarks bradysia plusiospina sp. nov. resembles b. pseudocampestris mohrig, 1978 and b. angustata tuomikoski, 1960 in the general form of the hypopygium and gonostylus, as well as in having the anal lobe of the wing poorly developed. bradysia angustata differs from the others in having an apical tooth and slightly stronger megasetae in its gonostylus, and in having its wing even narrower than the others. bradysia plusiospina sp. nov. can be distinguished from b. pseudocampestris in having stronger megasetae, some megasetae in a more lateral position than in the latter species and in having a more setose basal part of its hypopygium (see mohrig 1978: figs 4, 7). bradysia plusiospina sp. nov. is placed in the bradysia fungicola group of menzel & mohrig (2000). bradysia quercina menzel & köhler, 2014 fig. 7a–c material examined paratypes sweden: småland, gränna, lönnemålen, 58°02.935’ n, 14°34.382’ e, next to old cellar in norway spruce forest, malaise trap (trap id 17), 10–24 sep. 2003, swedish malaise trap project, 3 ♂♂, no. 3184, 3185, smnh, 4103, mzh; småland, kalmar län, bäckebo, grytsjön nature reserve, old aspen forest in boulder terrain, malaise trap, 15 jun. 2006, swedish malaise trap project, 1 ♂, smnh 6781. switzerland: sihlwald near zürich, window trap, 5 jun. 1996, schiegg, 1 ♂, pkhh 1805. heller k., hippa h. & vilkamaa p., taxonomy of bradysia 13 remarks bradysia quercina menzel & köhler, 2014 was described as belonging to the bradysia praecox group on the basis of a number of specimens from norway, italy, sweden and switzerland (köhler et al. 2014), and its affinities were thoroughly discussed. to compare to the figures of the holotype provided in the original description, we present drawings of the hypopygium and gonostylus of a paratype from sweden (fig. 7). discussion bradysia is by far the most species-rich genus of sciaridae, including about 400 species worldwide, most in the palaearctic region. with the present new species included, about 70 species are known from both finland and sweden. in the nearctic region, only 20 species are known (mohrig et al. 2013). this is a strongly biased figure and our current knowledge of the nearctic fauna is still very incomplete. even less is known of bradysia from other biogeographical regions (see vilkamaa et al. 2012). the newly described species show that even in the areas for which the sciarid fauna is best known, new species can be discovered. with many of the species morphologically very similar to each other, the genus is probably the most challenging of all sciaridae regarding its taxonomy. the holarctic fauna has been divided into sixteen species groups (menzel & mohrig 2000), but these have been formed mainly based on general similarity. a recent phylogeny based on molecular characters challenges even the monophyly of the genus (shin et al. 2013). fig. 7. bradysia quercina menzel & köhler, 2014 (from sweden). a. part of hypopygium, ventral view. b. gonostylus, ventral view. c. apical part of gonostylus, ventral view. scales = 0.1 mm. european journal of taxonomy 122: 1–15 (2015) 14 acknowledgements we thank rob deady (montréal, canada), werner mohrig (poseritz, germany) and jukka salmela (rovaniemi, finland) for materials and björn rulik (bonn, germany) for doing the dna analyses. this work was supported by the swedish taxonomy initiative. rob deady kindly checked the english. references enderlein g. 1911. die phyletischen beziehungen der lycoriiden (sciariden) zu den fungivoriden (mycetophiliden) und itonididen (cecidomyiiden) und ihre systematische gliederung. archiv für naturgeschichte 77, suppl. 3: 116–201. http://www.biodiversitylibrary.org/item/52219#page/908/ mode/1up [accessed 21 april 2015] hippa h. & vilkamaa p. 1991. the genus prosciara frey (diptera, sciaridae). entomologica fennica 2: 113–155. hippa h. & vilkamaa p. 1994. the genus camptochaeta gen. n. (diptera, sciaridae). acta zoologica fennica 194: 1–85. hippa h., vilkamaa p. & heller k. 2010. review of the holarctic corynoptera winnertz, 1867, s. str. (diptera, sciaridae). zootaxa 2695: 1–197. juslén a., kuusinen m, muona j., siitonen j. & toivonen h. 2008. puutteellisesti tunnettujen ja uhanalaisten metsälajien tutkimusohjelma [research programme of deficiently known and threatened forest species] (putte). suomen ympäristö 1: 1–146. köhler a., menzel f., thunes k.h. & søli g.e.e. 2014. black fungus gnats (diptera: sciaridae) in oak canopies: description of bradysia quercina menzel & köhler spec. nov. and new records for norway. studia dipterologica 20: 325–331. menzel f. & mohrig w. 2000. revision der paläarktischen trauermücken (diptera: sciaridae). studia dipterologica, suppl. 6 (for 1999): 1–761. mohrig w. 1978. zur kenntnis flügelreduzierter dipteren der bodenstreu. ix. beitrag: gattungen corynoptera, bradysia und plastosciara (sciaridae). zoologischer anzeiger 201: 424–432. mohrig w., menzel f. & kozánek m. 1992. neue trauermücken (diptera, sciaridae) aus nord-korea und japan. dipterological research 3: 17–32. mohrig w., heller k., hippa h., vilkamaa p. & menzel f. 2013. revision of the black fungus gnats (diptera: sciaridae) of north america. studia dipterologica 19: 141–286. rudzinski h.-g. 1994. neue mitteilungen zur trauermückenfauna österreichs (diptera nematocera: sciaridae). entomofauna, zeitschrift für entomologie 15: 281–292. shin s., jung s., menzel f., heller k., lee h. & lee s. 2013. molecular phylogeny of black fungus gnats (diptera: sciaroidea: sciaridae) and the evolution of larval habitats. molecular phylogenetics and evolution 66: 833–846. http://dx.doi.org/10.1016/j.ympev.2012.11.008 tuomikoski r. 1960. zur kenntnis der sciariden finnlands. annales zoologici societatis zoologiae botanicae fennicae ‚vanamo‘ 21 (4): 1–164. vilkamaa p. & hippa h. 1996. review of the genus prosciara frey (diptera, sciaridae) in the indomalayan region. acta zoologica fennica 203: 1–57. vilkamaa p. & hippa h. 2006. corynoptera vagula tuomikoski and allied species (diptera: sciaridae). zootaxa 1150: 31–42. http://www.biodiversitylibrary.org/item/52219#page/908/mode/1up http://www.biodiversitylibrary.org/item/52219#page/908/mode/1up http://dx.doi.org/10.1016/j.ympev.2012.11.008 heller k., hippa h. & vilkamaa p., taxonomy of bradysia 15 vilkamaa p., hippa h. & mohrig w. 2012. the genus bradysia winnertz (diptera, sciaridae) in new caledonia, with the description of thirteen new species. zootaxa 3489: 25–44. vilkamaa p., hippa h. & heller k. 2013a. review of the genus camptochaeta hippa & vilkamaa (diptera, sciaridae), with the description of nine new species. zootaxa 3636: 476–488. http://dx.doi. org/10.11646/zootaxa.3636.3.6 vilkamaa p., hippa h. & heller k. 2013b. notes on the taxonomy of the holarctic corynoptera winnertz sensu lato (diptera, sciaridae), with description of six new species. zootaxa 3710: 322–332. http:// dx.doi.org/10.11646/zootaxa.3710.4.2 vilkamaa p., hippa h. & heller k. 2013c. taxonomy of the sciaridae (diptera) of northern europe: description of eight new species. studia dipterologica 20: 47–58. winnertz j. 1867. beitrag zu einer monographie der sciarinen. zoologisch-botanischen gesellschaft in wien, vienna. manuscript received: 6 february 2015 manuscript accepted: 25 march 2015 published on: 8 may 2015 topic editor: koen martens desk editor: danny eibye-jacobsen printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark. http://dx.doi.org/10.11646/zootaxa.3636.3.6 http://dx.doi.org/10.11646/zootaxa.3636.3.6 http://dx.doi.org/10.11646/zootaxa.3710.4.2 http://dx.doi.org/10.11646/zootaxa.3710.4.2 european journal of taxonomy 299: 1–2 issn 2118-9773 https://doi.org/10.5852/ejt.2017.299 www.europeanjournaloftaxonomy.eu 2017 · galkowski c. et al. this work is licensed under a creative commons attribution 3.0 license. d n a l i b r a r y o f l i f e c o r r i g e n d u m 1 the following corrections have been made to paper no. 290 (https://doi.org/10.5852/ejt.2017.290) redescription of proformica nasuta (nylander, 1856) (hymenoptera, formicidae) using an integrative approach – corrigendum christophe galkowski 1, claude lebas 2, philippe wegnez 3, alain lenoir 4 & rumsaïs blatrix 5,* 1,2,3,4,5 antarea (www.antarea.fr), association for the study and mapping of ants from metropolitan france. 1 104 route de mounic, 33160 saint-aubin-de-medoc, france. 2 2 impasse del ribas, 66680 canohès, france. 3 walbru (www.fourmiswalbru.com), belgian association for the inventory of ant species in wallonia and brussels, and rue de la grotte 23, 4651 herve, belgium. 4 irbi, institut de recherche sur la biologie de l’insecte, umr cnrs 7261, université françois rabelais, faculté des sciences, parc grandmont, 37200 tours, france. 5 cefe umr 5175, cnrs – université de montpellier – université paul valéry montpellier – ephe, 1919 route de mende, 34293 montpellier cedex 5, france. * corresponding author: rumsais.blatrix@cefe.cnrs.fr 1 e-mail: chris.gal@wanadoo.fr 2 e-mail: cllebas@free.fr 3 e-mail: wegnez.phil@gmail.com 4 e-mail: alain.lenoir@univ-tours.fr galkowski c., lebas c., wegnez p., lenoir a. & blatrix r. 2017. redescription of proformica nasuta (nylander, 1856) (hymenoptera, formicidae) using an integrative approach – corrigendum. european journal of taxonomy 299: 1–2. https://doi.org/10.5852/ejt.2017.299 in the original publication, fig. 1 mistakenly cited proformica pilinotum sp. nov., which is a nomen nudum. the intended name was proformica longipilosa sp. nov., which is reflected in the updated version of the figure provided below, which also includes a correction of the year of publication cited in the figure legend. https://doi.org/10.5852/ejt.2017.299 www.europeanjournaloftaxonomy.eu https://creativecommons.org/licenses/by/3.0/ https://doi.org/10.5852/ejt.2017.290 www.antarea.fr http://www.fourmiswalbru.com mailto:rumsais.blatrix%40cefe.cnrs.fr?subject= mailto:chris.gal%40wanadoo.fr?subject= mailto:cllebas%40free.fr?subject= mailto:wegnez.phil%40gmail.com?subject= mailto:alain.lenoir%40univ-tours.fr?subject= https://doi.org/10.5852/ejt.2017.299 european journal of taxonomy 299: 1–2 (2017) 2 published on: 14 march 2017 printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands. fig. 1. distribution of the 22 localities where proformica nest samples were collected. the star indicates the type locality of proformica nasuta (nylander, 1856) (i.e., beaucaire). 0 50 100 km species proformica nasuta proformica longipilosa sp. nov. altitude 0 500 m 500 1000 m 1000 1500 m > 1500 m italy france spain a new genus for the lesser moorhen gallinula angulata sundevall, 1850 (aves, rallidae) george sangster1, juan carlos garcía-r2 & steve a. trewick3 1 department of bioinformatics and genetics, swedish museum of natural history, p.o. box 50007, se–104 05 stockholm, sweden. corresponding author, email: g.sangster@planet.nl 2 hopkirk institute, massey university, private bag 11-222, palmerston north, new zealand. email: j.c.garciaramirez@massey.ac.nz 3 phoenix lab, ecology group, institute of agriculture and environment, massey university, private bag 11-222, palmerston north, new zealand. email: s.trewick@massey.ac.nz 1 urn:lsid:zoobank.org:author:18bb564b-ca65-40af-93aa-932a61843535 2 urn:lsid:zoobank.org:author:df883332-5661-4d3d-aaba-1e738e95885d 3 urn:lsid:zoobank.org:author:ef574ef2-0aa8-4554-8f6e-d75657912974 abstract. molecular phylogenetic analysis has demonstrated that the genus gallinula is not monophyletic and comprises four major lineages. a review of the nomenclature of gallinula shows that generic names are available for three lineages but that a fourth is as yet unnamed. a new monotypic genus, paragallinula gen. nov., is described for lesser moorhen (gallinula angulata sundevall, 1850). key words. gallinules, morphology, nomenclature, paragallinula gen. nov., phylogenetics. sangster g., garcía-r j.c. & trewick s.a. 2015. a new genus for the lesser moorhen gallinula angulata sundevall, 1850 (aves, rallidae). european journal of taxonomy 153: 1–8. http://dx.doi.org/10.5852/ejt.2015.153 introduction recent molecular phylogenetic analysis has clarified the relationships among rails and has shown that several currently recognized genera are not monophyletic (garcía-r et al. 2014a). the study included nuclear and mitochondrial sequence data representing 70 species of rails in 22 of the 33 extant genera, including seven species traditionally placed in gallinula brisson, 1760. gallinula as traditionally circumscribed (e.g., sibley & monroe 1990; taylor & van perlo 1998) was found to be paraphyletic with respect to fulica linnaeus, 1758 and porzana vieillot, 1816. spot-flanked gallinule gallinula melanops (vieillot, 1819 ) was sister to three (of eleven) species of porzana, and two further species of gallinula (g. mortierii (du bus, 1840) and g. ventralis gould, 1837 were found to be their sister group. surprisingly, the lesser moorhen gallinula angulata sundevall, 1850, which until now has always been placed in the genus gallinula, was resolved as sister to a clade comprising fulica linnaeus, 1758 and all other species of gallinula. these relationships are well-supported and suggest that taxonomic revision is warranted. european journal of taxonomy 153: 1–8 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2015.153 www.europeanjournaloftaxonomy.eu 2015 · sangster g. et al. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:cf2e388c-9bc5-4a16-a598-3074142c3eab 1 mailto:g.sangster%40planet.nl?subject=g.sangster%40planet.nl mailto:j.c.garciaramirez%40massey.ac.nz?subject=j.c.garciaramirez%40massey.ac.nz mailto:s.trewick%40massey.ac.nz?subject=s.trewick%40massey.ac.nz http://zoobank.org/urn:lsid:zoobank.org:author:18bb564b-ca65-40af-93aa-932a61843535 http://zoobank.org/urn:lsid:zoobank.org:author:df883332-5661-4d3d-aaba-1e738e95885d http://zoobank.org/urn:lsid:zoobank.org:author:ef574ef2-0aa8-4554-8f6e-d75657912974 http://dx.doi.org/10.5852/ejt.2015.153 http://dx.doi.org/10.5852/ejt.2015.153 http://www.europeanjournaloftaxonomy.eu/index.php/ejt https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:cf2e388c-9bc5-4a16-a598-3074142c3eab current knowledge of the phylogenetic relationships among coots and gallinules leaves two major options for the generic assignment of g. angulata. fulica and several species of gallinula including g. angulata could be combined in a single genus for which fulica has nomenclatural priority. alternatively, g. angulata could be removed from gallinula and placed in a separate genus. merging gallinula with fulica would result in the change of at least six species names (g. chloropus (linnaeus, 1758), g. galeata (m.h.c. lichtenstein, 1818), g. comeri allen, 1892, g. nesiotis p.l. sclater, 1861, g. tenebrosa gould, 1846 and g. angulata), and it would fail to recognise the widely accepted ecological, morphological and phylogenetic distinction between gallinules (gallinula as defined below) and coots. placement of g. angulata in a separate genus minimises taxonomic instability and maximises the information content of genus limits because it allows gallinula and fulica to be retained as separate genera. however, this strategy requires additional adjustments (some of which have already been adopted) to reconcile the systematic status of the gallinules, which are not a natural group. in this paper, we provide an updated molecular phylogeny of the “fulica” clade (sensu garcía-r et al. 2014a), including a member of the putative genus “porphyriornis” (gallinula nesiotis) which was not previously sampled (garcía-r et al. 2014a), and a description of morphological characters to propose a new genus for the lesser moorhen. materials and methods we conducted a phylogenetic analysis of the fulica-clade using the same sequences and methods as garcía-r et al. (2014a), but also including an available cytochrome b sequence of g. nesiotis obtained from genbank (ef681998) which is the only marker shared between the datasets of groenenberg et al. (2008) and garcía-r et al. (2014a). thus, we conducted bayesian and maximum likelihood analyses of ~2900 bp of mitochondrial dna (partial sequences of the mitochondrial genes cytochrome b, cytochrome oxidase subunit i and 16s ribosomal rna) and ~1900 bp of nuclear dna (fragments of beta-fibrinogen intron 7 and recombination activating gene 1). we rooted the tree using an outgroup comprising species within the “aramides” clade (garcía-r et al. 2014a). we review the nomenclatural status of all generic names applied to gallinula (sensu sibley & monroe 1990; taylor & van perlo 1998) to assess if any of these apply to g. angulata. the diagnosis of the new genus is based on the morphological dataset of livezey (1998), which represents the most comprehensive and detailed comparison of morphological variation in rails. livezey’s (1998) study included all extant species of fulica and gallinula plus a number of extinct species, including gallinula nesiotis and gallinula (pareudiastes) pacifica (hartlaub & finsch, 1871). results phylogenetics a phylogeny of the fulica-clade is shown in fig. 1. the topology is identical to that of garcía-r et al. (2014a), but with g. nesiotis added as the sister of g. gallinula and g. galeata. gallinula (sensu sibley & monroe 1990; taylor & van perlo 1998) was resolved as paraphyletic and comprised four major lineages: (1) g. chloropus, g. galeata, g. nesiotis and g. tenebrosa; (2) g. ventralis and g. mortierii; (3) g. melanops; and (4) g. angulata (fig. 1). nomenclature the genus gallinula brisson, 1760 has g. chloropus as its type species (lineage 1 in fig. 1). the synonymy of gallinula (sensu sibley & monroe 1990; taylor & van perlo 1998) includes at least 10 names, although some of these have been reinstated as valid genera in recent classifications. however, none of these names can be applied to g. angulata. european journal of taxonomy 153: 1–8 (2015) 2 hydrogallina lacépède, 1799 (type species fulica chloropus linnaeus, 1758, currently gallinula chloropus) and stagnicola brehm, 1831 (type species fulica chloropus linnaeus, 1758, currently gallinula chloropus) are junior synonyms of gallinula. tribonyx du bus, 1840 (type species tribonyx mortierii du bus, 1840) has been used for two australian species, tasmanian native-hen g. mortierii and black-tailed native-hen g. ventralis by some authors (e.g., christidis & boles 2008; dickinson & remsen 2013). recognition of tribonyx as a separate genus from gallinula is supported by phylogenetic studies of morphology (livezey 1998) and dna sequences (garcía-r et al. 2014a), which showed that g. mortierii and g. ventralis are not closely related to other species of gallinula (lineage 2 in fig. 1). neither study indicated a close relationship between these species and lesser moorhen, and the name tribonyx is thus not applicable to g. angulata. brachyptrallus lafresnaye, 1840 (type species brachyptrallus ralloides lafresnaye, 1840, currently tribonyx mortierii du bus, 1840) is a junior synonym of tribonyx and not applicable to g. angulata. porphyriops pucheran, 1845 (type species fulica crassirostris j.e. gray, 1829; now gallinula melanops crassirostris) has been used for the spot-flanked gallinule (e.g., wolters 1975; blake 1977; livezey 1998). its recognition as a separate genus from gallinula is now supported by phylogenetic studies of morphology (livezey 1998) and dna sequences (garcía-r et al. 2014a) (lineage 3 in fig. 1). again, neither study indicated a close relationship between spot-flanked gallinule and lesser moorhen. therefore, porphyriops is not applicable to g. angulata. pareudiastes hartlaub & finsch, 1871 (type species pareudiastes pacificus hartlaub & finsch, 1871) and edithornis mayr, 1933 (type species edithornis silvestris mayr, 1933) have been proposed as the generic names of samoan moorhen and san cristobal moorhen, respectively. these two species have fig. 1. bayesian analysis of the fulica-clade (sensu garcía-r et al. 2014a). numbers above nodes indicate posterior probabilities from bayesian analysis. numbers below nodes are percent bootstrap values from maximum likelihood analysis. colours indicate relevant clades with current nomenclature at branch tips. coloured genus names are those adopted here to reconcile phylogeny and known morphological diversity. sangster g. et al., a new genus for gallinula angulata 3 both been placed in gallinula in the past (e.g., sibley & monroe 1990; taylor & van perlo 1998) but they share a number of unique character states and are now typically combined in the genus pareudiastes (olson 1973, 1975a; livezey 1998; dickinson & remsen 2013). a phylogenetic analysis of morphological characters indicates that g. angulata is not closely related to pareudiastes (livezey 1998). in addition, g. angulata does not share any unique apomorphies with pareudiastes and differs from the latter by at least 14 integumentary characters (listed below). thus, the name pareudiastes is not applicable to g. angulata. porphyriornis allen, 1892 (type species porphyriornis comeri allen, 1892) is sometimes used as the generic name of gough moorhen g. comeri and tristan moorhen g. nesiotis (e.g., peters 1934; livezey 1998). however, because both species are closely related to g. chloropus, these are typically placed in gallinula (wolters 1975; sibley & monroe 1990; taylor & van perlo 1998; dickinson & remsen 2013), an arrangement supported by a molecular phylogenetic study (groenenberg et al. 2008; see also fig. 1). porphyriornis represents a synonym of gallinula. microtribonyx sharpe, 1893 (type species gallinula ventralis gould, 1837) is sometimes used for a monotypic genus comprising t. ventralis (wolters 1975), but this name is now generally regarded as a synonym of tribonyx or gallinula, depending on whether the former is recognised as a separate genus (with two species), as we advocate here. pyramida oliver, 1955 (type species rallus hodgeni scarlett, 1955, currently tribonyx hodgenorum) was proposed for an extinct flightless species, now known from both north and south island, new zealand. subsequently, scarlett (1970) placed this species in the genus capellirallus falla, 1954, whereas olson (1975b) transferred it to tribonyx, which he considered a subgenus of gallinula. in livezey’s (1998) phylogenetic analysis of morphological characters, this species was included within a polytomy comprising tribonyx, pareudiastes, edithornis, porphyriornis, gallinula and fulica. in any case, pyramida is not applicable to g. angulata. as no generic name appears to be available for g. angulata, we propose a new genus taxon. class aves linnaeus, 1758 order gruiformes (bonaparte, 1854) family rallidae (rafinesque, 1815) paragallinula gen. nov. urn:lsid:zoobank.org:act:909ac100-559e-4ca3-bc7e-e6fd87002e30 type species gallinula angulata sundevall, 1850. monotypic. differential diagnosis paragallinula gen. nov. differs from gallinula, pareudiastes, tribonyx and porphyriops in that (i) the orange suffusion on the frontal shield does not encompass the entire shield (in gallinula encompassing entire frontal shield (fig. 2); in pareudiastes, tribonyx and porphyriops no suffusion of orange). in addition, paragallinula gen. nov. differs from most species in the fulica-clade (except t. ventralis) in (ii) showing marked sexual plumage dimorphism (taylor 1996; taylor & van perlo 1998). paragallinula gen. nov. further differs from gallinula in (iii) the lack of a contrasting reddish band (“garter”) on the legs immediately proximal to the ankle joint (present in gallinula) (fig. 2). european journal of taxonomy 153: 1–8 (2015) 4 http://zoobank.org/urn:lsid:zoobank.org:act:909ac100-559e-4ca3-bc7e-e6fd87002e30 paragallinula gen. nov. differs from fulica in (ii) gradual change in colouration on hindneck from dark grey head to paler lower neck and back (abrupt change in fulica); (iii) absence of broad cutaneous lobes on digits ii–iv of feet (present in fulica); and (iv) presence of narrow cutaneous folds on lateral and medial sides of digits i–iv of feet (absent in fulica). paragallinula also differs from fulica in at least 12 diagnostic osteological characters (characters 120, 124, 125, 127, 139, 216, 274, 298, 311, 312, 316, 343 in livezey 1998, to which is referred for descriptions of characters and their states). paragallinula gen. nov. further differs from the two species of pareudiastes in (iii) frontal shield with acuminate posterior margin (squared margin in pareudiastes); (iv) ventroanterior position of nasal depression (ventromedial in pareudiastes); (v) fully feathered lores (in pareudiastes, lores with sparsely feathered, yellow-orange area from base of maxilla to orbit); (vi) gradual change in colouration on hindneck from dark grey head to paler lower neck and back (very dark without noticeable change in colour from head to back in pareudiastes); (vii) flank feathers with large white streaks (no streaks in pareudiastes); (viii) undertail-coverts white dorsolaterally, black ventromedially (entirely black or brownish-black in pareudiastes); (ix) six pairs of rectrices (four in pareudiastes); (x) rectrices forming normally-shaped tail (in pareudiastes, rectrices very short barely protruding beyond uppertail and undertail-coverts); (xi) leading edge of distalmost primary with narrow white margin (absent in pareudiastes); (xii) underwing-coverts with white bars (not barred in pareudiastes); (xiii) bend of wing whitish (no white colouration in pareudiastes); and (xiv) presence of narrow cutaneous folds on lateral and medial sides of digits i–iv of feet (absent in pareudiastes). paragallinula gen. nov. further differs from tribonyx in (ii) broad naked area on crown in downy plumage (absent in tribonyx); (iii) long, filamentous white filoplumes in downy plumage (absent in tribonyx); (iv) frontal shield with acuminate posterior margin (rounded in tribonyx); (v) undertail-coverts white dorsolaterally, black ventromedially (entirely blackish in tribonyx); (vi) six pairs of rectrices (eight in tribonyx); (vii) underwing-coverts dark grey, blackish or black (medium brown in tribonyx); and (viii) fig. 2. lesser moorhen paragallinula angulata sundevall, 1850, kgomo kgomo, south africa, feb. 2011 (photo by mark tittley). this photograph illustrates two diagnostic character states differentiating paragallinula from the genus gallinula: the orange colouration on the frontal shield does not cover the entire shield, and the lack of a contrasting reddish band on the legs proximal to the ankle joint. sangster g. et al., a new genus for gallinula angulata 5 bend of wing whitish (no white colouration in tribonyx). paragallinula also differs from tribonyx in a single osteological character: (ix) ossa digitorum pedis, digiti tertius, phalanx intermedialis (secundi) is equal to or longer than phalanx quartus (in tribonyx modally shorter than phalanx distalis (tertius). paragallinula gen. nov. further differs from porphyriops (p. melanops) in (iii) broad naked area on crown in downy plumage (absent in porphyriops); (iv) long, filamentous white filoplumes in downy plumage (absent in porphyriops); (v) ventroanterior position of nasal depression (ventromedial in porphyriops); (vi) feathers at base of bill, lores and crown not contrasting with rest of head (in porphyriops contrastingly black); (vii) flank feathers mostly plain but with large white streaks at tips (flank feathers brown-grey, spotted white in porphyriops); (viii) central belly dark slate-grey (whitish in porphyriops); (xi) undertailcoverts white dorsolaterally, black ventromedially (white in porphyriops); and (x) rectrices dark grey or black (brown in porphyriops). in addition, there are two diagnostic osteological differences between paragallinula and porphyriops: (xi) os frontale, facies dorsalis moderately convex (flat or slightly concave in porphyriops), and (xii) os squamosum, fossa temporalis distinct, separated medially from each other by significantly broad, elevated expanse of os frontale (in porphyriops, indistinct, shallow, limited to region immediately ventral to processus postorbitalis). etymology the generic name is derived from the greek para (beside) and the latin gallinula (a little hen or chicken). it denotes the resemblance of p. angulata to species of gallinula but highlights that they are independent evolutionary lineages. the gender of the name is feminine. distribution paragallinula angulata is found in most of the african continent from senegal and gambia to ethiopia, namibia, botswana and south africa. discussion in the past, the validity and systematic position of many monotypic genera of birds were problematic. this is because the genera were not based on phylogenetic studies but on subjectively inferred evolutionary distinctiveness which in turn was based on phenotypic distinctiveness and commonly their insular location. indeed, some taxonomists have placed large numbers of species in monotypic genera for such reasons, but the introduction of molecular phylogenetic methods in avian taxonomy revealed that many of these were nested within other genera (e.g., johnson et al. 2001; gibson & baker 2012; alström et al. 2015). the resulting gradual elimination of monotypic genera in many bird groups has enhanced understanding of avian diversification. conversely, several species have been shown to occupy a phylogenetic position inconsistent with their traditional generic assignment, resulting in the recognition of new monotypic genera (e.g., chesser et al. 2009; slager & klicka 2014). it is for this latter reason that, to re-circumscribe gallinula as a monophyletic group and so eliminate its current paraphyly, we erect a new monotypic genus paragallinula gen. nov. for g. angulata. this reflects the well-supported phylogenetic position of g. angulata outside the gallinula-fulica clade and avoids the inclusion of five species of gallinula in the distinctive genus fulica, and accords with pleas for ‘economy of nomenclatural change’ (vences et al. 2013). the contents of the genus gallinula have been gradually revised by the transfer of phylogenetically divergent species to other genera. several species of rails which are still referred as “gallinules” have sometimes been placed in the genus gallinula, including allen’s (porphyrio alleni thomson, 1842), purple (p. martinicus (linnaeus, 1766)) and azure (p. flavirostris (j.f. gmelin, 1789)) gallinules (e.g., ripley 1977; howard & moore 1991). molecular phylogenetic evidence accords with plumage features showing a closer relationship to porphyrio swamphens and far removed from gallinula moorhens european journal of taxonomy 153: 1–8 (2015) 6 (e.g., typical g. chloropus) (trewick 1997; garcía-r et al. 2014a). such discoveries demonstrate the effect of convergence in body-form and coloration of separate rail lineages on taxonomic inference. reconciliation using molecular data helps us understand how phenotypic traits evolve in response to ecological conditions, and in this case shows that the informal composition of gallinules is not a natural grouping. analysis of the timing and pattern of diversification within rallidae established on fossils and complete mitochondrial genomes is revealing (garcía-r et al. 2014a, 2014b). the node that separates fulica from the most closely related group of gallinula (including g. chloropus) dates to 18 mya (95% highest posterior density interval of 10–26 mya), whereas the mean age of the fulica group is under 10 mya. the position of g. angulata as the sister of fulica and gallinula indicates that the time of divergence between lesser moorhen, gallinula and fulica is similar to or exceeds that of several widely accepted genera of rails, including amaurolimnas sharpe, 1893 / aramides pucheran, 1845, neocrex sclater & salvin, 1869 / pardirallus bonaparte, 1856, amaurornis reichenbach, 1853 / gallicrex blyth, 1852 and aramidopsis sharpe, 1893 / lewinia g.r. gray, 1855 (garcía-r et al. 2014a, 2014b). this further supports that gallinula and fulica are best retained as separate genera but that lesser moorhen should be placed in a different genus. our treatment reconciles paraphyly of gallinula as follows: gallinula comprising g. chloropus, g. galeata, g. comeri, g. nesiotis and g. tenebrosa; tribonyx comprising t. ventralis and t. mortierii; porphyriops melanops; and paragallinula angulata. acknowledgements we are grateful to mark tittley for permission to use his excellent photograph of the lesser moorhen. two anonymous referees made helpful suggestions, which improved the quality of he manuscript. references alström p., jønsson k.a., fjeldså j., ödeen a., ericson p.g.p. & irestedt m. 2015. dramatic niche shifts and morphological change in two insular bird species. royal society open science 2 (3), 140364. http://dx.doi.org/10.1098/rsos.140364 blake e.r. 1977. manual of neotropical birds. vol. 1. university of chicago press, chicago. chesser r.t., claramunt s., derryberry e. & brumfield r.t. 2009. geocerthia, a new genus of terrestrial ovenbird (aves: passeriformes: furnariidae). zootaxa 2213: 64–68. christidis l. & boles w.e. 2008. systematics and taxonomy of australian birds. csiro publishing, collingwood, vic. dickinson e.c. & remsen j.v. jr 2013. the howard and moore complete checklist of the birds of the world. fourth edition, vol. 1: non-passerines. aves press, london. garcía-r j.c., gibb g.c. & trewick s.a. 2014a. deep global evolutionary radiation in birds: diversification and trait evolution in the cosmopolitan bird family rallidae. molecular phylogenetics and evolution 81: 96–108. http://dx.doi.org/10.1016/j.ympev.2014.09.008 garcía-r j.c., gibb g.c. & trewick s.a. 2014b. eocene diversification of crown group rails (aves: gruiformes: rallidae). plos one 9 (10): e109635. http://dx.doi.org/10.1371/journal.pone.0109635 gibson r. & baker a.j. 2012. multiple gene sequences resolve phylogenetic relationships in the shorebird suborder scolopaci (aves: charadriiformes). molecular phylogenetics and evolution 64: 66–72. http://dx.doi.org/10.1016/j.ympev.2012.03.008 groenenberg d.s.j., beintema a.j., dekker r.w.r.j. & gittenberger e. 2008. ancient dna elucidates the controversy about the flightless island hens (gallinula sp.) of tristan da cunha. plos one 3(3): e1835. http://dx.doi.org/10.1371/journal.pone.0001835 sangster g. et al., a new genus for gallinula angulata 7 http://dx.doi.org/10.1098/rsos.140364 http://dx.doi.org/10.1016/j.ympev.2014.09.008 http://dx.doi.org/10.1371/journal.pone.0109635 http://dx.doi.org/10.1016/j.ympev.2012.03.008 http://dx.doi.org/10.1371/journal.pone.0001835 howard r. & moore a. 1991. a complete checklist of the birds of the world. second edition. academic press, london. johnson k.p., de kort s., dinwoodey k., mateman a.c., ten cate c., lessells c.m. & clayton d.h. 2001. a molecular phylogeny of the dove genera streptopelia and columba. the auk 118: 874– 887. http://dx.doi.org/10.1642/0004-8038(2001)118[0874:ampotd]2.0.co;2 livezey b.c. 1998. a phylogenetic analysis of the gruiformes (aves) based on morphological characters, with an emphasis on the rails (rallidae). philosophical transactions of the royal society of london b 353: 2077–2151. http://dx.doi.org/10.1098/rstb.1998.0353 olson s.l. 1973. a classification of the rallidae. wilson bulletin 85: 381–416. olson s.l. 1975a. the south pacific gallinules of the genus pareudiastes. wilson bulletin 87: 1–5. olson s.l. 1975b. a review of the extinct rails of the new zealand region (aves: rallidae). national museum of new zealand records 1: 63–79. peters j.l. 1934. check-list of birds of the world. vol. 2. museum of comparative zoology, cambridge, massachusetts. ripley s.d. 1977. rails of the world. dodine, boston. scarlett r.j. 1970. the genus capellirallus. notornis 17: 303–319. sibley c.g. & monroe b.l. 1990. distribution and taxonomy of birds of the world. yale university press, new haven. slager d.l. & klicka j. 2014. a new genus for the american tree sparrow (aves: passeriformes: passerellidae). zootaxa 3821: 398–400. http://dx.doi.org/10.11646/zootaxa.3821.3.9 taylor p.b. 1996. family rallidae (rails, gallinules and coots). in: del hoyo j., elliot a. & sargatal j. (eds) handbook of the birds of the world. vol. 3. hoatzin to auks: 108–209. lynx edicions, barcelona. taylor p.b. & van perlo b. 1998. rails, a guide to the rails, crakes, gallinules and coots of the world. pica press, sussex. trewick s.a. 1997. flightlessness and phylogeny amongst endemic rails (aves: rallidae) of the new zealand region. philosophical transactions of the royal society of london b 352: 429–446. vences m., guayasamin j.m., miralles a. & de la riva i. 2013. to name or not to name: criteria to promote economy of change in linnaean classification schemes. zootaxa 3636: 201–244. http://dx.doi. org/10.11646/zootaxa.3636.2.1 wolters h.e. 1975. die vogelarten der erde. eine systematische liste mit verbreitungsangaben sowie deutschen und englischen namen. part 1: 1–80. paul parey, hamburg. manuscript received: 3 september 2015 manuscript accepted: 8 october 2015 published on: 16 november 2015 topic editor: rudy jocqué desk editor: kristiaan hoedemakers printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark. european journal of taxonomy 153: 1–8 (2015) 8 http://dx.doi.org/10.1642/0004-8038(2001)118[0874:ampotd]2.0.co;2 http://dx.doi.org/10.1098/rstb.1998.0353 http://dx.doi.org/10.11646/zootaxa.3821.3.9 http://dx.doi.org/10.11646/zootaxa.3636.2.1 http://dx.doi.org/10.11646/zootaxa.3636.2.1 european journal of taxonomy 69: 1-23 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2013.69 www.europeanjournaloftaxonomy.eu 2013 · martin cheek & matthew jebb this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e 1 recircumscription of the nepenthes alata group (caryophyllales: nepenthaceae), in the philippines, with four new species martin cheek1 & matthew jebb2 1 herbarium, royal botanic gardens, kew, richmond, surrey, tw9 3ae, u.k. email: m.cheek@kew.org (corresponding author) 2 national botanic garden, glasnevin, dublin 9, ireland email: matthew.jebb@opw.ie abstract. an overview of nepenthes in the philippines is presented. four new species, nepenthes extincta sp. nov., n. kitanglad sp. nov., n. kurata sp. nov. and n. leyte sp. nov. are described and illustrated from the philippines and placed in the nepenthes alata group. an updated circumscription and key to the species of the group is provided. delimitation and comparison with the regiae group is given. all four of the newly described species are assessed as threatened using the international union for the conservation of nature 2012 standard, and one, n. extincta sp. nov. is considered likely to be already extinct due to open-cast mining. logging and conversion of forest habitat are thought to be the main threats to the other three species. key words. conservation, nepenthes alata group, mindanao, threatened, ultramafic. cheek m. & jebb m. 2013. recircumscription of the nepenthes alata group (caryophyllales: nepenthaceae), in the philippines, with four new species. european journal of taxonomy 69: 1-23. http://dx.doi.org/10.5852/ ejt.2013.69 introduction this paper forms part of studies towards a world monograph of nepenthes l. (linnaeus 1753) (nepenthaceae), building on a skeletal revision of nepenthes (jebb & cheek 1997) and the flora malesiana account (cheek & jebb 2001). nepenthes is the sole genus in its family. the species mainly occur in malesia, but with ca. 12 outlying species in madagascar, seychelles, sri lanka, ne india, indochina, solomon islands, new caledonia and australia. cheek & jebb (2001) recorded 87 species of nepenthes, but since 1st jan. 2001, 50 new specific names have been published in the genus (international plant names index continuously updated) and others have been resurrected from synonymy. the total number of species now accepted in the genus is estimated as140 and is set to rise further. within the philippines 12 species were recognised in cheek & jebb (2001). since 1st jan. 2001, 16 new names have been published at species-level for the country (international plant names index continuously updated). these derive mainly from exploration of previously unexplored areas by nepenthes enthusiasts (e.g. heinrich et al. 2009; cheek 2011) and are also due to the discovery of http://dx.doi.org/10.5852/ejt.2013.69 http://www.europeanjournaloftaxonomy.eu http://creativecommons.org/licenses/by/3.0/ http://www.europeanjournaloftaxonomy.eu mailto:matthew.jebb@opw.ie http://dx.doi.org/10.5852/ejt.2013.69 http://dx.doi.org/10.5852/ejt.2013.69 european journal of taxonomy 69: 1-23 (2013) 2 previously overlooked specimens in herbaria around the world (e.g. cheek & jebb 2013a, b, c, d). some of the latter, e.g. nepenthes alzapan jebb & cheek (cheek & jebb 2013b) are considered possibly already extinct, since habitat destruction in the philippines over the last 100 years has been so extensive. the philippines is thought to have remained two-thirds forested as recently as 1925 according to sohmer & davis (2007). myers et al. (2000) estimated that remaining primary vegetation in the philippines 75 years later amounted to only 3%. lowland primary forest (occurring below 500 m) has now all but gone from the philippines (sohmer & davis 2007). sohmer & davis (2007) estimate species extinction levels due to habitat destruction as between 9 and 28% in one representative, mainly forest genus, psychotria l. (linnaeus 1759) (rubiaceae) which they revised taxonomically for the philippines. since habitat destruction continues in the philippines, it is a race against time to discover, publish, assess and draw attention to the conservation needs of species before they become extinct, if they have not already been lost. of the species of nepenthes known for the philippines, about half occur in scrub or stunted habitats on ultramafic substrates (both lowland and submontane) and appear to be restricted to this substrate, while the remainder occur mainly in submontane forest (500–2000 m) on non-ultramafic, mainly volcanic substrates. species diversity is highest on mindanao island, the southernmost and second-largest island, followed by palawan island. all known philippine nepenthes species are endemic, apart from n. mirabilis lour. (druce) (druce 1916; basionym: loureiro 1790) which is known from tawi tawi island close to borneo, and from the northern coast of mindanao and its associated islands. phytogeographically, the nepenthes of palawan have closer affinities with borneo than with the rest of the philippines (cheek & jebb 1999). most of the endemic species can be assigned to one of three main groups: (1) the nepenthes villosa hook.f. (hooker 1851) group, with petiolate leaves, well-developed lid columns, and peristomes with blade-like teeth, and lids which lack a basal appendage. these are restricted to higher altitude ultramafic habitats and are most diverse in palawan, but extend to ne borneo (cheek & jebb 1999: 894). (2) the insignes or n. ventricosa blanco (blanco 1837) group which is most diverse in the philippines (but absent from palawan), extends to new guinea and to borneo and is characterised in cheek & jebb (2013b). in the philippines they occur in both ultramafic and non-ultramafic, mainly submontane forest habitats. its species lack a petiole, have broadly subcylindrical pitchers with a broad peristome which is conspicuously toothed on its inner surface, lids which lack appendages, and which have transversely elliptic nectar glands except along the midline band. (3) the nepenthes alata blanco (blanco 1837) group, characterised in cheek & jebb (2013d), is endemic to and extends throughout the philippines except palawan, is most diverse in mindanao, and occurs in the same range of habitats as the insignes. its species have winged petioles, lids with basal ridges, usually producing appendages and upper pitchers usually broadest at the base. outside of these three groups, three other species occur, namely n. philippinensis macfarl. (macfarlane 1908), n. abalata jebb & cheek (cheek & jebb 2013a: 153) and n. micramphora v.b.heinrich, s.mcpherson, gronem. & v.b.amoroso (heinrich et al. 2009). n. philippinensis is endemic to palawan, and part of the n. hirsuta hook.f. (hooker 1873) group otherwise restricted to borneo (cheek & jebb 1999: 890–891). the unusual n. abalata occurs only in the small low-lying, drier islands between palawan and panay islands. it appears to be restricted to grassland and is similar and possibly related to the grassland species of indo-china. finally, n. micramphora of the extreme se of mindanao fits none of these groups and has an isolated position. cheek m. & jebb m., nepenthes (caryophyllales: nepenthaceae) in the philippines 3 many philippine species of nepenthes are only known from a single, sometimes sterile, specimen. however, usually there is little doubt that they are distinct taxa, since most characters of use in delimiting species in the genus are found in the pitchers and not in the flowers or fruit. in nepenthes the large number of taxonomic characters in the pitcher morphology and micro-morphology, especially in the upper and intermediate pitchers, parallels the importance of characters present in the flowers and inflorescences of many other plant groups. in many other plant groups evolution of character diversity in floral characters appears to be correlated with assisting pollination, often by a single or narrow range of insect species. in nepenthes, cross-pollination appears from initial studies to be performed by a broad range of generalist flies and wasps (lian 1995; adam 1998). perhaps for this reason there is comparatively little specieslevel differentiation florally in the family. in contrast, evidence has been steadily accumulating in recent years that the pitchers of different nepenthes species are specialised at gathering nutrients from one of a variety of animal sources. they are not always pitfall trappers of any prey item that happens to fall in, as had previously been assumed. nepenthes species studied by jebb (1991) in new guinea predominantly trapped ants, but also a spectrum of other insects, while nepenthes albomarginata t.lobb ex lindley (lindley 1849) traps almost exclusively a species of termite (clarke 1997). nepenthes hemsleyana macfarl. (macfarlane 1908), the pitchers of which, providing day roosts for a species of bat, benefit from the resulting faecal deposits produced (grafe et al. 2011), while a group of nepenthes species in montane borneo are adapted to attract, detain and collect the faeces of a species of tree shrew and a rat (greenwood et al. 2011). in each of these cases modification in the morphology and micro-morphology of the pitchers appears to have been critical to collect nutrients from the target animals. no doubt in future, morphological and micromorphological character states seen in the pitchers of other nepenthes species or species groups, will also be shown to be important to targeting hitherto unsuspected specialised animal nutrient sources. in the last three cases, the morphological modifications of the pitcher each appear to target only one or two species of animal. however, it is possible that future studies will show that some or most species of nepenthes trap a wider range of insect species available in their environment. since the range of e.g. arthropod species can be expected to vary from one mountain (or other geographic location) to another, it is possible that speciation in nepenthes might be driven by modifications of the pitcher caused by selection for efficiency in trapping, or otherwise targeting, the spectra of animal species present at each. this might explain the common pattern seen in several nepenthes groups to have different species, as recognised by pitcher morphology and micromorphology, on different mountains or islands. an alternative hypothesis to explain this speciation pattern invokes genetic drift (stace 1991). in this hypothesis, the progeny of the few founding individuals at a new location might accumulate morphological changes over time independent of any selection pressures, for example as postulated for podostemaceae rich. ex kunth (kunth 1816) species at the same waterfall by van steenis (1981), exemplifying his theory of autonomous evolution. this scenario assumes that animal targeting is not strongly selected for in nepenthes. this might be the case if the threshold for the need for nutrients derived from animals in nepenthes is very low, or if there is no threshold at all, for example if nepenthes can survive without capturing animal nutrients through their pitchers. materials and methods the data in this study, one of a series, derives from herbarium specimens including their corresponding field notes, and if available, photographs. the location of the herbarium at which the specimens are located is indicated by the citation of the international code registered in index herbariorum (thiers continuously updated). european journal of taxonomy 69: 1-23 (2013) 4 the shape and dimensions of the upper and intermediate pitchers, including those of the body, mouth and lid of the pitcher, and of the fringed wings if present, are useful characters. the shape and features of the lower surface of the pitcher lid, and the shape of the pitcher mouth and its peristome are the largest source of such characters, as are the nectar glands on the lower surface of the lid (cheek & jebb 2001). the nectar gland size, shape, orientation, position, density and the absence or, if present, the extent of a nectar gland perimeter wall, can be especially useful in delimiting species. in some species nectar glands are dimorphic, and in a very few, trimorphic. indumentum is also frequently an important source of species characters. here the classification used in cheek & jebb (2001: 7-8) is maintained. since the mouth and the lid surface are often obscured or not presented in mounted herbarium specimens of nepenthes, it is frequently necessary, after prior permission from the herbarium manager of the specimens concerned, to soak the mouth area and lid, and sometimes entire pitchers, in warm soapy water so that they can be repositioned for study of the mouth and lower surface of the lid. by this means it is possible to view the three-dimensional shape of the pitcher, if this is not already known from photographs or observation of live material. measurements are made of stem, leaf and pitcher dimensions using a graduated ruler. the full range of measurements is given in the description, with extreme dimensions, disconnected with the normal range, indicated in brackets, e.g. “leaf-blade 12.5–22(–28) cm long”, indicating that a single leaf-blade was measured as 28 cm long, all others being in the 12.5–22 cm long range. observations and measurements of hair dimensions and of peristome and lower lid details are made with a leica wild m8 binocular microscope fitted with an eyepiece graticule graduated in units of 0.025 mm at x40 magnification. the drawings were prepared with the aid of a camera lucida fitted to the microscope. the four species described as new in this paper, n. extincta sp. nov., n. kitanglad sp. nov., n. kurata sp. nov. and n. leyte sp. nov. came to light when examining loans of previously unstudied specimens from e, bish and brit, which were compared with material from b, bm, bo, f, gh, k, kep, l, ndc, ny, ph, s, w, wag, u, uc, us, on which earlier studies have been published (jebb & cheek 1997; cheek & jebb 1999, 2001; cheek 2011; cheek & jebb 2013a, b, c, d). the material referred to matched no other species description, as represented by the both the specimens from the herbaria cited, and by protologues and other literature regarding the species for which material is only available in the philippines and not available for loan. the material best fitted the characters of nepenthes group (3) described above, that is the nepenthes alata group. however, unusual features for this group were present, which are considered in the discussion at the end of this paper. a key to the species of the group is provided below. the rapid increase in the number of new species from mindanao in recent years is likely to continue as specimens become available from new areas. many of the species of the n. alata group on this island seem narrowly geographically localised, although others, such as n. graciliflora are widespread. results key to the species of the nepenthes alata group updated from cheek & jebb (2013b) 1. lower surface of lid, including appendage (if present), densely and evenly covered in uniformly minute circular nectar glands (0.15–0.2 mm diam.) ..............................…….................……….. 2 – lower surface of lid with nectar glands either absent from the appendage (if present) and/or, sparse, large or dimorphic (larger glands 0.35–0.4 mm diam. or larger) ...................................……….….. 3 2. stems glabrous to glabrescent; upper pitchers lacking fringed wings in upper part; outer surface lacking stellate hairs entirely. s luzon to mindanao ........ n. graciliflora elmer (elmer 1912) cheek m. & jebb m., nepenthes (caryophyllales: nepenthaceae) in the philippines 5 – stems persistently pubescent; upper pitchers with fringed wings in upper part; outer pitcher surface > 50% covered in grey stellate hairs. n luzon ................................ n. alata blanco (blanco 1837) 3. stem internodes winged from the decurrent petiole bases. mindanao, sarangani prov. ..... ....................................................................................... n. saranganiensis sh.kurata (kurata 2003) – stem terete or angular, lacking wings .............................................................................................. 4 4 upper pitchers funnel-shaped or funneliform-cylindric ................................................................... 5 – upper pitchers not funnel-shaped, but subcylindric, widest at base, or equally at base and apex .... 6 5. stems and abaxial surface of midrib moderately densely covered in white appressed hairs 0.5– 1.5 mm long; lid of upper pitchers ovate, longer than broad. volcanic substrate. mindanao, mts apo & matutum ....................................................… n. copelandii macfarl.(macfarlane 1908) – stems and leaf-blades glabrous; lid of upper pitcher broader than long. ultramafic substrate. mindanao, mt kiamo ............................... n. ceciliae gronem. et al. (gronemeyer et al. 2012) 6 petiole appearing flat, at least distally (wings held flat) ................................................................... 7 – petiole appearing cylindrical (wings incurved) ............................................................................... 12 7. stems and lower surface of midrib conspicuously densely pubescent; lid appendage well-developed, hooked .............................................................................................................................................. 8 – stems and inner surface of midrib glabrous or inconspicuously and sparsely pubescent; lid appendage moderately or well-developed, never hooked .................................................................................. 9 8. upper pitchers widest at base, contracting slightly above into a narrower cylinder with fringed wings. negros & biliran isl. …………...…… n. negros jebb & cheek (cheek & jebb 2013d) – upper pitchers equally wide at base and apex, contracting slightly at the middle, lacking fringed wings. mindanao, surigao prov. ..............… n. ramos jebb & cheek (cheek & jebb 2013c) 9. lid apex with pocket. mindanao, s cotabato ...................... n. tboli jebb & cheek sp. ined. – lid apex lacking pocket or any appendage .................................................................................... 10 10. petiole canaliculate proximally (flat distally); blade hairy on upper surface; upper pitchers stout, length: breadth ratio 2–2.5:1; fringed wings absent. mindanao, mt hamiguitan .................. ............................................................ n. hamiguitanensis gronem. et al. (gronemeyer et al. 2010) – petiole flat proximally; blade glabrous on upper surface; upper pitchers slender, length: breadth ratio > 3:1; fringed wings present below peristome ……................…………………………………… 11 11. upper pitcher with lid about half as long as mouth; mouth not concave but flat; column absent; lid base truncate. mindanao, mt malindang …….........…… n. kurata jebb & cheek sp. nov. – upper pitcher with lid about as long as mouth; mouth highly concave; column present; lid base cordate. mindanao, kitanglad mts …….......……. n. kitanglad jebb & cheek sp. nov. 12. largest pitchers robust, 18–24 cm long; peristome 7–8 mm wide, curved in section but not cylindric; inner edge of peristome with small teeth visible, outer edge lobed. mindanao ………..……....13 – largest pitchers 12 cm long; peristome 2–3 mm wide, narrowly cylindrical, inner edge lacking visible teeth (unless dissected), outer edge not lobed. leyte isl. ..... n. leyte jebb & cheek sp. nov. european journal of taxonomy 69: 1-23 (2013) 6 13. largest pitchers without fringed wings; leaf midrib densely, minutely white stellate-hairy; lid base deeply cordate. mindanao, red mt ….........................…… n. extincta jebb & cheek sp. nov. – largest pitchers with fringed wings; leaf midrib with brown black bristle-like hairs 1–1.5 mm long; lid base truncate. ne mindanao ……..............…… n. mindanaoensis sh.kurata (kurata 2001) taxonomic descriptions order caryophyllales juss. ex bercht. & j.presl. sens. lat. (berchtold & presl 1820) family nepenthaceae dumort. (dumortier 1829) genus nepenthes l. (linnaeus 1753) nepenthes kurata jebb & cheek sp. nov. urn:lsid:ipni.org:names:77134486-1 fig. 1 diagnosis differs from n. mindanaoensis sh.kurata in the petiole wings patent (not involute), the hairs of stem, midrib and leaf-edge bushy, 0.1 mm long, not bristle-like 1–1.5 mm long; the lid about half as long as the mouth, lid base rounded or truncate (not about as long as the mouth, base cordate). etymology named as a noun in apposition for shigeo kurata, whose book on the nepenthes of mount kinabalu (kurata 1976) inspired interest in the genus among its many readers, and whose descriptions of nepenthes are models of detail, precision and clarity. type philippines. mindanao, “prov. of misamis, mount malindang”. may 1906, mearns & hutchinson in forest bureau 4632 (holotype k!; isotype ph!). fig 1. synonym nepenthes alata blanco var. ecristata macfarlane, nepenthaceae. in: engler a. (ed.) das pflanzenreich heft 36, 4, 3: 72 (1908). – type: lectotype, designated here: philippines, mindanao, “prov. of misamis, mount malindang”, may 1906, mearns & hutchinson in forest bureau 4632 (lecto-: k!; isolecto-: ph!). description terrestrial shrub-climber, height unknown. climbing stems terete to slightly angular, 4–6 mm diam.; internodes 30–50 mm long; axillary buds not evident; indumentum inconspicuous, persistent to the fifth internode from the apex, hairs translucent brown, simple or 2–3-armed from the base, hairs straight, variously angled from the horizontal, ca. 0.1 mm long, covering ca. 5% of the surface except the axils (100% coverage) surface brown-black, matt. leaves of rosette shoots thinly coriaceous, blade narrowly elliptic, 8–9 × 2–2.5 cm; apex and base acute; longitudinal nerves 1–2 pairs, within 2 mm of the margin, moderately conspicuous on both surfaces; pennate nerves at 90° from the midrib, numerous and moderately conspicuous; upper surface drying glossy pale brown, lower surface matt, mid-brown. leaves of climbing stems as the rosette leaves, but blades suboblong or oblong-lanceolate 10–12.5 × 3.2–3.8 cm; apex obtuse or acute; base obtuse; lower surface with sessile red glands ca. 0.5 mm diam.; midrib 40–60% covered in patent, brown, simple or basally bifurcate-trifurcate hairs 0.1–0.3(–0.5) mm long; margin fringed, in young leaves, with hairs 0.25 mm long, pale-brown, 1–4-armed from the base. petiole winged-canaliculate, 4–5 × 0.7 cm, wings patent; base clasping the stem for ⅓ to ½ its circumference, sometimes decurrent as an obtuse ridge to the node below. lower http://www.ipni.org/urn:lsid:ipni.org:names:77134486-1 cheek m. & jebb m., nepenthes (caryophyllales: nepenthaceae) in the philippines 7 fig. 1. nepenthes kurata sp. nov. a. habit, climbing stem with upper pitcher. b. indumentum of midrib, lower surface of leaf-blade. c. indumentum of tendril. d. indumentum of outer pitcher surface. e. lid, lower surface, showing nectar gland distribution on right (upper pitcher). f–g. profiles of basal lid ridges, without appendage (f), with appendage weakly developed (g), and moderately developed (h). i spur of intermediate pitcher. j. longitudinally elliptic nectar glands of lid midline. k. orbicular nectar glands of lid, outside the midline. l. peristome from above, short teeth and holes discernible. m. peristome viewed from inside pitcher. n. peristome transverse section, outer surface to right. a–e, j & k from gaerlan et al. ppi 10911; f–i, l–n from mearns & hutchinson 4632. scale bars: single = 1 mm; graduated single = 2 mm; double = 1 cm; graduated double = 5 cm. all drawn by andrew brown. european journal of taxonomy 69: 1-23 (2013) 8 pitchers unknown. intermediate pitchers (tendrils uncoiled: mearns & hutchinson 4632) 12.5–17.2 cm long, ellipsoid in the basal third to half, 4–5.7 cm wide, constricted, more or less abruptly, 5–7.5 cm from the base into the subcylindrical upper part, 2.1–3 cm diam. dilating slightly towards the apex 3–4 cm diam.; outer surface strongly reticulated with raised nerves when dry, 2–5% covered in hairs of two types (fig. 1d), (1) large erect hairs 0.3–0.75 mm long, with a single, major, curved arm, and 1–2 much smaller erect arms, and (2) minute, 3–6-armed stellate hairs 0.05–0.1 mm diam., which are more frequent, (ca. 4 per mm2); surface covered throughout (6–10 per mm2) with sessile, depressedglobose glands 0.1–0.2 mm diam.; fringed wings reduced to ridges except in the ca. 25 mm below the peristome, widening to 3 mm broad, with fringed elements 2.5 mm long, 2–5 mm apart; mouth oblique, suborbicular, ovate, 3–4.8 × 2.7–4.5 cm; apex with a column 9–10 mm long; peristome rounded to slightly flattened, 2–2.5 mm wide, more or less even in width, ribs 0.25–0.5 mm apart, conspicuous, about 0.1 mm high, outer edge lacking lobes, inner edge with very short teeth and conspicuous holes, teeth < 0.1 mm long. lid much smaller than the mouth, ovate, or broadly ovate, 25–35 × 25–30 mm, apex rounded to obtuse, base rounded to truncate; lower surface with a low basal ridge ca. 1 mm high, 7–10 mm long, either lacking a protruding appendage entirely (fig. 1f) or with a modestly developed appendage 1–2 mm high (fig. 1h); nectar glands only slightly dimorphic, (1) midline nectar glands sparse, longitudinally elliptic, 0.5–0.7 × 0.1–0.25 mm, with a thin marginal rim (fig. 1j), (2) outside the midline nectar glands circular (fig. 1k), sparse, < 1 per mm2, only 35–50 on each side of the midline, the largest scattered in the distal half, 0.5 mm diam., grading down to those of the marginal equatorial areas ca. 0.25 mm diam., and those at the attachment point with the peristome and the basal ridge and appendage, 0.15 mm diam.; sessile depressed-globose minute red glands 0.1–0.2 mm diam. are scattered over the surface at a density of 3–8 glands per mm2; minute inconspicuous stellate hairs ca. 0.075 mm diam. occur in an uneven, 0.5–1 mm wide band, near the margin widening to 1.5 mm wide at the lid apex. spur unbranched, curving downwards, stout at base and tapering to a slender apex, ca. 5 mm long, with scattered long, subpatent hairs 0.3–0.7 mm long (fig. 1i). upper pitchers (tendril coiled, gaerlan et al. in ppi 10914) resembling the intermediate pitchers, but fringed wings 1–2 mm wide, fringed elements 2.5 mm long, (2–)4–5 mm apart, dilating to 4.5 cm below the mouth; pitcher green, peristome maroon. lid broadly ovate to suborbicular 32 × 35 mm, lower surface with a basal ridge 9–10 mm long, ca. 2 mm high, bearing a central, symmetrical, protruding appendage 2 × 3 mm; nectar glands denser, ca. 110 on each side of the midline. male and female inflorescences and infructescences unknown. additional material philippines. mindanao, prov. misamis occidental, s.e. slopes of mt. malindang, lake duminagat, may 1993, gaerlan, sagcal & romero in ppi 10911 (brit!). distribution, habitat & phenology philippines, mindanao; evergreen forest, volcanic substrates. elevation: ca. 1400 m. conservation status nepenthes kurata sp. nov. is here assessed as critically endangered under criterion d of iucn (2012) since currently only two individuals, probably at a single location (as currently defined by iucn) are known. this site, the ca. 6 ha crater lake duminagat, is within the ca. 50,000 ha mt malindang range natural park of which at least 20,000 ha has been cleared for cultivation purposes, but which is a tentative world heritage site (http://whc.unesco.org/en/tentativelists/5029/, downloaded 16 july 2013). in 2012 the park was designated as an asean heritage park (http://news.pia.gov.ph/index. php?article=1451343449808, downloaded 16 july 2013). it is to be hoped that further investigation will discover additional individuals and locations for this species, decreasing its threat status, and increasing the likelihood that it can be protected. http://whc.unesco.org/en/tentativelists/5029/ http://news.pia.gov.ph/index.php?article=1451343449808 http://news.pia.gov.ph/index.php?article=1451343449808 cheek m. & jebb m., nepenthes (caryophyllales: nepenthaceae) in the philippines 9 since the terrain of mt malindang is reported as being rugged, with much forest surviving, there is every reason to hope that the species survives there, unlike nepenthes robcantleyi cheek (cheek 2011) also from mindanao, which is already suspected to be extinct in the wild due to the almost total clearance of forest habitat at the single known wild location due to logging (cheek 2011). remarks the first nepenthes taxa described from mindanao, both of the n. alata group, (cheek & jebb 2013d), were n. alata var. ecristata macfarl. (macfarlane 1908), based on mearns & hutchinson 4632 from mt malindang, and n. copelandii macfarl. (macfarlane 1908) from mt apo. the first of these we here elevate to species level as n. kurata jebb & cheek sp. nov. previously we had considered this taxon to be synonymous with n. mindanaoensis sh.kurata (kurata 2001) (cheek & jebb 2013d). the two taxa do have similarities in the overall shape of the upper pitchers, the weakly to moderately developed basal lid appendage and the sparse nectar glands of the lower lid surface. however they can be distinguished using the characters in table 1. the number and extent of these features merit elevation from varietal to specific-level recognition in our opinion. although the type specimen has rosette stems and intermediate pitchers only, a second specimen, with climbing stems and upper pitchers, gaerlan et al. in ppi 10911 came to light recently. it is from the type locality and matches the type in essential details. nepenthes kurata sp. nov. has the spot character within the nepenthes alata group of a small, more or less orbicular lid, only about half the length of the pitcher mouth. macfarlane (1908) characterised his n. alata var. ecristata by the lid appendage being either reduced or absent; the nectar glands being few, medium to large in size, and irregularly dispersed. of the single specimen cited (mearns & hutchinson 4632), only two sheets (ph and k) have been found, both annotated in macfarlane’s hand, each with two intermediate pitchers. although all four pitchers share a basal ridge (fig. 1f–h), only one of the four has an appendage, and that is only moderately developed as a convex emergence from the basal ridge (fig. 1h). however a recent collection (gaerlan et al. in ppi 10977) with upper pitchers, does show a developed appendage (fig. 1e), suggesting the epithet ecristata “lacking a crest” is inappropriate. in any case, the code demands priority only at one rank, so there is no requirement to adopt the varietal epithet at specific level, for which reason macfarlane’s taxon is renamed as n. kurata sp. nov. the upper pitchers also differ from the intermediate pitchers in the greater density of the nectar glands on the lower surface of the lid. however the shape, distribution and size of the nectar glands remain similar. this is the only known species of nepenthes from mt malindang at this time, and it is therefore the most westerly known species of the genus in mindanao. nepenthes kurata sp. nov. is still incompletely known, full details on its ecology, altitudinal range, population density, inflorescences and infructescences, and ethnobotany remain to be discovered. the type specimens were collected by major e.a. mearns and w.j. hutchinson in 1906 on the first recorded ascent of mt malindang, a volcanic mountain in the nw of mindanao. both sheets are annotated in the hand of macfarlane as “n. alata var. ecristata macfarlane”, and either could be selected as lectotype of that name. the k sheet is accordingly selected. european journal of taxonomy 69: 1-23 (2013) 10 nepenthes kitanglad jebb & cheek sp. nov. urn:lsid:ipni.org:names:77134487-1 fig. 2 diagnosis differs from n. saranganiensis sh.kurata in having angled, (not winged) stems, lower and upper pitchers strongly dimorphic, (not subdimorphic); in being a climbing epiphyte of forest (not a terrestrial shrub of open areas) and in having a strongly concave pitcher mouth with a long neck (not with the pitcher mouth flat or only slightly concave, lacking a neck). etymology named, as a noun in apposition, for mt kitanglad, the type and only known locality of the species. type philippines. mindanao, bukidnon province, intavas, impasug-ong, mt kitanglad, 18 jul. 1991, gaerlan, sagcal & fernando in ppi 3274 (holotype bish!; isotype brit!). description epiphytic climber, probably 1 m tall or more. short stems terete, 4–5 mm diam., internodes ca. 6 cm long, axillary buds not evident; surface glossy, appearing glabrous but with extremely sparse brown, simple hairs ca. 0.5 mm long, glabrescent. climbing stems strongly 4-angular, 7–9 mm diam., internodes 11–12 cm long; indumentum as short stems. leaves of short, and of climbing stems more or less identical, thickly papery; blade oblong-elliptic, 15.5–33 × 4.3–7 cm; apex acute, not peltate, tendril arising abruptly; base cuneate, decurrent to petiole; longitudinal nerves 3–4 pairs, conspicuous in the marginal half on the upper surface; pennate nerves arising at about 45° from the midrib, irregular, reticulate, branching in the marginal half; drying brown-black above, matt mid-brown below, appearing glabrous apart from margin but with indumentum as stem, densest on midrib but soon glabrescent; lower surface with sessile depressed-globose red-black glands ca. 0.05 mm diam.; margin densely fringed with soft fine orange-brown patent simple or bifurcate hairs 1 mm long. petiole evenly winged along its length, 4–5 × 0.4–0.9 cm, wings patent; at base clasping the stem for ⅔ to ¾ of its circumference, decurrent nepenthes kurata sp. nov. nepenthes mindanaoensis geography mt malindang, nw mindanao surigao prov. ne mindanao substrate volcanic ultramafic habitat m a.s.l. submontane forest; ca. 1400 m open scrub; 200–300 m indumentum (stem, leaf midrib and margin translucent-brown, simple or bushy, 0.1 mm long dark red-brown, bristle-like hairs 1–1.5 mm long lower surface leaf-blade hairs absent (except midrib) hairy petiole wings patent involute (petiole appearing cylindrical) longitudinal nerves of leafblade arising along length of midrib arising from petiole pitcher lid base truncate or rounded deeply cordate mouth:lid (length ratio) ca. 2:1 0.9–1.2:1 midline nectar glands of lower lid surface longitudinally elliptic (versus circular outside midline) circular, not differentiated from those of the rest of the lid table 1. diagnostic characters separating n. kurata sp. nov. and n. mindanaoensis. http://www.ipni.org/urn:lsid:ipni.org:names:77134487-1 cheek m. & jebb m., nepenthes (caryophyllales: nepenthaceae) in the philippines 11 fig. 2. nepenthes kitanglad sp. nov. a. habit, climbing stem with upper pitcher (from herbarium specimen). b. upper pitcher (from live plant). c. lower pitcher. d. indumentum from leaf margin. e. indumentum from tendril. f. indumentum from stem (extremely sparse hairs). g. indumentum of outer pitcher surface, plan view. h. ibid., profile view. i. lower surface of lid (upper pitcher). j. detail of glands from lower surface of lid. k. spur of upper pitcher. l. peristome, upper pitcher, view from above. m. peristome, view from inside of pitcher, showing minute teeth. n. peristome, transverse section (outer surface on right). a, d–n from gaerlan et al. in ppi 3274 (bish); b from mcpherson (2009: fig 417); c from gaerlan et al. in ppi 3274 (brit). scale bars: double = 1 cm; double graduated = 5 cm; single = 1 mm; single graduated = 2 mm. all drawn by andrew brown. european journal of taxonomy 69: 1-23 (2013) 12 diagonally as a narrow wing, in short stems 7 mm long, in climbing stems 18 mm long, and continuing as a ridge to the node below. lower pitchers narrowly ovoid-cylindric, 12.5 cm tall, 5 cm broad, widest in the basal half, narrowing steadily to ca. 3 cm wide below the peristome; outer surface 10–25% covered in pale brown hairs of two types, (1) bushy brown hairs 0.1–0.25 mm long and wide, with 4–8 arms ascending from a short central axis, 7–12 per mm2, (2) long brown straight erect hairs 1.5–1.75 (–2.5) mm long, with 2–4 short branches ascending from along the length of the main axis, sparse; fringed wings, 2–4 mm wide, running 3–4 cm from peristome towards base of pitcher, then diminished to slender ridges, wings extended over the peristome by two foliose flaps 3–4 × 3–4 mm, fringed elements 4–5 mm long, 2.5 mm apart (1.5 mm apart on foliose flaps); mouth ovate-lanceolate, highly oblique, concave, ca. 4.1 × 2.8 cm; column developed, tapering towards lid ca. 9 mm long, 2.5 mm wide at midpoint; peristome subcylindric, 1 mm wide at front of pitcher to 3 mm wide at sides, ridges ca. 2.5 per mm, ridges 0.1 mm high, inner edge lacking conspicuous teeth or holes, outer edge not lobed. lid narrowly ovate to rhombic 3.5 × 2.3 cm, apex rounded, base rounded to truncate, lower surface lacking a basal appendage, but with a low basal ridge 10 mm long, 0.5–1 mm high, extending from the junction with the peristome; nectar glands small and sparse, 6–8 on each side of the midline which mainly lacks glands, absent from basal ridge, nectar glands monomorphic, slightly perithecoid, orbicular or slightly elliptic, 0.25(–0.35) mm long, mixed with denser sessile depressed-globose, red-black glands, 0.05– 0.1 mm diam., 8 per mm2; marginal 2–3 mm of lower surface with minute stellate hairs densest near margin; upper surface with same indumentum as outer pitcher surface, but long hairs rarely seen. spur not seen. upper pitchers (tendril coiled) ovoid-cylindric, green, slightly maroon above, 21.5 × 6.5 cm, widest in the ovoid basal third, narrowing to ca. 5 cm wide in the cylindrical upper part; outer surface with same indumentum as lower pitcher; fringed wings reduced to ridges apart from two foliose flaps immediately below peristome, point of attachment 3–4 mm long, angular-elliptic, 9 × 6 mm, bearing fringes 2–7 mm long; mouth ovate-lanceolate 7 × 4 cm, oblique, concave, the frontal part straight; column ca. 1.5 × 0.8 cm; peristome rounded-flattened, 1.75–5.5 mm wide, widest at sides, ca. 1.75 ridges per mm, ridges 0.1 mm high, inner edge lacking conspicuous teeth or holes, outer edge not lobed. lid ovate-triangular, 5 × 3.8–4.2 cm, apex rounded, base truncate; lower surface with a weakly developed, convex basal appendage 1.5 mm high, arising from a low basal ridge 7 mm long; nectar glands ca. 16 on each side of the midline, sparsely scattered, more or less absent from midline, but present at appendage, nectar glands orbicular or slightly elliptic, slightly or strongly perithecoid, 0.25–0.5 × 0.25–0.45(–0.75) mm; sessile glands 0.05–0.1 mm diam., 8–20 per mm2; upper surface of lid with indumentum as outer surface of pitcher. spur inserted 2 mm below junction of lid and pitcher, pointing downwards, terete, 17 × 0.9–1 mm, dilating to the 1.8 mm wide rhombic-acute apex, indumentum moderately dense of long patent simple hairs as on the pitcher outer surface. male and female inflorescences unknown. distribution and habitat philippines, mindanao, bukidnon province, known only from mt kitanglad; epiphytic in mossy forest, geology volcanic, elevation 1800–2100 m. conservation here n. kitanglad sp. nov. is assessed as critically endangered since it is known from only a single location, mt kitanglad, on an island which has seen extensive forest clearance for logging and agricultural expansion in recent years (mcpherson 2009: 759). for these reasons one species, n. robcantleyi cheek is already suspected to be extinct in the wild (cheek 2011). nepenthes kitanglad sp. nov. is not a spectacular or especially bizarre species so is unlikely to come under pressure of collection for the horticultural trade which has brought several species of the genus close to extinction. cheek m. & jebb m., nepenthes (caryophyllales: nepenthaceae) in the philippines 13 remarks mcpherson (2009: 755–759; figs 417 & 418) depicts from volcanic mt kitanglad in n-central mindanao a plant under the name of n. saranganiensis sh.kurata (kurata 2003: 41). yet, the kitanglad plants he depicts differ from n. saranganiensis as depicted in its protologue in habit, habitat and in morphology. in 2013 sheets of gaerlan et al. in ppi 3274 (bish, brit) became available from kitanglad. these matched mcpherson’s (2009) depiction, enabling a detailed comparison to be made with n. saranganiensis. the conclusion is that the kitanglad material represents a different species from n. saranganiensis and is here described as n. kitanglad sp. nov. differences between the two taxa are given in table 2. n. kitanglad sp. nov. is unusual in the n. alata species group (cheek & jebb 2013d) in the strongly concave mouth of the upper pitchers, in which the base of the lid is held over the mouth. it is also unusual in that the rear of the peristome narrows to a neck, forming a moderately well-defined column for the lid. within the n. alata group, these two features are otherwise currently known only in n. hamiguitanensis gronem., wistuba, v.b.heinrich, s.mcpherson, mey & v.b.amoroso (gronemeyer et al. 2010), but that species is restricted to ultramafic mt hamiguitan in se mindanao and differs greatly in the shape of the upper pitchers which are stout, widest at the midpoint, with a funneliform lower half narrowing to a more slender, cylindrical upper half. it is possible that n. hamiguitanensis, n. kitanglad sp. nov. and n. saranganiensis are related since all have angled stems (or in the case of the last species, winged), a feature otherwise unknown in the otherwise terete-stemmed n. alata group. nepenthes kitanglad sp. nov. is unique in the n. alata group in its lid posture, as seen from photographs and herbarium specimens. the lid is held at ca. 45° below the horizontal, largely concealing the mouth. in other species it is usually elevated above the horizontal, sometimes by ca. 45° (n. saranganiensis) or as much as 90° or more (n. graciliflora elmer). so far, n. kitanglad sp. nov. is the only species of nepenthes recorded from mt kitanglad. table 2. diagnostic characters separating n. saranganiensis and n. kitanglad sp. nov. n. saranganiensis n. kitanglad sp. nov. habit terrestrial shrub climbing epiphyte habitat open, shrubby areas forest altitudinal range 700–800 m 1800–2100 m climbing stems terete, winged, wings 1.5–2 mm wide angled, wings absent lower & upper pitchers dimorphic? monomorphic-subdimorphic strongly dimorphic mouth ovate, flat to slightly concave lanceolate; strongly concave column absent present, 0.9 cm long lid posture elevated above horizontal by ca. 45º declinate below horizontal by ca. 45º lid nectar gland midline dense with large glands midline ± lacking glands lid basal ridge and appendage convex appendage present ridge only, appendage absent upper pitcher shape equally wide at base and apex, waisted at midpoint basal third widest, ovoid, tapering into the narrow, cylindric upper part european journal of taxonomy 69: 1-23 (2013) 14 nepenthes extincta jebb & cheek sp. nov. urn:lsid:ipni.org:names:77134488-1 fig. 3 diagnosis differs from n. mindanaoensis sh.kurata in the pitchers lacking fringed wings (versus with fringed wings), the lid base truncate (not cordate), the indumentum of the midrib of dense minute grey-white stellate hairs, (not of sparse black bristle-like hairs). etymology nepenthes extincta sp. nov. is named to signify that this species may already be extinct globally. type philippines. mindanao island “red hills (= 400 m alt.), se of claver, near the northeastern coast of the mindanao island. boundary of the surigao del sur and surigao del norte” 8 aug. 1978, fumihiro konta 12365 (holotype bish!). description terrestrial shrub, probably about 1 m tall. leaves elliptic to elliptic-lanceolate, 13–17 × 5.5–8 cm, thickly leathery, glossy above, matt mid brown below; apex obtuse to acute, not peltate; base rounded to obtuse, not decurrent; longitudinal nerves arising from base of blade, where 5–6 pairs arise on each side of the midrib, at blade midpoint 4–5 pairs occur in the outer third of the blade; pennate nerves arising at ca. 45° from the midrib, irregularly branching, ends traversing the inner longitudinal nerves; all nerves most conspicuous on upper surface; midrib deeply depressed on upper surface, lacking hairs, highly exserted on lower surface and densely (80–90% cover) grey-white stellate hairy, the hairs gathering dirt, hairs sessile, arms 5–8, 0.25–0.5 mm diam., fine, acute, appressed to surface; lower surface of blade sparsely hairy, densest towards midrib, ca. 15% cover, decreasing at margin to 5–10%, cover, hairs mainly stellate, as midrib, mixed with sparser erect, bristle-like hairs 1–2 mm long, of several types (1) with short branches arising along the length of the main axis; (2) with 2–6 ± equal erect arms; (3) with a single long erect bristle arising from a stellate hair, the “dagger-hair” of kurata (2003) more rarely (4) hairs with 2–3 erect, equal arms from the base; depressed-globose sessile red-black glands 0.03 mm diam., raised, dense, conspicuous; upper surface of blade with stellate hairs, as lower surface, scattered along the margins of the midrib. petiole 4.5 × 0.5–0.7cm, appearing cylindric due to the two involute wings, indumentum of appressed, stellate, fine 5–8-armed hairs, ca. 20% cover. lower and upper pitchers unknown, possibly not produced. intermediate pitchers (tendrils not coiled, fringed wings absent) ovoid-cylindric, 18–24 × 5.9–8.2 cm, widest in the basal half, narrowing gradually to the cylindric upper half (4.8–5.5 cm wide), not constricted or waisted; outer surface 10–20% covered in minute, white, 2–4-armed, bushy-stellate hairs 0.12–0.15 mm wide and high, the arms stout and raised, 10–15 hairs per mm2, mixed with sparser black depressed-globose sessile glands 0.07 mm diam., 4–5 per mm2, long simple and bristle-like hairs absent; fringed wings absent, reduced to ridges; mouth ovate 7–8 × 4.5–5.5 cm, oblique, concave, column weakly defined ca. 1.5 × 0.7 cm; peristome subcylindric, 7–8 mm wide, widest at sides, outer edge lobed, lobes 1–3 per side, 10–12 mm wide, inner side inconspicuously toothed, teeth 0.25 mm long; ridges 4 per mm, 0.1 mm high. lid ovate 5.2–6.5 × 4–5.2 cm, apex rounded, base truncate, lower surface with a basal ridge ca. 8 mm long, 1–2 mm high, bearing a pronounced straight convex appendage 4 × 2.5–5 mm; nectar glands of two distinct size classes (1) smaller, elliptic or orbicular, frequent, bordered glands 0.3–0.6(–0.7) × 0.25 mm, the border ca. 0.1 mm wide, glossy pale brown, dense, (1(–2) per mm2) along the midline these are longitudinally elliptic, elsewhere with their short axis orientated towards the base of the lid; the appendage completely covered in smaller type nectar glands; (2) larger glands elliptic to orbicular, (1–)1.25–2 × 1.25 mm, the http://www.ipni.org/urn:lsid:ipni.org:names:77134488-1 cheek m. & jebb m., nepenthes (caryophyllales: nepenthaceae) in the philippines 15 fig. 3. nepenthes extincta sp. nov. a. habit, showing intermediate pitcher. b. indumentum, lower surface of blade (midrib on left). c. branched bristle hairs (detail from lower surface of blade). d. stellate hairs and sessile glands (detail from lower surface of blade). e. branched erect hairs (detail from lower surface of blade). f. stellate hairs, with appressed arms, leaf-blade lower surface, near midrib. g. upper part of intermediate pitcher, lid posture as in life. h. lower surface of upper pitcher lid. i. basal lid appendage and spur. j. basal lid appendage and ridge, side view. k-m. details of nectar glands, lower lid surface. n. indumentum, outer surface of pitcher. o. peristome, view from inside pitcher showing minute teeth and holes at edge. p. peristome, view from above. q. transverse section of peristome (outer surface on right). a–f & h–q from konta 12365, g from sketch by m. cheek. scale bars: double, graduated = 5 cm; single = 1 mm; single graduated = 2 or 3 mm as indicated. all drawn by andrew brown. european journal of taxonomy 69: 1-23 (2013) 16 lumina often invaginated by a projection of the border, border 0.2 mm thick, 4–15 on each side of the lid, scattered around the margin and towards the apex of the midline; sessile, depressed-globose, red glands, 0.05 mm in diam., 1–2 per mm2; marginal 0.5–1 mm of lower surface with minute branched hairs 0.1 × 0.1 mm; upper surface with indumentum as outer pitcher. spur inserted 5–6 mm below junction of lid with peristome, cylindric 8–14 × 1–1.2 mm, apex shortly bifid, surface covered in minute appressed, matted, white-grey stellate hairs. male and female inflorescences and infructescence unknown. distribution & habitat philippines, mindanao, surigao del sur. open scrub habitats on ultramafic substrate with n. merrilliana macfarl. (macfarlane 1911) and n. graciliflora elmer (elmer 1912). elevation: ca. 400 m. conservation nepenthes exincta sp. nov. is here assessed as critically endangered under criterion d of iucn (2012) since only a single individual has ever been recorded (the type specimen collected in 1978). the locality data given corresponds with the large area of ultramafic known as ‘red mountain’ se of claver in ne mindanao. in fact, red mountain is a series of low red hills. the ne mindanao, probably due to its large areas of ultramafic substrate, supports several narrowly endemic and often spectacular nepenthes species, several of which are known from single locations. for this reason, it has been intensively visited in recent decades by devotees of the genus. despite this, no additional collections or observations of this taxon have come to light in the last 25 years and it is possible that it is restricted to the red mountain, and is now extinct, since this happens to be the largest nickel mining site in the world’s second largest nickel producing country (gallares 2013). the foundation for the philippine environment (pacudan 2013) recently reported on the environmental damage due to extensive and massive nickel mining “as far as your eyes can see.”, “the scene of endless open pit mining at the red mountain made our heart bleed.” (pacudan 2013). the biggest mining companies operating at the red mountain are taganito mining corporation, platinum group metals corporation (pgmc), taganito hpal nickel corporation, adnama mining resources inc. (amri) and zhen shou mining (almeda 2012). it is much to be hoped that this species is refound, and not proved to be extinct, and if found, that it can be protected and monitored. remarks nepenthes extincta sp. nov. is most likely to be confused with n. mindanaoensis and they may share a common origin. both species occur on open ultramafic substrates in ne mindanao, both have robust, ovoid-cylindric pitchers arising from thickly leathery blades in which the longitudinal nerves arise from the base of the blade. in both species the petiole has involute wings, so appear cylindric, and the blade is not decurrent to the petiole – unusual features in the n. alata group. the two can be distinguished using the characters in table 3. the two species are not sympatric so far as is known. fumihiro konta, collector of the only known material of this species, has not been traced by us. an internet search shows that he has published on the plants of s china and thailand, as well as japan, covering specialisms such as asclepiadaceae, ferns and vegetation mapping. “a list of the ferns and flowering plants of mt fuji, 1984” is his most referenced work. since “environmental impact studies” are listed among his interests, it may have been in that context that he collected the type specimen recorded here. cheek m. & jebb m., nepenthes (caryophyllales: nepenthaceae) in the philippines 17 nepenthes leyte jebb & cheek, sp. nov. urn:lsid:ipni.org:names:77134489-1 fig. 4 diagnosis differs from n. alata b lanco in having, except in the hairy axils, glabrous stems, (not densely white hairy); upper pitchers lacking fringed wings (versus with fringed wings); nectar glands on lower surface of lid dimorphic, concentrated around margin and appendage (not monomorphic, uniformly dense and distributed). etymology the specific epithet “leyte” is here used as a noun in apposition, to commemorate the island of that name, to which the species appears unique. type philippines. eastern visayas, leyte, mt lobi, near dagami, 7 nov. 1999, argent, mendum, fuentes, r., belonias, b.s. 99214 (holotype k!; isotypes e!, pnh). description terrestrial climber, height 2 m (probably), drying brown. climbing stems subterete, 4–6 mm diam., with a slight ridge below the leaf bases; the axil with a shallow groove containing a spike-like bud 1–2 mm long, inserted 5–6 mm above the axil; internodes 3–7 cm long; surface with scattered redblack, depressed-globose, sessile raised glands 0.05–0.08 mm diam.; hairs absent, except in the axillary grooves which have white, moderately dense, basally branched hairs with arms erect, ca. 1 mm long. rosette stems and leaves unknown. leaves of climbing stems spirally inserted, thinly leathery; blade narrowly oblong-elliptic, 13.5–16 × 2.5–3.8 cm; apex acute, not peltate; base cuneate, abruptly decurrent to the petiole; longitudinal nerves 1 pair, moderately close to the margin, inconspicuous; pennate nerves numerous, conspicuously raised on both surfaces, more or less patent, irregular; both surfaces drying brown, subglossy above, matt below; midrib on both surfaces 5–10% covered with fine white-translucent simple or 3–5-armed stellate hairs, on the upper surface 0.2–0.3 mm diam., on the lower surface 0.1–0.2 mm diam., the leaf-blade otherwise glabrous, apart from sessile red-black glands as the stem, 0.05–0.1 mm diam., 2–6 per mm2. petiole evenly winged along its length, the wings incurved (field notes); (2.5–)3–4.5 × 0.2–0.4 cm; clasping the stem for ½ its circumference, very shortly decurrent by table 3. diagnostic characters separating n. extincta sp. nov. from n. mindanaoensis. nepenthes extincta sp. nov. nepenthes mindanaoensis indumentum of leaf-blade (lower surface of midrib) densely covered in grey-white stellate hairs 0.25–0.5 mm diam., arms 5–8, fine, appressed to surface sparsely covered in brown or black, erect bristle-like hairs 1–1.5 mm long intermediate/upper pitchers shape ovoid basal half gradually tapering into cylindric upper half ovoid basal third abruptly narrowing to cylindric upper twothirds fringed wings absent from intermediate pitchers present on lower, intermediate and upper pitchers lid base ± truncate markedly cordate basal appendage of lid strongly developed absent http://www.ipni.org/urn:lsid:ipni.org:names:77134489-1 european journal of taxonomy 69: 1-23 (2013) 18 1–2 mm. lower and intermediate pitchers unknown. upper pitcher (tendril coiled) 12–15 × 4.5–5 cm; ovoid-ellipsoid in the lower half, upper half cylindrical, 3–3.5 cm diam., not constricted at any point; outer surface 10–30% covered in minute red stellate hairs, hairs ca. 0.1 mm diam., both sessile and shortly stalked, 4–6-armed, arms suberect or patent, density 3–5 per mm2, mixed with sessile red-black glands 0.05 mm diam. as the leaf-blade and stem, hairs denser on lid, and towards the peristome where they are mixed with sparse erect bushy-bristle hairs 0.2–0.3 mm long; “almost uniformly green with a few purple spots mainly on the ventricose base” (argent et al. 99214); fringed wings are absent, reduced to inconspicuous ridges; mouth ovate, 4–4.5 × 3–3.5 cm, oblique, slightly concave, “glaucous green inside with just a few red spots”; peristome (1–)2–3(–5) mm wide, narrowly subcylindrical, rounded at the front, becoming slightly flattened and widest at the sides, towards the lid, ca. 4 ridges per mm, ridges 0.075–0.15 mm high, inner edge inconspicuous, holes and teeth not visible (unless dissected: fig. 4p); outer edge not lobed; column weakly developed, ca. 7 × 3 mm. lid ovate 3.2 × 2.9(–3.2) cm; apex shallowly retuse, the sinus 3–7 mm wide; base cordate, the sinus 4 mm deep, 8–15 mm wide; green; margin undulate; lower surface with convex basal appendage, 0.4–0.7 × 1–2 mm, arising from near the midpoint of the 5–6 × 0.5 mm long basal midline ridge; nectar glands slightly dimorphic, each with a different distribution: (type 1) moderately dense on the basal ridge and appendage (fig. 4l) and in a ca. 2 mm band each side (but not extending along midline), glands with raised borders, shortly elliptic, 0.1– 0.2(–0.3) mm long; (type 2) slightly larger, (0.1–)0.2–0.3 mm long, moderately dense, in bands 2–4 mm wide along the lid margins, 25–40 glands on each side, one sheet (atypical?) with a few additional large elliptic glands, 0.7 × 0.4 mm, bordered, very sparsely scattered between the margins; sessile red-black glands, as stem, leaf and outer pitcher surface, 0.005–0.01 mm diam., scattered over surface ca. 3 per mm2; marginal part of lower surface with a few minute stellate hairs. spur inserted 2 mm below junction of lid and pitcher on ridge; simple, stout at base tapering to a long, acute apex; 7–9.5 × 0.5–0.7 mm; completely covered in long, grey appressed hairs, hairs (0.5–)0.7–1(–1.2) mm long. upper surface of lid with two prominent nerves, nerves densely (80–90% cover) white hairy, hairs of two types: (1) basally 1–2-branched hairs 0.3–0.4 mm long, (2) minute 3–5-armed stellate hairs 0.1 mm diam.; remainder of lid surface with type (2) hairs, but indumentum 30–40% cover, and with sparse perithecoid nectar glands 0.25 mm long. inflorescence and infructescence unknown. distribution & habitat philippines, visayas, leyte; volcanic geology; “climbing on fallen tree in submontane mossy forest”, elevation 900 m (argent et al. 99214). conservation nepenthes leyte sp. nov. is known currently from a single individual in an unprotected area, in a country, including specifically the island of leyte, where most of the original forest habitat has been cleared for timber and agricultural land and where forest degradation and clearance are ongoing (myers et al. 2000; googleearth viewed 2 oct. 2013). accordingly, it is here assessed as critically endangered under criterion d of iucn (2012). it is to be hoped that further exploration will reveal additional localities for the species, and that protection can be arranged before it becomes extinct. currently no protected areas are believed to be present on leyte apart from the 2193 ha lake danao national or natural park which is about 14 km to the w of the type location. however this seems to be mainly a recreational area, and within the reserve, illegal settlement, slash and burn agriculture and illegal logging are reported to be problems (http://en.wikipedia.org/wiki/lake_danao_(leyte)#threats, viewed 12 oct. 2013). viewing the area immediately around lake danao on googleearth (2007 imagery, viewed 12 oct 2013) confirms that large areas have been and were in 2007 in the process of being cleared and inhabited, and that these activities extend towards the e and the only known location for n. leyte sp. nov. some original forest still survives along the central high ridge of leyte, where terrain appears rugged, including the location indicated as the type location of the species, however the resolution of the imagery here is not http://en.wikipedia.org/wiki/lake_danao_(leyte)#threats cheek m. & jebb m., nepenthes (caryophyllales: nepenthaceae) in the philippines 19 fig. 4. nepenthes leyte sp. nov. a. habit, climbing stem with upper pitchers. b. stem section showing axillary hair patch and supra-axillary bud. c. lower surface of leaf-blade, with sessile glands. d. midrib of leaf-blade, lower surface, stellate to simple hairs. e. midrib of leaf-blade, upper surface with stellate hairs. f. upper lid of pitcher, upper surface. g. indumentum of upper surface of lid, over nerve. h. as g but distant from nerve. i. spur of upper pitcher and junction of lid with peristome (inverted). j. lid of upper pitcher, lower surface. k. detail of j showing large nectar glands. l. detail of j showing midline ridge with appendage and type (1) nectar glands. m. indumentum of outer pitcher surface. n. peristome of upper pitcher viewed from above. o. peristome viewed from inside pitcher. p. peristome dissected to expose the inner edge, with teeth. q. peristome, transverse section (outer surface on right). all drawn from argent et al. 99214 by andrew brown. european journal of taxonomy 69: 1-23 (2013) 20 sufficiently high to gauge how intact the habitat is. the eastern side of leyte has higher rainfall and the forest has extensively been replaced by intensive industrial oil palm plantations which extended in 2003 to within 4 km of the type location (googleearth imagery dating from 2003, viewed 2 oct. 2013). collection of nepenthes leyte sp. nov. from the wild to supply the horticultural trade is considered a low risk for this species since its pitchers are not as spectacular or as bizarre as those of other members of the genus in the philippines. remarks argent et al. 99214, here described as nepenthes leyte sp. nov., while superficially similar to n. graciliflora, the only other species of the genus known on leyte (wenzel 680, gh!; barbon et al. in ppi 8735, brit!; ibid. 8561, brit!), cannot be confused with it. this is due to the stellate hairs present on the outer pitcher surface of nepenthes leyte sp. nov. (versus absent in n. graciliflora), and the dimorphic nectar lid glands concentrated around the margin and appendage (not monomorphic, uniformly dense and distributed). it also has petioles that appear cylindrical since the wings are involute (not with patent wings). apart from n. graciliflora itself, only one other member of the n. alata group is, so far, known from the visayas: n. negros (negros and biliran islands). however, n. negros has densely hairy stems (versus glabrous), upper pitcher with fringed wings (not absent) and the inner peristome edge has conspicuous teeth and holes (versus conspicuous teeth absent). nepenthes leyte sp. nov. can be distinguished from other species of the nepenthes alata group using the key above. discussion in an earlier paper, the nepenthes alata group of species, confined to the philippines, was characterised and a key was provided to the then recognised species (cheek & jebb 2013d). in that paper the species of the n. alata group were characterised as possessing: (1) a basal appendage on the lower surface of the pitcher lid, (2) a terete stem, (3) a distinct, but winged petiole, the petiole wings wide, decurrent from the blade, (4) the peristome finely ridged, the outer edge not or only slightly lobed, (5) the inner surface lacking conspicuous teeth, (6) the mouth ovate, oblique, without a well-developed column (cheek & jebb 2013d). the four new species described here have characters that necessitate modification of the nepenthes alata group as circumscribed in cheek & jebb (2013d). all but n. leyte sp. nov. of the new species have stems that can be 4-angled in section rather than terete. in n. extincta sp. nov. and in n. leyte sp. nov., the petiole wings are incurved, the two edges overlapping, giving the petiole a cylindrical appearance. both n. extincta sp. nov. and n. kitanglad sp. nov. have broad, partly flattened peristomes (rather than narrowly cylindric) and pitcher mouths which are concave (rather than straight). in the case of n. kitanglad sp. nov., a moderately well-developed column is formed (rather than none, or a weakly developed column). several of these characters are more usual in the regiae group of species of danser (1928), which are considered by us to be closely related to the n. alata group. the two species groups share (1) petiolate leaves and, (2) on the basal part of the lower surface of the lid, appendages emerging from a keel-like ridge or crest, and (3) inflorescences with 2-flowered partial-peduncles. the two species groups can be both defined and separated from each other using the key below. it is possible that future discoveries will provide evidence that the two groups are a continuum, and should be united. – upper pitchers infundibuliform or infundibuliform-cylindrical, always widest at the peristome; petioles not or weakly winged, u-shaped in section, the wing width if present, shorter than the width of the central, non-winged part of the petiole; stem and leaf indumentum of persistent, red-brown bristle-like hairs. borneo, sulawesi & new guinea .............................................................. .................................................................................................... regiae species group (danser 1928) cheek m. & jebb m., nepenthes (caryophyllales: nepenthaceae) in the philippines 21 – upper pitchers usually ovoid-cylindrical, rarely subcylindrical, always widest in the basal half (except in n. ceciliae and n. copelandii ); petiole wing width wider than the width of the central, non-winged part of the petiole, wings either held flat or inrolled and then the petiole appearing cylindrical. philippines excluding palawan .................................................... ................................................................................ n. alata species group (cheek & jebb 2013d) acknowledgements janis shillito typed the manuscript. barbara kennedy at bish and tiana rehman at brit kindly provided the loan of the material cited. lee davies and yvette harvey digitised the images. references adam j.h. 1998. reproductive biology of bornean nepenthes (nepenthaceae). journal of tropical forest science 10(4): 456-471. almeda v.l. 2012. mining firms told to clean up claver bay, ‘chocolate river’ by 2013. minda news [online]. available from http://www.mindanews.com/environment/2012/03/30/mining-firms-told-toclean-up-claver-bay-chocolate-river-by-2013/ [accessed 15 aug. 2013] berchtold b.v. von & presl j.s. 1820. o přirozenosti rostlin, aneb rostlinár, obsahugjcj: gedanj on žiwobytj rostlinném pro sebe a z ohledu giných žiwoků, podlé stawu nyněgssjbo znánj; k rozssjřenj přirodnictwj; w potaženj na užitećnost w rolnictwj, hospodářstwj, řemestech, uměnj i obchodu a w wztahowánj obzwlásstnjm na lekařstwj. enders, prague. blanco m. 1837. flora de filipinas. en la inprenta de sto. thomas por d. candido lopez, manila. cheek m. 2011. nepenthes robcantleyi sp. nov. 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national botanic garden of belgium, meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark http://dx.doi.org/ 10.1038/35002501 http://fpe.ph/wp-content/uploads/2013/01/mru-fieldexposure-sarihay.pdf http://fpe.ph/wp-content/uploads/2013/01/mru-fieldexposure-sarihay.pdf http://sweetgum.nybg.org/ih/ http://sweetgum.nybg.org/ih/ european journal of taxonomy 103: 1–20 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2014.103 www.europeanjournaloftaxonomy.eu 2014 · grootaert p. & shamshev i. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:cc7f0646-9d66-4068-9336-c6dc12098c4e 1 new species of platypalpus (diptera: hybotidae) from the democratic republic of the congo patrick grootaert 1 & igor shamshev 2 1, 2 entomology, royal belgian institute of natural sciences, rue vautier 29, b-1000, brussels, belgium. email: patrick.grootaert@naturalsciences.be (corresponding author) 2 all-russian institute of plant protection, shosse podbel’skogo 3, 188620, st. petersburg, russia. temporary address: royal belgian institute of natural sciences, brussels. email: shamshev@mail.ru 1 urn:lsid:zoobank.org:author:b80bc556-9087-4d0d-9d69-7fa9be5779c4 2 urn:lsid:zoobank.org:author:569f41cc-ec2b-4cf0-802a-8d7056c72c93 abstract. six new species of platypalpus macquart, 1827 are described from tropical forest at yangambi (democratic republic of the congo): platypalpus bolikoi sp. nov., p. ikoso sp. nov., p. lokonda sp. nov., p. manjano sp. nov., p. saffradi sp. nov. and p. yangambensis sp. nov. all species are photographed and, except for p. saffradi sp. nov. known only from females, male terminalia are illustrated for all. a key is provided for the six species of dr congo. coi barcodes are available for all species at genbank. keywords. afrotropical region, diptera, platypalpus, new species. grootaert p. & shamshev i. 2014. new species of platypalpus (diptera: hybotidae) from the democratic republic of the congo. european journal of taxonomy 103: 1–20. http://dx.doi.org/10.5852/ejt.2014.103 introduction the genus platypalpus macquart, 1827 is the largest group of the hybotidae subfamily tachydromiinae and one of the most speciose genera of empidoidea on the whole. currently about 550 species of platypalpus are known worldwide but about 75% of them were described from the palaearctic and nearctic regions (yang et al. 2007). in the afrotropical region only 28 species are known. the first species was described by loew (1858) from south africa; becker (1914) described three species from kenya; collin (1922) described a single species from the seychelles; smith (1967a) described five species from tanzania and eighteen species from south africa (smith 1967b, 1969). only south african species of platypalpus have been keyed. in contrast to the palaearctic region where platypalpus is very common and diverse, few platypalpus species are found in tropical lowland forests in the afrotropical region. having a similar habitat and niche as elaphropeza macquart, 1827, platypalpus probably comes into competition with elaphropeza that is likely more adapted to tropical conditions. however above 500 m, platypalpus is more speciesrich and the number of elaphropeza species drops (grootaert & shamshev 2006). http://dx.doi.org/10.5852/ejt.2014.103 https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:cc7f0646-9d66-4068-9336-c6dc12098c4e http://zoobank.org/urn:lsid:zoobank.org:author:b80bc556-9087-4d0d-9d69-7fa9be5779c4 http://zoobank.org/urn:lsid:zoobank.org:author:569f41cc-ec2b-4cf0-802a-8d7056c72c93 http://dx.doi.org/10.5852/ejt.2014.103 european journal of taxonomy 103: 1–20 (2014) 2 the present paper is the result of a short sampling campaign at the end of may and first half of june 2013 in the surroundings of yangambi (dr congo, oriental province) at an altitude of about 400 m. the collection was made at the beginning of the ‘short dry season’ and indeed it did not rain at all during the campaign. collections were made in a patchwork of forest types ranging from garden patches, bamboo, over young forest to mixed primary and monodominant (gilbertiodendron j.léonard) forest. the study was in the scope of the podo project cobimfo (congo basin integrated monitoring for forest carbon mitigation and biodiversity) in which a relation is being sought between carbon fixation and biodiversity. the aim of this project is to get baseline reference data on the carbon balance and biodiversity in pristine and intervened dense tropical forests of the congo basin and to increase our understanding of the relationship between both variables as a function of forest management. only thirteen specimens of platypalpus were found out of a total of about 1600 tachydromiinae showing how rare platypalpus is. however, they belong to six species all new for science. moreover platypalpus is here reported for the first time from dr congo. the present study is also the continuation of a major review of the hybotidae of dr congo that was started in 2010 (grootaert & shamshev 2013). material and methods the holotypes will be housed in the collections of the centre de surveillance de la biodiversité in kisangani (dr congo) as soon as the facilities will be fully operational, but are now temporarily deposited in the royal belgian institute of natural sciences, brussels, belgium (rbins). in the descriptions we cite under ‘reg.’ the register number that is a unique identifier referring to the locality, date and microhabitat. when we cite the holotype we indicate also the ms name or code that was used throughout the description process e.g. ‘plat3’ is the third platypalpus species found in the congo samples. all specimens were collected by the first author (pg) by sweep netting and are conserved in 70% ethanol. the specimens from which dna was extracted are conserved in our tissue collection at rbins with the cuticular parts still being recognisable. we refer to the barcode of the tube in the tray in which it is conserved: e.g. ‘tube ab31515301’ stored in tray called ‘congo2’. the terminology and descriptive format used in the descriptions follow grootaert & shamshev (2012). in describing the sclerites of the thorax the terminology used by previous authors is often indicated in brackets. the scale bar represents 0.1 mm, but this is only an indicative measure because the drawings are projections of structures that are usually not in a horizontal plane. abbreviations the following abbreviations are used in the illustrations: a = apodeme lc = left cercus lel = left epandrial lamella rc = right cercus rel = right epandrial lamella rs = right surstylus. the species-groups of platypalpus are according to grootaert & chvála (1992). grootaert p. & shamshev i., new platypalpus (hybotidae) from dr congo 3 results class insecta linnaeus,1758 order diptera linnaeus,1758 superfamily empidoidea latreille, 1804 family hybotidae macquart, 1823 subfamily tachydromiinae meigen, 1822 genus platypalpus macquart, 1827 coryneta meigen, 1800: 27. type-species: musca cursitans fabricius, 1775, by subsequent designation of engel (1939: 43). suppressed by iczn (1963: 339; opinion 678). platypalpus macquart, 1827: 92. type-species: musca cursitans fabricius, 1775, by subsequent designation of westwood (1840: 132). platypalpus bolikoi sp. nov. urn:lsid:zoobank.org:act:f1fb7470-9502-4b74-b175-4c76bc47e5f4 figs 1–5 diagnosis small black species of the pallidiventris-cursitans group. one pair of verticals. postpedicel short, 1.5 times as long as wide. thorax faintly tomentose on scutum. legs including all coxae yellow, but mid femur brownish dorsally and hind femur dark brown on apical 2/3, apical half of tarsomere 5 of all legs brownish; large pointed apical spur on mid tibia. etymology the species is dedicated to mr. lola boliko, who guided the first author in the forest during the expedition in 2013. type material holotype ♂, dr congo, yangambi, 7 jun. 2013, young forest jeu2 (0°47’41.77” n 24°29’30.91” e; reg. 33053; leg. p. grootaert; plat4); mid leg in tray entomo 004 in position a2 with barcode ab42406107 figs 1–2. platypalpus bolikoi sp. nov., holotype ♂. 1. wing, abdomen and hind femur. 2. mid femur, posterior view. http://zoobank.org/urn:lsid:zoobank.org:act:f1fb7470-9502-4b74-b175-4c76bc47e5f4 european journal of taxonomy 103: 1–20 (2014) 4 was extracted for dna and the coi barcode can be accessed in genbank with the accession number: kj768325. description male length. body 2.0 mm; wing 1.8 mm. head. entirely black. occiput densely greyish pollinose; with 1 pair of long yellow verticals. ocellar tubercle greyish pollinose, with subequally very short anterior and posterior setae. frons broad, somewhat broadened toward ocellar tubercle, subshining. face narrow, greyish pollinose, clypeus subshining. antenna with scape and pedicel brown, postpedicel and stylus yellowish brown; postpedicel small, ovoid, 1.5 times as long as wide; stylus 1.8 times as long as postpedicel. proboscis pale yellowish brown, shorter than head is high. palpus pale yellow, large, elongate ovate, with scattered pale setulae. thorax. black to brown; faintly tomentose, katepisternum (= sternopleuron) shining; bristles yellowish to yellowish brown. postpronotal lobe large, with 1 long and several minute setae. mesonotum with 2 notopleurals (anterior seta very short, posterior seta long), 1 moderately long postalar and 4 scutellars (apical pair long, lateral pair short); additionally, notopleuron with several setulae; acrostichals short, sparse, arranged in 2 broad irregular rows; dorsocentrals arranged in 1-2 rows, nearly as long as acrostichals, 1 prescutellar long. legs. largely reddish yellow, but mid femur brownish dorsally, hind femur dark brown on about apical 2/3, tarsomere 5 of all legs yellowish brown apically. coxae and trochanters with ordinary yellowish to brownish yellow setae of different lengths. fore femur thickened; with rows of short anteroventral and posteroventral setae. fore tibia rather slender, clothed with ordinary setulae. mid femur very thickened; with double row of black ventral spinules and row of long, yellow posteroventral setae. mid tibia bearing figs 3–5. platypalpus bolikoi sp. nov., holotype ♂. 3. right epandrial lamella. 4. epandrium with cerci, dorsal view. 5. left epandrial lamella. scale bar = 0.1 mm. grootaert p. & shamshev i., new platypalpus (hybotidae) from dr congo 5 a row of black ventral spinules; with long pointed largely black apical spur. hind femur slender, with short anteroventral setae. hind tibia slender, clothed with ordinary setulae. tarsi of all legs unmodified. wing. hyaline, with brownish veins. one moderately long black costal seta. veins r4+5 and m1+2 somewhat convergent just before meeting wing margin. crossveins m-cu and r-m separated; cell bm longer and somewhat broader than cell br; vein cua2 straight. anal vein evanescent in apical part and lacking in basal part. calypter brown, with brown setae. halter brownish. abdomen. with tergites brownish, subshining, covered with faint very short setae; sternites brownish, with similar setae; segment 8 with several moderately long yellowish brown posteromarginal setae. terminalia rather small, brown (figs 3–5). both cerci digitiform, equally long, shorter than right epandrial lamella. right and left epandrial lamella equally long, set with unmodified setae. only short setae of left border of left epandrial lamella. female unknown. comparison in the key of smith (1969) the new species runs to p. vicina smith, 1969 that was described from south africa. in the latter species the fore femur is darkened above while in p. bolikoi sp. nov. the fore femur is entirely yellow. in p. vicina the mid and hind coxae are darkened. here they are yellow. platypalpus soror smith, 1969 has a black proboscis and yellow halteres. in the new species the proboscis is pale and halteres are brown. distribution dr congo. platypalpus ikoso sp. nov. urn:lsid:zoobank.org:act:9245014c-ecb6-4dd1-b1ea-8990af64c6bd figs 6–13 diagnosis a medium-sized species of the albiseta-group. postpedicel 5.3 times as long as wide; stylus white, 1.6 times as long as postpedicel. legs reddish-yellow. mid femur with a short flattened apical spur. etymology the specific epithet ‘ikoso’ means fly or mosquito in tumburu, the local language spoken at yangambi. type material holotype ♂, dr congo, yangambi, 3 jun. 2013, young forest jeu1 (0°47’22.77” n 24°31’03.76” e; reg. 33034; leg. p. grootaert; ms name plat3); mid leg in tray entomo 004 position a1 with barcode ab42406101 was extracted for dna and coi barcode is available in genbank with accession number: kj768325. description male length. body 3.7 mm; wing 2.9 mm. http://zoobank.org/urn:lsid:zoobank.org:act:9245014c-ecb6-4dd1-b1ea-8990af64c6bd european journal of taxonomy 103: 1–20 (2014) 6 head. entirely black. occiput densely greyish pollinose; with 1 pair of long black verticals. ocellar tubercle greyish pollinose, with 2 black, moderately long anterior and 2 very short posterior setae. frons narrow, somewhat broadened toward ocellar tubercle, above antennae slightly broader than anterior ocellus, greyish pollinose. face narrow, on middle as broad as anterior ocellus, somewhat broader below antennae and on lower part, greyish pollinose, clypeus subshining. antenna with scape, pedicel and postpedicel brown; postpedicel conical, long, 5.3 times as long as wide; stylus white, long pubescent, 1.6 times as long as postpedicel. proboscis brownish yellow, rather long, somewhat shorter than head is high. palpus brownish yellow, small, ovate, with scattered dark setulae. thorax. (fig. 6). with mesonotum brown, otherwise reddish brown; scutum shining (except prescutellar depression), katepisternum (= sternopleuron) with small shining spot, otherwise thorax faintly tomentose; figs 6–8. platypalpus ikoso sp. nov., holotype ♂. 6. head, thorax, lateral view. 7. mid legs, anterior view. 8. wing. grootaert p. & shamshev i., new platypalpus (hybotidae) from dr congo 7 bristles black. postpronotal lobe small, elongate, narrow, with several minute setulae. mesonotum with 2 notopleurals (anterior seta very short, posterior seta moderately long), 1 moderately long postalar and 4 scutellars (apical pair long and cruciate, lateral pair very short); additionally, notopleuron with several setulae; acrostichals minute, scattered, arranged in 2 irregular rows; dorsocentrals arranged in 1 row, nearly as long as acrostichals, 2 prescutellars long. legs. largely reddish yellow, fore and hind tibiae, entire fore tarsus, apical tarsomeres of mid and hind tarsi rather brownish yellow. coxae and trochanters with ordinary yellowish to brownish yellow setae of different lengths. fore femur slender; with rows of thin anteroventral and posteroventral setae shorter closer to apex, the latter including 2 longer brownish setae closer to base. fore tibia slender, clothed with ordinary setulae. mid femur (fig. 7) very thickened, long; with double row of black ventral spinules and row of long, black posteroventral setae. mid tibia bearing a row of black ventral spinules; with short flattened apical spur. hind femur slender, with short anteroventral setae. hind tibia slender, clothed with ordinary setulae. tarsi of all legs unmodified. wing. (fig. 8). uniformly brownish infuscate, with brownish veins. one moderately long costal seta. veins r4+5 and m1+2 somewhat convergent just before meeting wing margin. crossveins m-cu and r-m contiguous; cell bm longer and somewhat broader than cell br; vein cu not reaching wing border. vein cua2 somewhat sinuate. anal vein distinct in apical part and evanescent in basal part. calypter brown, with brown setae. halter with stem blackish brown, knob brown. figs 9–13. platypalpus ikoso sp. nov., holotype ♂. 9. hypandrium. 10. left cercus. 11. right epandrial lamella. 12. epandrium with cerci, dorsal view. 13. left epandrial lamella. scale bars = 0.1 mm. european journal of taxonomy 103: 1–20 (2014) 8 abdomen. with tergites brownish, subshining, covered with faint very short setae; sternites brownish, with similar setae; segment 8 with several long black posteromarginal setae. terminalia (figs 9–13) rather small, brown. left cercus (figs 10, 12) much longer than right cercus. left epandrial lamella (figs 12, 13) much smaller than right epandrial lamella (figs 11, 12). apex of left epandrial lamella with some long setae. female unknown. comparison in the key of smith (1969) platypalpus ikoso sp. nov. runs to p. inermifemur smith, 1969 and p. ngomensis smith, 1969 both described from south africa. in p. inermifemur the ratio of second and third sections of vein cu (fifth vein sensu smith 1969) is 3 : 3.5 and the hind femur is without distinct anteroventral setae. in p. ngomensis the ratio of second and third section of vein cu is 2 : 3.5 and the hind femur bears short, but distinct anteroventral setae. in p. ikoso sp. nov. the apical section of vein cu is twice as long as the second section being 2 : 4, which is comparable to p. ngomensis. the femur bears distinct anteroventral setae that are half as long as femur is wide. in p. ngomensis the left epandrial lamella is about as long as the right epandrial lamella. in p. ikoso sp. nov. the left lamella is very short (fig. 12). in addition vein cu does not reach the wing border in p. ikoso sp. nov. and p. ngomensis, but it does in p. inermifemur. distribution dr congo. platypalpus lokonda sp. nov. urn:lsid:zoobank.org:act:9d8bc13d-db91-423e-8a20-ab96db911e61 figs 14–20 diagnosis a medium-sized species of the albiseta-group. one pair of long vertical bristles. antenna black, postpedicel nearly 4 times as long as wide; with white stylus, almost 2.5 times as long as postpedicel. mesonotum dark brown; pleura brown. mid tibia with short pointed apical spur. etymology the specific epithet ‘lokonda’ means forest in tumburu language, the language spoken at yangambi. type material holotype ♂, dr congo, yangambi, 27 may 2013, primary mixed forest (0°48’04.69” n 24°31’37.87” e; reg. 33011; leg. p. grootaert; ms name plat2); mid leg in tray entomo 004 at position a6 with barcode ab42406131; coi barcode available in genbank with accession number: kj768328. paratype 1 ♂, dr congo, yangambi, 31 may 2013, monodominant gilbertiodendron forest gil3, sweeping (0°47’46.09” n 24°29’53.46” e; reg. 33025; leg. p. grootaert); in tray entomo 001 at position e2 with barcode ab48937147; coi barcode available in genbank with accession number: kj768327. description male length. body 2.5 mm; wing 2.2 mm. http://zoobank.org/urn:lsid:zoobank.org:act:9d8bc13d-db91-423e-8a20-ab96db911e61 grootaert p. & shamshev i., new platypalpus (hybotidae) from dr congo 9 head. entirely black. occiput densely greyish pollinose; with 1 pair of long black verticals. ocellar tubercle greyish pollinose, with long anterior and minute posterior setae. frons narrow, somewhat broadened toward ocellar tubercle, greyish pollinose. face very narrow, somewhat broadened below, greyish pollinose, clypeus subshining. antenna (fig. 15) with scape, pedicel and postpedicel brown; postpedicel conical, long, 3.7 times as long as wide; stylus white, densely pubescent, 2.4 times as long as postpedicel. proboscis brownish yellow, shorter than head is high. palpus pale yellow, short, elongate ovate, with scattered pale setulae. thorax. (figs 14, 15). with mesonotum dark brown, otherwise yellowish brown; faintly tomentose, scutum shining anteriorly; bristles black. pleura also finely tomentose including katepisternum (= sternopleuron). postpronotal lobe small, narrow, with several minute setulae. mesonotum with 2 notopleurals (anterior seta very short, posterior seta long), 1 moderately long postalar and 4 scutellars (apical pair long, cruciate, lateral pair very short); additionally, notopleuron with several setulae; acrostichals biserial, very short, few setae on anterior part of scutum; dorsocentrals uniserial, very short, 2 prescutellar pairs (anterior setae proclinate, posterior setae inclinate) long. legs. yellow. coxae and trochanters with ordinary yellowish to brownish yellow setae of different lengths. fore femur slender; with rows of anteroventral and posteroventral yellowish setae longer on basal part. fore tibia slender, clothed with ordinary setulae. mid femur (fig. 17) very thickened; with double row of black ventral spinules and row of long, brownish yellow posteroventral setae. mid tibia bearing a row of black ventral spinules; with short flattened apical spur. hind femur slender, with short anteroventral setae. hind tibia slender, clothed with ordinary setulae. tarsi of all legs unmodified. figs 14–17. platypalpus lokonda sp. nov., paratype ♂. 14. habitus, lateral view. 15. head and thorax, dorsal view. 16. wing. 17. mid legs, posterior view. european journal of taxonomy 103: 1–20 (2014) 10 wing. (fig. 16). faintly infuscate, with brownish veins. one moderately long costal seta. veins r4+5 and m1+2 convergent just before meeting wing margin. crossveins m-cu and r-m contiguous; cell bm somewhat longer and broader than cell br. vein cua2 sinuate. anal vein distinct in apical part and evanescent in basal part. calypter brown, with brown setae. halter brownish. abdomen. with tergites brownish, subshining, covered with faint very short setae; sternites brownish, with similar setae; segment 8 with several long black posteromarginal setae. terminalia (figs 14, 18–20) small, brown. left cercus much longer than right cercus, both widely separated. cerci clothed by unmodified hairs and setae. left epandrial lamella (figs 19, 20) much shorter than right epandrial lamella (figs 18, 19). left epandrial lamella with two long apical setae. female unknown. comparison platypalpus lokonda sp. nov. is very closely related to p. ngomensis smith, 1969 described from natal, south africa. it differs in having brown prothoracic sterna and propleura which are yellow in p. ngomensis. in the latter species, the left epandrial lamella is about as long as the right epandrial lamella, in p. lokonda sp. nov., the left lamella is very short. distribution dr congo. figs 18–20. platypalpus lokonda sp. nov., holotype ♂. 18. right epandrial lamella. 19. epandrium with cerci, dorsal view. 20. left epandrial lamella. scale bars = 0.1 mm. grootaert p. & shamshev i., new platypalpus (hybotidae) from dr congo 11 platypalpus manjano sp. nov. urn:lsid:zoobank.org:act:85ee94e0-1364-4011-834e-2e421cb4e83a figs 21–27 diagnosis a medium-sized yellow species of the albiseta-group. postpedicel brown, a little more than 5 times as long as wide; stylus white, about as long as postpedicel. mid tibia with short yellow apical sharp spur, shorter than tibia is wide. etymology the word ‘manjano’ means yellow in turumbu, language that is spoken in yangambi. it refers to the yellow body colour of the new species. type material holotype ♂, dr congo, yangambi, 27 may 2013, primary mixed forest (0°48’04.69” n 24°31’37.87” e; reg. 33015; leg. p. grootaert; ms name plat5), mid leg in position ab42406125 was extracted for dna. a coi barcode can be found in genbank with accession number: kj768329. description male length. body 2.4 mm; wing 2.7 mm. figs 21–24. platypalpus manjano sp. nov., holotype ♂. 21. habitus, lateral view. 22. wing. 23. head and thorax, dorsal view. 24. mid leg, anterior view. http://zoobank.org/urn:lsid:zoobank.org:act:85ee94e0-1364-4011-834e-2e421cb4e83a european journal of taxonomy 103: 1–20 (2014) 12 head. entirely black. occiput greyish pollinose; with 1 pair of long black verticals. ocellar tubercle greyish pollinose, with 2 black, short anterior and 2 minute posterior setae. frons very narrow, somewhat widened toward ocellar tubercle, above antennae nearly as broad as anterior ocellus, greyish pollinose. face very narrow, narrower than anterior ocellus, greyish pollinose, clypeus subshining. antenna with scape, pedicel and postpedicel brown; postpedicel conical, long, 5.3 times as long as wide; stylus white (except basal segment), long pubescent, about as long as postpedicel. proboscis brownish, long, nearly as long as head is high. palpus yellow, small, ovate, with scattered setulae and a long brownish subapical seta. thorax. almost entirely yellow, only postpronotal lobe, notopleuron narrowly along lower margin, anepisternum (= mesopleuron) narrowly along upper margin and anterior spiracle yellowish brown; scutum shining, otherwise thorax subshining; bristles brownish. postpronotal lobe elongate narrow, with 1 minute seta. mesonotum with 2 notopleurals (anterior seta very short, posterior seta long, strong), 1 long strong postalar and 4 scutellars (apical pair long and cruciate, lateral pair very short); acrostichals very short, scattered, arranged in 2 broadly spaced rows; dorsocentrals uniserial, sparse, nearly as long as acrostichals, 2 prescutellar pairs somewhat longer. legs. almost entirely yellow, except for fore tibia and apical four tarsomeres more brownish than fore femur; mid and hind leg with only apical tarsomere brownish. coxae and trochanters with ordinary yellowish to brownish setae of different lengths. fore femur slender; with very short anteroventrals and 4 fine moderately long brownish posteroventral setae on about basal half. fore tibia slender, clothed with ordinary setulae. mid femur very thickened; with double row of black ventral spinules and row of long, black posteroventral setae (fig. 24). mid tibia bearing a row of black ventral spinules; with very short flattened apical spur. hind femur slender, with short anteroventral setae. hind tibia slender, clothed with ordinary setulae. tarsi of all legs unmodified. figs 25–27. platypalpus manjano sp. nov., holotype ♂. 25. right epandrial lamella. 26. epandrium with cerci, dorsal view. 27. left epandrial lamella. scale bars = 0.1 mm. grootaert p. & shamshev i., new platypalpus (hybotidae) from dr congo 13 wing. (fig. 22) finely, uniformly infuscate, with brownish veins. costa between r1 and r2+3 darkened. one moderately long costal seta. veins r4+5 and m1+2 almost parallel just before meeting wing margin. crossveins m-cu and r-m contiguous; cell bm longer and distinctly broader than cell br; vein cua2 somewhat sinuate reaching wing border. anal vein distinct in apical part and evanescent in basal part. calypter brown, with about 10 brown setae. halter darkened. abdomen. with brownish tergites and yellowish sternites (except segment 8); tergites covered with short brownish setae; sternites with similar but somewhat longer setae; segment 8 bearing numerous very long black strong setae. terminalia (figs 25–27) moderately large, brown. left cercus (fig. 26) much longer and broader than right cercus, clothed with long unmodified setae. left epandrial lamella much shorter than right epandrial lamella bearing about 4 long apical setae. female unknown. comparison in smith’s (1969) key the new species runs to p. rhodesiensis smith, 1969 that was described from females found in rhodes-inyanya national park in zimbabwe. however the latter has scape and pedicel yellow. in p. manjano sp. nov. the scape and pedicel are brown. in p. rhodesiensis the scutellum is reddish yellow, but blackish on disc and sometimes completely black. in the new species the scutellum is yellow. distribution dr congo. platypalpus saffradi sp. nov. urn:lsid:zoobank.org:act:3dff850a-3bf1-4855-8109-cd7c5bc60696 figs 28–31 diagnosis a medium-sized yellow species of the minutus-group with thorax and legs yellow, but all coxae white; abdomen with tergites 1-5 brownish. no postpronotal (= humeral) seta present. one pair of vertical bristles. scape and pedicel black; postpedicel brown, 2.5 times as long as broad; stylus brown. mid tibia with long, but blunt apical spur. etymology the word ‘saffradi’ means yellow in swahili language and it refers to the yellow body colour of the new species. type material holotype ♀, dr congo, yangambi, 26 may 2013, primary mixed forest mix4 (reg. 33007, leg p. grootaert; ms name plat6). paratype 1 ♀, dr congo, 4 jun. 2013, mixed primary forest mix3, sweeping (0°47’12.80” n 24°31’24.85” e; reg. 33040; leg. p. grootaert) (rbins), mid leg extracted for dna in tube with position code ab42406119. http://zoobank.org/urn:lsid:zoobank.org:act:3dff850a-3bf1-4855-8109-cd7c5bc60696 european journal of taxonomy 103: 1–20 (2014) 14 description female length. body 2.8 mm; wing 2.8 mm. head. entirely black. occiput densely greyish pollinose; with 1 pair of long black verticals. ocellar tubercle greyish pollinose, with 2 black, very short anterior and 2 minute posterior setae. frons narrow, parallel-sided, above antennae nearly 1.5 times as broad as anterior ocellus, greyish pollinose. face very narrow, on middle narrower than anterior ocellus, somewhat broader below antennae and on lower part, greyish pollinose, clypeus subshining. antenna (fig. 30) with scape and pedicel black, postpedicel and stylus brownish; postpedicel conical, 3 times as long as wide; stylus 1.5 times as long as postpedicel. proboscis brownish yellow, rather long, somewhat shorter than head is high. palpus (fig. 29) yellow, small, ovate, with scattered pale setulae. thorax. (figs 29, 30). almost entirely reddish yellow, only metanotum brownish yellow; posterior spiracle brownish; postpronotal lobe, scutum behind postpronotal lobe, notopleuron and scutellum with some tomentosity; otherwise scutum shining, entire pleuron subshining; bristles brown. postpronotal lobe elongate ovate, with several minute setulae. mesonotum with 2 notopleurals (anterior seta very short, posterior seta moderately long), 1–2 moderately long postalars and 4 scutellars (apical pair long and cruciate, lateral pair very short); additionally, notopleuron with several setulae; acrostichals short, figs 28–31. platypalpus saffradi sp. nov., holotype ♀. 28. habitus, dorsal view. 29. head and thorax, lateral view. 30. head and thorax, dorsal view. 31. mid leg, anterior view. grootaert p. & shamshev i., new platypalpus (hybotidae) from dr congo 15 arranged in 2 narrow irregular rows, lacking on prescutellar depression; dorsocentrals arranged in 1–2 rows, nearly as long as acrostichals, 2 prescutellars somewhat longer. legs. almost entirely yellow, only last tarsomere brownish yellow. coxae and trochanters almost white, with ordinary yellowish to brownish yellow setae of different lengths. fore femur thickened; with uniformly very short anteroventral and posteroventral setae. fore tibia slender, clothed with ordinary setulae somewhat longer dorsally. mid femur (fig. 31) very thickened, long; with double row of black ventral spinules and row of long, yellow posteroventral setae, longest near middle as long as femur is wide. mid tibia bearing a row of black ventral spinules; with long blunt-tipped apical spur bearing short seta. hind femur slender, with short anteroventral setae. hind tibia slender, clothed with ordinary setulae. tarsi of all legs unmodified. wing. finely, uniformly infuscate, with brownish veins. one moderately long costal seta. veins r4+5 and m1+2 parallel just before meeting wing margin. crossveins m-cu and r-m narrowly separated; cell bm longer and distinctly broader than cell br; vein cua2 recurrent, somewhat sinuate. anal vein distinct in apical part and evanescent in basal part. calypter darkened, with brown setae. halter darkened. abdomen. with tergites 1–5 brownish, tergites 6–8 brownish yellow, subshining, covered with short brownish setae; sternites yellowish to brownish yellow, sternite 8 darker; with scattered short setae; cercus slender, brown. male unknown. comparison in the key of smith (1969), p. saffradi sp. nov. runs to p. eshowensis smith, 1969 described from zululand (south africa). in the latter species the frons is wider (2.5 times ocellar width), face below antenna about 1.5 times ocellus and there is a distinct fine humeral seta. in p. saffradi sp. nov. the frons is narrower (only 1.5 times the width of anterior ocellus), face is almost linear and there is no distinct humeral seta. platypalpus eshowensis has a long sharp apical spur on mid tibia, while it is long, but blunt in p. saffradi sp. nov. with a black bristle at tip. finally the apical two tarsomeres of all legs are darkened while in p. saffradi sp. nov. only the apical half of the apical tarsomere is brownish. platypalpus saffradi sp. nov. is provisionally placed in the minutus-group because of the presence of a well-developed postpronotal lobe lacking however a distinct postpronotal seta. mesonotum is polished and mid tibia bears a long apical spur. distribution dr congo. platypalpus yangambensis sp. nov. urn:lsid:zoobank.org:act:c6bf7e87-a8e1-4d0e-8ec0-65366caebc2f figs 32–38 diagnosis a medium-sized black species of the minutus-group with two pairs of indistinct vertical bristles. scutum shining black; pleura brownish black dusted but katepisternum largely polished. legs yellow but mid femur with narrow ventral brown stripe that is shortly interrupted near middle and hind femur a black ring in middle. mid tibia half as long as femur, with a strong sharp spur. http://zoobank.org/urn:lsid:zoobank.org:act:c6bf7e87-a8e1-4d0e-8ec0-65366caebc2f european journal of taxonomy 103: 1–20 (2014) 16 etymology the specific epithet refers to the type locality, yangambi. type material holotype ♂, dr congo, yangambi, 27 may 2013, primary mixed forest, path to gil4 sweeping from 9.05 am to 9.45 am (0°47’32.65” n 24°31’42.33” e; reg. 33010; leg. p. grootaert; ms name plat1). paratypes 1 ♂ from same provenance as holotype (rbins); 1 ♂ with same provenance as holotype in tray entomo001 at position e4, completely extracted with barcode ab48937135 (rbins); 1 ♂ with same provenance as holotype with mid leg in tray entomo004 at position a3 with barcode ab42406113 has been extracted for dna (rbins); 1 ♀, yangambi, 26 may 2013, primary mixed forest, mix4 (0°47’12.80” n 24°31’24.85” e; reg. 33007; leg. p. grootaert) (rbins); 1 ♂, yangambi, 27 may 2013, primary mixed forest, sweeping along path to mix4 from 2.15 pm to 3.05 pm (0°48’04.69” n 24°31’37.87” e; reg. 33015, leg. p. grootaert) male completely extracted in tray entomo001 at position e3 with barcode ab48937141 (rbins). the coi barcodes are available in genbank with accession numbers: kj768330, kj768331 and kj768332. figs 32–35. platypalpus yangambensis sp. nov., paratype ♂. 32. habitus, lateral view. 33. head and thorax, lateral view. 34. wing. 35. mid leg, anterior view. grootaert p. & shamshev i., new platypalpus (hybotidae) from dr congo 17 description male length. body 3.0 mm; wing 2.4 mm. head. entirely black in ground-colour. occiput entirely faintly greyish pollinose; with 2 pairs of dark very short verticals (hardly distinguished from other occipital setae), further clothed with short bristly hairs, longer setae on lower part. ocellar tubercle greyish pollinose, with 2 dark, short anterior and 2 minute posterior setae. frons widened toward ocellar tubercle, above antennae nearly 1.5 times as broad as anterior ocellus, shining. face narrow, greyish pollinose, clypeus subshining. antenna (fig. 33) with scape and pedicel black, postpedicel brown; postpedicel conical, about 3.0 times as long as wide; stylus dirty white, nearly 1.5 times as long as postpedicel. proboscis brownish, long, nearly as long as head is high. palpus (fig. 33) yellow, ovate, short, with scattered setulae and a long, brown subapical seta. thorax. (fig. 33). largely brown, postalar ridge yellow, anepisternum (= mesopleuron) along posterior margin, katepisternum (= sternopleuron) on upper posterior corner and anepimeron (= pteropleuron) and meron (= hypopleuron) yellowish brown; scutum (except notopleural depression) shining, sternopleuron with large shining spot; long bristles brownish. postpronotal lobe moderately large, with 1 short, fine seta and several setulae. mesonotum with 2 notopleurals (posterior longer), 1 short postalar and 4 scutellars (apical pair long and cruciate, lateral pair very short); acrostichals very short, arranged in 2 almost regular, very broadly spaced rows; dorsocentrals uniserial, nearly as long as acrostichals. legs. almost entirely yellow; fore and hind coxae and trochanters almost white; mid coxae yellowish; mid femur with narrow ventral brown stripe (interrupted before middle), hind femur with broad brownish ring on apical part leaving narrow subapical space yellow, knees of mid and hind legs brownish. coxae and trochanters with ordinary yellowish to brownish setae of different lengths. fore femur moderately thickened, with minute pale brown anteroventral and posteroventral setae; a strong brown posterior seta near base of femur. fore tibia very thickened, spindle-shaped, clothed with ordinary setulae. mid femur figs 36–38. platypalpus yangambensis sp. nov., paratype ♂. 36. right epandrial lamella. 37. epandrium with cerci, dorsal view. 38. left epandrial lamella. scale bars = 0.1 mm. european journal of taxonomy 103: 1–20 (2014) 18 (fig. 35) about 1.5 times stouter and almost twice longer than fore femur; with double row of black ventral spinules and row of long, brown posteroventral setae. mid tibia about half as long as femur, with long pointed apical spur bearing 2 subapical setulae of different lengths. hind femur slender, with short anteroventral setae. hind tibia slender, clothed with ordinary setulae. tarsi of all legs unmodified. wing. (fig. 34). finely, uniformly faintly infuscate, with brownish veins. one short black costal seta. veins r4+5 and m1+2 convergent just before meeting wing margin. crossveins m-cu and r-m separated; bm longer and slightly broader than br. vein cua2 somewhat sinuate, not recurrent. anal vein distinct in apical part and evanescent in basal part. calypter brown, with brown setae. halter darkened. abdomen. with tergites brown, subshining, covered with very short setae; sternites somewhat paler, with similar setation; segment 8 with several moderately long yellowish brown posteromarginal setae. terminalia (figs 32, 36–38) moderately large, brown. both cerci equally long, fine digitiform, not protruding from epandrial lamellae. right and left epandrial lamellae equally long. female palpus elongate, about half as long as labrum, pointed. fore tibia almost slender, somewhat stouter basally. abdomen with tergites 1–5 brown, subshining, covered with scattered setulae, tergites 6–8 paler, sternites pale; cercus narrow, brownish. comparison this species does not resemble any of smith’s species (1967a, 1967b, 1969). it might be compared with p. gracilipes smith, 1967 described from tanzania (smith 1967a) but in the latter the proboscis is short, less than half head height while in the new species the proboscis is as long as head is high. distribution dr congo. key to platypalpus of dr congo 1. thorax with at least pleura yellow or reddish yellow ....................................................................... 2 – thorax dark brownish or black ......................................................................................................... 3 2. stylus white (albiseta-group) with postpedicel 5.3 times as long as broad; mid tibia with pointed apical spur shorter than tibia is wide ....................................................................p. manjano sp. nov. – stylus brown with postpedicel 2.5 times as long as broad; mid tibia with long blunt apical spur longer than tibia is wide ........................................................................................ p. saffradi sp. nov. 3. mid tibia with long sharp apical spur ................................................................................................ 4 – apical spur on mid tibia shorter than tibia is wide. species from the albiseta-group ...................... 5 4. mid femur darkened above ..................................................................................... p. bolikoi sp. nov. – mid femur yellow, ventrally with dark brown black stripe along the double row of spinules ............ ..................................................................................................................... p. yangambensis sp. nov. 5. postpedicel 5.3 times as long as wide; stylus 1.6 times as long as postpedicel .........p. ikoso sp. nov. – postpedicel nearly 4 times as long as wide; stylus almost 2.5 times as long as postpedicel ............... ............................................................................................................................... p. lokonda sp. nov. grootaert p. & shamshev i., new platypalpus (hybotidae) from dr congo 19 acknowledgements we thank dr erik verheyen coordinator of rbins’s part of cobimfo as well as prof. benjamin dudu akaibe (university of kisangani) for their support. further the first author thanks mr thierry bouyer (rbins), dr jean lambert wetsi lofete (university of kisangani) and mr lola boliko (inera) for their help in the field. last but not least, we thank dr adrian plant and dr andreas stark for their many useful comments. references becker t. 1914. diptères nouveaux récoltés par mm. ch. alluaud et r. jeannel en afrique orientale 1911-1912. annales de la société entomologique de france 83:120–130. collin j.e. 1922. empididae from the seychelles. the entomologist’s monthly magazine 58: 184–189. engel e.o. 1939. empididae. in: lindner e. (ed.) die fliegen der palaearktischen region. bd. iv: 41–104. schweizerbart’sche verlagsbuchhandlung, stuttgart. grootaert p. & chvála m. 1992. monograph of the genus platypalpus (diptera: empidoidea, hybotidae) of the mediterranean region and the canary islands. acta universitatis carolinae biologica 36, univerzita karlova, praha. grootaert p. & shamshev i. 2006. the genus platypalpus macquart (diptera: hybotidae) from northeast thailand with comments on the species groups in the oriental region. journal of natural history 39 (47): 4031–4065. http://dx.doi.org/10.1080/00222930500533781 grootaert p. & shamshev i.v. 2012. the fast-running flies (diptera, hybotidae, tachydromiinae) of singapore and adjacent regions. european journal of taxonomy 5: 1–162. http://dx.doi.org/10.5852/ ejt.2012.5 grootaert p. & i. shamshev 2013. the flies of the family hybotidae (diptera, empidoidea) collected during the boyekoli ebale congo 2010 expedition in democratic republic of congo. zootaxa 3603: 1–61. http://dx.doi.org/10.11646/zootaxa.3603.1.1 international commission on zoological nomenclature (iczn) 1963. opinion 678. the suppression under the plenary powers of the pamphlet published by meigen, 1800. bulletin of zoological nomenclature 20: 339–342. loew h. 1858. bidrag till kännedomen om afrikas diptera. öfversigt af kongliga vetenskaps-akademiens förhandlingar 14 (9): 337–383. macquart j. 1827. insectes diptères du nord de la france. vol. 3. platypézines, dolichopodes, empides, hybotides. imprimerie de l. danel, lille. http://dx.doi.org/10.5962/bhl.title.8146 meigen j.w. 1800. nouvelle classification des mouches à deux ailes (diptera l.) d’après un plan tout nouveau. perronneau, paris. smith k.v.g. 1967a. chapter ix. diptera (brachycera): empididae. in: hanström b., brinck p. & rudebeck g. (eds) south african animal life: results of the lund university expedition in 1950-1951. vol. 13: 215–233. almqvist & wiksell, stockholm. smith k.g.v. 1967b. afrikanische empididae (dipt.). stuttgarter beiträge zur naturkunde 179: 1–16. smith k.v.g. 1969. the empididae of southern africa (diptera). annals of the natal museum 19, natal museum, pietermaritzburg. westwood j.o. 1840. order xiii. diptera aristotle. (antliata fabricius. halteriptera clairv.). in: westwood j.o. (ed.) an introduction to the modern classification of insects; founded on the natural habits and corresponding organisation of the different families. vol. 2. synopsis of the genera of british http://dx.doi.org/10.1080/00222930500533781 http://dx.doi.org/10.5852/ejt.2012.5 http://dx.doi.org/10.5852/ejt.2012.5 http://dx.doi.org/10.11646/zootaxa.3603.1.1 http://dx.doi.org/10.5962/bhl.title.8146 european journal of taxonomy 103: 1–20 (2014) 20 insects: 125–154. longman, orme, brown, green and longmans, london. http://dx.doi.org/10.5962/ bhl.title.12455 yang d., zhang k., yao g. & zhang j. 2007. world catalog of empididae (insecta: diptera). china agricultural university press, beijing. manuscript received: 3 february 2014 manuscript accepted: 29 august 2014 published on: 25 november 2014 topic editor: koen martens desk editor: charlotte thionois printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark. http://dx.doi.org/10.5962/bhl.title.12455 http://dx.doi.org/10.5962/bhl.title.12455 the primary types of some species of centris bees described by european entomologists in the 18th and 20th centuries (hymenoptera: apidae) felipe vivallo hymn laboratório de hymenoptera, departamento de entomologia, museu nacional, universidade federal do rio de janeiro, quinta da boa vista, são cristóvão 20940‒040 rio de janeiro, rj, brazil. email: fvivallo@yahoo.com urn:lsid:zoobank.org:author:ac109712-1474-4b5d-897b-1ee51459e792 abstract. in this paper are presented notes on the primary types of some species of the oil-collecting bees of the genus centris described by the european naturalists and entomologists amédée louis michel lepeletier de saint-fargeau, anders christian jensen-haarup, arthur louis marie joseph vachal, charles émile blanchard, embrik strand, félix édouard guérin-méneville, guillaume-antoine olivier, jean guillaume audinet-serville, jean pérez, johann christoph friedrich klug, johann ludwig christ, john obadiah westwood, josef anton maximilian perty, jules dominique, karl hermann konrad burmeister, karl wilhelm von dalla torre, massimiliano spinola, peter cameron, and wilhelm ferdinand erichson. information on the type status, type locality and depository are provided. in order to stabilize some names, lectotype designations were made for centris rhodophthalma, c. sponsa var. asuncionis, c. transversa, hemisia byssina and ptilotopus americanus. centris sponsa var. asuncionis is withdrawn from the synonymy of c. sponsa, revalidated and raised to species level. centris byssina is proposed as nomen oblitum and as a new junior synonym of c. trigonoides, nomen protectum. keywords. anthophila, biodiversity, oil-collecting bees, solitary bees, systematics. vivallo f. 2023. the primary types of some species of centris bees described by european entomologists in the 18th and 20th centuries (hymenoptera: apidae). european journal of taxonomy 864: 1–27. https://doi.org/10.5852/ejt.2023.864.2083 introduction centris fabricius, 1804 is one of the new world solitary bee lineages with the highest species richness and one of the largest distribution ranges (moure et al. 2007). this great specific diversity results in a complex taxonomic history, both of its subgenera and of the species that have been described. in recent years, a series of taxonomic works on the species of the genus have been published (vivallo 2016, 2019a, 2019b, 2019c, 2020a, 2020c, 2020e, 2020f, 2020g, 2020h, 2020i, 2020j) which allowed the stabilization of the nomenclature of the group and the proposition of new synonyms and revalidations. 1 european journal of taxonomy 864: 1–27 issn 2118-9773 https://doi.org/10.5852/ejt.2023.864.2083 www.europeanjournaloftaxonomy.eu 2023 · vivallo f. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:4f73ab5f-a4b7-43ad-949e-a40ffeeabf4f mailto:fvivallo%40yahoo.com?subject= https://zoobank.org/urn:lsid:zoobank.org:author:ac109712-1474-4b5d-897b-1ee51459e792 https://doi.org/10.5852/ejt.2023.864.2083 https://orcid.org/0000-0002-4487-0804 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ https://zoobank.org/urn:lsid:zoobank.org:pub:4f73ab5f-a4b7-43ad-949e-a40ffeeabf4f the vast majority of the species in this group was described by european entomologists, who had access to the material that was collected in america and taken to europe through scientific expeditions or by private collectors who later marketed their specimens (vivallo 2020c). this material was deposited in the most important museums of the continent, being available to specialists such as frederick smith (1805‒1879), amédée louis michel lepeletier de saint-fargeau (1770‒1845), sándor mocsáry (1841‒1915), heinrich friese (1860‒1948), johan christian fabricius (1745‒1808), adolf ducke (1876‒1959), giovanni gribodo (1846‒1924), curt schrottky (1874‒1937), and theodore dru alison cockerell (1866‒1948), among others. the primary types of centris described by those authors were studied by vivallo (2019a, 2019b, 2020a, 2020c, 2020e, 2020g, 2020h, 2020i, 2020j), providing taxonomic stability to the group. continuing this line, the primary types of the species described by the european entomologists anders christian jensen-haarup, arthur louis marie joseph vachal, charles émile blanchard, embrik strand, félix édouard guérin-méneville, guillaume-antoine olivier, jean pérez, johann christoph friedrich klug, johann ludwig christ, john obadiah westwood, josef anton maximilian perty, jules dominique, karl hermann konrad burmeister, karl wilhelm von dalla torre, massimiliano spinola, peter cameron, wilhelm ferdinand erichson, and jean guillaume audinet-serville with his friend amédée louis michel lepeletier de saint-fargeau are here studied, providing notes on the type status and depository of the type material. material and methods all labels are here considered whitish and rectangular and the data contained on them is black, handwritten or printed unless otherwise indicated. the specific features of the labels, like coloration or type of writing, are given in square brackets ([]). the backward slash (\) indicates other labels on the pin of the same specimen, and two backward slashes (\\) indicate the information on the reverse of the label, all quoted verbatim. as a result of aging, some originally white labels now have a slightly yellowish coloration. to differentiate them from current white labels, they are indicated as “yellowish white”. photographs were enhanced and organized in plates using photoshop (ver. cs2). the specimens cited are housed in the following collections: macn = museo argentino de ciencias naturales, buenos aires, argentina mhnn = muséum d’histoire naturelle de nantes, nantes, france mnhn = muséum national d’histoire naturelle, paris, france mnrj = museu nacional, universidade federal do rio de janeiro, rio de janeiro, brazil msnt = museo regionale di scienze naturali di torino, torino, italy nhmd = natural history museum of denmark, copenhagen, denmark nhmuk = natural history museum, london, united kingdom oumnh = oxford university museum of natural history, oxford, united kingdom zmb = museum für naturkunde, berlin, germany zsm = zoologische staatssammlung münchen, munich, germany. in most cases the authors cited here did not explicitly indicate how many specimens they used in the species descriptions and this cannot be inferred objectively from the original descriptions. following the recommendation 73f of the international code of zoological nomenclature (1999, iczn henceforth), i have considered all specimens as from a syntype series, even if only a single exemplar was found in the collections studied. exemplars designated lectotypes will be properly labeled as such, as well as eventual paralectotypes. results recognition of the type specimens the primary types of the species studied were recognized following the information indicated in their respective original descriptions, place of deposit, as well as the data on labels of the specimens. both european journal of taxonomy 864: 1–27 (2023) 2 pieces of information were contrasted in order to verify the existence of inconsistencies that could interfere in the recognition of the primary types. charles émile blanchard charles émile blanchard (1819‒1900) was one of the most famous french zoologists and entomologists of the 19 th century. in 1833, when he was only 13 years old, blanchard began to frequent the laboratory of the mnhn, thanks to the help of the famous french naturalist and entomologist jean victoire audouin (1797–1841), who taught entomology at that institution. at the museum, blanchard became technician in 1838, and a few years later assistant-naturalist. he published valuable works in various zoological groups, which allowed him to occupy the chair of natural history of crustaceans, arachnids and insects. in 1860, he began to lose his sight and became blind in 1890. during this period, he gradually restricted access to the collections to amateurs, triggering a general decline in the museum’s activities and, simultaneously, the dispersion of the collections. blanchard died in paris at the age of 80. blanchard’s centris bee blanchard described only one taxon in the genus centris, the species c. langsdorfii blanchard, 1840 in homage to the german naturalist, explorer and russian diplomat georg heinrich von langsdorff (1774–1852). blanchard did not indicate the sex or the number of specimens studied for the description of this species, although based on the data mentioned, they must correspond to one or more females collected in brazil. the specific locality where the material came from, as well as its collector, are unknown. it is possible that it was collected by langsdorff himself during his stay as a russian consul in rio de janeiro, brazil. from there, he organized expeditions to minas gerais state, between 1813 and 1820, and to the amazon between 1825 and 1829. considering the current known distribution range of this species, its type material was probably collected during the first expedition in southeastern brazil. unfortunately, the current whereabouts of the type material of c. langsdorfii are unknown. possibly, it was lost during the dispersal of the collections of the mnhn between 1860 and 1892. class insecta linnaeus, 1758 order hymenoptera linnaeus, 1758 family apidae latreille, 1802 genus centris fabricius, 1804 centris langsdorfii blanchard, 1840 centris langsdorfii blanchard, 1840: 405 (spelled langsdorsii [sic] in the text and langsdorfii in the figure legend). type data syntypes female, whereabouts unknown. type locality brazil. karl hermann konrad burmeister karl hermann konrad burmeister (1807–1892) was a german naturalist, zoologist, entomologist, herpetologist, botanist and geologist. he was nationalized argentinean and developed most of his career in that country. in 1850, burmeister traveled to brazil, visiting rio de janeiro and minas gerais states. vivallo f., the centris bees described by european entomologists 3 european journal of taxonomy 864: 1–27 (2023) 4 in the latter, he went to lagoa santa spending a season in company of the danish naturalist and father of brazilian paleontology and archeology peter wilhelm lund (1801–1880). from 1862 to 1892, burmeister was director of the argentinean museum of natural sciences, currently known as museo argentino de ciencias naturales (macn). he died in buenos aires in 1892, at the age of 85. burmeister’s centris bees burmeister described numerous species of flora and fauna from argentina, four of them being bees of the genus centris. three of these species were described based on series of specimens of both sexes collected mainly in mendoza and buenos aires. although burmeister did not indicate the name of the collector of those specimens, it is very likely that they were collected by him during his many trips into the interior of the country. some specimens of the primary series bear labels of holotype, paratype or allotype. however, they were labeled as such after the description of the species and not by burmeister, since the handwriting is different, and mainly because the concept of paratype and allotype was not established in burmeister’s time. all type specimens are housed in the macn. centris muralis burmeister, 1876 centris muralis burmeister, 1876: 162–163. type data this species was described based on specimens of both sexes collected in mendoza and rio negro, argentina. the lectotype male was designated by moure (1960a) and it has the following data label: [light green label] carm. patag. [printed]\ [red-rimmed white yellowish label] 223 [handwritten]\ [light blue label] 100 [printed]\ [black-rimmed pink label] centris [handwritten] holotypus [printed] muralis burm [handwritten]\ [white yellowish label] lectotipo desig moure, 1960 [handwritten in blue] (macn). paralectotype female with the following data label: [red-rimmed white yellowish label] 223 [handwritten]\ [light green label] mendo-za. [printed]\ [black-rimmed white yellowish label] muralis burm. mendozae [handwritten]\ [black-rimmed pink label] centris [handwritten] allotypus [printed] muralis burm [handwritten] (macn). in the macn there is a conspecific (metander) male without collection locality but with the same number “223” present in the lectotype and the paralectotype specimens. this possible additional paralectotype has the following data label: [red-rimmed white yellowish label] 223 [handwritten]\ [black-rimmed light yellowish orange label] centris [handwritten] paratypus [printed] muralis burm. [handwritten] (macn). type locality argentina: buenos aires province, carmen de patagones. centris nigriventris burmeister, 1876 centris nigriventris burmeister, 1876: 165. vivallo f., the centris bees described by european entomologists 5 type data this species was proposed based on an undetermined number of specimens of both sexes collected in mendoza and buenos aires. two males and one female belonging to the type series were examined, the latter being here designated as lectotype. the chosen specimen has the following data label: [red-rimmed white yellowish label] 224 [handwritten]\ [light green label] buen. ayres. [printed]\ [black-rimmed white yellowish label] nigriventris burm. rep. arg. [handwritten]\ [blackrimmed pink label] centris [handwritten] allotypus [printed] nigriventris burm. [handwritten] (macn). paralectotype male with the following data label: [red-rimmed white yellowish label] 224. [handwritten]\ [light green label] mendo-za. [printed]\ [black-rimmed pink label] centris [handwritten] holotypus [printed] nigriventris burm. [handwritten] (macn). paralectotype male with the following data label: [red-rimmed white yellowish label] 224. [handwritten]\ [light green label] mendo-za. [printed]\ [black-rimmed light yellowish orange label] centris [handwritten] paratypus [printed] nigriventris burm. [handwritten] (macn). type locality argentina: buenos aires province, buenos aires. centris pectoralis burmeister, 1876 centris pectoralis burmeister, 1876: 161–162 (junior synonym of c. obsoleta lepeletier, 1841). type data this species was described apparently based on a single female collected in corrientes province, argentina. the holotype bears the following data label: [red-rimmed white yellowish label] 221. [handwritten]\ [light green label] döcong [undecipherable handwriting]\ [black-rimmed white yellowish label] pectoralis burm. corrient. [handwritten]\ [black-rimmed pink label] centris [handwritten] holotypus [printed] pectoralis burm [handwritten] (macn). type locality argentina: corrientes province, río guayquiraró. centris vulpecula burmeister, 1876 centris vulpecula burmeister, 1876: 164. type data this species was described based on a series composed of specimens of both sexes collected in uruguay, brazil and in the argentinean cities of mendoza and paraná. according to roig-alsina (2000), the specimen from brazil corresponds to c. tarsata smith, 1874 and those from uruguay and paraná to c. trigonoides lepeletier, 1841. the male from mendoza was designated by him as lectotype. the specimen has the following data label: [red-rimmed white yellowish label] 231. [handwritten]\ [light green label] mendo-za. [printed]\ vulpecula nobis [handwritten]\ [black-rimmed pink label] centris vulpecula burm. ♂ [handwritten] lectotypus [printed] a. roig alsina 2000 [handwritten] (macn). the paralectotypes are also housed in the same collection. european journal of taxonomy 864: 1–27 (2023) 6 type locality argentina: mendoza province, mendoza. peter cameron peter cameron (1847‒1912) was a very enthusiastic english amateur entomologist and specialist of hymenoptera. he was a very prolific, and by some considered chaotic, descriptor of species (morley 1913). his collection is currently housed at nhmuk; his type specimens are also deposited in this collection, as well as at oumnh. cameron died in new mills, england, aged 65. cameron’s centris bee cameron described only one species of centris under the name paracentris fulvohirta cameron, 1903. the description was based on an undetermined number of males, collected by the english mountaineer, explorer and illustrator edward whymper (1840‒1911). in 1880, whymper organized an expedition to ecuador, designed primarily to collect data for the study of altitude sickness and the effect of reduced pressure on the human body (chisholm 1911). the results of his journey were published in 1892, in a volume entitled “travels amongst the great andes of the equator” (bonney 1892). during his travel, whymper made a collection of amphibians and reptiles that he handed over to the belgian-british zoologist george albert boulenger (1858‒1937) at the british museum (boulenger 1882). he also made a very large collection of insects that were housed in the same institution and later studied by cameron. centris fulvohirta (cameron, 1903) paracentris fulvohirta cameron, 1903: 236 (junior synonym of centris caelebs friese, 1899). type data lectotype male (nhmuk, revised). subsequent designation: moure (1999). data label: [light yellow label] paracentris fulvohirta cam. type ecuador [handwritten]\ [red-rimmed circular label] type [printed]\ b.m. type hym. [printed] 17b.907. [handwritten]\ cameron coll. 1904‒29. [printed]\ michachi ecuador 9-10,000 feet. ed. whymper. [printed]\ [black-rimmed] lectotype fulvohirta cm. [handwritten] det. j.s. moure 19 [printed] 57 [handwritten]\ [black-rimmed] centris (paracentris) caelebs friese, 1900 f. zanella det., 1998 [printed] (nhmuk). paralectotype with the following data label: paracentris fulvohirta cam. type ecuador [handwritten]\ [red-rimmed circular label] type [printed]\ b.m. type hym. [printed] 17b.907. [handwritten]\ cameron coll. 1904‒29. [printed]\ quito [handwritten] ecuador [printed] 9000 [handwritten] feet. ed. whymper. [printed]\ c. fulvo-hirtaparatype in the collection [handwritten]\ [white label with yellow lateral margins] paralectotype [printed] paracentris fulvohirta cameron, 1903 [handwritten] (nhmuk). type locality ecuador: pichincha province, machachi (9–10 000 feet). johann ludwig christ johann ludwig christ (1739‒1813) was a german naturalist, gardener and pastor. despite being a specialist in hymenoptera, he also became interested in fruit growing, agriculture and beekeeping while he was a pastor, first in wetterau and later in kronberg, germany (wilhelm 1957). christ died in his country of origin, aged 74. vivallo f., the centris bees described by european entomologists 7 christ’s centris bee christ only contributed to the knowledge of centris describing a variety of c. flavifrons (fabricius, 1775) from brazil. currently, it is considered a junior synonym of that species. centris flavifrons brasiliana (christ, 1791) apis flavifrons brasiliana christ, 1791: 140 (junior synonym of c. flavifrons). type data this variety was proposed based on an undetermined number of females collected in brazil. unfortunately, the whereabouts of the type material are currently unknown. type locality brazil. karl wilhelm von dalla torre karl wilhelm von dalla torre (1850–1928) was an austrian naturalist. he studied mathematics and natural sciences at the universität innsbruck, where he was later hired as professor of zoology (clément 1928). he studied actively the taxonomy of hymenoptera, where he published several classic works on specific groups as well as the first catalogues of the order. dalla torre died in innsbruck, aged 77. dalla torre’s centris bee dalla torre did not describe any species of centris, but instead proposed a new name for c. thoracica smith, 1874, a junior homonym of a species previously described by lepeletier (1841). the primary type of c. thoracica smith is currently housed at nhmuk and it bears the following data label: [redrimmed circular label] type h.t. [printed]\ b.m. type hym. [printed] 17b.904. [handwritten]\ st. dom. 55.1. [printed]\ centris thoracica smith (type) [handwritten]\ nhmuk 01081396 [qr code] (nhmuk). centris domingensis dalla torre, 1896 centris domingensis dalla torre, 1896: 304 (nom. nov. for c. thoracica smith, 1874). centris thoracica – smith 1874: 370 (junior primary homonym of c. thoracica lepeletier, 1841). jules dominique jules dominique (1838–1902) was a french abbé and naturalist, specialist of lichens and insects. he was in charge of organizing the entomological collection of nantes, france, which houses a large part of his collection (bureau 1903). dominique died in his country of origin, aged 64. dominique’s centris bees dominique described three species in centris, unfortunately, all currently considered junior synonyms. the specimens he studied were collected along the maroni river by the family of constant bar (1817‒1884) a french entomologist who lived in french guiana. european journal of taxonomy 864: 1–27 (2023) 8 centris debilis dominique, 1898 fig. 1a centris debilis dominique, 1898: 59 (junior synonym of c. analis (fabricius, 1804)). type data this species was proposed based on an unknown number of males. the lectotype was designated by rasmussen et al. (2007) and it bears the following data label: [black-rimmed white yellowish label] maroni guyane-française legs e. bar [printed]\ centris (heterocentris) analis fabricius, 1804 det. rasmussen & mahé [printed]\ [red label] lecotype centris debilis dominique, 1898 des. rasmussen, 2006 [printed] (mhnn) (fig. 1a). type locality french guiana: maroni river. centris dominiquella dominique, 1898 fig. 1b centris dominiquella dominique, 1898: 59 (junior synonym of c. nitens lepeletier, 1841). type data this species was described based on an undetermined number of females. the lectotype was designated by rasmussen et al. (2007) and it has the following data label: [black-rimmed white yellowish label] maroni guyane-française legs. e. bar [printed]\ centris (c.) nitens lepeletier de s.f., 1841 det. rasmussen & mahé [printed]\ [red label] lecotype centris dominiquella dominique, 1898 des. rasmussen, 2006 [printed] (mhnn) (fig. 1b). fig. 1. data labels. a. centris debilis dominique, 1898 (lectotype ♂). b. centris dominiquella dominique, 1898 (lectotype ♀︎). vivallo f., the centris bees described by european entomologists 9 type locality french guiana: maroni river. centris zonalis dominique, 1898 fig. 2a centris zonalis dominique, 1898: 59 (junior synonym of c. laticincta (spinola, 1841)). type data dominique described this species using an undetermined number of female specimens. the lectotype was designated by rasmussen et al. (2007) and it has the following data label: [black-rimmed white yellowish label] maroni guyane-française [printed]\ centris (melacentris) insignis smith, 1854 det. rasmussen & mahé [printed]\ [red label] lectotype centris zonalis dominique, 1898 des. rasmussen, 2006 [printed] (mhnn) (fig. 2 a). paralectotype female with the following data label: [black-rimmed white yellowish label] maroni guyane-française [printed]\ centris ( melacentris) insignis smith, 1854 det. rasmussen & mahé [printed]\ [yellow label] paralectotype centris zonalis dominique, 1898 des. rasmussen, 2006 [printed] (mhnn). type locality french guiana: maroni river. wilhelm ferdinand erichson wilhelm ferdinand erichson (1809‒1848) was a german doctor and entomologist. he worked actively in different areas of zoology, mainly with beetles, which led him to be appointed curator of the coleoptera collection of the zmb (klug 1850). erichson died very young, only 39 years old. erichson’s centris bee erichson described several new species of insects, but only two in centris. when he proposed them, hemisia klug, 1807 was considered the valid name of the genus, so he placed his species in it. later, the genus name was updated to centris. centris clitelligera (erichson, 1848) hemisia clitelligera erichson, 1848: 591 (junior synonym of c. flavifrons). type data this species was proposed based on an undetermined number of females collected in guyana. unfortunately, the whereabouts of the syntypes are unknown. type locality guyana. centris varia (erichson, 1848) hemisia varia erichson, 1848: 591. european journal of taxonomy 864: 1–27 (2023) 10 type data as the previous species, erichson described this new taxon based on an undetermined number of females. the whereabouts of the syntypes are unknown. type locality guyana. félix édouard guérin-méneville félix édouard guérin-méneville (1799‒1874) was a french entomologist, famous for his work “iconographie du règne animal de g. cuvier”, a complement to the work published by the french zoologists georges cuvier (1769‒1832) and pierre andré latreille (1762‒1833). guérin-méneville founded several magazines of zoology and he was an active member of the société entomologique de france. he passed away in paris, aged 74. guérin-méneville’s centris bee guérin-méneville described a variety in the genus anthophora latreille, 1803 that actually belongs to centris. unfortunately, his contribution was unsuccessful, since the variety he proposed corresponds to a species previously described by lepeletier in 1841. centris versicolor apicalis (guérin-méneville, 1844) anthophora versicolor apicalis guérin-méneville, 1844: 455 (junior synonym of c. poecila lepeletier, 1841). type data this variety was described based on an undetermined number of females collected in cuba. the whereabouts of the syntypes are unknown. type locality cuba. anders christian jensen-haarup anders christian jensen-haarup (1863–1934) was a danish entomologist specialized in hymenoptera. in 1904, he traveled to argentina on an insect collecting expedition and spent most of his time in the province of mendoza. in 1906, he returned to argentina accompanied by his compatriot, colleague and friend peter jörgensen (1870–1937). both naturalists made extensive collections of insects, which they had to sell to cover travel costs or send to specialists for identification (rasmussen 2007). the specimens of his collection that were not sold or retained by some of their colleagues are currently housed at nhmd. jensen-haarup died in denmark, aged 71. jensen-haarup’s centris bee during his stay in mendoza, jensen-haarup collected some specimens including a new species of centris that he dedicated to laureano lyngbye, his host in the city. although he did not indicate the date of collection, the specimens were probably collected during his second visit to argentina. vivallo f., the centris bees described by european entomologists 11 centris lyngbyei jensen-haarup, 1908 fig. 2b centris lyngbyei jensen-haarup, 1908: 107–108. type data to describe this species, jensen-haarup mentioned that he studied four females collected in chacras de coria, argentina. however, the characters cited in the description belong to males, not females. the lectotype specimen was designated by zanella (2002) and it bears the following data label: [black-rimmed] centris lyngbyei n. sp. j.hrp [handwriten] jensen-haarup det. [printed]\ [black-rimmed] chacr. de coria prov. de mendoza rep. argentina jensen-haarup [handwritten]\ [black-rimmed] type coll. j = hrp. [handwritten]\ [red label] holo [handwritten] type [printed]\ [white label with black discontinuous strokes] centris tricolor ♀︎ [handwritten] 1907 friese det. [printed] fr. [handwritten]\ [pink label] lectotype centris lyngbyei jensen-haarup, 1908 f. zanella, 1999 [printed]\ zmuc 00240257 [printed] (nhmd) (fig. 2b). type locality argentina: mendoza province, chacras de coria. johann christoph friedrich klug johann christoph friedrich klug (1775‒1856) was a german entomologist. he taught medicine and entomology at the universität zu berlin (currently the humboldt-universität zu berlin) where he also worked as curator of the insect collection (gerstaecker 1856). he died in berlin, aged 80. klug’s centris bees in 1810, klug proposed ptilotopus as a new genus, including in it a single species that he described simultaneously. the species is currently considered valid, but the genus was later downgraded and included as a subgenus of centris. fig 2. data labels. a. centris zonalis dominique, 1898 (lectotype ♀︎). b. centris lyngbyei jensenhaarup, 1908 (lectotype ♂). european journal of taxonomy 864: 1–27 (2023) 12 a couple of years before that publication, klug (1808) published an article on sexual dimorphism in hymenoptera, citing some examples in several genera of wasps, ants and bees. among his comments on species of centris, he cited “byssina”, a species of hemisia that had previously been named by the german entomologist and zoologist johann karl wilhelm illiger (1775–1813) but that he never formally described. klug provided some morphological features that allow the identification of this species and the sex of the specimens he studied, resulting in its formal description. the species remained unnoticed during all these years, until a visit to the zmb where i studied the specimens described by klug, which had previously been recognized as such by another entomologist. the researcher designated one of the specimens as lectotype, but unfortunately, the nomenclatural act was never published. to formalize this action and avoid future taxonomic problems, this designation is published here, acknowledging the previous work carried out by him. centris americana (klug, 1810) ptilotopus americanus klug, 1810: 31. type data this species was described based on an undetermined number of males from an unknown locality, according to moure et al. (2007), “america”. a male studied by klug from cayenne was found at zmb being here designated lectotype. the specimen bears the following data label: [green label] cajen. dy [handwritten]\ [red label] type [printed]\ centris americana [handwritten] 1907 friese det. [printed]\ [white label with lateral red margins] lectotype [printed] ptilotopus americanus klug, 1810 [handwritten] desig. melo, 2016 [printed\ http://coll.mfn-berlin.de/u/58fa3d [qr code] (zmb). as can be seen, this specimen was already labeled as lectotype, however, this designation has remained unpublished until now. type locality french guiana: cayenne. centris byssina (klug, 1808) fig. 3 megilla byssina illiger, 1806: 142 (nom. nud.). hemisia byssina klug, 1808: 57 (nomen oblitum, new junior synonym of c. trigonoides lepeletier, 1841 (nomen protectum)). type data this species was described based on an undetermined number of males. at least part of the type series is housed at zmb. the lectotype is here designated bearing the following data label (fig. 3): [green label with black lower margin] bah. gom [handwritten]\ [white yellowish label] 1600 [printed]\ [white yellowish label] zool. mus. berlin [printed in blue ink]\ [white label with red lateral margins] lectotype [printed] hemisia byssina klug, 1808 [handwritten] desig. melo, 2016 [printed]\ http://coll.mfn-berlin.de/u/58fa36 [qr code] (zmb). type locality brazil. http://coll.mfn-berlin.de/u/58fa3d http://coll.mfn-berlin.de/u/58fa36 vivallo f., the centris bees described by european entomologists 13 comment this species was described several years earlier than centris trigonoides. however, in this case the priority principle applies (iczn 1999: article 23.9.1). this allows for prevailing usage of names when the “senior synonym has not been used as a valid name after 1899” (condition 23.9.1.1) and the junior synonym has been used “in at least 25 works, published by at least 10 authors in the immediately preceding 50 years and encompassing a span of not less than 10 years” (condition 23.9.1.2). under this circumstance, both conditions are met because the name ‘byssina’ was cited only once by illiger (1906) as “megilla byssina” (nom. nud .) and then it was completely forgotten, while c. trigonoides is by far one of the most cited centridine bees (vivallo 2019a). considering this, and according to the article 23.9, c. trigonoides must be considered the valid name of the species (nomen protectum), and c. byssina one of its junior synonyms (nomen oblitum). amédée louis michel lepeletier de saint-fargeau and jean guillaume audinet-serville amédée louis michel lepeletier de saint-fargeau (1770‒1845) was a famous french entomologist, specialist of hymenoptera and one of the pioneers in the study of centris bees (vivallo 2019a). after his death in 1845, his collection began to be fragmented (casolari & casolari moreno 1980). one part was sent to the mnhn, and the other one was split and subsequently absorbed by the collections of the english entomologist and archaeologist john obadiah westwood (1805‒1893) in the united kingdom, and of the french-italian entomologist and naturalist massimiliano spinola (1780‒1857) in italy (baker 1994). lepeletier died in france, aged 74. jean guillaume audinet-serville (1775‒1858) was a french entomologist and specialist of orthoptera. audinet-serville was a friend of lepeletier and they both contributed in the publication of the “encyclopédie méthodique. histoire naturelle. entomologie, ou histoire naturelle des crustacés, des arachnides et des insectes”. he passed away in france, aged 82. fig. 3. hemisia byssina klug, 1808 (lectotype ♂). a. frontal view. b. habitus, lateral view. scale bars: a = 0.5 mm; b = 2 mm. european journal of taxonomy 864: 1–27 (2023) 14 lepeletier & serville’s centris bee although lepeletier described several dozen species of centris, only one of them was done in partnership with serville. they included their new species in ptilotopus, considered at that time to be a different group from centris and not one of its subgenera as it is today. centris americanorum (lepeletier & serville, 1825) ptilotopus americanorum lepeletier & serville, 1825: 239 (junior synonym of c. americana). type data this species was proposed based on at least two males collected at an undetermined locality. moure & seabra (1960) found two of those specimens in the spinola collection at msnt, designating one of them as lectotype, but without providing additional information about its data labels. unfortunately, the primary type of this species was not available during the preparation of this research. type locality unknown. guillaume-antoine olivier guillaume-antoine olivier was a french naturalist and entomologist. he was a very active collector and made a lot of expeditions, he was a very close friend of the danish naturalist johan christian fabricius, who in 1804 described the genus centris. olivier died in france, aged 58. olivier’s centris bee olivier proposed several new species of bees, but only one in centris. the identity of this species was misunderstood, and it was only fixed recently by vivallo (2016). centris dimidiata (olivier, 1789) apis dimidiata olivier, 1789: 64. type data olivier described this species based on an undetermined number of females collected in french guiana. the material studied by him is currently lost; therefore vivallo (2016) designated a neotype in order to resolve taxonomic issues related to a couple of species that were proposed as junior synonyms. unfortunately, the neotype female was subsequently destroyed. it had the following data label: hymnrj 000312 [printed]\ [black-rimmed white yellowish label] coleção campos seabra [printed]\ [black-rimmed white yellowish label] porto velho guaporé brasil xi-1954 m. alvarenga, dente, pereira e werner [printed]\ [red label] neotype apis dimidiata olivier, 1789 f. vivallo des., 2016 [printed] (mnrj†). type localities original type locality: french guiana, cayenne. neotype locality: brazil, rondônia state, porto velho. jean pérez jean pérez (1833‒1914) was a french zoologist and entomologist. he worked as professor of zoology at the faculty of sciences of the university of bordeaux, france, and was honorary member of the société vivallo f., the centris bees described by european entomologists 15 entomologique de france (anonymous 1916). during his professional career, he focused on the study of solitary and social bees. pérez passed away in france, aged 80. pérez’s centris bees pérez published more than 100 articles and notes, two of them containing centris bees. the first paper was published in 1905 and it contained the description of two new species from mexico. the second article was published in 1911 and contained the description of a new species from northern chile. centris confinis pérez, 1905 centris confinis pérez, 1905: 40 (junior synonym of c. nitida smith, 1874). type data this species was described based on an undetermined number of females. a single specimen of the type series was found at mnhn that was interpreted as “type” by snelling (1984). this assumption can be considered a valid lectotype designation (iczn 1999: article 74.6). the specimen bears the following data label: type [printed in red]\ centris confinis j. p. [handwritten]\ type [handwritten with blue ink] (mnhn). type locality “mexique?”. centris rhodophthalma pérez, 1911 centris rhodophthalma pérez, 1911: 55–59. type data this species was described based on an undetermined number of females collected in chañarcillo, northern chile by the chilean naturalist carlos emilio porter (1867‒1942). a single female was found at mnhn that was cited by moure et al. (2007) as holotype. however, i here consider it as syntype, following the recommendation 73f of the iczn (1999). this specimen is here designated lectotype, and it bears the following data label: chañarcillo [handwritten]\ type [printed in red]\ centris rhodophthalma j p [handwritten] (mnhn) (fig. 4a). type locality chile: atacama region, chañarcillo. centris transversa pérez, 1905 fig. 4b centris transversa pérez, 1905: 39–40. type data this species was proposed based on specimens of both sexes collected by the french naturalist léon diguet (1859‒1926) in tehuacán, puebla state, mexico. diguet traveled to mexico where he worked as a chemical engineer. during this stay, as well as following six trips he made to this country, he collected specimens for the mnhn. one male and two females of the type series are found in that collection being european journal of taxonomy 864: 1–27 (2023) 16 one of these latter designated here lectotype. the female has the following data label: [black-rimmed light green label] museum paris mexique état de puebla env. de tehuacan l. diguet 1903 [printed]\ [black-rimmed] centris [printed] (hemisiella) transversa pérez ♀︎ [handwritten] det. snelling [printed] ’83 [handwritten]\ [light blue label] lectotype centris transversa pérez, 1905 des. f. vivallo, 2017 [printed] (mnhn) (fig. 4 b). paralectotype female with the following data label: [black-rimmed light green label] museum paris mexique état de puebla env. de tehuacan l. diguet 1903 [printed]\ centris transversa typus perez [handwritten]\ centris nitida sm. r. du burpron det. [handwritten]\ type [printed in red] (mnhn). paralectotype male with the following data label: [black-rimmed light green label] museum paris mexique état de puebla env. de tehuacan l. diguet 1903 [printed]\ [black-rimmed] centris [printed] (hemisiella) transversa pérez ♂ [handwritten] det. snelling [printed] ’83 [handwritten]\ [light blue label] paralectotype centris transversa pérez, 1905 des. f. vivallo, 2017 [printed] (mnhn). at nhmuk was found a female which apparently belonged to the type species of this species. however, it was collected in 1904 and not in 1903, as pérez cited explicitly the year of collection of his specimens; therefore, here it is not interpreted as a syntype. that exemplar has the following data label: mexique, tehuacan l. diguet 1904 [handwritten]\ schulz coll. 1908-157. [printed]\ centris transversa typus ♀︎ pérez [handwritten]\ [black-rimmed red label] spec. typ. [printed]\ [red-rimmed circular label] type [printed]\ b.m. type hym. [printed] 17b905 [handwritten] (nhmuk). fig. 4. data labels. a. centris rhodophthalma pérez, 1911 (lectotype ♀︎). b. centris transversa pérez, 1905 (lectotype ♀︎). vivallo f., the centris bees described by european entomologists 17 type locality mexico: puebla state, tehuacán. josef anton maximilian perty josef anton maximilian perty (1804–1884) was a german naturalist and entomologist. he was a professor of zoology at ludwig-maximilian-universität münchen, and at universität bern, switzerland (hess 1887). throughout his professional life, perty published numerous articles on a wide variety of topics. all of them testify of a strictly scientific education, a wide knowledge of literature and a keen capacity for observation; but unfortunately, especially in his later years, also of a tendency towards miraculous and spiritualistic visions (hess 1887). perty passed away in bern, aged 80. perty’s centris bees perty described a single species in centris that was later transferred to eufriesea cockerell, 1908 (apidae: euglossini), and three species in xylocopa latreille, 1802 (apidae: xylocopini) that were later transferred to centris. the type specimens of these species were collected by the german naturalist johann baptist von spix (1781–1826) and the german doctor, botanist and anthropologist carl friedrich philipp von martius (1794–1868) during an expedition to brazil. centris flavicrus (perty, 1833) xylocopa flavicrus perty, 1833: 150 (junior synonym of c. xanthocnemis). type data perty described this species based on an unknown number of specimens of an undetermined sex. the type material of this species, which corresponds to females, is currently lost. type locality brazil: piauí state. centris moerens (perty, 1833) xylocopa moerens perty, 1833: 150. type data as in the previous case, perty did not indicate how many specimens he studied to describe this species. the lectotype female was designated by moure (1960b, by inference of a holotype, iczn 1999: article 74.6) and it has the following data label: [black-rimmed green label] brasilien coll. perty [handwritten]\ [green-rimmed white yellowish label with green horizontal lines] 4. brasil. x. moerens perty [handwritten]\ [pink label] holotypus [printed] xylocopa moerens perty ♀︎ diller 1989 [handwritten] zoologische staatssammig. münchen [printed] (zsm) (fig. 4 a). type locality brazil: minas gerais state. centris xanthocnemis (perty, 1833) xylocopa xanthocnemis perty, 1833: 150. european journal of taxonomy 864: 1–27 (2023) 18 type data this species was proposed based on an undetermined number of males collected in piauí state. the lectotype was designated by moure (1960b, by inference of a holotype, iczn 1999: article 74.6) and it bears the following data label: [black-rimmed green label] piauhy. brasilien. coll. perty. [handwritten]\ [green-rimmed white yellowish label with green horizontal lines] 5. bras. piauhy x. xanthocnemis pty. [handwritten]\ [pink label] lectotypus [printed] xylocopa xanthocnemis perty ♀︎ diller 1989 [handwritten] zoologische staatssammlg. münchen [printed] (zsm) (fig. 4b). type locality brazil: piauí state. massimiliano spinola massimiliano spinola was a french-italian doctor, naturalist and entomologist. he descended from a rich and powerful family from italy, with lands in europe and south america, from which he received many insects. thanks to his wealth, spinola made extensive purchases of insect collections (gestro 1915), which allowed him to make important contributions to the taxonomy of the orders coleoptera, hemiptera and hymenoptera. he passed away in tassarolo, italy, aged 77. spinola’s centris bees spinola proposed six new species in hemisia, at that time considered the valid name for centris. according to moure et al. (2007) the primary types of four of those species are housed at msnt, but unfortunately, they were not available during the preparation of this article. centris bombiformis (spinola, 1841) hemisia bombiformis spinola, 1841: 148–149 (junior synonym of c. americana). type data this species was described based on a female specimen. moure & seabra (1960) mentioned a male from pará state, brazil, housed at mnhn as the probable type of this species, which disagrees with the information provided by spinola. however, that specimen was not found at mnhn, despite active search. the current condition and depository of the primary type is unknown. type locality french guiana: cayenne. centris chilensis (spinola, 1851) hemisia chilensis spinola, 1851: 167. type data spinola described this species using an undetermined number of females collected in the coquimbo mountain range, chile. the whereabouts of the syntypes are unknown. according to zanella (2002), they are also probably housed at msnt (not examined). type locality chile: coquimbo region. vivallo f., the centris bees described by european entomologists 19 centris laticincta (spinola, 1841) hemisia laticincta spinola, 1841: 148. type data spinola proposed this species using two females from french guiana. the syntypes are at msnt (not examined). type locality french guiana: cayenne. centris leprieuri (spinola, 1841) hemisia leprieuri spinola, 1841: 146–147 (junior synonym of c. decolorata lepeletier, 1841). type data this species was based on a single female collected in cayenne, french guiana. the holotype is housed at msnt (not examined). type locality french guiana: cayenne. centris nigerrima (spinola, 1851) hemisia nigerrima spinola, 1851: 167. fig. 5. data labels. a. xylocopa moerens perty, 1833 (lectotype ♀︎). b. xylocopa xanthocnemis perty, 1833 (lectotype ♂). european journal of taxonomy 864: 1–27 (2023) 20 type data this species was described based on specimens of both sexes collected in northern chile. the syntypes are at msnt (not examined). type locality chile: coquimbo region, coquimbo. centris pyropyga (spinola, 1841) hemisia pyropyga spinola, 1841: 148 (junior synonym of c. nobilis westwood, 1840). type data spinola described this species using a single female specimen. the holotype is currently housed at msnt (not examined). type locality french guiana: cayenne. embrik strand embrik strand (1876–1947) was a norwegian botanist, entomologist, arachnologist and naturalist. between 1901 and 1903, he worked as curator in the museum of zoology of the universitetet i kristiania, currently universitetet i oslo (university of oslo), norway. later he moved to germany to continue his studies in zoology. in norway, strand worked at different museums and universities until 1923, when he moved to latvia. there he worked as professor of zoology at the rīgas universitāte, currently latvijas universitāte (university of latvia) (natvig 1944). specimens of the strand’s collection can be found in the zoological museum of the university of oslo, and in the zmb. strand passed away in riga, latvia, aged 71. strand’s centris bee strand described a single taxon in centris, the variety c. sponsaasuncionis strand, 1910. the description was based on specimens collected by the hungarian pharmacist, biologist, botanist and researcher jános dániel anisits (1856‒1911). anisits migrated to paraguay after he graduated as a pharmacist in budapest (vivallo 2020d). in this country he worked actively, collecting biological material and discovering several new species, mainly plants (magyarország és latin-amerika 2019). this variety remained forgotten until it was cited by moure et al. (2007) as junior synonym of c. sponsa smith, 1854. despite the similarity between them, they seem to be different species. here, it is proposed to withdraw c. sponsa asuncionis from the synonymy of c. sponsa, revalidate it and raise it to species level. centris asuncionis strand, 1910 nom. rev. centris sponsa var. asuncionis strand, 1910: 521–522. type data strand proposed this variety based on several specimens of both sexes collected in the cities of asunción and sapucai, in se paraguay. two males and one female of the type series were found at zmb, the latter vivallo f., the centris bees described by european entomologists 21 being designated lectotype. the female specimen bears the following data label: [black-rimmed white yellowish label] asuncion, paraguay j.d. anisits [printed] villa morra 19.ii.06 [handwritten]\ centris sponsa var. asuncionis m. [handwritten] strand det. [printed]\ [black-rimmed red label] type [printed]\ [white label with lateral red margins] lectotype [printed] centris sponsa asuncionis strand, 1910 [handwritten] desig. melo, 2016 [printed]\ http://coll.mfn-berlin.de/u/58fa33 [qr code] (zmb). paralectotype male with the following data label: asuncion, paraguay j.d. anisits [printed] 10.ii.06 [handwritten]\ centris sponsa v. asuncionis m. [handwritten] strand det. [printed]\ [white label with lateral yellow margins] paralectotype [printed] centris sponsa asuncionis strand, 1910 [handwritten] desig. melo, 2016 [printed] (zmb). paralectotype male with the following data label: asuncion, paraguay j. d. anisits [printed] sapucay xii.04 [handwritten]\ [black-rimmed red label] type [printed]\ centris sponsa v. asuncionis m. [handwritten] strand det. [printed] (zmb). the lectotype and one paralectotype were previously labeled as such, but those designations remained unpublished. i agree with that interpretation and maintain both specimens as such. type locality paraguay: distrito capital, asunción (villa mora). arthur louis marie joseph vachal best known as joseph vachal (1838–1911), he was a french notary, politician, entomologist, and the 5 th most prolific describer ever of new bee taxa (rasmussen 2012). thanks to his contact with the french entomologist and carcinologist eugène louis bouvier (1856‒1944), former chair of entomology at the mnhn, vachal started working at the museum, deciding that after his death, his entire entomological collection would be donated to that institution (rasmussen 2012). he passed away in argentat, france, at the age of 72. vachal’s centris bee vachal described centris autrani vachal, 1904 based on specimens collected by the french naturalist gustave-adolphe baer (1839‒1918) between 1902 and 1903 in northern argentina. the name of this species was in homage of his friend and colleague the austrian botanist eugène john benjamin autran (1855‒1912), at that time, a senior official of the ministry of agriculture of argentina. centris autrani vachal, 1904 centris autrani vachal, 1904: 16 (junior synonym of c. flavohirta friese, 1899). type data vachal described this species based on two females and one male from argentina and one female from arica, chile. apparently, this type series was composed by at least two species, because c. autrani (= c. flavohirta) does not occur in chile (vivallo 2020b). unfortunately, the female from chile was not found during the preparation of this paper. the rest of the type series, including the lectotype male designated by zanella (2002) is housed at mnhn. the specimen bears the following data label: [yellowish label] argentine prov. tucuman lara 4000m g. a. baer, 2-1903 [printed]\ [yellowish label] museum paris [printed] tucuman a. baer 1903 [handwritten]\ [yellowish label] centris autrani 1903 n. sp j. vachal [handwritten]\ [red label] type [printed]\ [white label] lectotype centris autrani vachal f. zanella, 2002 [printed] (mnhn). european journal of taxonomy 864: 1–27 (2023) 22 despite vachal mentioning only four specimens in the original description, another six males with the same original label of baer were found at mnhn. those specimens were labeled as paralectotypes by zanella (2002). type locality argentina: tucumán province, lara. john obadiah westwood john obadiah westwood (1805‒1893) was an english entomologist, archaeologist and the first appointed curator of the hope entomological collections at the oxford university museum of natural history (anonymous 1893). westwood was one of the first entomologists with an academic position at the university of oxford. he donated to that institution his insect collection which was later incorporated to the hope collection (wandolleck 1894). westwood died in oxford, aged 87. westwood’s centris bee westwood described a single species in centris, based on an undetermined number of specimens that belonged to the collection of the british entomologist and naturalist frederick william hope (1797‒1862). in 1849, hope gave his entire collection of insects to the university of oxford (baker 1994), which included the specimens studied by westwood. currently, it is formally known as the hope entomological collections and it is held by the oxford university museum of natural history (oumnh). centris nobilis westwood, 1840 centris nobilis westwood, 1840: 263–264. type data westwood proposed this new species based on an undetermined number of females from an unknown locality in south america. moure & seabra (1960) studied a single female housed at oumnh that they recognized as “type”. according to the article 74.6 (iczn 1999), this assumption can be treated as a valid lectotype designation. the specimen bears the following data label: centris nobilis westw nat lib. bees p 264 pl 20 fi [handwritten] (oumnh). type locality “locality doubtful; but in all probability south america”. discussion the detailed transcription of the labels of the type material is essential to unequivocally recognize the specimen or specimens that were used to describe a certain species. in many cases, only through this procedure is it possible to detect inconsistencies between the information contained in the original descriptions and in the labels of the specimens considered primary types. the careful and detailed execution of this procedure allows confirmation of the status of the material, as well as the identity of a particular species. as is often the case with old descriptions, it is not always possible to know the number of specimens studied with only the information indicated in the original description, since often they only mention male and female characteristics in a general way. in these cases, it is prudent to consider such specimens as part of a type series and designate a lectotype. this procedure brings taxonomic stability by fixing the vivallo f., the centris bees described by european entomologists 23 name of the species. otherwise, complex problems may arise where the same name is being applied to more than one species as a result of a lack of extensive taxonomic work. this point is especially important in those cases in which the species were described from syntypes or composed of type series that ended up distributed in different collections. among the old melittologists, perhaps the most exemplary case is that of heinrich friese, who described a large number of species of centris bees mostly based on series of specimens deposited in several american and european collections (vivallo 2019b, 2020c) using a very confusing and inaccurate labeling system (rasmussen & ascher 2008). acknowledgments i want to thank arturo roig-alsina (macn), françois meurgey (mhnn), agnièle touret-alby (mnhn), lars vilhelmsen (nhmd), david notton (nhmuk), james hogan (oumnh), michael ohl (zmb), and stefan schmidt (zsm) for allowing me to visit and work in the collections under their care. i also thank michael engel for his help. this article is part of the sigma project nº21565 mn/ufrj and the contribution number 64 from the hymn. references anonymous 1893. professor john o. westwood, m.a., f.l.s., &c. geological magazine 10 (3): 143. https://doi.org/10.1017/s0016756800168736 anonymous 1916. liste chronologique des travaux scientifiques de jean pérez (1833‒1914). annales de la société entomologique de france 85: 355–366. baker d.b. 1994. type material in the university museum, oxford, of bees described by comte amédée lepeletier de saint-fargeau and pierre andré latreille (hymenoptera: apoidea). journal of natural history 28: 1189–1204. https://doi.org/10.1080/00222939400770591 blanchard e. 1840. orthoptères, neuroptères, hémiptères, hymenoptères, lépidoptères et diptères. in: laporte f.l. 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(ed.) the natural history of bees. in jardine w. (ed.) the naturalist’s library: 260–301. lizars, edinburgh. wilhelm s. 1957. christ, johann ludwig. neue deutsche biographie 3: 217. zanella f.c.v. 2002. sistemática, filogenia e distribuição geográfica das espécies sul-americanas de centris (paracentris) cameron, 1903 e de centris (penthemisia) moure, 1950, incluindo uma análise filogenética do “grupo centris” sensu ayala 1998 (hymenoptera, apoidea, centridini). revista brasileira de entomologia 46 (4): 435–488. https://doi.org/10.1590/s0085-56262002000400001 manuscript received: 12 september 2022 manuscript accepted: 28 november 2022 published on: 27 march 2023 topic editor: tony robillard section editor: gavin broad desk editor: marianne salaün printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; leibniz institute for the analysis of biodiversity change, bonn – hamburg, germany; national museum of the czech republic, prague, czech republic. https://doi.org/10.13102/sociobiology.v67i3.4968 https://doi.org/10.1590/1678-4766e2020020 https://doi.org/10.11606/1807-0205/2020.60.33 https://doi.org/10.5852/ejt.2020.618 https://doi.org/10.1590/s0085-56262002000400001 two new species of the genus deuteraphorura absolon, 1901 (hexapoda, collembola, onychiuridae) from georgian caves with remarks on the subterranean biodiversity of the caucasus mountains andrea parimuchová 1,*, shalva barjadze 2, eter maghradze 3 & ľubomír kováč 4 1,4 institute of biology and ecology, faculty of science, pavol jozef šafárik university in košice, šrobárova 2, 04154 košice, slovakia. 2,3 institute of zoology, ilia state university, giorgi tsereteli 3, 0162, tbilisi, republic of georgia. * corresponding author: andrea.parimuchova@upjs.sk 2 email: shalva.barjadze@yahoo.com 3 email: eter.magradze.1@iliauni.edu.ge 4 email: lubomir.kovac@upjs.sk 1 urn:lsid:zoobank.org:author:6533715d-0789-401a-bffd-91abdb36428b 2 urn:lsid:zoobank.org:author:ab36bef1-c006-41a3-861e-e0b79ee35fbf 3 urn:lsid:zoobank.org:author:a5ce76d6-214b-40d8-b06d-181c0ab4d7c9 4 urn:lsid:zoobank.org:author:64c6117e-db5c-4569-bf89-5c812c118760 abstract. specimens of deuteraphorura collected in 11 georgian caves were analysed morphologically and molecularly based on the coi gene barcode region. two molecular delimitation methods revealed four species (motus); however, only two of them were distinguished morphologically and are described in this paper as new to science. both new species, d. colchisi sp. nov. and d. kozmani sp. nov., belong to the group with a pseudocellus on the first thoracic tergum; the differential diagnosis table to this species group is provided. the potential of the caucasus as a hotspot region of subterranean biodiversity and evolution centre of subterranean animals is discussed. keywords. cave fauna, species delimitation, motus, hotspot, evolution centre. parimuchová a., barjadze s., maghradze e. & kováč ľ. 2023. two new species of the genus deuteraphorura absolon, 1901 (hexapoda, collembola, onychiuridae) from georgian caves with remarks on the subterranean biodiversity of the caucasus mountains. european journal of taxonomy 879: 64–82. https://doi.org/10.5852/ejt.2023.879.2161 introduction the biodiversity of caves has been one of the main topics of subterranean biology for decades. within the palearctic region, “hotspot areas” were defined in the mountains of southern europe based on the diversity of obligate cave-dwellers (troglobionts and stygobionts) (culver et al. 2004, 2006; reboleira et al. 2011). europe is traditionally a centre of speleobiological research, resulting in rich data on karst european journal of taxonomy 879: 64–82 issn 2118-9773 https://doi.org/10.5852/ejt.2023.879.2161 www.europeanjournaloftaxonomy.eu 2023 · parimuchová a. et al. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:b8be4779-ef8d-4b63-8643-e1468f306c5b 64 https://orcid.org/0000-0002-8977-9991 https://orcid.org/0000-0001-8992-4987 https://orcid.org/0000-0002-4796-9439 https://orcid.org/0000-0001-8194-2128 mailto:andrea.parimuchova%40upjs.sk?subject= mailto:shalva.barjadze@yahoo.com mailto:eter.magradze.1%40iliauni.edu.ge?subject= mailto:lubomir.kovac%40upjs.sk?subject= https://zoobank.org/urn:lsid:zoobank.org:author:6533715d-0789-401a-bffd-91abdb36428b https://zoobank.org/urn:lsid:zoobank.org:author:ab36bef1-c006-41a3-861e-e0b79ee35fbf https://zoobank.org/urn:lsid:zoobank.org:author:a5ce76d6-214b-40d8-b06d-181c0ab4d7c9 https://zoobank.org/urn:lsid:zoobank.org:author:64c6117e-db5c-4569-bf89-5c812c118760 https://doi.org/10.5852/ejt.2023.879.2161 https://creativecommons.org/licenses/by/4.0/ https://zoobank.org/urn:lsid:zoobank.org:pub:b8be4779-ef8d-4b63-8643-e1468f306c5b regions and their subterranean fauna. it has been observed that areas of maximum diversity occur in the range of ca 42°–46° in europe and 34° in the north america, but potential hotspots were also expected in the karst regions of the western caucasus in georgia (culver et al. 2006) and in south-east asia, with a high level of endemism in troglobiotic taxa (deharveng & bedos 2000). the last mentioned has been confirmed, and the region of southern china is recently the richest in troglobiotic trechini beetles (e.g., tian et al. 2016, 2017). biodiversity-rich areas were also identified in brazil (souza-silva & ferreira 2016). caves in the caspian territory of the palearctic have been almost neglected in terms of biospeleology. recent studies have made a significant contribution to revealing the subterranean diversity and documenting the great potential of this area for the discovery of the new taxa. caucasian subterranean fauna has been intensively studied mostly in georgia, resulting in regularly annotated lists of cave fauna (barjadze & djanashvili 2008; barjadze et al. 2012, 2015) with a total of 86 troglobiotic and stygobiotic species as of 2019 (barjadze et al. 2019). in the last decade, new species of collembola have been described in the genera deuteraphorura absolon, 1901 (jordana et al. 2012), arrhopalites börner, 1906 and pygmarrhopalites vargovitsh, 2009 (vargovitsh 2012, 2013, 2017, 2022), and plutomurus yosii, 1956 (barjadze et al. 2022) and the new genus troglaphorura, with highly troglomorphic species (vargovitsh 2019). unexpected diversity was documented in diplopoda (antić & makarov 2016; antić & reip 2020), and new troglomorphic taxa were described in isopoda (gongalski & taiti 2014) and opilionida, with the highly specialised species in the genus nemaspela (martens et al. 2021) and pseudoscorpionida of the genus globochtonius (zaragoza et al. 2021) . in the present contribution, we describe two new collembolan species of the genus deuteraphorura discovered during recent biological investigations in the caves of western georgia, and discuss the diversity indicated by molecular analyses. we emphasise the significance of caves in the caucasian mountains and consider this important mountain range to be a hotspot area of subterranean biodiversity. material and methods study area in georgia, the total area of the karstic rock outcrop occupies 4475 km2 with more than 1500 caves (asanidze et al. 2019). specimens of deuteraphorura from 11 relatively warm horizontal caves of low altitudes in georgia were morphologically studied and molecularly analysed (table 1). we called the cave close to prometheus and datvi cave as sakadzhia cave; however, there are doubts about the real location of this cave (barbakadze pers. com.). in this study, populations were defined as individuals collected in caves located in different karst areas of georgia with a minimum distance of ca 2.5 km between the closest caves (motena–inchkhuri) and maximum distance of ca 85 km (motena–kozmani) (fig. 1). morphological examination for morphological study, specimens were separately mounted on permanent slides in swann medium (liquido de swann) modified after rusek (1975) and studied in phase-contrast carl zeiss axio 5 microscope and leica dm 2500 microscope equipped with dil optics (differential interference contrast), a measuring eyepiece (micrometric ocular) and a drawing arm. the images were taken with an axiocam 208 color (carl zeiss) camera with zen imaging software. drawings were edited using adobe photoshop cs6. chaetotaxy of the tibiotarsus is presented after deharveng (1983), and of the labium after fjellberg (1999). parimuchová a. et al., two new cave deuteraphorura species from georgia 65 molecular data analysis and species delimitation methods one to three specimens from each population were analysed in the molecular laboratory of the department of zoology, ibe fs upjs, košice, slovakia. to prevent contamination, all dna laboratory work was conducted under sterile conditions with the use of barrier tips. total dna was extracted with the machrey-nagel nucleospin tissue kit according to the modified manufacturer´s protocol with 50 μl of elution buffer twice. a polymerase chain reaction (pcr) (saiki et al. 1988) was carried out using a 12.5 μl reaction volume consisting of 1 μl of template dna (not quantified), 10× pcr buffer (topbio), 12.5 mm of dntp mix, 5 μm of each primer and 0.125 units of taq polymerase (topbio) on a genepro (bioer co. ltd, china) thermal cycler. a fragment of the coi gene (588 bp) was amplified using the primers lco1490_jj (5’ cha cwa ayc ata aag ata tyg g-3’) and hco2198_jj (5’awa ctt cvg grt gvc caa ara atc a -3’; astrin & stüben 2008). thermal cycling conditions were as follows: 94°c for 3 min followed by 5 cycles of 94°c for 30 sec., 45°c for 1 min 30 sec. and 72°c for 1 min, followed by 35 cycles of 94°c for 30 sec., 51°c for 1 min 30 sec. and 72°c for 1 min followed by 1 min in 72°c. after verification on agarose electrophoresis, reaction products were purified using exo i/fastap (thermo fisher scientific). the sequencing of the purified products was performed using lco1490_jj by the sanger method (eurofins genomics, ebersberg, germany). in cases when the primer failed to produce high quality chromatogram, reverse primer sequencing was employed. sequences were edited and trimmed of unreadable short stretches (ca 30 bp at the 5’ and 3’ ends) with geneious prime ver. 2022.1.1 (copyright © 2005–2022 biomatters ltd). since none of the sequences contained stop codons or indels in orf, all were considered to be true mitochondrial and not nuclear copies. all the sequences were verified as consistent with onychiuridae congeners using the genbank blastn search (the mega blast algorithm with the default setting). sequences were aligned with the geneious prime ver. 2022.1.1 (copyright © 2005–2022 biomatters ltd) software by muscle (codons) algorithm using the invertebrate mitochondrial genecode and fig. 1. locations of studied caves in georgia. for cave abbreviations see table 1. european journal of taxonomy 879: 64–82 (2023) 66 default parameters. standard dna barcoding distance analysis was conducted in mega x (kumar et al. 2018) f using the tamura-3 parameter method (tamura 1992). a neighbour-joining tree (saitou & nei 1987) with tamura-3 parameter method (tamura 1992) was constructed and the robustness of the tree nodes was assessed by bootstrap analysis with 1000 replications, values under 60 are not shown. both barcoding gapand evolutionary models were applied for coi marker. assemble species by automatic partitioning (asap) method (puillandre et al. 2021) used genetic distances to propose species hypotheses. the kimura (k2p) model with default parameters was used to merge sequences into groups. the poisson tree processes (ptp) model, used for species delimitation based on the number of substitutions, was performed using on-line software (zhang et al. 2013). a maximum likelihood (ml) tree was inferred using auto substitution model and 1000 ultrafast bootstrap analysis (hoang et al. 2018) in iq-tree software (nguyen et al. 2015). correlation between geographical and genetic distances (tamura-3 parameter model, pairwise deletion option) of populations was evaluated by mantel test (999 permutations) using the genalex 6.5 program. all new sequences are available in genbank (accession numbers: oq271838–oq271861). abbreviations ant. = antennal segment abd. = abdominal tergum aoiii = antennal organ of the third antennal segment table 1. list of caves with their characteristics, and administrative and geographic location in georgia. cave characteristics according to tatashidze et al. (2009) and tsikarishvili & bolashvili (2013). names of tourist caves are bolded. abbreviations: ak = askhi karst massif; ch = chiatura; ee = entrance elevation; et = eutrophic; im = imereti; kg = kharagauli; kh = khoni; ma = martvili; mt = mesotrophic; op = odishi plateau; ot = oligotrophic; sam = samegrelo; st = sataplia-tskaltubo; ts = tskaltubo; x = site of air temperature measurements in caves is unclear; y = elevation between the entrance and the deepest site of the cave; zi = zemo-imereti plateau; * trophic level estimation is based on presence of organic material in internal parts of the cave; – = not measured. abb. cave name ee (m a.s.l.) cave length (m) cave temp. x (°c) cave depth y (m) trophic level* region district karst area xom khomuli 95 70 13.5–14 2 mt im ts st mel melouri 424 5300 12–13 15 ot im ts st sak sakadzhia 141 – – – mt im ts st ssb satsurblia 305 125 11.7 20 mt im ts st pro prometheus 147 2900 13.5–14.5 – et im ts st dat datvi 140 56 – – mt im ts st sat satevzia 215 250 – 0 mt im kh st koz kozmani 652 200 – – mt im kg zi shv shvilobisa 730 1000 12.3 10 ot im ch zi mot motena 570 95 13–13.6 14 ot sam ma ak ink inchkhuri 380 65 – – et sam ma op parimuchová a. et al., two new cave deuteraphorura species from georgia 67 ibe fs upjs = institute of biology and ecology, faculty of science, p.j. šafárik university, košice, slovakia izisu = institute of zoology, ilia state university, tbilisi, georgia ms = microsensillum mvo = male ventral organ pao = postantennal organ pso = pseudocellus psx = parapseudocellus tita = tibiotarsus th. = thoracic tergum vt = ventral tube results molecular species delimitation we employed delimitation methods based on the coi mitochondrial gene to define the molecular operational taxonomic units (motus) and assess their congruence with the current species level based on morphology and geographic distribution. we obtained alignment of 24 coi sequences with a length of 605 bp. fig. 2. a neighbour-joining tree (nj) with species delimitation of georgian cave populations of deuteraphorura absolon, 1901 based on coi molecular marker, morphology and geographic location in karst areas. numbers and coloured columns indicate groups (species) identified by particular methods asap (assemble species by automatic partitioning) and bptp (bayesian poisson tree processes). the question mark (?) indicates ambiguous result in shvilobisa cave due to low number of studied specimens. for abbreviations of caves in the nj tree see table 1. european journal of taxonomy 879: 64–82 (2023) 68 the asap method delimited four species, and the best partition had an asap score of 2.0 (p <0.05). the bptp method estimated four groups (species) with support from 0.519 to 1.0, thus corresponding to the asap delimitation (fig. 2). the distribution of k2p distances revealed a clear barcode gap. as determined with asap, specimens diverging at a k2p distance above 3% belong to different species (fig. 3). species 1 comprised specimens from most of the sataplia-tskaltubo karst caves: khomuli, satsurblia, melouri, prometheus, datvi and sakadzhia. species 2 contained specimens from caves of three karst areas represented by satevzia cave, motena cave and inchkhuri cave. species 3 comprised only a single specimen from shvilobisa cave, and species 4 consisted exclusively of specimens from kozmani cave (fig. 2). morphological character analysis was able to confirm the species status of only two of the four motus revealed by molecular delimitation methods; these two species have been given scientific names and are described taxonomically below. fig. 3. histogram of coi k2p distances between specimens of deuteraphorura absolon, 1901. the red line indicates the threshold distance above which specimens are considered to belong to different species, according to asap method. parimuchová a. et al., two new cave deuteraphorura species from georgia 69 taxonomy phylum arthropoda von siebold, 1848 subphylum hexapoda blainville, 1816 class collembola lubbock, 1870 order poduromorpha börner, 1913 family onychiuridae lubbock (in börner, 1913) subfamily onychiurinae börner, 1901 genus deuteraphorura absolon, 1901 deuteraphorura colchisi parimuchová, barjadze & kováč sp. nov. urn:lsid:zoobank.org:act:aac4848a-79d3-4a5c-8458-a8cb7bb2ea8f fig. 4, table 2 deuteraphorura sp. – zaragoza et al. 2021. etymology the name is derived from ‘colchis’ – the historical geographical, ethnical and political entity of georgia which today is located in the west of the country. type material holotype georgia • ♀; imereti, tskaltubo, satsurblia cave; 42.38805000° n, 42.60626700° e; 12 mar. 2020; eter maghradze leg.; hand collecting on wood; ibe fs upjs. paratypes georgia – imereti, tskaltubo • 3 ♀♀; khomuli cave; 42.31562° n, 42.63613° e; 11 apr. 2020; eter maghradze leg.; hand collecting on wood, guano, water surface • 1 ♀, 3 ♂♂; melouri cave; 42.38752° n, 42.62819° e; 28 may 2019; eter maghradze leg.; pitfall traps with pork liver, hand collecting on guano and speleothems; ibe fs upjs • 3 ♀♀; prometheus cave; 42.37716° n, 42.60086° e; 13 feb. 2018; eter maghradze leg.; pitfall traps with pork liver, hand collecting on wood, guano and speleothems; ibe fs upjs • 1 ♀; datvi cave; 42.37444° n, 42.59583° e; 27 dec. 2019; eter maghradze leg.; pitfall traps with pork liver; izisu • 1 ♀; same collection data as preceding; ibe fs upjs • 2 ♀♀; satsurblia cave; same collection data as for holotype; ibe fs upjs • 1 ♂; same collection data as preceding; izisu • 1 ♀, 1 ♂; sakadzhia cave; 42.36756387° n, 42.59123348° e; 28 dec. 2020; eter maghradze leg.; hand collecting on guano and detritus • 3 ♀♀, 3 ♂♂; imereti, khoni, satevzia cave; 42.43153377° n, 42.56590444° e; 10 feb. 2020; eter maghradze leg.; hand collecting on guano and water surface; • 10 ♀♀, 2 ♂♂; same collection data as preceding; izisu ibe fs upjs • 1 ♀, 1 ♂; imereti, chiatura, shvilobisa cave; 42.3254° n, 43.26786° e; 8 oct. 2021; eter magradze, shalva barjadze, lado shavadze, mariam gogshelidze leg.; hand collecting on guano, wood and detritus; ibe fs upjs • 3 ♀♀; samegrelo, martvili, inchkhuri cave; 42.45678637°n, 42.40425674°e; 18 jul. 2020; 10 jul. 2021; eter maghradze, shalva barjadze, lado shavadze, mariam gogshelidze leg.; hand collecting guano, wood, water surface and walls; izisu • 5 ♀♀; same collection data as preceding; ibe fs upjs • 1 ♀; motena cave; 42.47657295° n, 42.39126228° e; 10 jul. 2021; shalva barjadze, lado shavadze, mariam gogshelidze leg.; hand collecting on walls; ibe fs upjs . description body length 1.3–2.3 mm in females, 1.85–2.1 in males (average 1.78 mm; n = 46), shape cylindrical (fig. 4a). colour white to pale brownish in ethyl alcohol. cuticular granulation fine and uniform, slightly dense around pseudocelli. antennae almost as long as head, area antennalis relatively well marked. pao european journal of taxonomy 879: 64–82 (2023) 70 https://zoobank.org/urn:lsid:zoobank.org:act:aac4848a-79d3-4a5c-8458-a8cb7bb2ea8f fig. 4. deuteraphorura colchisi parimuchová, barjadze & kováč sp. nov. a. dorsal chaetotaxy (the same scale as in fig. 4b). b. ventral chaetotaxy of abdomen. c. pao. d. mvo in adult specimen (other than in fig. 4b). parimuchová a. et al., two new cave deuteraphorura species from georgia 71 with 10–14 compound vesicles (fig. 4c). ant. i with 8–9 chaetae in one row, ant. ii with 14–15 chaetae. aoiii with 5 papillae, 5 guard chaetae, 2 sensory rods almost as long as papillae, 2 rough sensory clubs and lateral ms (as in fig. 5b). lateral ms on ant. iv placed basally at the level of second row of chaetae. apical organite simple in unprotected cavity. maxillary outer lobe simple with 1 basal chaeta and 2 sublobal hairs. labium of ab-type, with 6 proximal chaetae. basomedian field with 4 chaetae, basolateral field with 5 chaetae. head ventrally with 4 postlabial chaetae. pso formula dorsally as 33/133/3(4)3(4)4(3)5(6)3-4 (fig. 4a) (2 pso on th. i sometimes appear); ventrally as 12/011/3212 (fig. 4b for abdominal ventral pso formula); head ventrally with 1 anterior 1 posteromedial and 1 postero-lateral pso. psx weakly visible. subcoxae 1 of i–iii pairs leg with 2,2,2 pso. dorsal body chaetae only weakly differentiated into macro and mesochaetae. th. i with 7 chaetae per half. thii–abdiii with 3 + 3 medial chaetae respectively. vt with 5–7 chaetae per half, basal chaetae table 2. list of species with 3 pso on hind margin of the head and 1 pso on th. i. abbreviations: abs = absent; f = forked; l = long; mvo = number of setae in male ventral organ; s = simple; t = thick. species distribution habitat body length (mm) pao dorsal pso ventral pso subcoxae 1 pso mvo d. akelaris jordana & beruete, 1983 spain (navarra) cave 1.2 12–14 33/133/45454 3/011/ 2211 ? 2t/8t d. arminiaria (gisin, 1961) austria cave 1.5–2.2 12 33/133/33354 3/011/ 2112 2,2,2 ? 4t/6t d. bizkaiensis beruete, arbea & jordana in beruete et al. 2021 spain (basque) cave 0.8–1.06 12–13 33/133/33353 3/011/ 3111 2,2,2 2t/8t d. closanica gruia, 1965 romania caves 1.25–1.8 12 334/133/33353 3/011/4121-22 1,1,1 4s/35-40s d. dashtenazensis arbea, yahyapour & shayanmehr in yahyapour et al. 2020 iran soil, litter 1.4–1.9 13-15 33/133/33353 3/000/ 1221 2,2,2 -/8t d. galani beruete, arbea & jordana, 2001 spain (navarra) cave 1.0–1.2 12–14 33/133/3-443454 3/011/ 4212 ? 2t/8t d. harrobiensis beruete, arbea & jordana, 2001 spain (navarra) cave 1.1.–1.3 12–15 33/133/344464 3/011/ 4111 ? 2t/8t d. jitkae (rusek, 1964) slovakia soil, forest 2.1 21 33/133/33342 1/???/2321-22 1,1,1 2/ numerous s,f d. kosarovi (zonev, 1973) ? ? ? 15 33/133/33354 3/011/ 4222 ? ? d. trisilvaria (gisin, 1962) austria cave 1.6–2.4 18 33/133/33354 3/011/ 3211 2,2,2 abs d. colchisi sp. nov. georgia cave 1.1–2.3 10–14 33/133/3(4) 3(4)4(3)5(6)3-4 3/011/ 3212 2,2,2 -/20-25 t, s d. kozmani sp. nov georgia cave 1.5–2.6 14–16 33/133/4(3)4 (5)3-45(6)3(4) 3/011/ 3222 2,2,2 -/ 20-25 l, t, f european journal of taxonomy 879: 64–82 (2023) 72 fig. 5. deuteraphorura kozmani parimuchová, barjadze & kováč sp. nov. a. dorsal chaetotaxy. b. aoiii. c. mvo (enlargment of modified chaeta). d. tita and claw of leg iii (dic contrast image; chaeta in c-row not visible from this view). parimuchová a. et al., two new cave deuteraphorura species from georgia 73 absent. chaetae on th. i–iii sterna absent. furca remnant with 2 + 2 thin chaetae in one row. mvo present only in fully adult males in form of 10–20 thickened, short and bent spine-like chaetae only on abd. iii sternum (fig. 4d). subcoxae 1 of legs i–iii 4,4,4 chaetae, subcoxae 2 with 3, 13–14, 14–16 chaetae, trochanters with 8–9 chaetae each and femora with 15, 13–15, 13–15 chaetae, respectively. tita i–iii with 18, 19, 17 chaetae including 9 chaetae in distal whorl. tita i with 6 + m chaetae in b row and 2 chaetae in c row, tita ii with 7 + m chaetae in b-row and 2 chaetae in c-row, tita iii with 6 + m chaetae in b row and 1 chaeta in c row. claw without teeth. empodium with basal lamella, tip of filament reaching two-thirds of the claw length (as in fig. 5d). ecology and distribution the species is known only from caves in western georgia where it inhabits warm caves at low altitudes. by its morphology, it does not display any obvious troglomorphic adaptations. remarks see remarks for d. kozmani sp. nov. deuteraphorura kozmani parimuchová, barjadze & kováč sp. nov. urn:lsid:zoobank.org:act:bd48c182-8833-4537-9c44-024938dc47cd fig. 5, table 2 etymology the species was named after the type locality, the kozmani cave in georgia. type material holotype georgia • ♂; imereti, kharagauli, kozmani cave; 42.10092528° n, 43.28852625° e; 14 sept. 2021; eter maghradze leg.; hand collecting on detritus; ibe fs upjs. paratypes georgia • 6 ♀♀, 2 ♂♂; same collection data as for holotype; izisu • 8 ♀♀, 1 ♂; same collection data as for holotype; ibe fs upjs. description body length 1.8–2.6 mm in females, 1.5–2.0 in males (average 2.0 mm; n = 18), shape cylindrical (fig. 5a). colour white to pale brownish in ethyl alcohol. cuticular granulation fine and uniform, slightly dense around pseudocelli. antennae almost as long as head, area antennalis relatively well marked. pao with 14–16 compound vesicles. ant. i with 8 chaetae in one row, ant. ii with 14–15 chaetae. aoiii with 5 papillae, 5 guard chaetae, 2 sensory rods almost as long as papillae, 2 rough sensory clubs and lateral ms (fig. 5b). lateral ms on ant. iv placed basally at the level of second row of chaetae. apical organite simple in unprotected cavity. maxillary outer lobe simple with 1 basal chaeta and 2 sublobal hairs. labium of ab-type, with 6 proximal chaetae. basomedian field with 4 chaetae, basolateral field with 5 chaetae. head ventrally with 5 postlabial chaetae. pso formula dorsally as 33/133/4(3)4(5)3-45(6)3(4) (fig. 5a); ventrally as 12/011/3222; head ventrally with 1 anterior, 1 postero-medial and 1 postero-lateral pso. psx weakly visible. subcoxae 1 of i–iii pairs leg with 2,2,2 pso. european journal of taxonomy 879: 64–82 (2023) 74 https://zoobank.org/urn:lsid:zoobank.org:act:bd48c182-8833-4537-9c44-024938dc47cd dorsal body chaetae only weakly differentiated into macro and mesochaetae. th. i with 6–7 chaetae per half. thii–abdiii with 3 + 3 medial chaetae respectively. vt with 5–6 chaetae per half, basal chaetae mostly absent. chaetae on th. i–iii sterna absent. furca remnant with 2 + 2 thin chaetae in one row. mvo present only in fully adult males in form of 20–25 thickened, long and forked chaetae only on abd. iii sternum (fig. 5c). subcoxae 1 of legs i–iii with 4, 4, 4 chaetae, subcoxae 2 with 3,14–17, 15–17 chaetae, trochanters with 8–10 chaetae each and femora with 14–15, 13–15, 13–15 chaetae, respectively. tita i–iii with 18, 19, 17 chaetae including 9 chaetae in distal whorl. tita i with 6 + m chaetae in b row and 2 chaetae in c row, tita ii with 7 + m chaetae in b-row and 2 chaetae in c-row, tita iii with 6 + m chaetae in b row and 1 chaeta in c row. claw without teeth. empodium with basal lamella, tip of filament reaching two-thirds of the claw length (fig. 5d). ecology and distribution the species is known only from the type locality, occurring on guano and decaying organic material. it does not display any obvious troglomorphic adaptations. remarks both species belong to the species group of deuteraphorura with 3 pso on hind margin of the head and possessing the pso on the first thoracic tergum. the vast majority of these species occupy caves in southern and central europe. as morphological characters vary within both new species, reliable distinguishing from each other is possible only by ventral pseudocellar formula and shape of mvo in matured males. deuteraphoruracolchisi sp. nov. has simple thickened chaetae in mvo, while modified chaetae in d.kozmani sp. nov. are longer and weakly forked at the tip. similar to the new species, d. dashtenazensis arbea, yahyapour & shayanmehr, 2020 has mvo only on abd. iii, but it differs in number of chaetae on this organ. diagnostic morphological characters of both new species and other species of this group are listed in table 2. discussion only a few species of deuteraphorura have been registered in georgia to date: d. variabilis (stach, 1954), d. kruberaensis jordana & baquero, 2012 (barjadze et al. 2012, 2015) and d. inermis (tullberg, 1869); however, the occurrence of the last species in georgian caves is doubtful (barjadze et al. 2012). intensive cave sampling using an integrative approach reveals a greater diversity of cave deuteraphorura in georgia than previously thought. delimitation methods indicated the presence of several distinct molecular lineages (motus) within the (morpho)species colchisi, while only one in kozmani. three groups within the species of colchisi point to recent speciation of deuteraphorura in georgian caves, as revealed in deuteraphorura and protaphorura from the western carpathians (parimuchová et al. 2017, 2020). three approaches to species delimitation (morphological, molecular and geographical) contradict one another in the population from shvilobisa cave. molecularly, specimens from this cave represent unique species; however, it is located in the same karst area as kozmani cave–zemo-imereti plateau. geography was considered a reliable delimitation tool in onychiuridae (sun et al. 2017), but the in the case of such a complex karst area as zemo-imereti, particular structural plateaus have a different geological history (lezhava et al. 2019; tielidze et al. 2019), which may have a decisive impact on the isolation of subterranean populations within this area and the evolution of independent phyletic lineages of deuteraphorura. morphologically, specimens from shvilobisa cave are similar to those from the geologically similar sataplia-tskaltubo karst area. but due to the relatively small number of specimens for morphological and molecular study, the population from shvilobisa cave needs further examination. the satevzia, motena and inchkhuri caves are located in different karst areas, but they are geographically relatively close; they share the parimuchová a. et al., two new cave deuteraphorura species from georgia 75 same cryptic species, which is documented by a positive correlation between geographic and genetic distance. as the family onychiuridae reflects a high level of morphological variability and the left-right asymmetry in chaetotaxy and pseudocellar patterns (e.g., jordana et al. 2012; kaprus' et al. 2014; sun & wu 2014; parimuchová et al. 2017, 2020; vargovitsh 2019), identification of species-specific characters, corresponding to molecular delimitation, is very problematic. inadequacies in the morphological taxonomy of onychiuridae are caused by a lack of sufficient morphological characters and thus a high level of cryptic diversity in this family (sun et al. 2017). subterranean biodiversity of the caucasus mts – a hotspot area areas of the highest subterranean biodiversity (hotspots) were defined based on the number of species adapted to subterranean life per cave (culver & sket 2000; reboleira et al. 2011). regarding the number of troglobiotic species, we have to return to the definition of troglobiont/troglobite as a species that exclusively inhabits a subterranean environment with a preference for its deep parts, and eventually also showing morphological adaptations to subterranean life (sket 2008; trajano & carvalho 2017; howarth & moldovan 2018). the degree of troglomorphism is not correlated with occupied cave depths, as documented by non-adapted animals occurring in the deepest parts of caves (sendra & reboleira 2012). to distinguish troglobionts based only on the level of morphological adaptations to the cave environment is very ambiguous, particularly in pre-adapted groups of invertebrates living in deeper soil horizons. moreover, recent speciation could be a reason for the low development of troglomorphic characters in onychiuridae (e.g., fiera et al. 2021). thus, the real species richness of a given area could be underestimated in this family when considering only morphological traits. it is known that troglomorphic adaptations are not universal in all cave-adapted species. the level of troglomorphy could be correlated with evolutionary age, showing up well in ‛old’ troglobionts and weakly to moderately in ‛young’ ones (kováč et al. 2016). guanobionts regularly contradict the generally accepted morphological traits of obligate cave-dwellers (e.g., culver & pipan 2009, 2015). in troglobiotic onychiuridae, a rather edaphomorphic appearance without progressive troglomorphic adaptations has been documented in a large number of cave-dwelling species from the romanian carpathians (fiera et al. 2021). on the other hand, the highly troglomorphic troglaphorura gladiator vargovitsh, 2019, from georgia in the caucasus, and deuteraphorura muranensis parimuchová & kováč, 2020, distributed at the northernmost distribution limit of troglobionts in europe (parimuchová et al. 2020), show an extremely high level of troglomorphy, as much as the species of the genus ongulonychiurus from spain and croatia, respectively (thibaud & massoud 1986; sun et al. 2019), and pilonychiurus from algeria (pomorski 2007). in contrast, absolonia gigantea (absolon, 1901) from dinarides, of an unusually large size, lacks distinct troglomorphy similar to protaphorura janosik weiner, 1990, and p. cykini parimuchová & kováč in parimuchová et al., 2017a, from the western carpathians and siberia, respectively, and megaphorura arctica (tullberg, 1877), which is abundant in the substrate at the foot of bird cliffs in the arctic. these discrepancies suggest that the microhabitat (or the trophic niche that a species occupies) determines the level of troglomorphy to a greater extent than geographic distribution in a biodiversity hotspot or evolutionary origin in terms of young and old troglobionts. based on the category of troglomorphisms (length of antennae, claws and furca), four life forms were distinguished in troglobiotic arrhopalitidae, including neustonic, atmobiotic, intermediate and intrasubstrate troglobionts (vargovitsh 2022). a similar approach could be applied to representatives of the family of onychiuridae, whose species living exclusively on the water surface have a clearly elongated claw, while species living in guano and sediments generally have a short claw. the occurrence of highly troglomorphic species is thus possible everywhere in the subterranean environment where the microhabitat character requires specific adaptations; however, there are european journal of taxonomy 879: 64–82 (2023) 76 environmental indicators to a higher incidence. aside from hypotheses considering habitat heterogeneity, historical circumstances and habitat productivity, a high terrestrial species richness is also enhanced by the west-east orientation of mountains, which historically reduced the migration potential of invertebrates and increased their invasion rate via subterranean habitats (culver et al. 2006; deharveng et al. 2012). similar to the geographic characteristics seen in mountain ranges in southern europe, the caucasus is predestined to be a hotspot of subterranean biodiversity and potential evolution centre. the relatively large extent and connectivity of the karst, especially in abkhazia, georgia, may enable subterranean species to disperse more widely and access various microhabitats inside the karst, the pattern revealed in dinarides (bregović & zagmajster 2016). the length of passages and surface productivity as a variable of subterranean species richness (culver et al. 2004, 2006) point in favour of the caucasus as a hotspot, as well. the long-term stable areas of high precipitation are important for subterranean terrestrial diversity, while productive energy is important only on a global scale (bregović & zagmajster 2016). the fauna of the western caucasian caves importantly contributes to global subterranean diversity, as was documented across multiple arthropod taxa (e.g., sendra & reboleira 2012; antić & makarov 2016; barjadze et al. 2019; antić & reip 2020; martens et al. 2021; zaragoza et al. 2021). the caucasus as a significant hotspot of subterranean biodiversity is well documented in diplopoda, with a high level of endemism and a high proportion of troglobiotic species, especially in the orders julida and chordeumatida (antić & makarov 2016; antić & reip 2020). altogether, the 19 troglobiotic species known from a single cave in georgia (fiera et al. 2021) are almost equal to the threshold of 20 troglobiotic and stygobiotic species in the hotspot caves (culver & sket 2000). thus, it is herein documented that the western caucasus is a centre of speciation in the collembola genera of arrhopalites, pygmarrhopalietes (arrhopalitidae) and plutomurus (tomoceridae) (fiera et al. 2021), and based on the present results, potentially also in the genus deuteraphorura (onychiuridae). acknowledgements this article is published in the frame of the grant: “complex morphological and molecular investigations of cave dwelling collembolans (hexapoda) in georgia and slovakia” supported by the national scholarship programme of the slovak republic. the study was financially supported by grant apvv21-0379 (slovak research and development agency), grant vega 1/0438/22 (slovak scientific grant agency) and grant “conservation actions and invertebrates’ investigations in sataplia-tskaltubo karst caves, georgia” supported by the conservation leadership programme (clp-04125220). we would like to thank speleologist valeri barbakadze (imereti cave protected areas, georgia) for assistance during the expeditions. we are thankful to david lee mclean for linguistic correction of the manuscript. references absolon k. 1901. weitere nachricht über europäische höhlen collembolen und über die gattung aphorura a.d. macgillivary. zoologischer anzeiger 24 (646): 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(collembola) aus bulgarien. godishnik na sofiiskiya universitet sv. kliment okhridski biologicheski fakultet kniga 1 zoologiya 65 (1): 157–161. yahyapour e., shayanmehr m., vafaei-shoushtari r. & arbea j.i. 2020. a review of the iranian species of the family onychiuridae (collembola, poduromorpha), with description of five new species from hyrcanian forests in iran. zootaxa 4861 (1): 1–22. https://doi.org/10.11646/zootaxa.4861.1.1 manuscript received: 21 october 2022 manuscript accepted: 13 february 2023 published on: 10 july 2023 topic editor: tony robillard section editor: javier i. arbea desk editor: marianne salaün printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; leibniz institute for the analysis of biodiversity change, bonn – hamburg, germany; national museum of the czech republic, prague, czech republic. european journal of taxonomy 879: 64–82 (2023) 82 https://doi.org/10.11646/zootaxa.4250.1.2 https://doi.org/10.11646/zootaxa.4250.1.2 https://doi.org/10.11646/zootaxa.4619.2.4 https://doi.org/10.3390/d14080678 https://doi.org/10.1093/bioinformatics/btt499 https://doi.org/10.1093/bioinformatics/btt499 https://doi.org/10.11646/zootaxa.4861.1.1 european journal of taxonomy 672: 1–18 issn 2118-9773 https://doi.org/10.5852/ejt.2020.672 www.europeanjournaloftaxonomy.eu 2020 · dierickx k. & snoeks j. this work is licensed under a creative commons attribution licence (cc by 4.0). r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:1b1d9e9b-a5b5-4e9a-976c-6409f81dad76 1 protomelas krampus, a new paedophagous cichlid from lake malawi (teleostei, cichlidae) katrien dierickx 1,* & jos snoeks 2 1,2 ichthyology, section vertebrates, department of biology, royal museum for central africa, tervuursesteenweg 13, 3080 tervuren, belgium. 1 bioarch, department of archaeology, university of york, england, united kingdom. 2 laboratory of biodiversity and evolutionary genomics, leuven university (ku leuven), charles deberiotstraat 32, 3000 leuven, belgium. * corresponding author: katrien.dierickx.icht@gmail.com 2 email: jos.snoeks@africamuseum.be 1 urn:lsid:zoobank.org:author:390dd3dc-a240-4359-8643-c76ffca68c4f 2 urn:lsid:zoobank.org:author:8841919c-2a14-4deb-9cc4-69e3c8ca3295 abstract. a new paedophagous species of protomelas, p. krampus sp. nov., is described from lake malawi. it has been found in lukoma bay in tanzania, near mara point in mozambique, and at otter point, chizumulu, the likoma islands and mazinzi reef in malawi. this species is placed in the genus protomelas based on its melanin pattern, which comprises a continuous midlateral stripe. a morphometric study was done to compare this species with its congeners and similar species of hemitaeniochromis and caprichromis. it differs from most congeners by having only one inner tooth row. furthermore, p. krampus sp. nov. differs from p. insignis, p. spilopterus, h. brachyrhynchus, h. urotaenia, caprichromis liemi and c. orthognathus by its shorter premaxillary pedicel, shorter prepectoral distances and dentition. it also differs largely in its melanin pattern from the paedophagous species c. liemi, c. orthognathus, diplotaxodon greenwoodi and naevochromis chrysogaster, as well as h. brachyrhynchus and h. urotaenia. protomelas krampus sp. nov. has been observed to ram mouth-brooding cichlids from above to feed on their eggs or larvae. keywords. cichliformes, east africa, mouth-brooding cichlids, mozambique, trophic ecology. dierickx k. & snoeks j. 2020. protomelas krampus, a new paedophagous cichlid from lake malawi (teleostei, cichlidae). european journal of taxonomy 672: 1–18. https://doi.org/10.5852/ejt.2020.672 introduction lake malawi is home to about 800 to 1000 species of endemic cichlids (snoeks 2000; konings 2016). because of their explosive speciation, species can be difficult to delineate and many species complexes and genera still need to be taxonomically resolved. in lake malawi, haplochromines constitute the major part of the cichlids. the endemic genus protomelas eccles &trewavas, 1989 currently encompasses 14 to 16 valid species (konings 2016; froese & pauly https://doi.org/10.5852/ejt.2020.672 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licences/by/4.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:1b1d9e9b-a5b5-4e9a-976c-6409f81dad76 mailto:katrien.dierickx.icht%40gmail.com?subject= mailto:jos.snoeks%40africamuseum.be?subject= http://zoobank.org/urn:lsid:zoobank.org:author:390dd3dc-a240-4359-8643-c76ffca68c4f http://zoobank.org/urn:lsid:zoobank.org:author:8841919c-2a14-4deb-9cc4-69e3c8ca3295 https://doi.org/10.5852/ejt.2020.672 european journal of taxonomy 672: 1–18 (2020) 2 2019; fricke et al. 2020). the genus is characterized by a typical melanin pattern with a well-developed continuous midlateral band from one eye length behind the opercle to the caudal fin and a dorsolateral band that may be spotted. jaws are short and the length of the premaxillary pedicel is less than one third that of the head. representatives of this genus have bicuspid anterior outer teeth that may be replaced by unicuspid teeth in adults; posterior outer teeth are simple (eccles & trewavas 1989). one species, haplochromis urotaenia regan, 1922, was placed in a newly described genus, hemitaeniochromis eccles & trewavas, 1989, based on its melanin pattern having a typical discontinuous midlateral band (eccles & trewavas 1989). for a long time, it remained the sole species within this genus. several undescribed species were attributed to the genus and later protomelas spilopterus (trewavas, 1935) was transferred by some authors to hemitaeniochromis (see, e.g., turner 1996; snoeks & hanssens 2004; oliver 2012; konings 2016). in addition, oliver (2012) described a new species based on two specimens with a similar melanin pattern, h. brachyrhynchus oliver, 2012. there is, however, major confusion on the precise delimitation of the genera protomelas and hemitaeniochromis (snoeks & hanssens 2004; oliver 2012). already in 1983, mckaye & kocher observed an undescribed paedophagous species near otter point, lake malawi. this species seemed to differ from caprichromis orthognathus (trewavas, 1935) by its melanin pattern and behaviour (mckaye & kocher 1983). eccles & trewavas (1989) suggested that this undescribed cichlid might be p. spilopterus, which is a suspected paedophage based on the resemblance of the jaws and pharyngeal teeth with those of known paedophagous species in lake victoria. based on its head shape and behaviour, as observed by konings (1989, 2016), it seems, however, to be an undescribed species, which was referred to as protomelas sp. ‘paedophage’ and later hemitaeniochromis sp. ‘paedophage’ (see, e.g., konings 1989, 2016). snoeks & hanssens (2004) also mentioned a possibly paedophagous species, protomelas sp. ‘paedophage’, stating that this may be the same species as observed by konings (1989). these authors placed the only two available specimens in the genus protomelas because of its continuous midlateral stripe. mckaye & kocher (1983) noted that paedophagous species, i.e., caprichromis liemi (mckaye & mackenzie, 1982) and c. orthognathus, may change colour depending on their prey species. therefore, as turner (1996) already noted, it is not possible to rule out that this undescribed species may be conspecific to either of the species in caprichromis eccles & trewavas, 1989 or congeneric based solely on its melanin pattern. a morphological analysis was needed to compare this undescribed paedophagous species with similar taxa. three confirmed paedophagous species have currently been described from lake malawi. protomelas spilopterus has short, oblique jaws with thick gums and one or two inner tooth rows. both caprichromis liemi and c. orthognathus have a melanin pattern with a diagonal stripe running from the nape to the base of the caudal fin. the teeth on the lower jaw are embedded in thick gums. especially c. orthognathus has a steeply inclined mouth (eccles & trewavas 1989; konings 2016). in addition to p. spilopterus, c. liemi and c. orthognathus, two more species are suspected of having a paedophagous diet (stauffer & mckaye 1986; eccles & trewavas 1989). diplotaxodon greenwoodi stauffer & mckaye, 1986 is a widespread deep-water species. the genus diplotaxodon trewavas, 1935 is characterized by the lack of distinct markings such as bars or spots on the body. its lower jaw protrudes and the teeth on the lower jaw are embedded in thick gums. diplotaxodon greenwoodi has a steeply inclined mouth, large eyes and one to three inner tooth rows (stauffer & mckaye 1986; trewavas & eccles 1989; snoeks 2004). naevochromis chrysogaster (trewavas, 1935) has a suprapectoral spot, a midlateral spot covering the lateral line, a supranal spot positioned between the lateral lines and a precaudal spot. the chin is prominent and thick. it has a broad mouth and fleshy lips that embed the teeth. there is one inner row of teeth in the upper and two in the lower jaw (eccles & trewavas 1989; konings 2016). as is clear from various discussions (e.g., eccles & trewavas 1989; snoeks 2004; oliver 2012; konings 2016) there is still some confusion regarding the classification of species within the genera protomelas dierickx k. & snoeks j., a new paedophagous cichlid 3 and hemitaeniochromis. in this study, we follow the classification provided by eccles & trewavas (1989), expanded by oliver (2012). it is, however, clear that a taxonomical revision of the species in the genera protomelas and hemitaeniochromis is necessary. for now, we suggest including the newly described paedophagous species in the genus protomelas based on its continuous midlateral band and short jaws, following snoeks & hanssens (2004). we await a further revision of the complex for a final generic allocation. below, the species that is new to science is described as protomelas krampus sp. nov. material and methods in total, 44 specimens of protomelas, hemitaeniochromis and caprichromis in the collections at the royal museum of central africa, tervuren, and the natural history museum, london, were examined: five specimens of c. liemi, five of c. orthognathus, one of h. brachyrhynchus, five of h. urotaenia, eleven of h. sp. ‘insignis like’ (sensu snoeks & hanssens 2004), four of p. insignis, two of p. krampus sp. nov. and eleven of p. spilopterus. the measurements and counts performed here follow snoeks (2004). two additional measurements, as described by oliver (2012), were taken: the snout width and belly length. also, the gape inclination, the angle between the midlateral line and the anteriormost tip of the upper jaw, was measured. in total, 21 length measurements, 13 counts (including the number of vertebrae via x-rays), one angle (gape inclination) (see tables 4 and 5) and some qualitative observations on the body and head shape, dentition, and the colour pattern in alcohol were made. principal component analysis (pca) was performed in past3 (hammer et al. 2001) to explore the multivariate data set. measurements were log-transformed and the covariance matrix was used. when using log-transformed measurements, the individual loadings of all variables on the first principal component (pc 1) are of the same magnitude and sign, and pc 1 can therefore be regarded as a proxy for multivariate size (jolicoeur 1963; snoeks 2004; van steenberge et al. 2015). the correlation matrix was used for the raw meristic data. repositories mrac = musée royal de l’afrique centrale, tervuren, belgium (used traditionally for collections of the royal museum of central africa) bmnh = british museum of natural history, london, uk (used traditionally for collections of the natural history museum) abbreviations used in text hl = head length pc = principal component pca = principal component analysis sl = standard length results comparative morphometrics a pca on 21 log-transformed measurements including all specimens showed a clear separation of protomelas krampus sp. nov. from all other species in the negative part of pc 2 on the second principal axis (fig. 1). in general morphology, it seems to be most similar to c. orthognathus. the most important loadings on pc 2 are of the premaxillary pedicel length, cheek depth, belly length, caudal peduncle length, interorbital width and dorsal fin base (table 1). in a comparison with both species of caprichromis, p. krampus sp. nov. remains clearly separated on the third principal axis (fig. 2). the most important european journal of taxonomy 672: 1–18 (2020) 4 table 1. loadings of pc 1, pc 2 and pc 3 of the principal component analysis of 21 log-transformed measurements of all specimens in the analysis. fig. 1. scatter plot of pc 1 against pc 2 for a principal component analysis of 21 log-transformed measurements of all specimens studied (n = 44). protomelas krampus sp. nov. = blue dot; hemitaeniochromis brachyrhynchus oliver, 2012 = purple square; h. urotaenia (regan, 1922) = green square; hemitaeniochromis sp. ‘insignis like’ = red triangle; p. spilopterus (trewavas, 1935) = grey inverted triangle; p. insignis (trewavas, 1935) = golden diamond; caprichromis liemi (mckaye & mackenzie, 1982) = light brown dash; c. orthognathus (trewavas, 1935) = dark brown rectangle. measurement pc1 pc 2 pc 3 standard length 0.21 -0.16 0.03 body depth 0.23 -0.05 0.18 prepectoral distance 0.20 0.08 -0.05 predorsal distance 0.21 0.16 0.01 preventral distance 0.21 -0.05 0.05 preanal distance 0.21 -0.11 -0.02 dorsal fin base length 0.22 -0.22 0.07 anal fin base length 0.20 -0.18 0.05 caudal peduncle length 0.22 -0.25 0.12 caudal peduncle depth 0.24 -0.17 0.11 belly length 0.21 -0.27 -0.08 head width 0.22 0.16 0.29 head length 0.20 0.10 -0.10 premaxillary pedicel length 0.18 0.61 -0.14 snout length 0.24 0.19 -0.31 lacrimal depth 0.26 0.14 -0.55 cheek depth 0.26 -0.33 -0.19 eye diameter 0.14 0.19 0.33 inter orbital width 0.24 0.24 0.48 snout width 0.24 0.13 0.07 lower jaw length 0.22 -0.04 -0.17 % variance 90.77% 3.04% 2.13% dierickx k. & snoeks j., a new paedophagous cichlid 5 table 2. loadings of pc 1, pc 2 and pc 3 of the principal component analysis of 21 log-transformed measurements of both species of caprichromis eccles & trewavas, 1989 and p. krampus sp. nov. measurement pc1 pc 2 pc 3 standard length 0.21 -0.10 -0.12 body depth 0.20 0.08 -0.21 prepectoral distance 0.20 0.17 -0.04 predorsal distance 0.20 0.19 0.07 preventral distance 0.22 0.04 -0.24 preanal distance 0.23 0.01 -0.09 dorsal fin base length 0.22 -0.06 -0.26 anal fin base length 0.19 -0.08 -0.08 caudal peduncle length 0.23 -0.32 0.21 caudal peduncle depth 0.25 -0.15 0.02 belly length 0.22 -0.20 0.10 head width 0.22 -0.11 0.29 head length 0.20 0.13 -0.09 premaxillary pedicel length 0.15 0.61 0.52 snout length 0.23 0.35 -0.34 lacrimal depth 0.22 0.33 -0.16 cheek depth 0.30 -0.29 -0.10 eye diameter 0.15 -0.10 -0.04 inter orbital width 0.25 -0.13 0.39 snout width 0.23 -0.03 0.26 lower jaw length 0.22 0.02 -0.00 % variance 86.45% 9.34% 1.90% fig. 2. scatter plot of pc 1 against pc 3 for a principal component analysis of 21 log-transformed measurements of protomelas krampus sp. nov., caprichromis liemi (mckaye & mackenzie, 1982) and c. orthognathus (trewavas, 1935) (n = 12). p. krampus sp. nov. = blue dot; c. liemi = light brown dash; c. orthognathus = dark brown rectangle. european journal of taxonomy 672: 1–18 (2020) 6 loadings on pc 3 are for the premaxillary pedicel length, the interorbital width, the snout length, the head width, the dorsal fin base and the snout width (table 2). in a pca on the raw meristics, the picture was less clear. still, p. krampus sp. nov. was separated from all other species (fig. 3), mostly on pc 1. the most important loadings on pc 1 are of the number of vertebrae, the number of both upper and lower jaw teeth, the number of longitudinal line scales, the table 3. loadings of pc 1 and pc 2 of the principal component analysis of 13 counts. counts pc 1 pc 2 longitudinal line 0.33 -0.37 upper lateral line 0.26 -0.06 lower lateral line 0.24 -0.37 lower gill rakers 0.05 0.08 upper gill rakers 0.08 0.09 dorsal fin spines 0.31 -0.20 dorsal fin rays 0.25 0.57 anal fin rays 0.29 0.45 pectoral fin rays 0.07 -0.26 vertebrae 0.43 -0.05 inner teeth rows 0.22 0.05 upper teeth 0.40 -0.10 lower teeth 0.34 0.22 % variance 29.05% 12.70% fig. 3. scatter plot of pc 1 against pc 2 for a principal component analysis of 13 counts of all specimens studied (n = 44). protomelas krampus sp. nov. = blue dot; hemitaeniochromis brachyrhynchus oliver, 2012 = purple square; h. urotaenia (regan, 1922) = green square; hemitaeniochromis sp. ‘insignis like’ = red triangle; p. spilopterus (trewavas, 1935) = grey inverted triangle; p. insignis (trewavas, 1935) = golden diamond; caprichromis liemi (mckaye & mackenzie, 1982) = light brown dash; c. orthognathus (trewavas, 1935) = dark brown rectangle. dierickx k. & snoeks j., a new paedophagous cichlid 7 number of dorsal fin spines and the number of anal fin rays. on pc 2, the highest loadings are the number of dorsal fin rays, the number of anal fin rays, the number of scales on the lower lateral line, the number of longitudinal line scales, the number of pectoral fin rays and the number of outer lower jaw teeth (table 3). taxonomy phylum chordata haeckel, 1874 class actinopterygii klein, 1885 order cichliformes betancur et al., 2013 family cichlidae bonaparte, 1840 subfamily pseudocrenilabrinae fowler, 1934 tribe haplochromini poll, 1986 genus protomelas eccles & trewavas, 1989 protomelas eccles & trewavas, 1989: 40 (type species: chromis kirkii günther, 1894, by original designation). protomelas krampus sp. nov. urn:lsid:zoobank.org:act:5c1913c2-e6c9-43cd-bb1c-5522dfc60b8c figs 4–5; tables 4–5 diagnosis protomelas krampus sp. nov. differs from most species of protomelas, i.e., p. annectens (regan, 1922), p. fenestratus (trewavas, 1935), p. kirkii (günther, 1894), p. labridens (trewavas, 1935), p. macrodon eccles, 1989, p. marginatus (trewavas, 1935), p. pleurotaenia (boulenger, 1901), p. similis (regan, 1922), p. spilonotus (trewavas, 1935), p. taeniolatus (trewavas, 1935), p. triaenodon (trewavas, 1935) and p. virgatus (trewavas, 1935), by having only one inner tooth row, whereas the other species have two rows. it can be distinguished from p. spilopterus by a shorter premaxillary pedicel (17.0–19.2 vs 21.8–28.4% hl), a larger gape inclination (70–80 vs 40–60°), a shallower body (33.3–34.3 vs 36.0–42.8% sl), a smaller predorsal (27.4–29.1 vs 33.4–37.4% sl) and prepectoral distance (29.4–29.8 vs 31.8–40.0% sl), a shorter and more slender head (length 27.8–29.2 vs 30.4–34.6% sl and width 39.3–40.3 vs 41.4– 50.7% hl, respectively), and a smaller interorbital width (22.9–23.4 vs 25.0–33.6% hl). in addition, p. krampus sp. nov. has more gill rakers on the outer epibranchial (further mentioned as upper gill rakers) than p. spilopterus (5 vs 3–4) and more vertebrae (31–32 vs 29). the inner teeth are tricuspid in p. krampus sp. nov. while mixed unicuspid/tricuspid or unicuspid in p. spilopterus. the outer teeth of the lower jaw of p. krampus sp. nov. are oriented straight up, whereas those of p. spilopterus are angled forward, except for in one specimen which has slightly inwards curved teeth. protomelas krampus sp. nov. differs from p. insignis by a shorter premaxillary pedicel (17.0–19.2 vs 27.9–30.0% hl), a deeper cheek (31.6–40.2 vs 23.2–28.5% hl), the larger gape inclination (70–80 vs 30°), a smaller predorsal (27.4–29.1 vs 34.2–35.9% sl) and prepectoral distance (29.4–29.8 vs 32.7– 35.4% sl), a shorter and more slender head (length 27.8–29.2 vs 31.6–34.3% sl and width 39.3– 40.3 vs 41.3–44.8% hl, respectively), a shallower lacrimal (19.6–20.4 vs 21.4–22.4% hl), a smaller interorbital width (22.9–23.4 vs 26.9–32.8% hl), and a longer lower jaw (37.7–38.8 vs 31.2–37.3% hl). the upper jaw has fewer outer teeth in p. krampus sp. nov. than in p. insignis (37–40 vs 43–50). outer lower jaw teeth are straight in p. krampus sp. nov. but curved inwards in p. insignis. http://zoobank.org/urn:lsid:zoobank.org:act:5c1913c2-e6c9-43cd-bb1c-5522dfc60b8c european journal of taxonomy 672: 1–18 (2020) 8 protomelas krampus sp. nov. differs from the undescribed species mentioned by snoeks & hanssens (2004) as hemitaeniochromis. sp. ‘insignis like’ by a shorter premaxillary pedicel (17.0–19.2 vs 24.7– 30.7% hl), a deeper cheek depth (31.6–40.2 vs 22.5–28.8% hl), a larger gape inclination (70–80 vs 40–60°), a shorter predorsal (27.4–29.1 vs 33.5–38.3% sl) and prepectoral distance (29.4–29.8 vs 31.1–38.1% sl), a shorter and more slender head (length 27.8–29.2 vs 31.3–34.5% sl and width 39.3–40.3 vs 40.8–46.8% hl, respectively), and a longer lower jaw (37.7–38.8 vs 28.9–35.0% hl). protomelas krampus sp. nov. has more gill rakers on the outer epibranchial than h. sp. ‘insignis like’ (5 vs 3–4). there are more dorsal fin rays in p. krampus sp. nov. than in h. sp. ‘insignis like’ (11–12 vs 9–10). the outer teeth of the lower jaw of p. krampus sp. nov. are oriented straight up, whereas those of h. sp. ‘insignis like’ are angled forward. protomelas krampus sp. nov. differs from both species of the genus hemitaeniochromis by its continuous midlateral stripe, which places it in protomelas, whereas h. urotaenia and h. brachyrhynchus have an anteriorly spotted stripe. in addition, p. krampus sp. nov. differs from h. urotaenia by a shorter premaxillary pedicel (17.0–19.2 vs 27.5–28.9% hl), a deeper cheek (31.6–40.2 vs 27.3–30.6% hl), a larger gape inclination (70–80 vs 30–50°), a longer dorsal fin base (53.6–58.3 vs 49.3–50.5% sl), a shorter predorsal (27.4–29.1 vs 37.0–38.7% sl) and prepectoral distance (29.4–29.8 vs 35.2–36.7% sl), a shorter and more slender head (length 27.8–29.2 vs 34.9–37.3% sl and width 39.3–40.3 vs 40.5–42.5% hl respectively), and a shallower lacrimal (19.6–20.4 vs 22.7–23.3% hl). protomelas krampus sp. nov. has more vertebrae (31–32 vs 30), more gill rakers on the outer epibranchial (5 vs 2–4) and more dorsal fin rays (11–12 vs 9–10) than h. urotaenia. protomelas krampus sp. nov. has only one inner tooth row with tricuspid teeth, whereas h. urotaenia has two rows of mostly unicuspid teeth. the outer lower jaw teeth are straight in p. krampus sp. nov., whereas they are curved inwards in h. urotaenia. protomelas krampus sp. nov. differs from h. brachyrhynchus by a shorter premaxillary pedicel (17.0– 19.2 vs 27.1% hl), a deeper cheek (31.6–40.2 vs 21.4% hl), a larger gape inclination (70–80 vs 45°), a shallower body (33.3–34.3 vs 37.7% sl), a shorter predorsal (27.4–29.1 vs 37.8% sl), preventral (39.0–44.4 vs 46.1% sl) and prepectoral distance (29.4–29.8 vs 37.0% sl), a shorter anal fin base length (18.3–18.8 vs 21.2% sl), a shorter and more slender head (27.8–29.2 vs 35.3% sl and 39.3–40.3 vs 43.8% hl, respectively), a longer snout (33.3–34.2 vs 30.1% hl), a deeper lacrimal (19.6–20.4 vs 17.2% hl), a smaller eye (22.5–27.0 vs 32.2% hl), and a longer lower jaw (37.7–38.8 vs 32.9% hl). protomelas krampus sp. nov. has more vertebrae (31–32 vs 30), upper gill rakers (5 vs 4) and dorsal fin rays (11–12 vs 10) than h. brachyrhynchus. the outer teeth of the lower jaw of p. krampus sp. nov. are oriented straight up, whereas those of h. brachyrhynchus are angled forward and slightly outwards. protomelas krampus sp. nov. differs from the paedophagous species of the genus caprichromis by its melanin pattern: it has a midlateral band from behind the opercle to the base of the caudal fin, whereas species of caprichromis have a diagonal stripe from the nape to the base of the caudal fin. protomelas krampus sp. nov. further differs from c. orthognathus by a shorter premaxillary pedicel (17.0–19.2 vs 21.9–25.4% hl) and a shorter prepectoral distance (29.4–29.8 vs 30.2–33.4% sl). there are more soft dorsal fin rays in p. krampus sp. nov. than in c. orthognathus (11–12 vs 10) and more gill rakers on the outer epibranchial (5 vs 3–4). protomelas krampus sp. nov. has fewer outer teeth in the lower jaw (31–36 vs 42–50). protomelas krampus sp. nov. differs from c. liemi by a shorter premaxillary pedicel (17.0–19.2 vs 24.7–28.0% hl), a deeper cheek (31.6–40.2 vs 25.5–29.5% hl), a larger gape inclination (70–80 vs dierickx k. & snoeks j., a new paedophagous cichlid 9 30–60°), a shallower body (33.3–34.3 vs 36.1–38.6% sl), shorter predorsal (27.4–29.1 vs 35.0–36.7% sl) and prepectoral distances (29.4–29.8 vs 34.2–36.5% sl), a shorter and wider head (length 27.8– 29.2 vs 31.9–33.6% sl and width 39.3–40.3 vs 35.9–38.2% hl, respectively), and a shallower lacrimal (19.6–20.4 vs 20.8–23.9% hl). there are fewer scales along the upper lateral line in p. krampus sp. nov. than in c. liemi (20–23 vs 25–27). protomelas krampus sp. nov. has fewer outer teeth in the upper jaw (37–40 vs 44–47) and lower jaw (31–36 vs 44–59) than c. liemi. protomelas krampus sp. nov. differs from diplotaxodon greenwoodi by the lack of a melanin pattern in the latter species. it has isognathous jaws, whereas d. greenwoodi has a protruding lower jaw. protomelas krampus sp. nov. differs from naevochromis chrysogaster by its melanin pattern, which consists of three large spots on the lateral sides in the latter instead of a continuous midlateral line. it has a more strongly inclined gape than n. chrysogaster. protomelas krampus sp. nov. has only one inner tooth row on the lower jaw, whereas n. chrysogaster has two. etymology the specific name, ʻkrampusʼ, is a noun in apposition and was chosen in reference to the european folklore character krampus. this demon puts naughty children in a bag and takes them away, which is reminiscent of a paedophagous behaviour. the goat-like appearance of krampus also implicitly refers to the head-butting behaviour of the species. the same implicit reference to this behaviour is also found in the genus name caprichromis of other paedophagous species of lake malawi. material examined holotype tanzania • ♀; lake malawi, lukoma bay; 11°22.50′ s, 34°52.00′ e; 11 jan. 1998; sadc/gef taxonomy team leg.; 116.1 mm sl; mrac 99-041-p-4768. paratype mozambique • 1 ♂; lake malawi, mara rocks; 12°11.34′ s, 34°41.73′ e; 22 may 1998; sadc/ gef taxonomy team leg.; 181.1 mm sl; mrac 99-041-p-4767. comparative material hemitaeniochromis brachyrhynchus (oliver, 2012) malawi • 1 paratype; lake malawi, nkhata bay, south bay; 11°36.22′ s, 34°19.16′ e; 27 sep. 1997; sadc/gef taxonomy team leg.; 81.6 mm sl; mrac 99-041-p-1746. hemitaeniochromis urotaenia (regan, 1922) malawi • 2 specs; lake malawi, kande bay, s of bandawe; 11°56.47′ s, 34°09.41′ e; 3 jun. 1997; sadc/gef taxonomy team leg.; 142.2–182.7 mm sl; mrac 99-041-p-1738-1739 • 3 specs [of 5 in lot]; lake malawi, nkhotakota; 12°52.95′ s, 34°19.33′ e; 22 sep. 1997; sadc/gef taxonomy team leg.; 87.8–119.0 mm sl; mrac 99-041-p-1741-1745. hemitaeniochromis sp. “insignis like” malawi • 3 specs; lake malawi, chipoka to makanjila, sw arm and se arm; 14°00.62′ s, 34°37.29′ e; 18 nov. 1997; sadc/gef taxonomy team leg.; 122.0–148.5 mm sl; mrac 99-041-p-1747-1749 • 1 spec.; lake malawi, nkhotakota; 12°52.95′ s, 34°19.34′ e; 22 sep. 1997; sadc/gef taxonomy team leg.; 76.9 mm sl; mrac 99-041-p-1750 • 1 spec.; lake malawi, senga bay; 13°45.13′ s, 34°29.22′ e; 8 jun. 1997; sadc/gef taxonomy team leg.; 145.2 mm sl; mrac 99-041-p-1773 • 2 specs; lake malawi, chipoka to makanjila, sw arm and se arm; 13°52.45′ s, 34°54.74′ e; 9 oct. 1997; sadc/ european journal of taxonomy 672: 1–18 (2020) 10 gef taxonomy team leg.; 139.7–172.4 mm sl; mrac 99-041-p-2625-2626 • 1 spec.; lake malawi, chipoka to makanjila, sw arm and se arm; 13°57.77′ s, 34°43.37′ e; 10 oct. 1997; sadc/gef taxonomy team leg.; 142.7 mm sl; mrac 99-041-p-2627 • 1 spec.; lake malawi, sw arm; 14°07.00′ s, 34°44.30′ e; 18 dec. 1996; sadc/gef taxonomy team leg.; 169.0 mm sl; mrac 99-041-p-1755. mozambique • 1 spec.; lake malawi, chiwanga bay; 12°39.03′ s, 34°46.56′ e; 11 apr. 1998; sadc/gef taxonomy team leg.; 111.9 mm sl; mrac 99-041-p-1753 • 1 spec.; lake malawi, chiwanga bay; 12°39.03′ s, 34°46.56′ e; 11 apr. 1998; sadc/gef taxonomy team leg.; 117.8 mm sl; mrac 99-041-p-1754. protomelas spilopterus (trewavas, 1935) malawi • lectotype, 3 paralectotypes; lake malawi, south end; 1925; 147.1–166.0 mm sl; bmnh 1935.6.14.644-647 • 2 paralectotypes [of 3 in lot]; lake malawi, mwaya; 1925; 149.1–153.4 mm sl; bmnh 1935.6.14.649-651 (2 of 3) • 5 paralectotypes [of 6 in lot]; lake malawi, monkey bay; 1925; 85.5–160.4 mm sl; bmnh 1935.6.14.652-657. protomelas insignis (trewavas, 1935) malawi • lectotype, 3 paralectotypes; lake malawi, monkey bay; 1925; 81.7–161.0 mm sl; bmnh 1935.6.14.839-843. caprichromis liemi (mckaye & mackenzie, 1982) malawi • 1 spec.; lake malawi, chipoka to makanjila, sw arm and se arm; 14°00.62′ s, 34°37.29′ e; 18 nov. 1997; sadc/gef taxonomy team leg.; 129.2 mm sl; mrac 99-041-p-2647. mozambique • 1 spec.; lake malawi, chilola bay; 12°06.45′ s, 34°46.79′ e; 7 apr. 1998; sadc/ gef taxonomy team leg.; 117.9 mm sl; mrac 99-041-p-2648 • 1 spec.; lake malawi, tchulutcha reef, metangula; 12°42.19′ s, 34°47.46′ e; 25 may 1998; sadc/gef taxonomy team leg.; 173.2 mm sl; mrac 99-041-p-2649 • 2 specs; lake malawi, namisse, s of cobue; 12°10.08′ s, 34°42.97′ e; 24 may 1998; sadc/gef taxonomy team leg.; 103.2–144.4 mm sl; mrac 99-041-p-5018-5019. caprichromis orthognathus (trewavas, 1935) malawi • 1 spec.; lake malawi, mazinzi bay, se arm; 4 sep. 1980; d.s.c. lewis leg.; 146.7 mm sl; mrac 81-02-p-33 • 1 spec.; lake malawi; 25 jun. 1962; 154.1 mm sl; mrac 191849. mozambique • 1 spec.; lake malawi, chiwanga bay; 12°38.52′ s, 34°46.37′ e; 10 apr.1998; sadc/ gef taxonomy team leg.; 122.0 mm sl; mrac 99-041-p-2644 • 2 specs; lake malawi, chiwanga bay; 12°38.81′ s, 34°46.68′ e; 10 apr. 1998; sadc/gef taxonomy team leg.; 128.3–131.4 mm sl; mrac 99-041-p-2645-2646. diplotaxodon greenwoodi (stauffer & mckaye, 1986) malawi • 1 spec.; lake malawi, senga bay; 13°45.25′ s, 34°40′30″ e; 8 jun. 1997; sadc/gef taxonomy team leg.; mrac 1999-041-p-10765 • 1 spec.; lake malawi, chipoka to makanjila, sw arm and se arm; 13°57′46.19″ s, 34°43′22.19″ e; 10 oct. 1997; sadc/gef taxonomy team leg.; mrac 1999-041-p-10766. mozambique • 1 spec.; lake malawi, chilola bay, 2nd bay of cobwe; 12°00′44.39″ s, 34°47′16.80″ e; 8 apr. 1998; sadc/gef taxonomy team leg.; mrac 1999-041-p-10768. tanzania • 1 spec.; lake malawi, weismann bay; 9°30′46.80″ s, 34°00′19.80″ e; 14 jan. 1998; sadc/gef taxonomy team leg.; mrac 1999-041-p-10767. dierickx k. & snoeks j., a new paedophagous cichlid 11 naevochromis chrystogaster (trewavas, 1935) malawi • 1 spec.; lake malawi, mdoka; 10°19′00.01″ s, 34°12′00″ e; may 1989; a. konings leg.; mrac 1991-095-p-0037 • 1 spec.; lake malawi, cape ngombo near makanjila point; 13°44′07.19″ s, 34°51′16.20″ e; 13 nov. 1997; sadc/gef taxonomy team leg.; mrac 1999-041-p-8524. tanzania • 1 spec.; lake malawi, lutara, n of bay; 10°25′57.61″ s, 34°33′33.59″ e; 11 nov. 1998; sadc/gef taxonomy team leg.; mrac 1999-041-p-5205. description based on holotype and one paratype (see figs 4–5 and tables 4–5). qualitative observations are described in the context of lake malawi haplochromine cichlids as conducted by snoeks (2004). body. moderately elongate. head. profile somewhat steep, clearly concave at eye level. snout above upper jaw convex. mouth very steep, vertically orientated with a gape inclination of 70–80°. premaxillary pedicel small. jaws fig. 4. photographs of preserved specimens of protomelas krampus sp. nov. a. holotype (rmca 99041-p-4768). b. paratype (rmca 99-041-p-4767). melanin pattern strongly faded. european journal of taxonomy 672: 1–18 (2020) 12 ta bl e 4. m ea su re m en ts f or p ro to m el as k ra m pu s sp . n ov . ( ho lo ty pe a nd p ar at yp e) , c om pa re d w ith th e ra ng es a nd m ea ns o f th e ex am in ed s pe ci m en s of p . i ns ig ni s (t re w av as , 1 93 5) , h em ita en io ch ro m is s p. ‘ in si gn is li ke ’, p. s pi lo pt er us ( tr ew av as , 1 93 5) , h em ita en io ch ro m is b ra ch yr hy nc hu s o liv er , 20 12 , h . u ro ta en ia (r eg an , 1 92 2) , c ap ri ch ro m is li em i ( m ck ay e & m ac ke nz ie , 1 98 2) a nd c . o rt ho gn at hu s (t re w av as , 1 93 5) . p. k ra m pu s sp . n ov . p. in si gn is h . s p. ‘i ns ig ni s lik e’ p. s pi lo pt er us h . b ra ch yr hy nc hu s h . u ro ta en ia c . o rt ho gn at hu s c . l ie m i h ol ot yp e p ar at yp e m or ph om et ri cs r an ge , m ea n (n = 4 ) r an ge , m ea n (n = 11 ) r an ge , m ea n (n = 11 ) (n = 1 ) r an ge , m ea n (n = 5 ) r an ge , m ea n (n = 4 ) r an ge , m ea n (n = 5 ) st an da rd le ng th (s l ) i n m m 11 6. 1 18 1. 1 81 .7 –1 61 .0 (1 28 .9 ) 76 .9 –1 72 .4 (1 35 .8 ) 85 .5 –1 66 .0 (1 37 .8 ) 81 .6 87 .8 –1 82 .7 (1 25 .6 ) 12 2. 0– 15 4. 1 (1 36 .5 ) 10 3. 2– 17 3. 2 (1 33 .6 ) g ap e in cl in at io n (i n de gr ee s) 70 80 30 (3 0) 35 –6 0 (4 7. 3) 40 –6 0 (4 5. 9) 45 30 –5 0 (4 1) 70 –9 0 (8 0. 5) 30 –6 0 (4 8) a s % sl : b od y de pt h 33 .3 34 .3 33 .8 –3 7. 0 (3 5. 6) 34 .2 –4 0. 3 (3 7. 2) 36 .0 –4 2. 8 (3 8. 9) 37 .7 31 .5 –3 5. 5 (3 3. 6) 32 .5 –3 4. 8 (3 3. 8) 36 .1 –3 8. 6 (3 7. 4) h ea d le ng th 27 .8 29 .2 31 .6 –3 4. 3 (3 2. 5) 31 .3 –3 4. 6 (3 3. 0) 30 .4 –3 4. 6 (3 2. 9) 35 .3 34 .9 –3 7. 3 (3 6. 1) 29 .3 –3 1. 1 (3 0. 3) 31 .9 –3 3. 6 (3 2. 8) pr ep ec to ra l d is ta nc e 29 .4 29 .8 32 .7 –3 5. 4 (3 3. 7) 31 .1 –3 8. 1 (3 4. 5) 31 .8 –4 0. 0 (3 5. 3) 37 .0 35 .2 –3 6. 7 (3 6. 2) 30 .2 –3 3. 4 (3 2. 0) 34 .2 –3 6. 5 (3 5. 2) pr ed or sa l d is ta nc e 29 .0 27 .4 34 .2 –3 5. 9 (3 4. 9) 33 .5 –3 8. 3 (3 5. 7) 33 .4 –3 7. 4 (3 6. 2) 37 .8 37 .0 –3 8. 7 (3 7. 6) 31 .3 –3 3. 4 (3 2. 5) 35 .0 –3 6. 7 (3 5. 6) pr ev en tr al d is ta nc e 39 .0 44 .4 40 .1 –4 2. 9 (4 1. 7) 38 .6 –4 8. 3 (4 3. 6) 41 .2 –5 1. 0 (4 6. 0) 46 .1 41 .7 –4 5. 4 (4 3. 4) 39 .7 –4 4. 1 (4 2. 2) 42 .8 –4 8. 7 (4 4. 5) pr ea na l d is ta nc e 62 .3 69 .8 67 .5 –7 0. 1 (6 8. 8) 63 .7 –7 2. 7 (6 9. 3) 64 .7 –7 2. 1 (6 9. 1) 70 .0 66 .0 –6 9. 9 (6 7. 7) 67 .2 –7 0. 3 (6 8. 4) 68 .5 –7 2. 6 (7 0. 7) d or sa l fi n ba se 53 .6 58 .3 52 .0 –5 8. 7 (5 6. 3) 50 .5 –5 6. 6 (5 3. 3) 49 .8 –5 7. 6 (5 4. 1) 54 .3 49 .3 –5 0. 5 (5 0. 1) 52 .9 –5 4. 3 (5 3. 7) 55 .3 –5 6. 6 (5 5. 7) a na l fi n ba se 18 .2 18 .8 17 .9 –1 9. 3 (1 8. 6) 16 .6 –2 0. 1 (1 8. 1) 17 .8 –2 0. 7 (1 9. 4) 21 .2 17 .4 –2 0. 2 (1 9. 0) 18 .4 –1 9. 3 (1 8. 9) 18 .3 –1 9. 9 (1 9. 2) c au da l p ed un cl e le ng th 17 .2 19 .6 17 .5 –1 7. 8 (1 7. 7) 15 .9 –1 9. 5 (1 8. 1) 17 .5 –1 9. 3 (1 8. 4) 15 .7 17 .0 –1 9. 4 (1 8. 2) 18 .5 –2 0. 9 (1 9. 4) 15 .4 –1 8. 1 (1 7. 0) c au da l p ed un cl e de pt h 11 .6 12 .6 12 .3 –1 3. 2 (1 2. 7) 11 .3 –1 3. 7 (1 2. 5) 11 .4 –1 4. 1 (1 2. 6) 12 .6 11 .3 –1 2. 8 (1 1. 9) 12 .3 –1 3. 2 (1 2. 6) 11 .2 –1 3. 4 (1 2. 2) b el ly le ng th 27 .8 29 .5 28 .3 –2 9. 8 (2 9. 2) 27 .3 –3 2. 3 (2 9. 7) 23 .4 –2 9. 0 (2 6. 1) 28 .5 26 .6 –2 8. 7 (2 7. 8) 29 .2 –3 2. 6 (3 0. 5) 24 .5 –3 1. 4 (2 8. 5) a s % h l : h ea d w id th 39 .3 40 .3 41 .3 –4 4. 8 (4 3. 4) 40 .8 –4 6. 8 (4 3. 2) 41 .4 –5 0. 7 (4 6. 6) 43 .8 40 .5 –4 2. 5 (4 1. 9) 40 .7 –4 5. 5 (4 3. 3) 35 .9 –3 8. 2 (3 7. 0) pr em ax ill ar y pe di ce l l en gt h 19 .2 17 .0 27 .9 –3 0. 0 (2 9. 3) 24 .7 –3 0. 7 (2 9. 0) 21 .8 –2 8. 4 (2 6. 3) 27 .1 27 .5 –2 8. 9 (2 8. 0) 21 .9 –2 5. 4 (2 3. 7) 24 .7 –2 8. 0 (2 7. 0) sn ou t l en gt h 34 .2 33 .3 32 .7 –3 9. 3 (3 6. 4) 30 .6 –3 6. 4 (3 3. 6) 30 .8 –3 7. 8 (3 3. 9) 30 .1 34 .1 –3 7. 5 (3 5. 6) 31 .4 –3 3. 7 (3 2. 2) 34 .8 –3 9. 6 (3 7. 2) l ac hr ym al d ep th 20 .4 19 .6 21 .4 –2 2. 4 (2 1. 9) 17 .4 –2 5. 3 (2 0. 6) 13 .6 –2 2. 7 (1 9. 0) 17 .2 22 .7 –2 3. 3 (2 3. 0) 18 .6 –2 2. 4 (2 0. 3) 20 .8 –2 3. 9 (2 2. 3) c he ek d ep th 31 .6 40 .2 23 .2 –2 8. 5 (2 5. 7) 22 .5 –2 8. 8 (2 6. 0) 22 .6 –3 1. 9 (2 7. 4) 21 .4 27 .3 –3 0. 6 (2 9. 0) 29 .2 –3 7. 0 (3 3. 1) 25 .5 –2 9. 5 (2 7. 7) e ye d ia m et er 27 .0 22 .5 23 .7 –2 9. 9 (2 5. 6) 21 .6 –3 0. 5 (2 5. 8) 24 .9 –3 0. 4 (2 7. 2) 32 .2 21 .6 –2 7. 6 (2 4. 4) 22 .7 –2 5. 4 (2 3. 9) 19 .5 –2 4. 6 (2 1. 8) sn ou t w id th 30 .8 32 .2 27 .7 –3 4. 5 (3 1. 5) 27 .9 –3 4. 8 (3 2. 3) 30 .2 –4 0. 8 (3 4. 9) 29 .2 31 .8 –3 7. 2 (3 4. 2) 32 .9 –3 6. 0 (3 4. 2) 28 .4 –3 1. 8 (3 0. 0) in te ro rb ita l w id th 22 .9 23 .4 26 .9 –3 2. 8 (2 9. 3) 23 .3 –3 0. 6 (2 5. 8) 25 .0 –3 3. 6 (2 9. 8) 22 .1 23 .2 –2 6. 0 (2 4. 9) 23 .2 –2 8. 2 (2 5. 8) 20 .4 –2 3. 6 (2 1. 6) l ow er ja w le ng th 37 .7 38 .8 31 .2 –3 7. 3 (3 4. 4) 28 .9 –3 5. 0 (3 2. 7) 29 .6 –3 9. 1 (3 6. 1) 32 .9 37 .6 –4 0. 6 (3 9. 0) 37 .8 –3 9. 2 (3 8. 4) 33 .6 –3 8. 4 (3 6. 2) dierickx k. & snoeks j., a new paedophagous cichlid 13 ta bl e 5. m er is tic s fo r p ro to m el as k ra m pu s sp . no v. ( ho lo ty pe a nd p ar at yp e) , co m pa re d w ith t he r an ge s of t he e xa m in ed s pe ci m en s of p . in si gn is (t re w av as , 19 35 ), h em ita en io ch ro m is s p. ‘ in si gn is l ik e’ , p. s pi lo pt er us ( tr ew av as , 19 35 ), h em ita en io ch ro m is b ra ch yr hy nc hu s o liv er , 20 12 , h . u ro ta en ia (r eg an , 1 92 2) , c ap ri ch ro m is li em i ( m ck ay e & m ac ke nz ie , 1 98 2) a nd c . o rt ho gn at hu s (t re w av as , 1 93 5) . p. k ra m pu s sp . n ov . p. in si gn is h . s p. ‘i ns ig ni s lik e’ p. s pi lo pt er us h . b ra ch yr hy nc hu s h . u ro ta en ia c . o rt ho gn at hu s c . l ie m i h ol ot yp e p ar at yp e m er is tic s (n = 4 ) (n = 11 ) (n = 11 ) (n = 1 ) (n = 5 ) (n = 4 ) (n = 5 ) l on gi tu di na l s er ie s 32 31 31 –3 4 30 –3 4 29 –3 1 33 30 –3 4 31 –3 3 32 –3 3 u pp er la te ra l l in e 23 20 23 –2 6 21 –2 7 19 –2 6 22 21 –2 4 21 –2 5 25 –2 7 l ow er la te ra l l in e 14 17 13 –1 7 13 –1 6 12 –1 5 14 13 –1 7 13 –1 5 13 –1 6 u pp er g ill ra ke rs 5 5 4– 5 3– 4 3– 4 4 2– 4 3– 4 3– 5 l ow er g ill ra ke rs 11 10 9– 13 11 –1 2 9– 12 10 9– 11 9– 12 11 –1 2 d or sa l fi n sp in es 17 17 16 –1 8 16 –1 8 15 –1 6 17 16 –1 7 15 –1 7 17 d or sa l fi n ra ys 11 12 10 –1 1 9– 10 10 –1 1 10 9– 10 10 10 –1 1 a na l fi n ra ys 9 10 9– 10 8– 9 8– 10 9 9 9 9– 10 pe ct or al fi n ra ys 15 13 14 –1 5 13 –1 5 12 –1 5 14 14 –1 5 13 –1 4 11 –1 4 u pp er o ra l j aw te et h 37 40 43 –5 0 24 –4 6 21 –3 8 37 35 –4 8 33 –4 2 44 –4 7 l ow er o ra l j aw te et h 31 36 34 –4 2 24 –3 4 23 –3 6 32 25 –3 7 42 –5 0 44 –5 9 in ne r t ee th ro w s 1 1 2 1– 2 1– 2 1 1– 2 1– 3 1– 2 v er te br ae 31 32 31 29 –3 1 29 30 30 30 –3 1 31 european journal of taxonomy 672: 1–18 (2020) 14 isognathous. lower jaw long. posterior side of lower jaw protrudes anteriorly. anterior side of lower jaw slightly wider than posterior side. deep chin. larger specimen with larger mouth inclination and deeper chin. lip longer than half length of lower jaw. lips normal. lower lip with mucosa embedding teeth. preopercle also inclined. long and slender gill rakers. teeth. outer row of teeth on upper and lower jaws mostly unequally bicuspid and some posterior teeth unicuspid in smaller specimen; anterior teeth exclusively unicuspid on both jaws of larger specimen. teeth straight and not curved inwards. anterior cusps of teeth on outer row in both jaws larger than posterior cusps in smaller specimen. anterior teeth larger than posterior teeth in smaller specimen. one inner row of irregularly placed tricuspid teeth on both jaws in smaller specimen. posterior and inner teeth in larger specimen not readily observable, being to a large extent or fully covered by fleshy gums. teeth closely set (space between teeth about half to one tooth width). fins. pectoral fin origin behind level of dorsal fin origin in smaller specimen. position of pectoral fin origin unknown in larger specimen because of damage. pelvic fin origin positioned slightly more backwards than level of dorsal fin origin. pectoral fin of holotype and pelvic fin in both types almost to level of anus. anal fin anterior to level of first soft dorsal fin ray. small scales on base of caudal fin rays. fig. 5. photograph of preserved paratype (rmca 99-041-p-4767) of protomelas krampus sp. nov., shortly after preservation. fig. 6. photograph of live specimen of protomelas krampus sp. nov. (specimen not preserved; length unknown) (copyright ad konings). dierickx k. & snoeks j., a new paedophagous cichlid 15 colour pattern in life based on a photograph by konings (2016) (fig. 6). body generally greyish. head and pectoral fin base yellowish. pelvic fin with white distal part of leading edge. five orange-brownish eggspots on anal fin. continuous dark midlateral band from about one eye length behind opercle to caudal fin. supralateral row of darks spots. dark spots also present just below dorsal fin. some spots are arranged in an interrupted vertical bar pattern. fig. 7. distribution map of protomelas krampus sp. nov. red star = holotype; red dot = paratype; green triangles = possible sightings by konings (2016); orange square = possible sighting by mckaye & kocher (1983); yellow diamond = possible sighting by stauffer (pers. comm.). inset: map of africa with indication of area of lake malawi. european journal of taxonomy 672: 1–18 (2020) 16 colour pattern in preserved specimens based on photographs by mckaye & kocher (1983) and snoeks (2004) (fig. 5). body generally brown or greyish. dorsum somewhat darker than belly. darker on dorsal parts of head and body contiguous with dorsal fin base. clear dark maculae on spiny part of dorsal fin, possibly also on soft dorsal fin part and caudal fin. continuous dark midlateral band from one eye length or about three scales behind opercle to caudal fin. one supralateral and one subdorsal row of darks spots. some spots as subtle, incomplete vertical bars. both types currently pale-coloured, probably due to light exposure (fig. 4). geographical distribution (fig. 7) the specimens of p. krampus sp. nov. were found in lukoma bay (11°22.50′ s, 34°52.00′ e), south of mbamba bay, lake malawi (tanzania) and at rocks just south of mara point (12°11.34′ s, 34°41.73′ e), aldeia mala, lake malawi (mozambique), at depths of 32.5–33.2 m. three others specimens that also may belong to this species were caught near otter point (14°03′ s, 34°49′ e), mangochi, lake malawi (malawi), at a depth of 7–20 m (mckaye & kocher 1983). it has also been observed near chizumulu and likoma islands (konings 2016) and mazinzi reef (stauffer, pers. comm.). hence the species has a confirmed distribution in the central-eastern part of the lake, but it may also occur in the southern part. ecology the behaviour and ecology of specimens most likely belonging to this species have been observed by mckaye & kocher (1983) and konings (1989, 2016). they observed this species feeding on eggs and fry while stealing them from mouth-brooding females of other cichlid species. the paedophage rams these females from above on the snout and the brooding females may release some eggs or larvae upon this impact. the brood can then be snatched by p. krampus sp. nov. the inclined position of the mouth enables the fish to immediately grab the brood since it is already in a good position relative to the prey after ramming from above (konings 1989, 2016). discussion based on its melanin pattern, the new species was assigned to the genus protomelas. eccles & trewavas (1989) conducted an extensive review of the characteristics of all species of protomelas. based on these descriptions, only those species that are similar to the new paedophage species, p. krampus sp. nov., were included in the morphometric part of this study. since p. krampus sp. nov. has only one inner tooth row, all twelve species with two or more inner rows (see diagnosis) were excluded from the detailed morphometric analyses. eclectochromis ornatus (regan, 1922), which according to oliver (2012) and konings (2016) belongs to protomelas, was also excluded based on the number of inner tooth rows and melanin pattern as described by eccles & trewavas (1989). only p. spilopterus, which was suggested by eccles & trewavas (1989) to be the undescribed paedophage species observed by mckaye & kocher (1983), and p. insignis were retained for the morphometric comparison, as well as an undescribed species from this genus mentioned by snoeks & hanssens (2004) with some similar traits, h. sp. ‘insignis like’. in the morphometric analyses it was found that p. krampus sp. nov. is distinguished from its congeners among others by its large gape inclination, very short premaxillary pedicel, deep cheek, long lower jaw, deep chin, short and slender head, small interorbital width, and shorter predorsal and prepectoral distance. both c. orthognathus and p. krampus sp. nov. have been observed showing a similar ramming behaviour (mckaye & kocher 1983; konings 2016), though the number of observations is limited; therefore, their strikingly similar morphology is probably linked to their food acquisition. the shorter head, the deep chin and short premaxillary pedicel of p. krampus sp. nov. enhance the aspect of an enlarged ventral part of the head. this may be an adaptation to the ramming behaviour typical of this species. caprichromis orthognathus rams from beneath, whereas p. krampus sp. nov. rams from above with the ventral part dierickx k. & snoeks j., a new paedophagous cichlid 17 of the head. when the chin is enlarged, it is easier to use it as a ram from this orientation and possibly to have more impact on the snout of the prey. this may also explain the large gape inclination. the different angle of attack and slight differences in morphology may be explained by niche partitioning (mckaye & kocher 1983). the precise phylogenetic relationships between the paedophagous species should, however, be further studied with genetic analyses and a thorough taxonomic revision. during our analyses, it became clear that there may be an additional undescribed species within the genus protomelas. the lectotype of p. spilopterus and one paralectotype (tag 2583 within bmnh 1935.6.14.652-657) differ from the other paralectotypes in several features. the head of these two specimens is steeper and the neurocranial crest appears to be more rounded. they show a clear chin, whereas the other paralectotypes lack a chin. the jaws of the lectotype and some paralectotypes are rounded, while the others have wide and anteriorly flattened jaws. the lacrimal appears to be somewhat larger and has a more rounded anterior side in the lectotype and paralectotype compared to the other paralectotypes. the caudal fin of the lectotype and paralectotype is emarginate, whereas the other paralectotypes of p. spilopterus have a truncate caudal fin with a longer dorsal lobe than ventral lobe. furthermore, the lectotype has bicuspid teeth, the second cusp of which is very small, in the outer row on both jaws, and a mixture of uniand tricuspid teeth on the two inner rows. its teeth are curved inwards. the teeth of all paralectotypes are angled forwards and are uniquely unicuspid and conical, and there is only one inner tooth row on both jaws. the deepest point of the body occurs at the level of the pectoral fin origin in the lectotype and posterior to the pectoral fin origin in most paralectotypes. the eggspots on the anal fin of the lectotype are placed in three rows: the distal row with seven spots, the middle row with three spots and one spot in the proximal row. only one other paralectotype has eggspots in two rows with five spots on the distal row and four or five spots on the proximal row. a more detailed study is necessary to assess whether or not the type series is polyspecific. acknowledgements we acknowledge the project members of the sadc/gef taxonomy team and the crew of the r/v usipa for their logistical support. we are grateful to oliver crimmen and james macclaine (natural history museum, london) for the loan of some specimens under their care and to ad konings for the use of his photograph of a live specimen. references eccles d.h. & trewavas e. 1989. malawian cichlid fishes: the classification of some haplochromine genera. lake fish movies, herten, west germany. fricke r., eschmeyer w.n. & van der laan r. 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(ed.) the cichlid diversity of lake malawi/nyassa/ niassa: identification, distribution and taxonomy: 12–19. cichlid press, el paso, tx. snoeks j. & hanssens m. 2004. identification guidelines to other non-mbuna. in: snoeks j. (ed.) the cichlid diversity of lake malawi/nyassa/niassa: identification, distribution and taxonomy: 266–310. cichlid press, el paso, tx. stauffer j.r. jr. & mckaye k.r. 1986. description of a paedophagous deep-water cichlid (teleostei: cichlidae) from lake malawi, africa. proceedings of the biological society of washington 99: 29–33. turner g.f. 1996. offshore cichlids of lake malawi. cichlid press, lauenau, germany. van steenberge m., vanhove m.p.m., breman f.c. & snoeks j. 2015. complex geographical variation patterns in tropheus duboisi marlier, 1959 (perciformes, cichlidae) from lake tanganyika. hydrobiologia 748: 39–60. https://doi.org/10.1007/s10750-013-1692-5 manuscript received: 1 february 2020 manuscript accepted: 11 may 2020 published on: 23 june 2020 topic editor: rudy jocqué section editor: felipe polivanov ottoni desk editor: danny eibye-jacobsen printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany; national museum, prague, czech republic. https://doi.org/10.11646/zootaxa.3410.1.3 https://doi.org/10.1016/s0065-2504(00)31005-4 https://doi.org/10.1007/s10750-013-1692-5 european journal of taxonomy 266: 1–9 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2017.266 www.europeanjournaloftaxonomy.eu 2017 · le gall l. et al. this work is licensed under a creative commons attribution 3.0 license. d n a l i b r a r y o f l i f e 1 toward the dna library of life line le gall 1, frédéric delsuc 2, stéphane hourdez 3, guillaume lecointre 4 & jean-yves rasplus 5 1, 4 institut de systématique, évolution, biodiversité, isyeb – umr 7205 – cnrs, mnhn, upmc, ephe, muséum national d’histoire naturelle, sorbonne universités, 57 rue cuvier, cp 39 75005, paris, france. 2 institut des sciences de l’evolution, umr 5554, cnrs, ird, ephe, université de montpellier, montpellier, france. 3 station biologique de roscoff, cnrs – upmc, umr 7144, adaptation et biologie des invertebrés en conditions extrêmes, 29680 roscoff, france. 5 inra, umr 1062, cbgp centre de biologie pour la gestion des populations, 34988 montferrier-sur-lez, france. 1 email: legall@mnhn.fr 2 email: frederic.delsuc@umontpellier.fr 3 email: hourdez@sb-roscoff.fr 4 email: guillaume.lecointre@mnhn.fr 5 email: jean-yves.rasplus@inra.fr abstract. the special set of papers entitled “dna library of life” constitutes an outcome of the project “bibliothèque du vivant” (bdv), which aims to promote the molecular taxonomy of eukaryotes by offering research teams the possibility to produce and manage a molecular library linked with specimens deposited in natural history museums. the project was funded by three french institutions (the cnrs, inra and mnhn), and provided access to the sequencing power offered by the genoscope for 105 teams between 2011 and 2013. it was subsequently supported by the cnrs through the “groupement de recherche génomique environnementale”. the scientific objectives of this programme were threefold: 1) species delimitation among species complexes; 2) phylogenetic reconstruction (including phylogenomics); and 3) metabarcoding and improving ngs methods for systematic purposes. within the present collection, 19 papers contribute to these objectives across a large taxonomic range and a worldwide geographic coverage. these papers propose taxonomic novelties (22 new species and 3 new genera) in both animal and plant taxa. keywords. census of life, eukaryotic biodiversity, molecular assisted systematics, museum life, species description. le gall l., delsuc f., hourdez s., lecointre g. & rasplus j.-y. 2017. toward the dna library of life. european journal of taxonomy 266: 1–9. http://dx.doi.org/10.5852/ejt.2017.266 http://dx.doi.org/10.5852/ejt.2017.266 www.europeanjournaloftaxonomy.eu https://creativecommons.org/licenses/by/3.0/ mailto:legall%40mnhn.fr?subject= mailto:frederic.delsuc%40umontpellier.fr?subject= mailto:hourdez%40sb-roscoff.fr?subject= mailto:guillaume.lecointre%40mnhn.fr?subject= mailto:jean-yves.rasplus%40inra.fr?subject= http://dx.doi.org/10.5852/ejt.2017.266 european journal of taxonomy 266: 1–9 (2017) 2 a brief history of the dna library of life long before the discovery of dna (watson & crick 1953), humans had already started to distinguish, name and classify organisms. the first inventories could be found within the written texts left by ancient greek philosophers, especially aristotle and his pupil theophrastus (4th–3rd century ad) who greatly contributed to organizing a census of existing species, classifying them into “animal” for organisms with the ability to move and “vegetal” for “unanimated” ones (for an in-depth review, see mayr 1982). while the usage of some organisms (especially plants) had been well documented in the middle ages, almost no progress was made on the classification of organisms before the renaissance and its technological advents. the construction of the microscope by a. van leeuwenhoek (hamarneh 1960) fostered the observation of tiny organisms and unveiled a previously invisible world that accommodates a wealth of species. a formal system of naming species was subsequently proposed by carl von linné in the species plantarum (linnaeus 1753). swiftly, the binomial system gained recognition in the scientific world and almost two million species have since been described, which nevertheless may represent only half (costello et al. 2013) – or even as little as one fifth (mora et al. 2011) – of the overall eukaryotic diversity on earth. whereas only morpho-anatomical characters were initially used to describe species, more recently the use of molecular characters has allowed biologists to estimate phylogenetic relationships more reliably and to discriminate between morphologically cryptic species. in providing a comprehensive phylogenetic framework, molecular systematics has enhanced our understanding of species diversity by unravelling cryptic diversity, morphological convergence, heteromorphic life cycles, sexual dimorphism and other evolutionary phenomena. molecular data consist of universal and quantifiable characters, which present the advantage of being both objective and abundant compared to morphological ones. the comparison of homologous characters facilitates the reconstruction of historical relationships among biological entities, as well as the estimation of their divergence times. even if molecular characters are not completely devoid of pitfalls, especially when inferring species relationships from gene genealogies (e.g., collins & cruickshank 2012), they have proven to be amenable in establishing libraries of life for prokaryotes and eukaryotes (e.g., hebert et al. 2003; blaxter 2004; quast et al. 2013), albeit with a few limitations. identifying what level of molecular difference defines a species is also a challenging task, as it refers to the debated plethora of species concepts and ultimately relates to underlying speciation mechanisms (roux et al. 2016). nonetheless, several methods for species delimitation using dna barcoding data have been proposed (pons et al. 2006; puillandre et al. 2012; zhang et al. 2013). moreover, the addition of novel data has in some cases challenged previous species delimitations, highlighting the need for automated and time-proficient algorithms to delimit species (ratnasingham & hebert 2013), ideally relying on the analysis of multiple loci in a coalescent framework (yang & rannala 2014). despite the apparent simplicity of using dna sequences to build a library of life, the number of species collected in the core database is far from representative of the diversity on earth. for instance, on the genbank taxonomy page (anonymous 2017), it is stated that all the organisms in the public sequence databases currently represent no more than 10% of the total number of described species. in the context of the on-going 6th global biodiversity crisis (mostly referred to as a mass extinction), the major challenge that taxonomists are currently facing is to describe species diversity before it actually becomes extinct. whereas some taxa and some regions on earth have been thoroughly studied (i.e., more than 70% of plants are presumed to have been described already), the biodiversity of certain communities (e.g., soil) and some of the smallest organisms (e.g., microbial eukaryotes (de vargas et al. 2015), fungi (hawksworth 2001) and small inconspicuous animals (appeltans et al. 2012)) remain poorly known, rendering the conservation status of those species impossible to assess (régnier et al. 2015). dna-assisted taxonomy is often seen as the holy grail in accelerated species discovery and description, as molecular data are abundant and amenable to quantitative statistical treatment. for instance, in the le gall l. et al., toward the dna library of life 3 marine realm, assuming that the current rhythm of species description is maintained, most of the species should be discovered before the end of this century (appeltans et al. 2012). nevertheless, the time-lapse between species discovery and species description is still 21 years on average (fontaine et al. 2012), which reminds us that the process leading to a formal description is long and leaves a gap between uncovered species and named species (yahr et al. 2016). the objective of this special set of papers in the european journal of taxonomy is to make a contribution, no matter how slim, toward the construction of a comprehensive dna library of life. contributions in this special set of papers to the dna library of life this special collection of articles is comprised of the present introduction, 18 research papers spanning the fields of biogeography, phylogeny and molecular systematics, as well as an opinion paper on the pitfalls in supermatrix phylogenomics (philippe et al. in press). the geographic coverage of the sampling conducted for the 18 research papers spans the entire planet and includes organisms from marine (castelin et al. in press a, castelin et al. in press b, fedosov et al. in press, galindo et al. in press, manghisi et al. in press, rousseau et al. in press, sabroux et al. in press) and terrestrial biomes (azofeifa-bolaños et al. in press, galkowski et al. in press, legendre et al. in press, le ru et al. in press, ohler & nicolas in press, prigent et al. in press, rabeau et al. in press, ramage et al. in press, soldati et al. in press, tu et al. in press, veron et al. in press). it is noteworthy that the freshwater biome is not covered by any of these studies, except for the paper by ohler & nicolas (in press) focusing on frogs that occupy freshwater habitats for at least part of their life cycle. interestingly, although the funding intended for this project originates from french institutions, only a single study (galkowski et al. in press) is limited to metropolitan france. most of the sampling for this project was performed in tropical areas, including some overseas french territories (fedosov et al. in press, rousseau et al. in press, ramage et al. in press, castelin et al. in press b, sabroux et al. in press, legendre et al. in press). last but not least, the majority of the papers deal with animals; the remainder deals with plants. there is no contribution on fungi or on unicellular eukaryote lineages. within this special set of papers for the dna library of life, ten papers propose taxonomic novelties, consisting of 22 new species (fedosov et al. in press, le ru et al. in press, tu et al. in press, galindo et al. in press, soldati et al. in press, azofeifa-bolaños et al. in press, galkowski et al. in press) and three new genera (rousseau et al. in press, rabeau et al. in press, sabroux et al. in press). the european journal of taxonomy “library of life” initiative includes the following papers (in order of publication, which will take place from the 30th of january to the 3rd of march, 2017): • le gall l., delsuc f., hourdez s., lecointre g. & rasplus j.-y. toward the dna library of life. • manghisi a., le gall l., bonillo c., gargiulo g.m., ribeira m.a. & morabito m. an assessment of the taxonomic status of the mediterranean endemic genus acrodiscus zanardini (halymeniales, rhodophyta). • fedosov a.e., stahlschmidt p., puillandre n., aznar-cormano l. & bouchet p. not all spotted cats are leopards: evidence for a hemilienardia ocellata species complex (gastropoda: conoidea: raphitomidae). • rousseau f., gey d., kurihara a., maggs c.a., martin-lescanne j., payri c., reviers b. de, sherwood a.r. & le gall l. molecular phylogenies support taxonomic revision of three species of laurencia (rhodomelaceae, rhodophyta), with the description of a new genus. • le ru b., capdevielle-dulac c., musyoka b.k., pallangyo b., njaku m., mubenga o., chipabika g., ndemah r., bani g., molo r. ong’amo g. & kergoat g.j. phylogenetic analysis and systematics of european journal of taxonomy 266: 1–9 (2017) 4 the acrapex unicolora hampson species complex (lepidoptera, noctuidae, noctuinae, apameini), with the description of five new species from the afrotropics. • ohler a. & nicolas v. which frog’s legs do froggies eat? the use of dna barcoding for identification of deep frozen frog legs (dicroglossidae, amphibia) commercialized in france. • ramage t., martins-simoes p., mialdea g., allemand r., duplouy a., rousse p., davies n., roderick g.k. & charlat s. a dna barcode-based survey of terrestrial arthropods in the society islands of french polynesia: host diversity within the symbiocode project. • rabeau l., gradstein s.r., dubuisson j.-y., nebel m., quandt d. & reeb c. new insights into the phylogeny and relationships within the worldwide genus riccardia (aneuraceae, marchantiophytina). • tu v.t., csorba g., ruedi m., furey n.m., son n.t., thong v.d., bonillo c. & hassanin a. comparative phylogeography of bamboo bats of the genus tylonycteris (chiroptera, vespertilionidae) in southeast asia. • galindo l.a., kool h.h. & dekker h. review of the nassarius pauperus (gould, 1850) complex (nassariidae). part 3, reinstatement of the genus reticunassa with the description of six new species. • soldati l., condamine f.l., clamens a.-l. & kergoat g.j. documenting tenebrionid diversity: progress on blaps fabricius (coleoptera, tenebrionidae, tenebrioninae, blaptini) systematics, with the description of five new species. • philippe h., de vienne d.m., ranwez v., roure b., baurain d. & delsuc f. pitfalls in supermatrix phylogenomics. • azofeifa-bolaños j.b., gigant l.r., nicolás-garcía m., pignal m., tavares-gonzález f.b., hágsater e., salazar-chávez g.a., reyes-lópez d., archila-morales f.l., garcía-garcía j.a., da silva d., allibert a., solano-campos f., rodríguez-jimenes g.d.c., paniagua-vásquez a., besse p., pérez-silva a. & grisoni m. a new vanilla species from costa rica closely related to v. planifolia (orchidaceae). • veron g., bonillo c., hassanin a. & jennings a.p. molecular systematics and biogeography of the hemigalinae civets (mammalia, carnivora). • sabroux r., corbari l., krapp f., bonillo c., le prieur s. & hassanin a. biodiversity and phylogeny of ammotheidae (arthropoda: pycnogonida). • pringent s.r., suwalski a. & veuille m. connecting systematic and ecological studies using dna barcoding in a population survey of drosophilidae (diptera) from mt oku (cameroon). • castelin m., williams s., buge b., maestrati p., lambourdière j., ozawa t., utge j., couloux a., alf a. & samadi s. untangling species identity in gastropods with polymorphic shells in the genus bolma (mollusca, vetigastropoda). • castelin m., de mazancourt v., marquet g., zimmerman g. & keith p. genetic and morphological evidence for cryptic species in macrobrachium australe and resurrection of m. ustulatum (crustacea, palaemonidae). • galkowski c., lebas c., wegnez p., lenoir a. & blatrix r. re-description of proformica nasuta (nylander, 1856) (hymenoptera, formicidae) using an integrative approach. • legendre f., grandcolas p. & thouzé f. molecular phylogeny of blaberidae (dictyoptera, blattodea) with implications for taxonomy and evolutionary scenarios. le gall l. et al., toward the dna library of life 5 perspectives toward a comprehensive dna library of life high-throughput dna sequencing has widened the field of possibilities for automated and accelerated taxon identification (coissac et al. 2016). this technological advancement has enhanced the use of molecular data to study functional ecology, organism interactions, community ecology and biodiversity, plus the dynamics and evolution of past and present interactions. nevertheless, further efforts are needed to aggregate a comprehensive dna library of life. the well-known taxonomic impediment is of course relevant, even if some authors have questioned it (appeltans et al. 2012). in addition, one of the major current challenges consists of improving our sampling methods and instrumentation to discover organisms that live in extreme environments. finally, species description is governed by the international code for zoological nomenclature (iczn 1999) and the international code of nomenclature for algae, fungi, and plants (mcneill & international association for plant taxonomy 2012), which leaves many microbial eukaryotic lineages orphaned and results in species belonging to the same phylogenetic lineage being described according to both codes (yilmaz et al. 2014). in order to perform effective biodiversity monitoring, upon which conservation depends, we are counting upon new technologies to further our understanding of biodiversity as a whole. this, however, will not be possible without a proper and well-curated taxonomic framework. we therefore urge our founding agencies to continue their financial support to taxonomy, with the aim of achieving a comprehensive dna library of life. acknowledgments we would like to express our sincere thanks to koen martens for accepting to host this special set of papers in the european journal of taxonomy. we are indebted to charlotte thionois and danny eibye-jacobsen who took care of most of the production process. the entire editorial team of european journal of taxonomy has always been most helpful. this special set of papers would not have been possible without the encouragement, the scientific support and the financial contributions of the institut ecologie et environnement (cnrs), the muséum national d’histoire naturelle, and the institut national de recherche agronomique. the sanger sequencing has been 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http://dx.doi.org/10.1093/bioinformatics/btt499 http://dx.doi.org/10.1093/bioinformatics/btt499 european journal of taxonomy 17: 1-31 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2012.17 www.europeanjournaloftaxonomy.eu 2012 · michael karner this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e 1 a revision of african psammoecus (coleoptera, silvanidae) and descriptions of two new species from the collection of the musée royal de l’afrique centrale michael karner dr. michael karner, c/o dr. damir kovac, sektion entomologie i, senckenbergisches naturforschendes institut, senckenberganlage 25, d-60325 frankfurt am main, germany email: mka_fra@yahoo.de abstract. a revision of the known african species of psammoecus is given, including redescriptions and illustrations of diagnostic characters. extensive material from the musée royal de l’afrique centrale (tervuren) is studied. two new species are described: psammoecus leleupi sp. nov., and ps. luchti sp. nov. four specific names are synonymized: psammoecus excellens grouvelle, 1908 = ps. trimaculatus motschulsky, 1858; ps. alluaudi grouvelle, 1912 = ps. trimaculatus motschulsky, 1858; ps. longulus grouvelle, 1878 = ps. longicornis schaufuss, 1872; ps. nitescens grouvelle, 1914 = ps. laetulus grouvelle, 1914. a key to the african species is provided. key words. coleoptera, africa, psammoecus, silvanidae, taxonomy. karner m. 2012. a revision of african psammoecus (coleoptera, silvanidae) and descriptions of two new species from the collection of the musée royal de l’afrique centrale. european journal of taxonomy 17: 1-31. http://dx.doi.org/10.5852/ejt.2012.17 introduction while examining extensive material of african psammoecus latreille, 1829 from the collection of the musée royal de l’afrique centrale (tervuren, belgium) it came clear that any attempt to determine the material at hand required studies of type material. since the studies of grouvelle (see references), no work was done on african species of this genus. the available papers, however, were found to be of little practical help, reflecting the taxonomic standards of the late 19th century. they frequently show a lack of detail in descriptions and diagnostic content. the present paper gives redescriptions, illustrations, and diagnoses for the known african species of psammoecus. two species contained in the material from the musée royal de l’afrique centrale are new to science. http://creativecommons.org/licenses/by/3.0/ www.europeanjournaloftaxonomy.eu european journal of taxonomy 17: 1-31 (2012) 2 material and methods specific identification of psammoecus is often possible by use of external characters, viz. general body shape, the pattern and form of pronotal teeth, and the puncturation of the body surface. the male genitalia provide diagnostic characters especially in the shape of the parameres. determination of single female specimens was sometimes found impossible, since female genitalia were not found to provide useful characters. an overview considering the general structure of the genitalia of psammoecus is given by pal (1985). although the genital structures are small and delicate, preparation of the genitalia is not very difficult. the animal is softened by soaking it in water overnight. the abdomen is separated from the body by use of a very fine insect pin, and macerated in warm, 5% solution of caustic potash (koh) for few minutes. after rinsing in water, the abdomen is dissected in a drop of 5% acetic acid to isolate the genitalia. it should never be attempted to pull the genitalia out of the abdomen, since this would involve a high risk of the structures, especially the setae, being destroyed. the isolated genitalia are embedded in a flat drop of embedding medium (either canada balsam or a water-soluble medium, e.g. as described by franzen & karner 1998) on a piece of transparent cellulose acetate film that is put on the specimen’s insect pin. drawings were made by aid of microscopes and a camera lucida at magnifications of 50 times and 400 times, respectively. the microsculpture of the body surface was studied at a magnification of 80 times. the lateral teeth bordering the pronotum were numbered, using roman numerals, beginning anteriorly, behind the group of very small teeth that are located at the anterior angle of the pronotum. measurements were taken as follows: length, from apical margin of clypeus to apex of elytra; head width, across eyes; head length, from apical margin of clypeus to imaginary line between hind margins of eyes; eye length, from anterior to posterior margin; antennal length, from base of 1st antennomere to apex of 11th antennomere; pronotal width, across maximum width, excluding spines; pronotal length, from anterior to posterior margin; elytral width, across maximum joint width; elytral length, along suture including scutellum. material is listed according to the localities, sorted from north-east to south-west. all label data are given. the labels are cited beginning with the uppermost one, the respective lines are separated by ‘|’. if words or single characters could not be deciphered, the characters are represented by a corresponding number of ‘?’ within squared brackets. also, comments on label colors, label shapes, and other regarding the respective specimen are included in squared brackets. specimens from the following collections were studied: bmnh = the natural history museum, london i.r.s.a.c. = institut pour la recherche scientifique en afrique centrale (the material studied here is stored in the collection of the musée royal de l’afrique centrale, tervuren) mhnaf = museu de história natural e aquário, funchal mkf = michael karner, frankfurt mnhn = muséum national d’histoire naturelle, paris mrac = musée royal de l’afrique centrale, tervuren zmhb = museum für naturkunde der humboldt-universität, berlin karner m., revision of african psammoecus 3 results classis hexapoda blainville, 1816 ordo coleoptera linnaeus, 1758 superfamilia cucujoidea latreille, 1802 familia silvanidae kirby, 1837 subfamilia brontinae erichson, 1845 tribus telephanini leconte, 1861 genus psammoecus latreille, 1829 psammoecus personatus grouvelle, 1919 (fig. 1) material examined holotype ♀: ‘funchal | madère’, ‘type’ [red label], ‘museum paris | 1917 | coll. grouvelle’, ‘psammoecus | personatus grl.’ [not grouvelle’s hand] (mnhn). other material 1♀ ‘musée du congo | haut-uele: moto | 1923 | l. burgeon’ (mrac). 1♂ ‘congo belge: p.n.a. | 7-9-iv-1955 | p. vanschuytbroeck | 12.743-58’, ‘secteur nord | riv. mukandwe, | affl. talya, 1.200 m’ (mrac). 1♂ ‘musée du congo | mombassa (36 km s. | lubero viii-1932 | l. burgeon’ (mrac). 2 spms ‘coll. mus. congo | uganda: 1400 m. | savane boisée | 5/8-viii-1957’, ‘mission zoolog. i.r.s.a.c. | en afrique orientale | p. basilewsky et | n. leleup’ (mrac, mkf). 1♀ ‘musée du congo | haut-uele: abimva | -v1925 | l. burgeon’ (mrac). 3♀♀ ‘musée du congo | haut-uele | watsa, xi-1919 | l. burgeon’ (mrac). 1♂ ‘musée du congo | ruvenzori: kalonge | (2050 m.) 9-viii-1932 | l. burgeon’ (mkf). 1♂ ‘coll. mus. congo | kivu: ibanda | 1952 | m. vandelanoite’ (mrac). 1♂ ‘i.r.s.a.c.-mus.congo | kivu: uvira | xi-1949 | n. leleup’ (mrac). 1♀ ‘i.r.s.a.c.-mus.congo | kivu: marais de la basse | kalimabenge (uvira) | n. leleup xi – 1955’ (mrac). 1♀ ‘i.r.s.a.c.-mus.congo | maniema: kabambare | (à la lumière) 16/18-x-54 | n. leleup’ (mrac). 1♀ ‘musée du congo | ifuri: adia | 1935 | 1.a | (r. belot.)’ (mrac). 1♀ ‘coll. mus. congo | elisabethville ii/iv-1949 | ch. seydel”, ‘a la lumière’ (mrac). 1♂ ‘coll. mus. congo | elisabethville, a la | lumière xi-50/vi-51 | ch. seydel’ (mrac). 1♂ ‘madeira 19-24.4.1978 | garajau | th. palm’, ‘psammoecus | personatus | grouvelle | det. g.israelson’ (mhnaf). differential diagnosis ps. personatus differs from ps. marginicollis grouvelle, 1908, ps. parallelus grouvelle, 1919, ps. laetulus grouvelle, 1914 and ps. luchti sp. nov. by the longer lateral teeth of the pronotum. it differs from ps. leleupi sp. nov. by the narrower elytral striae, narrower bases of lateral teeth of pronotum, and slender parameres with shorter and fewer setae; from ps. hacquardi grouvelle, 1889 by the 1st antennomere being less than two times as long as 2nd; from ps. laetulus by the maximum pronotal width being closer to the middle, narrower elytral striae, and less extensive dark elytral markings. redescription body. elongate oval, total length 2.50-3.00 mm (fig. 1a). surface yellowish brown. elytra with round brown or blackish-brown maculae on disc and narrow macula on posterior part of suture; some specimens without elytral maculae. antennae as in fig. 1b, yellowish brown, 7th to 10th and apex of 6th antennomere brown or blackish brown, 11th antennomere lighter than basal antennomeres. european journal of taxonomy 17: 1-31 (2012) 4 head. broad, temples rounded, immediately narrowed behind eyes; width 0.69-0.83 mm, length 0.360.45 mm, 1.71-1.90 times as wide as long. eyes protuberant, 0.21-0.24 mm long, distance of inner margins 0.43-0.54 mm. puncturation on vertex variable, moderate to very dense, distance between punctures irregular, slightly increasing anteriorly. pubescence composed of long, semierect setae, directed anteriorly; microsculpture absent. longitudinal impressions on vertex distinct, attaining anterior third of eyes. antennae 1.23-1.43 mm long; antennomere proportions of holotype as follows: 2.5 : 1.4 : 1.4 : 1.6 : 1.6 : 1.4 : 1.1 : 1.1 :1.1 : 1.0 : 2.3. pronotum. broad, width 0.71-0.88 mm, length 0.48-0.60 mm, 1.35-1.50 times as wide as long. anterior angles with distinct group of small, short teeth; lateral margins with five teeth, tooth i small, tooth ii larger, tooth iii the largest, tooth iv a little smaller than tooth ii, tooth v very small. posterior fig. 1. psammoecus personatus grouvelle, 1919. a. habitus of holotype. b. left antenna of holotype. c. parameres of specimen from kivu, ibanda. scale line a, b: 1 mm; c: 0.2 mm. karner m., revision of african psammoecus 5 angle diminutive. puncturation on disc coarser than on vertex, punctures sometimes hardly separated. pubescence as on vertex; microsculpture mostly not visible (holotypus), some specimens showing a very shallow reticulation. elytra. oval, length 1.63-2.00 mm, combined width 1.08-1.38 mm, 1.35-1.56 times as long as combined width. rows of punctures on disc somewhat narrower than interstices. pubescense composed of long, semierect setae, directed posteriorly; microsculpture absent. parameres. slender, curved basally, with small setae along the inner margins and single long seta at their apices (fig. 1c). psammoecus hacquardi grouvelle, 1889 (fig. 2) psammoecus haequardi [sic.] grouvelle, 1908b: 185. material examined lectotype ♀, by present designation: ‘zanzibar | raffray’ [blue label, grouvelle’s hand], ‘type’ [red label], ‘museum paris | 1917 | coll. grouvelle’ [blue label], ‘hacquardi | a. grou’ [not grouvelle’s hand] (mnhn). paralectotype ♀, by present designation, with identical data as lectotype (mnhn). other material 2 spms ‘tanzanie: mts. uluguru | kiroka, for. héliophile | alt. 725 m sous écorces | 27-31/v/71’, ‘coll. mus. tervuren | mission mts. uluguru | l. berger, n. leleup | j. debecker v/viii/71’ (mrac). 3 spms ‘tanzanie: mts. uluguru | kiroka, for. héliophile | alt. 600 m 24-30/vii/71 | dans arbres morts’, ‘coll. mus. tervuren | mission mts. uluguru | l. berger, n. leleup | j. debecker v/viii/71’ (mrac, 1 spm mkf). 1 spm ‘tanzanie: mts. uluguru | kiroka, for. héliophile | alt. 600 m 24-30/vii/71’, ‘coll. mus. tervuren | mission mts. uluguru | l. berger, n. leleup | j. debecker v/viii/71’ (mrac). 2 spms ‘a la lumière’ [blue label], ‘i.r.s.a.c.-mus.congo | kivu: terr masisi, 720 m. | walikale (rives loa) ix53 | n. leleup’ (mrac, mkf). 3 spms ‘dans | humus | en forêt’ [blue rectangular label], ‘i.r.s.a.c.mus.congo | kivu: terr masisi, 800 m. | mutakato ix-53 | n. leleup’, (mrac). 1♀ ‘musée du congo | eala | xi-1934 | j. ghesquiére’ (mrac). 3 spms ‘musée du congo | eala | 10-vii-1936 | j. ghesquiére | 2873’ (mrac). 1♂ ‘musée du congo | eala | vii-1936 | j. ghesquiére | 2875’, (mrac). 1♀ ‘i.r.s.a.c.-mus.congo | bolobo (fleuve congo | 5-x-1949 | n. leleup’, (mrac). 1 spm ‘coll. mus. tervuren | côte d‘ivoire: bingerville | xi. 1961 | j. decelle’ , (mrac). 4 spms ‘coll. mus. tervuren | côte d‘ivoire: bingerville | viii. 1962 | j. decelle’ (mrac, 1 spm mkf). 2 spms ‘coll. mus. tervuren | côte d’ivoire: bingerville | ix. 1962 | j. decelle’ (mrac). 1 spms ‘coll. mus. tervuren | côte d’ivoire: bingerville | x. 1962 | j. decelle’, (mrac). 1 spm ‘coll. mus. tervuren | côte d’ivoire: bingerville | xi. 1962 | j. decelle’ (mrac). 1 spm ‘coll. mus. tervuren | côte d’ivoire: bingerville | ix. 1963 | j. decelle’ (mrac). differential diagnosis ps. hacquardi differs from ps. marginicollis by the shape of the parameres and shorter antennae; from ps. personatus by the 1st antennomere being more than two times longer as 2nd; from ps. leleupi sp. nov. by the narrower elytral striae; from ps. laetulus by the maximum pronotal width being closer to the middle; from ps. parallelus by more extensive dark maculae on elytra, darkened elytral basis and slender shape european journal of taxonomy 17: 1-31 (2012) 6 of parameres; from ps. luchti sp. nov. by narrow bases of lateral teeth of pronotum, narrower elytral striae and darkened elytral bases. redescription body. elongate-oval, total length 2.40-3.05 mm (fig. 2a). surface yellowish to reddish brown, elytra with variable patterns of maculae; the basic pattern consists of brown or blackish brown maculae at the elytral bases, sutures, lateral margins and apices. specimens of lighter colour show no darker coloration on anterior half of elytral suture and posterior half of lateral margins; very dark specimens have dark elytra with light brown maculae on the anterior and posterior aspects of the disk. antennae as in fig. 2b, yellowish brown; 6th to 10th and apical half of 5th antennomere brown to blackish-brown, 11th antennomere yellowish brown, sometimes a little lighter than basal antennomeres. fig. 2. psammoecus hacquardi grouvelle, 1889. a. habitus of lectotype. b. left antenna of lectotype. c. parameres of specimen from tanzanie: uluguru mountains. scale line a, b: 1 mm; c: 0.2 mm. karner m., revision of african psammoecus 7 head. broad, temples immediately narrowed behind eyes; width 0.65-0.74 mm, length 0.38-0.44 mm, 1.61-1.80 times as wide as long. eyes moderately protuberant, 0.19-0.23 mm long, distance of inner margins 0.40-0.48 mm. puncturation on vertex moderate, density of punctures in some specimens irregular, varying from sparse to very dense. pubescence composed of long, semierect setae, directed anteriorly; microsculpture distinct, reticulate. longitudinal impressions on vertex distinct, almost attaining the middle of the eyes. antennae comparatively short, 1.30-1.43 mm long; antennomere proportions of lectotype as follows: 2.8 : 1.0 : 1.7 : 1.1 : 1.5 : 1.3 : 1.2 : 1.2 :1.2 : 1.2 : 2.7. pronotum. broad, some specimens with shallow impressions close to the apical margin; width 0.650.78 mm, length 0.50-0.60 mm, 1.27-1.33 times as wide as long. anterior angles with distinct groups of small, short teeth; lateral margins with five teeth, tooth i very short and slender, tooth ii longer and wider than tooth i, tooth iii longer than tooth ii, tooth iv a little shorter than tooth iii, tooth v very small; posterior angle with a very small, almost obtuse tooth. puncturation on disc denser than on vertex, punctures sometimes hardly separated. pubescence as on vertex; microsculpture distinct, reticulate. elytra. elongate-oval, length 1.50-1.90 mm, combined width 0.98-1.30 mm, 1.46-1.67 times as long as combined width. rows of punctures on disc narrower than interstices. pubescence composed of long, semierect setae; microsculpture absent. parameres. slender, short, with dense rows of small setae at the inner margins of their bases and two larger setae at their apices. (fig. 2c). remarks the determination of single female specimens, especially if they are lightly colored or immature, can be difficult. grouvelle (1908b) spells the name ‘haequardi’, which is to be considered a misspelling. psammoecus laetulus grouvelle, 1914 (fig. 3) psammoecus nitescens grouvelle, 1914: 149. syn. nov. material examined lectotype ♀, by present designation: ‘25’ [hand written on specimen’s mounting label], ‘type | h.t.’ [round label with red border], ‘seychelle islands. | percy sladen | trust expedition | 1913.-170’, ‘psammoecus | laetulus grouv. | type | figured specimen’ [not grouvelle’s hand; last line on reddish piece of paper glued to the label], ‘psammoecus | laetulus | type grouv.’ [grouvelle’s hand, ‘type’ printed on blue piece of paper glued on the label, mounted facing down], ‘silhouette, 1908. | seychelles exp.’ [mounted facing down], ‘syn| type’ [round label with blue border] (bmnh) other material 1 spm ‘25’ [hand written on specimen’s mounting label], ‘type | h.t.’ [round label with red border], ‘seychelle islands. | percy sladen | trust expedition | 1913.-170’, ‘silhouette, 1908. | seychelles exp.’ [mounted facing down], ‘psammoecus | nitescens | type grouv.’ [grouvelle’s hand, ‘type’ printed on blue piece of paper glued on the label], ‘psammoecus | nitescens | grouvelle | type figured’ [not grouvelle’s hand; ‘figured’ printed on reddish piece of paper glued to the label, mounted facing down], ‘syn| type’ [round label with blue border] (bmnh). this specimen is hereby designated to be the lectotypus of ps. nitescens. 5 spms ‘seychelle islands. | percy sladen trust | expedition | 1913.170’, ‘psammoecus | nitescens | a. grouvelle | paratype’, ‘seychelle islands. | percy sladen | trust expedition | 1913.-170’, ‘syn| type’ [round label with blue border] (bmnh). these specimens are european journal of taxonomy 17: 1-31 (2012) 8 hereby designated to be paralectotypes of ps. nitescens. 96 spms ‘iles séchelles – mahé | centre: morne séch| ellois, for. endémique, | 750 m. 13/17.vii.1972’, ‘coll. mus. tervuren | miss. zool. belge aux | séchelles – plg benoit | et. j. j. van mol’ (mrac, 4 spms in coll. mkf). differential diagnosis ps. laetulus differs from ps. hacquardi, ps. personatus, and ps. leleupi sp. nov. by the maximum pronotal width being located in the anterior third. it differs from ps. personatus also by shorter lateral teeth of pronotum, wider elytral striae, and more extensive dark elytral markings; from ps. leleupi sp. nov. also by lateral margin of pronotum less curved, slender parameres with shorter setae; from ps. marginicollis fig. 3. psammoecus laetulus grouvelle, 1914, specimen from mahé. a. habitus. b. left antenna. c. parameres. scale line a, b: 1 mm; c: 0.2 mm. karner m., revision of african psammoecus 9 by 1st antennomere being more than two times as long as 2nd; from ps. parallelus by more extensive dark maculae on elytra, darkened elytral basis and slender shape of parameres; from ps. luchti sp. nov. by less rounded lateral margin of pronotum, darkened elytral basis, and more slender parameres. redescription body. oval, total length 2.35-2.95 mm (fig. 3a), head and pronotum blackish-brown with the head in most specimens somewhat brighter; elytra yellowish brown with darkened humeral area, dark lateral margin, a wide transversal band behind the middle and apex dark. in many specimens the darkened areas on the elytra are enlarged, leaving only bright maculae on the anterior and posterior portions of the elytra. antennae as in fig. 3b, bright reddish-brown, 7th to 10th antennomeres and apex of 6th antennomere blackish-brown, 11th antennomere bright, almost white. head. transverse, 0.69-0.75 mm wide, 0.39-0.48 mm long, 1.53-1.77 times as wide as long. eyes large, protuberant, 0.21-0.23 mm long, distance of inner margins 0.43-0.48 mm. puncturation on vertex coarse, moderately dense; pubescence composed of long, semierect setae, directed anteriorly; microsculpture absent. longitudinal impressions on vertex distinct, almost parallel, attaining middle of eyes. antennae long and slender, 1.45-1.63 mm long, antennomere proportions as follows: 2.7 : 1.0 : 1.2 : 1.7 : 1.7 : 1.6 : 1.4 : 1.2 : 1.2 : 1.1 : 2.3. pronotum. subquadrate, posterior margin with shallow impressions; 0.66-0.78 mm wide, 0.53-0.63 mm long, 1.15-1.33 times as wide as long. anterior angles with small group of teeth; margins with five lateral teeth; tooth i small, only slightly bigger than teeth of anterior group; tooth ii somewhat bigger, tooth iii biggest, teeth iv and v small; posterior angle with very small tooth. puncturation and pubescence on pronotal disc in most specimens as on vertex, sometimes a little coarser. microsculpture variable, mostly absent, in few specimens shallow, reticulate. elytra. oval, length 1.54-1.88 mm, combined width 1.10-1.18 mm, 1.40-1.52 times as long as their combined width. rows of punctures on disc slightly narrower than interstices; pubescence composed of long, semi-erect setae, directed anteriorly; microsculpture absent. parameres. slender, narrowing gradually towards apex; inner margins with numerous small setae, few setae on outer margins of apical struts; apex with single large seta (fig. 3c). remarks both ps. laetulus and ps. nitescens are described in the same paper by grouvelle (1914). due to the name ps. laetulus preceding in the original article, it is considered to be the valid name for this taxon, with ps. nitescens being a synonym. psammoecus parallelus grouvelle, 1919 (fig. 4) material examined holotype ♀: ‘salisbury | mashonaland | aug. 1900 | gakm’ [on backside of mounting label], ‘type’ [round label with orange border], ‘salisbury, | mashonaland. | marshall. | 1902-245’, ‘psammoecus | parallelus | typ | grouv’ [grouvelle’s hand] (bmnh). other material 1 spm ‘coll. mus. congo | kasenyi | 19-viii-1937 | h.j. brédo’ (mrac). 1♀ ‘dans marais | roselier’, ‘i.r.s.a.c. – mus. congo | kibali-ituri: blukwa (tsa) | 1820 m. i-1954 | n.leleup’ (mrac). european journal of taxonomy 17: 1-31 (2012) 10 1♀ ‘i.r.s.a.c. mus. congo | bendera, terr. albertville | 1000 m. (à la lumière) | n. leleup x-1958’ (mrac). 1♂ ‘i.r.s.a.c. mus. congo | bendera, terr. albertville | riv. kiymbi 1000 m. | b. 18 n, leleup vii-1958’, ‘biot. no 18 | dans détritus | et humus’ (mrac). 1♂ ‘coll. mus. congo | urundi: bugesera, rives | lac tshohoha yx-1957 | n. leleup’ (mkf). 1 spm ‘musée du congo | haut-uelè: watsa | 1922 | l. burgeon’ (mrac). 1♀ ‘récolté dans | panicule de | riz’, ‘coll. mus. congo | 15 kn e. yangambi | 21 1952 | j .decelle’ (mrac). 2 spms ‘coll. mus. congo | kivu: sanghe, pl. ruzizi | (à la lumière) xii-1951 | h. bomans’ (mrac). 1♀ ‘i.r.s.a.c. – mus. congo | kivu: lemera, terr. uvira | 1600 m. xii-1956 | n. leleup’ (mrac). 2♀♀ ‘biot. no 25 | racines de | plantes basses’, ‘i.r.s.a.c. mus. congo | mont kabobo, terr. albert| ville, hte kiymbi 1700 m. | b.25 n. leleup ix-1958’ (mrac). 1 spm ‘coll. mus. congo | nioka | vii-1937 | j. ghesquière’ (mrac). 1 spm ‘coll. mus. congo | sankuru: gandajika | v-1955 | j. decelle’ (mrac). 1 spm fig. 4. psammoecus parallelus grouvelle, 1919. a. habitus of holotype. b. left antenna of holotype. c. parameres of specimen from bendera. scale line a, b: 1 mm; c: 0.2 mm. karner m., revision of african psammoecus 11 ‘coll. mus. congo | tanganika: musosa, 980 m. | (à la lumière) xi-1953 | h. bomans’ (mrac). 1♀ ‘coll. mus. congo | elisabethville (a la | lumière) x-1959 | ch. seydel’ (mkf). 1♂ ‘i.r.s.a.c. mus. congo | bas-congo: rives de | la n’tadi 9-ix-1949 | n. leleup’ (mrac). 1 spm ‘musée du congo | kisantu | -1932 | r.p. vanderyst’ (mrac). 1♀ ‘coll. mus. congo | bas congo: thysville | 1-2/xii-1952 | p. basilewsky’ (mrac). 1 spm ‘coll. mus. congo | côte d’ivoire: adiopo| doumé 1957 | p. dessart’ (mrac). differential diagnosis differs from ps. personatus and ps. leleupi sp. nov. by the shorter lateral teeth of pronotum; from ps. leleupi sp. nov. also by narrower elytral striae, round dark maculae in posterior third of elytra, darkened suture near elytral apex, and the absence of a darkened area at bases of elytra; from ps. hacquardi and ps. laetulus by round dark maculae in posterior third of elytra, darkened suture near elytral apex, the absence of a darkened area at bases of elytra and very wide bases and club-shaped extensions of parameres; from ps. marginicollis by 1st antennomere being almost three times as long as 2nd and from ps. marginicollis and ps. luchti sp. nov. by very wide bases and club-shaped extensions of parameres. redescription body. slender, total length 2.85-3.30 mm (fig. 4a), surface yellowish-brown, elytra with round dark maculae at beginning of posterior third near lateral margins, suture darkened along posterior third. antennae yellowish brown, 7th-10th and apex of 6th antennomere brown, 11th antennomere lighter reddishbrown, but somewhat darker than basis of antennae. head. wide, temples curved; width 0.48-0.78 mm, length 0.40-0.45 mm, 1.41-1.82 times as wide as long. eyes protuberant, 0.21-0.23 mm long, distance of inner margins 0.48-0.53 mm. puncturation on vertex coarse, dense, density of punctures variable. pubescence composed of long, semierect setae, directed anteriorly; microsculpture visible only in few specimens, very shallow, reticulate. longitudinal impressions on vertex distinct, extending slightly behind middle of eyes. antennae as in fig. 4b, slender, 1.35-1.63 mm long; antennomere proportions of holotype as follows: 2.9 : 1.2 : 1.4 : 1.5 : 1.5 : 1.3 : 1.2 : 1.1 : 1.0 : 1.0 : 2.2. pronotum. subquadrate, 0.73-0.81 mm wide, 0.60-0.68 mm long, 1.19-1.26 times as wide as long. anterior angles with distinct group of small teeth; lateral margins with five teeth, tooth i small, teeth ii and iii larger, of similar size, teeth iv and v as big as tooth i; posterior angles with small tooth. puncturation and pubescence on disc as on vertex, microsculpture absent. elytra. oval, length 1.80-2.10 mm, combined width 1.03-1.25 mm, 1.67-1.76 times as long as their combined width. rows of punctures on disc slightly wider than interstices; pubescense composed of long, semierect setae, directed posteriorly; microsculpture absent. parameres. with very large base and club-shaped apical extension. inner margins of base and apical extension with numerous setae of variable size, apex with one large seta (fig. 4c). psammoecus leleupi sp. nov. (fig. 5) etymology the name is dedicated to narcisse leleup (1912-2001), who collected many of the psammoecus stored in the the musée royal de l’afrique centrale. european journal of taxonomy 17: 1-31 (2012) 12 material examined holotype ♂: ‘coll. mus. congo | kivu: t. lubero, mt kibat| siro, visiki, 2080 m. xii-54 | r.p.m.j. célis’, ‘dans | terreau, au | berlese’, ‘i.r.s.a.c. – mus. congo | n. rhodesia: mpika | (muchinga mts.) 1700m. | vii.1960 n. leleup’, ‘dans | l’humus’ [blue label], ‘holotypus | psammoecus | leleupi | des. karner’ [red label] (mrac). paratypes 1♂ ‘coll. mus. tervuren | uganda: distr. busonga | env. jinja, i/iii.1968 | j-j. rwabuneza’, ‘paratypus | psammoecus | leleupi | des. karner’ [red label] (mrac). 1 spm ‘19 km sud | tapili’, ‘à la lampe | iv-1956’, ‘collection | j.g. pantos’, ‘622’, ‘coll. mus. congo | 622 | ex coll. j. pantos’, ‘paratypus | psammoecus | leleupi | des. karner’ [red label] (mrac). 1♀ ‘musée du congo | kivu: mabuita | xii.1935 | boutakoff’, ‘paratypus | psammoecus | leleupi | des. karner’ [red label] (mrac). 1♀ ‘musée du congo | rutshuru | iv-1937 | j. ghesquière’, ‘paratypus | psammoecus | leleupi | des. karner’ [red label] (mrac). 1 spm ‘ coll. mus. congo | ruanda: kayove, | 2000 m. terr. kisenyi, | p. basilewsky 14/ii-53’, ‘paratypus | psammoecus | leleupi | des. karner’ [red label] (mrac). 1♀ ‘ i.r.s.a.c.-mus. congo | kivu: terr. masisi, 700 m. | walikale ix1953 | n. leleup’, ‘paratypus | psammoecus | leleupi | des. karner’ [red label] (mrac). 1♀ ‘coll. mus. tervuren | burindi: mugera | vi/vii.1965 | j.j. rwabuneza’, ‘paratypus | psammoecus | leleupi | des. karner’ [red label] (mrac). 1♀ ‘musée du congo | k. 300. de kindu | 16-iv-1911 | l. burgeon’, ‘paratypus | psammoecus | leleupi | des. karner’ [red label] (mrac). 1♀ ‘musée du congo | manyema 1918 | katangar. kasa | dr. gérard’, ‘paratypus | psammoecus | leleupi | des. karner’ [red label] (mrac). 1♀ ‘i.r.s.a.c.-mus. congo | maniema: t. kasongo, | mwanakussu | n. leleup 21/22-x-54’, ‘a la lumière’, ‘paratypus | psammoecus | leleupi | des. karner’ [red label] (mrac). 1♀ ‘ musée du congo | lulouga: befale | -ix-1927 | l. [sic!] ghesquière’, ‘paratypus | psammoecus | leleupi | des. karner’ [red label] (mrac). 1♂ ‘i.r.s.a.c.-mus. congo | kwango: terr. de feshi, | rive g. kwenge ii-1959 | b.64 mme. j. leleup’, ‘biot. no 64 | llôt de forêt | marécag. inondée’, ‘paratypus | psammoecus | leleupi | des. karner’ [red label] (mkf). 1♂ ‘i.r.s.a.c.-mus. congo | kwango: kianza, terr. de | feshi, riv. sengi iii-1959 | b.76 mme j. leleup’, ‘biot. no 76 | tête de source | humus en forêt’, ‘paratypus | psammoecus | leleupi | des. karner’ [red label] (mrac). 1♀ ‘i.r.s.a.c.-mus. congo | thysville: grotte | b 15 d. 22-vii-1949 | n. leleup’, ‘paratypus | psammoecus | leleupi | des. karner’ [red label] (mrac). diagnosis maximum pronotal width closely behind anterior third; margin of pronotum rounded, bearing four triangular lateral teeth with wide bases. elytra with dark brown to blackish maculae at base, on the disc and with a dark apex. the large maculae on the disc often form a transverse band, connected to the apical maculae by the darkened suture; striae on elytral disc wider than interstices. parameres with wide bases and club-shaped extensions, bearing numerous long setae. differential diagnosis ps. leleupi sp. nov. differs from ps. hacquardi, ps. parallelus, ps. marginicollis, and ps. personatus by wider elytral striae; from ps. personatus also by wider bases of lateral teeth of pronotum, and broad parameres with numerous long setae. it differs from ps. laetulus by maximum pronotal width being closer to middle, lateral margin of pronotum more curved, parameres with wide basis, club-shaped extensions, and longer setae; from ps. marginicollis by longer lateral teeth of pronotum, and by parameres with wide basis and club-shaped extensions; from ps. parallelus by longer lateral teeth of pronotum, and extensive dark maculae on elytra with darkened area at elytral basis; from ps. luchti sp. nov. by darkened elytral bases and parameres with wide basis, club-shaped extensions, and numerous longer setae. karner m., revision of african psammoecus 13 description body. elongate oval, total length 2.80-3.20 mm (fig. 5a). elytra with dark brown to blackish maculae at base, on the disc and with a dark apex. the large maculae on the disc often form a transverse band, connected to the apical maculae by the darkened suture. head and pronotum darker than the bright areas of elytra, mostly even darker than the elytral maculae. antennal coloration variable, 7th to 9th and basis of 6th antennomere darkened, 10th and 11th antennomere very bright, often almost white. some specimens with darkened 10th antennomere. head. with large, protuberant eyes, temples narrowed immediately behind eyes; head width 0.760.85 mm, length 0.44-0.48 mm, 1.61-1.79 times as wide as long. puncturation on vertex dense, coarse; pubescence composed of long, semierect setae, directed anteriorly; microsculpture absent. longitudinal impressions on vertex distinct, slightly curved, attaining middle of eyes. eyes protuberant, 0.23-0.25 mm long, distance of inner margins 0.48-0.53 mm. antennae as in fig. 5b, comparatively robust, 1.381.53 mm long; antennomere proportions of holotypus as follows: 2.5 : 1.5 : 1.5 : 1.6 : 1.6 : 1.5 : 1.4 : 1.2 : 1.0 : 1.0 : 2.2. fig. 5. psammoecus leleupi sp. nov., holotype. a. habitus. b. left antenna. c. parameres. scale line a, b: 1 mm; c: 0.2 mm. european journal of taxonomy 17: 1-31 (2012) 14 pronotum. broad, near apical margin and apical angles with shallow impressions; width 0.830.91 mm, length 0.59-0.66 mm, 1.33-1.51 times as wide as long. anterior angles with distinct group of comparatively big teeth, lateral margins with four distinct triangular teeth; tooth i short, tooth ii somewhat longer, tooth iii the longest, tooth iv a bit shorter than tooth iii, but longer than tooth ii; posterior angle with distinct angular tooth. puncturation and pubescence on pronotal disc as on vertex. microsculpture absent. elytra. oval, length 1.88-2.13 mm, combined width 1.20-1.43 mm, 1.39-1.58 times as long as combined width. rows of punctures on disc wider than interstices; pubescense composed of long, semierect setae; microsculpture absent. parameres. with wide, stout base and club-shaped apical extension. inner margin of base with 3-4 longer and numerous short setae; inner margin of extension with numerous small setae, apical portion of extension with 2-3 long setae (fig. 5c). psammoecus luchti sp. nov. (fig. 6) etymology the name is dedicated to wilhelm lucht (1922-2000), who provided the author with much helpful advice in the course of his first steps into entomology. material examined holotype ♂: ‘i.r.s.a.c. – mus. congo | n. rhodesia: mpika | (muchinga mts.) 1700 m. | vii.1960 n. leleup’, ‘dans | l’humus’ [blue label], ‘holotypus | psammoecus | luchti | des. karner’ [red label] (mrac). paratypes 1♂ ‘coll. mus. tervuren | kivu: butembo | alt. 1740 m. iii.1965 | m.j. celis et collab.’, ‘sous | dendrocalamus’, ‘paratypus | psammoecus | luchti | des. karner’ [red label] (mkf). 1♂ ‘coll. mus. congo | katanga: gal. forest. | de la kisanga ix| n. leleup 1948 | (env. d’ elisabethville)’, ‘récolté dans | humus’ [blue label], ‘paratypus | psammoecus | luchti | des. karner’ [red label] (mkf). 1 spm ‘i.r.s.a.c. – mus. congo | n. rhodesia: abercorn, | 1800m. | vii.1960 n. leleup’, ‘galerie forestière | de la mwengo, | dans l’humus’, ‘paratypus | psammoecus | luchti | des. karner’ [red label] (mrac). 1 spm ‘i.r.s.a.c. – mus. congo | n. rhodesia: danger hill | à 45 km. n. de mpika | vii.1960 n. leleup’, ‘humus | de galerie | forestière’ [blue label], ‘paratypus | psammoecus | luchti | des. karner’ [red label] (mrac). 3 spms ‘i.r.s.a.c. – mus. congo | n. rhodesia: mpika | (muchinga mts.) 1700 m. | vii.1960 n. leleup’, ‘dans | l’humus’ [blue label], ‘paratypus | psammoecus | luchti | des. karner’ [red label] (mrac). diagnosis lateral margins of pronotum rounded, bearing five distinct teeth with wide bases. elytra elongate-oval, bright reddish-brown with dark, transverse band at beginning of posterior portion, sometimes with darkened suture near elytral apex; striae on elytral disc slightly wider than interstices. parameres small, elongate, spatula-shaped. differential diagnosis ps. luchti sp. nov.differs from ps. simoni grouvelle, 1892 by the elytral striae being only slightly wider than the interstices, elytra being more elongate, elytral basis not darkened. it differs from ps. hacquardi and ps. marginicollis by wider bases of lateral teeth of pronotum; from ps. hacquardi also by wider karner m., revision of african psammoecus 15 elytral striae and elytral basis not darkened; from ps. marginicollis, ps. parallelus, ps. leleupi sp. nov., ps. simoni, and ps. laetulus by spatula-shaped parameres. it further differs from ps. leleupi sp. nov. and ps. laetulus by elytral basis not darkened; from ps. personatus by shorter lateral teeth of pronotum with wider bases; and from ps. laetulus by more rounded lateral margin of pronotum. description body. elongate oval, total length 2.80-3.19 mm (fig. 6a). bright reddish-brown, elytra with dark, transverse band at beginning of posterior portion, some specimens also with darkened suture near elytral fig. 6. psammoecus luchti sp. nov., holotype. a. habitus. b. left antenna. c. parameres. scale line a, b: 1 mm; c: 0.2 mm. european journal of taxonomy 17: 1-31 (2012) 16 apex. antennae with 7th-10th antennomere blackish brown, some specimens with darkened apex of 6th antennomere. 11th antennomere, sometimes also 10th antennomere, bright yellowish brown, brighter than basal antennomeres. head. very wide, with large, protuberant eyes, temples narrowed immediately behind eyes. head width 0.79-0.88 mm, length 0.44-0.48 mm, 1.75-1.96 times as wide as long. eyes 0.20-0.23 mm long, distance of inner margins 0.50-0.60 mm. puncturation on vertex coarse, relatively dense, distance between punctures irregular; pubescence composed of long erect setae, directed anteriorly; microsculpture absent. longitudinal impressions on vertex distinct, short, attaining second third of eyes. antennae as in fig. 6b, long and slender, 1.45-1.55 mm long, antennomere proportions of holotype as follows: 2.6 : 1.1 : 1.4 : 1.3 : 1.5 : 1.4 : 1.3 : 1.1 : 1.0 : 1.1 : 2.1. pronotum. broad, shallowly impressed near apical margin and apical angles, width 0.80-0.96 mm, length 0.60-0.73 mm, 1.28-1.40 times as wide as long. anterior angles with group of very small teeth, lateral margins with five distinct, triangular teeth; tooth i small, tooth ii larger, tooth iii largest, tooth iv slightly smaller than tooth iii, tooth v smallest; posterior angle with very small angular tooth. puncturation and pubescence on disc as on vertex; microsculpture absent. elytra. elongate oval, length 1.85-2.05 mm, combined width 1.15-1.40 mm, 1.44-1.65 times as long as their combined width. rows of punctures on disc slightly wider than interstices, pubescence composed of semierect setae that are somewhat shorter as on head and pronotum; microsculpture absent. parameres. small, elongate; base less than 2 times as wide as apical extension, inner margins with few short setae, inner portion of apex with four longer setae, outer portion with one shorter and one longer seta (fig 6c). psammoecus grandis grouvelle, 1908 (fig. 7) material examined holotype ♀: ‘af. or. all. | kwai. weise’ [blue label, grouvelle’s hand], ‘type’ [grouvelle’s hand], ‘type’ [red label, printed], ‘museum paris | 1917 | coll. grouvelle’ [blue label], ‘psammoecus | grandis | g. grouv’ [grouvelle’s hand] (mnhn). other material 1 spm ‘coll. mus. congo | ruanda: gitarama, | 1850 m. terr. nyanza, | p. basilewsky i-1953’ (mrac). 1♂ ‘i.r.s.a.c.-mus. congo | kivu: terr. lubero | 2.200 m. 2-xii-1951 | n. leleup | (for mont. avec bamb.)’, ‘récolté dans l’humus’ [blue label] (mrac). 1 spm ‘coll. mus. congo | urundi: kitega 1600| 1700 m. 3/4-iii-1953 | p. basilewsky’ (mkf). 4 spms ‘coll. mus. congo | urundi: bururi, 1800 | 2000 m. 5/12-iii-1953 | p. basilewsky’ (mrac). differential diagnosis ps. grandis resembles ps. longicornis schaufuss, 1872 and ps. lateralis (grouvelle, 1899), but differs by the presence of small lateral teeth of pronotum. it differs from ps. longicornis also by the very large and curved parameres with very short extensions. redescription body. slender, narrow, total length 3.4-4.2 mm (fig. 7a), surface yellowish-brown, elytra with dark maculae of variable shape; some immature specimens only with a darkish macula in the posterior half karner m., revision of african psammoecus 17 of elytra, other specimens with posterior half of elytral suture and elytral maculae blackish-brown. 7th to 10th antennomere and apex of 6th antennomere darkened, 11th antennomere brighter than basis of antennae. head. with comparatively small eyes and broad, curved temples; width 0.80-0.89 mm, length 0.450.48 mm; 1.71-1.85 times as wide as long. eye length 0.23-0.25 mm, distance of inner margins 0.480.56 mm. puncturation on vertex irregular, pubescence composed of short, recumbent setae, directed anteriorly; microsculpture well defined, reticulate. longitudinal impressions on vertex short, curved, almost attaining the anterior third of eyes. antennae as in fig. 7b, short, length 1.58-2.00 mm, 1st antennomere very long and slender. antennomere proportions of holotype as follows: 3.33 : 1 : 1.17 : 1.5 : 1.2 : 1.7 : 1 : 1 : 1.8 : 1: 2.17. fig. 7. psammoecus grandis grouvelle, 1908. a. habitus of holotype. b. left antenna of holotype. c. parameres of specimen from kivu. scale line a, b: 1 mm; c: 0.2 mm. european journal of taxonomy 17: 1-31 (2012) 18 pronotum. subquadrate, width 0.89-1.03 mm, length 0.75-0.90 mm, 1.14-1.23 times as wide as long. surface uneven with flat impressions near the posterior margin close to the posterior angles as well as in the middle of the posterior margin. less distinct impressions also close to the anterior margin, in some specimens also in the middle of the pronotal disc. posterior angle with a very small group of teeth, lateral margins with four to five very short lateral teeth, posterior angle almost rudimentary. the holotype shows an asymmetrical tooth pattern with different numbers and location of lateral teeth on the two sides of the pronotum. puncturation slightly coarser and denser than on vertex, pubescence as on vertex; microsculpture well defined, reticulate. elytra. narrow, slender, length 2.10-2.67 mm, combined width 1.35-1.60 mm, 1.50-1.81 times as long as their combined width. rows of punctures on the disc a little narrower than interstices. pubescence rather flat, consisting of long setae. microsculpture visible only in few specimens, very weak, reticulate. parameres. with broad base, narrowing and curving inwards toward apex (fig. 7c). psammoecus marginicollis grouvelle, 1908 (fig. 8) material examined holotype ♂: ‘af or. all | kwai weise’ [blue label, grouvelle’s hand], ‘type’ [grouvelle’s hand], ‘type’ [red label], ‘museum paris | 1917 | coll. grouvelle’ [blue label], psammoecus | marginicollis | g. grouv’ [grouvelle’s hand]; (mnhn). other material 1♂ ‘mus. roy. afr. centr. | tanganyika: kilimandjaro | vers. o. (buchberger) | ex. coll. dr breuning’ (mrac). 1 spm ‘musée du congo | haut-uele: yebo moto | vi-1926 | l. burgeon’ (mrac). 1 spm ‘musée du congo | ituri: medje | fin viii-1925 | dr h. schouteden’ (mrac). 3 spms ‘coll.mus.tervuren | congo: dorsale de lubero | mt. muleke, vi/vii.1963 | m.j. celis’ (mrac). 1♂ ‚a la lumiére’ [blue label], ‘i.r.s.a.c.-mus.congo | kivu: terr. masisi, 720 m. | walikale (rives loa) ix-53 | n. leleup’ (mkf). 1♂ ‘coll. mus. congo | kivu: kalonge | 15-xii-1938 | hendrickx’ (mkf). 1 spm ‘forêt | transition’ [blue label], ‘i.r.s.a.c. – mus. congo | kivu: riv. nyakagera, terr. | kabare 1600/1700 m. | n. leleup xi-1955’ (mrac). 1 spm ‘musée du congo | kivu: burunga | 9-xii-1925 | dr h.schouteden’ (mrac). 1 spm ‘musée du congo | vuhovi | vii-1925 | h. j. bredo’ (mrac). 3 spms ‘musée du congo | mombassa (36 km.s. | lubero) viii-1932 | l. burgeon’(mrac). 1 spm ‘musée du congo | aruwimi: panga | 9/10-ix-1925 | dr h. schoudeten’ (mrac). 1 spm ‘coll. mus. tervuren | côte d’ivoire: bingerville | vi. 1962 | j. decelle’ (mrac). 1 spm ‘coll. mus. tervuren | côte d’ivoire: bingerville | vii. 1962 | j. decelle’ (mrac). 2 spms ‘coll. mus. tervuren | côte d’ivoire: bingerville | v.1964 | j. decelle’ (mkf) differential diagnosis ps. marginicollis differs from ps. personatus and ps. leleupi sp. nov. by the shorter lateral teeth of pronotum; from ps. laetulus and ps. parallelus by 1st antennomere being two times as long as 2nd; from ps. hacquardi by wider, less elongate parameres and longer antennae; from ps. leleupi sp. nov. by narrower elytral striae and parameres without club-shaped extension; from ps. parallelus by wide parameres without club-shaped extension; from ps. luchti sp. nov. by narrower bases of lateral teeth of pronotum and wider, less elongate parameres. karner m., revision of african psammoecus 19 redescription body. elongate oval, total length 2.50-3.50 mm (fig. 8a). surface yellowish to reddish brown, some specimens considerably darker with lateral margins of pronotum blackish brown. elytra with variable pattern of dark maculae; darker specimens with wide, horizontal maculae in the middle of the elytra and dark apical portion of elytral suture; the basal portion of the eltral suture not darkened. specimens of lighter colour with reduced maculae on the elytral discs and only slightly darkened or light apical portion of elytral suture. few specimens without elytral maculae. antennae yellowish or reddish brown, 7th to 10th antennomere dark brown or black, 11th antennomere yellowish brown, lighter than basal antennomeres. fig. 8. pammoecus marginicollis grouvelle, 1908, holotype. a. habitus. b. right antenna. c. parameres. scale line a, b: 1 mm; c: 0.2 mm. european journal of taxonomy 17: 1-31 (2012) 20 head. broad, temples rounded; width 0.65-0.76 mm, length 0.40-0.46 mm, 1.58-1.83 times as wide as long. eyes moderately prominent, 0.20-0.23 mm long, distance of inner margins 0.43-0.53 mm. puncturation on vertex variable, moderately dense, sometimes very dense. pubescence composed of long, semierect setae, directed anteriorly; microsculpture hardly visible, very shallow, reticulate. longitudinal impressions on vertex deep, attaining the anterior third of eyes. antennae as in fig. 8b, 1.58-1.70 mm long. antennomere proportions of holotype as follows: 2.0 : 1.1 : 1.3 : 1.4 : 1.5 : 1.2 : 1,1 : 1.0 :1.0 : 1.1 : 2.1. pronotum. subquadrate, width 0.70-0.85 mm, length 0.56-0.70 mm, 1.16-1.42 times as wide as long. surface even. anterior angle with distinct group of small teeth. lateral margin with five distinct, short teeth; tooth i short with wide base, tooth ii a little larger, teeth iii and iv still a little larger, triangular with wide bases, tooth v about as big as tooth ii. posterior angle with very small tooth. puncturation on pronotal disc coarser and denser as on vertex, pubescence as on vertex. microsculpture variable, in most specimens well defined, reticulate, in some specimens hardly visible or absent. elytra. elongate oval, length 1.63-2.30 mm, width 1.13-1.50 mm, 1.32-1.71 times as long as their combined width. rows of punctures on the disc a little narrower than interstices. pubescence composed of long, semierect setae; microsculpture absent. parameres. short, broadest in the middle, with one large apical seta (fig. 8c). remarks the shape of the parameres was found to vary to a small degree. so far, the material at hand was not sufficient to decide wether this variability has specific importance, hence all specimens mentioned here were determined to be conspecific with ps. marginicollis. psammoecus lateralis (grouvelle, 1899) (fig. 9) cryptamorpha lateralis grouvelle, 1899: 179. psammoecus lateralis grouvelle, 1919: 46. material examined holotype ♀: ‘andrangoloaka | 1600 m. o. s. o. de | tananarive.’, ‘type’ [red label], ‘museum paris | 1917 | coll. grouvelle’, ‘cryptamorpha | lateralis | g. grouv’ [grouvelle’s hand] (mnhn). differential diagnosis ps. lateralis resembles ps. longicornis, but differs by the smaller and less dense puncturation on head, pronotum, and elytral discs, and by the longer antennae. it differs from ps. grandis by absence of lateral teeth of pronotum. redescription body. elongate-oval, total length 3.60 mm (fig. 9a). surface yellowish brown with elytral margin and a round macula on posterior third of elytra dark brown. head. with long, slightly rounded temples; width 0.79 mm, length 0.48 mm, 1.65 times as wide as long. eyes moderately protuberant, 0.4 mm long, distance of inner margins 0.53 mm. puncturation on vertex of moderate density, punctures small, distance between punctures variable. pubescence composed of short karner m., revision of african psammoecus 21 semierect setae, directed anteriorly; microsculpture very distinct, reticulate. longitudinal impressions on vertex short, distinct, attaining the anterior quarter of the eyes. antennae as in fig. 9b, antennal length 1.78 mm; antennomere proportions of holotype as follows: 2.6 : 1.3 : 1.2 : 1.1 : 1.4 : 1.6 : 1.4 : 1.1 :1.1 : 1.0 : 1.9. pronotum. subquadrate, width 0.86 mm, length 0.75 mm, 1.15 times as wide as long. anterior and posterior angles with distinct groups of small teeth, lateral teeth absent. puncturation on pronotal disc sparser as on vertex, punctures of same size as on vertex. pubescence shorter and less erect as on vertex; microsculpture distinct, reticulate. elytra. elongate, length 2.35 mm, width 1.28 mm, 1.84 times as long as combined width. rows of punctures on disc distinctly narrower than interstices. pubescence composed of only slightly erect setae; microsculpture hardly visible, very shallow, reticulate. fig. 9. pammoecus lateralis (grouvelle, 1899), holotype. a. habitus. b. left antenna. scale line: 1 mm. european journal of taxonomy 17: 1-31 (2012) 22 psammoecus longicornis schaufuss, 1872 (fig. 10) psammoecus longulus grouvelle, 1878: 265. syn. nov. material examined holotype ‘type’, ‘coll. l. w. | schaufuß’, ‘type’ [red label], ‘longicornis | schauf | n.germany | caffr’, ‘longicornis | schauf | caffraria.’ (zmhb). other material 2♀♀, mounted on single card label ‘port natal | h. deysolle’ [grouvelle’s hand; second line hardly readable], ‘type’, ‘type’ [red label], ‘museum paris | 1917 | coll. grouvelle’ [blue label], ‘identique au | longicornis | sch. | comp. au ty.’ [grouvelle’s hand], ‘psammoecus | longulus | grouv’ [grouvelle’s hand] (mnhn). the specimen that is mounted on the right side is hereby designated to be the lectotype of ps longulus. 1 spm ‘coll. mus. congo | urundi: kitega 1600| 1700 m. 3/4-iii-1953 | p. basilewsky’ (mrac). 1 spm ‘recolté dans | l’humus’ [blue label], ‘i.r.s.a.c.-mus. congo | kivu : terr. kalehe | contr. s. du kahuzi, | 2200 m. 27-vii-51 | n. leleup’ (mrac). 1 spm ‘coll. mus. congo | kivu: ibanda | 1952 | m. vandelannoite’ (mrac). 1 spm ‘i.r.s.a.c.-mus. congo | kivu: lemera, terr. uvira | 1600 m. xii-1956 | n. leleup’ (mrac). 1 spm ‘recolté sous | détritus | végétaux’ [blue label]’, ‘i.r.s.a.c.-mus. congo | kivu : terr. uvira, | mulenge 1950 m. | (mavaisroselier) 4-ix-51 | n. leleup’, (mrac). 1♂ ‘i.r.s.a.c.-mus. congo | kivu : terr. uvira, mu| lenge 1880-2010 m. | (vert. for. ombroph.) v-1951 | n. leleup’, ‘recolté dans | l’humus’ [blue label] (mrac). 1 spm ‘recolté dans | racines | (plantes basses)’ [blue label], ‘i.r.s.a.c.-mus. congo | kivu : mwenga, | s.-o. itombwe, luiko | 1900 m (sav.herb.) i-1952 | n. leleup’ (mrac). 2 spms ‘coll. mus. congo | elisabethville | ii-1940 | h. j. bredo’ (mkf). differential diagnosis ps. longicornis resembles ps. lateralis, but differs by coarser puncturation of head, pronotum, and elytral discs and by the shorter antennae. it differs from ps. grandis by absence of lateral teeth of pronotum and by parameres with wide bases and distinct, slender extension. redescription body. elongate oval, total length 2.60-3.40 mm (fig. 10a). surface bright yellowish or reddish brown, elytra with dark brown maculae at the beginning of posterior half, near margin; the elytral suture and margins in apical third often darkened; some specimens lacking elytral maculae. antennae yellowish brown; 8th to 10th antennomeres slightly to moderately darkened, in some specimens also the 7th antennomere darkened; 11th antennomere of same color as basal antennomeres or only slightly brighter. head. broad, temples curved, head width 0.65-0.75 mm, length 0.38-0.45 mm, 1.67-1.83 times as wide as long. eyes only moderately protuberant, 0.18-0.20 mm long, distance of inner margins 0.44-0.53 mm. puncturation on vertex dense and coarse. pubescence composed of short, recumbent setae, directed anteriorly; microsculpture well defined, reticulate. longitudinal impressions on vertex distinct, attaining second fourth of eyes. antennae as in fig. 10b, 1.30-1.50 mm long. antennae of holotype missing; antennomere proportions of lectotype of ps. longulus as follows: 2.5 : 1.2 : 1.5 : 1.5 : 1.2 : 1.2 : 1.1 : 1.1 :1.0 : 1.2 : 2.3. pronotum. almost as long as wide; width 0.64-0.78 mm, length 0.58-0.73 mm, 1.01-1.15 times as wide as long. lateral teeth absent, anterior and posterior angle with small groups of teeth. puncturation and pubescence as on vertex; microsculpture well defined, reticulate. karner m., revision of african psammoecus 23 elytra. oval, length 1.68-2.10 mm, width 0.95-1.50 mm, 1.40-1.80 times as long as wide. rows of puctures somewhat wider than interstices. the pubescence of the studied specimens almost completely damaged, as far as visible, the setae are short and recumbent; microsculpture absent. parameres. with wide base and slender apical extension. inner margin with numerous short setae, apex with single long seta (fig. 10c). fig. 10. pammoecus longicornis schaufuss, 1872. a. habitus of specimen from port natal (= lectotype of ps longulus grouvelle, 1878). b. right antenna of specimen from port natal. c. parameres of specimen from kivu. scale line a, b: 1 mm; c: 0.2 mm. european journal of taxonomy 17: 1-31 (2012) 24 psammoecus trimaculatus motschulsky, 1858 psammoecus excellens grouvelle, 1908a: 115. syn. nov. psammoecus alluaudi grouvelle, 1912: 409. syn. nov. material examined 1♀ [round, red label]; [yellow label]; ‘psammecus [sic!] | trimaculatus | motch. | ceylon’ [motschulsky’s hand] (zmum). 3 spms ‘af. or. all. | eichelbaum’ [grouvelle’s hand], ‘type’ [red label], ‘museum paris | 1917 | coll. grouvelle’ [blue label], ‘psammoecus | excellens | g grouv’ [grouvelle’s hand] (mnhn). one ♂ specimen was designated as lectotpyus of ps excellens grouvelle and labelled accordingly. holotype of ps alluaudi. ♂: ‘madagascar | su[???]ieville’ [blue label, grouvelle’s hand], ‘type’ [red label], ‘museum paris | 1917 | coll. grouvelle’ [blue label], ‘psammoecus | alluaudi | g. grouv’ [grouvelle’s hand] (mnhn). remarks pal (1985) gave a redescription of ps. trimaculatus based on material from motschulsky’s collection (museum moscow) that he accepted as type material. he did not designate a lectotype. the material consisted of five specimens mounted on a single label (pal 1996, personal communication). it was not possible for the present author to see this material, but he had the opportunity to study a single, female specimen from motschulsky’s collection (zmum). this specimen is not assumed to be a type, while the material mentioned by pal may be discovered again. pal (1985) provided detailed figures of ps. trimaculatus, including the male genitalia, and gives an overview on the taxonomy. he states that it occurs in india, nepal, bhutan, sri lanka, myanmar, malaysia, australia, japan, and madagascar. recently, it has also been recorded as being established in brasil (thomas & yamamoto 2007). both ps. excellens grouvelle and ps. alluaudi grouvelle are conspecific and have to be synonymised with ps. trimaculatus motschulsky. psammoecus simoni grouvelle, 1892 (fig. 11) psammoecus simonis grouvelle, 1892: 287. psammoecus simoni – grouvelle 1908c: 476.— pal 1985. material examined lectotype ♂, by present designation: ‘antipolo | e. simon’, ‘type’ [red label], ‘museum paris | 1917 | coll. grouvelle’ [yellow label], ‘psammoecus | simoni | ty. a. grouv’ [grouvelle’s hand] (mnhn). paralectotype 1 spm with identical data as lectotype (mnhn). other material 5 spms ‘coll: mus. congo | madagascar: maroansetra | (à la lumière) ii/iv-1950 | j. vadon’ (mrac). 1spm ‘coll. mus. tervuren | n.e. madagascar: | ambodivoangy 1959 | j. vadon’ (mrac). karner m., revision of african psammoecus 25 differential diagnosis ps. simoni differs by its short oval habitus and the short, stout parameres that are fused with the basal piece from all other african psammoecus. the wide-based pronotal teeth resemble ps. luchti sp. nov., it differs by the elytral striae being considerably wider than interstices, elytra being shorter, darkened basis of elytra, parameres short, stout and fused with basal piece. redescription body. oval, total length 2.13-3.00 mm (fig. 11a). surface yellowish-brown, sometimes reddish-brown, elytra with brown or blackish-brown maculae: humeral swelling, a transverse band in the middle of the elytra, the elytral suture along the posterior two thirds and the elytral apex are dark. base of antennae yellowish or reddish brown, 6th to 10th antennomere darkened, 11th antennomere yellowish-white, some specimens with light apex of 10th antennomere. head. broad, temples narrowed immediately behind eyes; width 0.64-0.71 mm, length 0.33-0.44 mm, 1.67-1.73 times as wide as long. eyes protuberant, rounded, 0.17-0.20 mm long, distance of inner fig. 11. psammoecus simoni grouvelle, 1892, lectotype. a. habitus. b. right antenna. c. parameres. scale line a, b: 1 mm; c: 0.2 mm. european journal of taxonomy 17: 1-31 (2012) 26 margins 0.38-0.45 mm. puncturation on vertex coarse, density of punctures variable, pubescence composed of long, semierect setae, directed anteriorly; microsculpture absent. longitudinal impressions on vertex very shallow, attaining the middle of the eyes, sometimes shorter. antennae as in fig. 11b, 1.17-1.40 mm long, stout, antennomere proportions of lectotype as follows: 2.9 : 1.3 : 1.8 : 1.5 : 1.8 : 1.6 : 1.4 : 1.0 :1.2 : 1.4 : 2.8. pronotum. broad; width 0.62-0.74 mm, length 0.48-0.56 mm, 1.22-1.35 times as wide as long. surface smooth, without impressions. anterior angles with distinct groups of small teeth; lateral margins with four distinct teeth; tooth i very small, tooth ii a little larger, teeth iii and iv largest. posterior group of teeth consisting of a larger anterior tooth and a very small, almost obtuse posterior tooth. puncturation coarser than on vertex, punctures sometimes adjoining. pubescence as on vertex; microsculpture absent. elytra. oval, short, length 1.35-1.70 mm, combined width 1.00-1.23 mm, 1.27-1.43 times as long as their combined width. rows of punctures on disc wider than interstices. pubescence consists of long, semierect setae. microsculpture absent. parameres. short, stout, fused with basal piece; with distinct pattern of three large setae (fig. 11 c). remarks in his original description, grouvelle (1892) spells the name ‘simonis’. however, on the labels that grouvelle added to the syntypes as well as in a later paper (grouvelle, 1908c), he spells the name ‘simoni’. pal (1985) also uses the latter spelling. hence the present author considers ‘simonis’ to be a misprint and proposes to spell the name in accordance with grouvelle (1908c) and pal (1985). psammoecus spinosus grouvelle, 1882 (fig. 12) material examined 1 spm ‘i.r.s.a.c.-mus.congo | kivu: kavimvira (uvira) | (à la lumière) i-1956 | g. marlier’ (mrac). 1 spm ‘i.r.s.a.c.-mus.congo | kivu: kavimvira (uvira) | (à la lumière) ii/iii-1956 | g. marlier’ (mrac). 1 spm ‘coll. mus. congo | tanganika: moba, 780 m. | (à la lumière) xi-1953 | h. bomans’ (mrac). 1 spm ‘angola: cazombo, | alto zambeze ii-1955 | 4987.1 | e. luna de carvalho’ (mrac). 1♂ ‘nigeria vdofr | m state | iv 1975 | j. t. nedler coll.’ (mrac). 1 spm ‘coll. mus. tervuren | côte d’ivoire: bingerville | ii.1962 | j. decelle’, ‘psammoecus sp. | d. g. h. halstead | det. 1978’ (mkf). 1♂ ‘coll. mus. tervuren | côte d’ivoire: bingerville | i/12.iii.1962 | j. decelle’ (mrac). differential diagnosis ps. spinosus differs by the very characteristic shape of the pronotum and the distinctly toothed elytral margins from all other african psammoecus. redescription body. oval, total length 2.40-3.15 mm (fig. 12a), reddish-brown, elytra with darkened basis and with dark lateral macula near their middle, connected to the darkened suture in posterior half which is connected to another lateral macula close to the apex; the maculae forming a roughly x-shaped pattern on posterior two thirds of elytra. antennae reddish-brown, 7th to 10th antennomeres darker brown, 11th antennomere bright yellowish, almost white. head. wide, temples strongly narrowed immediately behind eyes; width 0.60-0.75 mm, length 0.380.45 mm, 1.58-1.71 times as wide as long. eyes very large and protuberant, length 0.18-0.23 mm, karner m., revision of african psammoecus 27 distance of inner margins 0.36-0.48 mm. puncturation on vertex irregular, punctures comparatively small and sparse, pubescence composed of moderately long, semierect setae, microsculpture absent. longitudinal impressions on vertex very distinct, deep, slightly curved, attaining slightly beyond the middle of eyes. antennae as in fig. 12b, long and slender, 1.23-1.30 mm long; antennomere proportions as follows: 2.8 : 1.0 : 1.3 : 1.2 : 1.3 : 1.4 : 1.3 : 1.0 : 1.0 : 1.1 : 2.2. pronotum. transverse, posterior and lateral margins with distinct impressions, close to the middle of the anterior margin with shallow impression; width 0.65-0.83 mm, length 0.45-0.58 mm, 1.38-1.44 times as wide as long. anterior angle with very distinct group or large teeth; lateral margin characteristic pattern of four teeth; tooth i large with wide basis, teeth ii and iii almost fused, forming a large forked tooth, fig. 12. psammoecus spinosus grouvelle,1882, specimen from bingerville. a. habitus. b. right antenna. c. parameres. scale line a, b: 1 mm; c: 0.2 mm. european journal of taxonomy 17: 1-31 (2012) 28 tooth iv as large as tooth i; posterior angle with small group of teeth. anterior margin with a small short tooth right between anterior group of teeth and middle of anterior margin. puncturation on disc somewhat coarser as on vertex, pubescence as on vertex, microsculpture absent. elytra. oval, length 1.55-2.08 mm, combined width 0.98-1.40 mm, 1.46-1.59 times as long as combined width. rows of punctures on the disc about as wide as interstices; pubescence composed of long semierect setae, directed posteriorly; interstices of lateral rows of punctures with tubercles that are enlarged to form distinct teeth; towards the elytral disc, these teeth become successively smaller. microsculpture absent. parameres. simple, narrowed towards apex, without well-defined basal part. inner portion of parameres with numerous small setae; apex with one large seta (fig. 12c). remarks it has not been possible to locate and study type material of ps. spinosus. however, grouvelle (1882) provides an illustration that leaves little doubt regarding the identity of this species, given its peculiar habitus. key to species 1. lateral margins of pronotum without teeth; only anterior and posterior groups of small teeth present ............................................................................................................................................................. 2 – lateral margins of pronotum with various number of large or small teeth ......................................... 3 2. 3rd antennomere shorter than 2nd; elytral rows of punctures narrower than interstices .......................... .................................................................................................................... lateralis (grouvelle, 1899) – 3rd antennomere longer than 2nd; elytral rows of punctures somewhat wider than interstices ............... ..................................................................................................................longicornis schaufuss, 1872 3. body length 3.40-4.20 mm; lateral margins of pronotum with very small teeth; 1st antennomere more than three times as long as 2nd ......................................................................... grandis grouvelle, 1908 – body length not more than 3.30 mm; lateral margins of pronotum with distinct teeth, 1st antennomere less than three times as long as 2nd ....................................................................................................... 4 4. interstices of lateral striae of elytra with small, distinct spines; pronotum with large lateral teeth, bases of tooth ii and iii connate ............................................................................. spinosus grouvelle, 1882 – interstices of lateral striae of elytra without spines, often with tubercles; lateral teeth of pronotum always well separated .......................................................................................................................... 5 5. 1st antennomere less than two times as long as 2nd ............................................................................... 6 – 1st antennomere more than two times as long as 2nd ............................................................................ 8 6. 3nd antennomere longer than 2nd ............................................................marginicollis grouvelle, 1908 – 3rd antennomere as long as 2nd ............................................................................................................. 7 7. striae on elytral disc narrower than interstices; lateral teeth of pronotum with narrow bases .............. .................................................................................................................. personatus grouvelle, 1919 – striae on elytral disc wider than interstices, lateral teeth of pronotum with wide bases ....................... ....................................................................................................................................... leleupi sp. nov. karner m., revision of african psammoecus 29 8. maximum pronotal width near middle (excluding lateral teeth) ......................................................... 9 – maximum pronotal width in anterior third (excluding lateral teeth) ..................................................11 9. lateral teeth of pronotum slender ...............................................................hacquardi grouvelle, 1889 – lateral teeth of pronotum triangular, with wide bases ...................................................................... 10 10. elytral striae distinctly wider than interstices; humeral swelling, a transverse band in the middle of elytra, elytral suture along posterior two thirds and elytral apex dark; parameres short, stout, fused with basal piece ................................................................................................ simoni grouvelle, 1892 – elytral striae slightly wider than interstices; elytra with dark, transversal band at beginning of posterior portion, sometimes with darkened suture near elytral apex; parameres spatula-shaped, distinctly separated from basal piece ...............................................................................luchti sp. nov. 11. elytrae with darkened humeral area, wide dark band across middle and darkened suture near apex; paramera slender ............................................................................................ laetulus grouvelle, 1914 – elytrae with round macula and darkened suture near apex .............................................................. 12 12. lateral margin of pronotum with 4 distinct teeth between small posterior tooth and anterior group of small teeth .......................................................................................... trimaculatus motschulsky, 1858 – lateral margin of pronotum with 5 distinct teeth between small posterior tooth and anterior group of small teeth ................................................................................................... parallelus grouvelle, 1919 discussion the genus psammoecus latreille, 1829 (coleoptera, silvanidae, brontinae, telephanini) consists of about 80 old world species, with one species recently found to be established also in brazil (thomas & yamamoto 2007). the vast majority of species inhabits tropical regions, where they are found in plant detritus and sometimes at light sources. after the studies of antoine henri grouvelle (1843-1917), little research on the taxonomy of this genus was done, until pal (1985) provided a revision of the indian psammoecus. hetschko (1930) lists 14 species of psammoecus for africa, including ps. bipunctatus (f.) var. algiricus pic, 1920 and ps. oblitus grouvelle, 1908. psammoecus bipunctatus is a common species in europe. the author was not able to retrieve the material studied by pic and has not seen other specimens of ps. bipunctatus from africa. hetschko lists psammoecus oblitus grouvelle, with reference to grouvelle (1908a: 116), where silvanus oblitus is described. the description itself is sufficient to conclude that grouvelle did indeed not describe a species belonging to the genus psammoecus. halstead (1973) includes the species in the genus parasilvanus grouvelle, 1912. since no original description is available, psammoecus oblitus is to be considered a nomen nudum. acknowledgements the author wishes to thank the following persons and institutions for their support and/or generous loan of material: dr nicole berti (museum national d’histoire naturelle), dr roger g. booth (the natural history museum, london), dr daniel burckhardt (naturhistorisches museum, basel), dr johannes frisch (museum für naturkunde der humboldt-universität, berlin), dr damir kovac (senckenbergisches naturforschendes institut, frankfurt am main), dr marc de meyer (musée royal de l’afrique centrale, tervuren), dr nikolay borisovich nikitsky (zoological museum of m.v. lomonosov state university), dr tarun k. pal (zoological survey of india, calcutta). european journal of taxonomy 17: 1-31 (2012) 30 references franzen b. & karner m. 1998. zur genitalpräparation bei kleinund kleinstkäfern. in: lucht w. & klausnitzer b. (eds.) die käfer mitteleuropas. band 15. 4. supplementband: 20-22. goecke & evers, krefeld, im gustav fischer verlag, jena. grouvelle a. 1878. cucujides nouveaux ou peu connus, 4e mémoire. annales de la société entomologique de france 8: 261-268, pl. 8. grouvelle a. 1882. cucujides nouveaux du musée civique de gènes. annali del museo civico di storia naturale giacomo doria 18: 275-296, pl. 7. grouvelle a. 1892. voyage de m.e. simon a l’île de luzon, philippines (mars et avril 1890), 5e mémoire. cucujides. annales de la société entomologique de france 61: 285-288. grouvelle a. 1899. descriptions de clavicornes d’afrique et de la région malgache, 2e mémoire. annales de la société entomologique de france 68: 136-185. grouvelle a. 1908a. clavicornes récoltés dans l’afrique orientale allemande par le dr. f. eichelbaum (2e liste). revue d’entomologie 27: 97-126. grouvelle a. 1908b. coléoptères clavicornes de l’afrique australe et orientale. annales de la société entomologique de france 77: 127-206. grouvelle a. 1908c. coléoptères de la région indienne. annales de la société entomologique de france 77: 315-495, pl. 8-9. grouvelle a. 1912. coléoptères cucujides nouveaux du genre psammoecus faisant partie des collections du muséum. bulletin du museum d’histoire naturelle de paris 18: 409-414. grouvelle a. & de peyerimhoff p. 1914. no. ii.—coleoptera: cucujidæ, cryptophagidæ: avec une description de la larve et de la nymphe de prostominia convexiuscula grouvelle [cucujidæ]. transactions of the linnean society of london. 2nd series: zoology 17: 141-159. http://dx.doi. org/10.1111/j.1096-3642.1914.tb00591.x grouvelle a. 1919. descriptions d’espèces nouvelles du genre psammoecus latr. in: mémoires entomologiques. études sur les coléoptères 2: 5-38. société entomologique de france, paris. halstead d.g.h. 1973. a revision of the genus silvanus latreille (s.l.) (coleoptera: silvanidae). bulletin of the british museum (natural history). entomology. 29 (2): 39-112. hetschko a. 1930. fam. cucujidae. coleopterorum catalogus 109: 3-122. w. junk, berlin. pal t.k. 1985. a revision of indian psammoecus latreille (coleoptera, silvanidae). records of the zoological survey of india, miscellaneous publication, occasional paper 71: 1-54. thomas m.c. & yamamoto p.t. 2007. new records of old world silvanidae in the new world (coleoptera: cucujoidea). coleopterists bulletin 61 (4): 612-613. http://dx.doi.org/10.1649/0010-065x( 2007)61[612:nroows]2.0.co;2 manuscript received on: 27 november 2011 manuscript accepted on: 4 june 2012 published on: 5 july 2012 topic editor: koen martens http://dx.doi.org/10.1111/j.1096-3642.1914.tb00591.x http://dx.doi.org/10.1111/j.1096-3642.1914.tb00591.x http://dx.doi.org/10.1649/0010-065x(2007)61[612:nroows]2.0.co;2 http://dx.doi.org/10.1649/0010-065x(2007)61[612:nroows]2.0.co;2 karner m., revision of african psammoecus 31 in compliance with the iczn, printed versions of all papers are deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; national botanic garden of belgium, meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark. 77 european journal of taxonomy 888: 77–96 issn 2118-9773 https://doi.org/10.5852/ejt.2023.888.2215 www.europeanjournaloftaxonomy.eu 2023 · asif m. et al. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e molecular and morphological studies reveal a new species of panaeolus (agaricales, basidiomycota) from punjab, pakistan muhammad asif 1,*, qudsia firdous 2, aiman izhar 3, abdul rehman niazi 4, samina sarwar 5 & abdul nasir khalid 6 1,2,3,4,6 fungal biology and systematics research laboratory, institute of botany, university of the punjab, quaid-e-azam campus 54590, lahore, pakistan. 1 plant mycology research laboratory, department of plant sciences, quaid-i-azam university, islamabad, pakistan. 5 department of botany, lahore college for women university, lahore, 54590, pakistan. * corresponding author: asifgondal101@gmail.com 2 email: qudsiafirdous26@gmail.com 3 email: aimanizhar25@gmail.com 4 email: mushroomniazi@gmail.com 5 email: samina_boletus@yahoo.com 6 email: drankhalid@gmail.com abstract. panaeolus punjabensis m. asif, q. firdous, a. izhar, niazi & khalid sp. nov. was collected from three different localities (bahawalnagar, kasur, and lahore) in punjab, pakistan. morphological observations and phylogenetic analyses based on nuclear encoded internal transcribed spacers (its15.8s-its2 = its) and d1/d2 domain of large subunit (28s) rdna confirmed the taxonomic distinctness of this species. the new species is potentially hallucinogenic and characterized by a parabolic pileus with a light brown center, broadly fusiform basidiospores, presence of cheilocystidia, pileocystidia, and caulocystidia, and absence of pleurocystidia and clamp connections. the dna sequences of the species clustered together in a well-supported distinct clade. we present a detailed description, photographs, and line drawings, and elucidate and discuss the phylogenetic position of the new species. morphological comparisons with phylogenetically and morphologically allied species are discussed. keywords. agarics, hallucinogenic, molecular systematics, saprotrophic, taxonomy. asif m., firdous q., izhar a., niazi a.r., sarwar s. & khalid a.n. 2023. molecular and morphological studies reveal a new species of panaeolus (agaricales, basidiomycota) from punjab, pakistan. european journal of taxonomy 888: 77–96. https://doi.org/10.5852/ejt.2023.888.2215 introduction the genus panaeolus (fr.) quél (1872: 151) has been placed in different families, including coprinaceae overeem & weese (doveri 2011), bolbitiaceae singer (tóth et al. 2013), and psathyrellaceae vilgalys, moncalvo & redhead (amandeep et al. 2014), or has been treated as incertae sedis (he et al. 2019). more recently, panaeolus along with copelandia bres., panaeolina pers., and panaeolopsis singer have been https://doi.org/10.5852/ejt.2023.888.2215 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ https://orcid.org/0000-0001-7327-2072 mailto:asifgondal101%40gmail.com?subject= mailto:qudsiafirdous26%40gmail.com?subject= mailto:aimanizhar25%40gmail.com?subject= mailto:mushroomniazi%40gmail.com?subject= mailto:samina_boletus%40yahoo.com?subject= mailto:drankhalid%40gmail.com?subject= https://doi.org/10.5852/ejt.2023.888.2215 https://orcid.org/0000-0002-3096-0198 https://orcid.org/0000-0002-1118-1148 https://orcid.org/0000-0002-6377-4498 https://orcid.org/0000-0002-5635-8031 https://orcid.org/0000-0002-8739-2881 european journal of taxonomy 888: 77–96 (2023) 78 placed in a distinct family galeropsidaceae singer (kalichman et al. 2020). these genera share several taxonomically important features such as the pileus covering, spore color, and coprophilous ecology (singer 1986; tóth et al. 2013; kalichman et al. 2020). the members of the family galeropsidaceae are mostly found growing on open lawns, in steppes and prairies, in mountain deserts and are characterized by a cutis that is rarely and slightly gelatinized, partly enclosed or ovoid pileus, lacunar hymenophore or in some cases regularly developed lamellae often connected by anastomoses, passive spore discharge, ochre-brown spores with germ pore, pileipellis with pileocystidia, the presence in some species of hymenial cystidia, and clamped hyphae (zeller 1943; singer & de leon 1982; malysheva et al. 2019). species in the genus are characterized by their typically coprophilous or nitrophilous habitat, slender fruiting body with the hemispherical pileus, cartilaginous and relatively long stipe, bluing context, epithelial pileipellis, and a black spore print that does not fade in concentrated sulphuric acid (watling & gregory 1987; gerhardt 1996; stamets 1996; strauss et al. 2022). panaeolus includes some of the most hallucinogenic species after psilocybe (fr.) p.kumm., for example, p. subbalteatus (berk. & broome) sacc. and p. cambodginiensis ola’h & r.heim whose basidiomata contain two hallucinatory compounds, i.e., psilocybin and psilocin (stamets 1996; andersson et al. 2009). there are sixteen species in the genus panaeolus reported in literature (he et al. 2019; hu et al. 2020) however, in index fungorum, 189 records are associated with this genus (accessed on 18 september 2022). species of panaeolus are mostly reported from asia and europe in tropical to sub-tropical and temperate habitats (senn-irlet et al. 1999; halama et al. 2014; kaur 2014; wang & tzean 2015; desjardin & perry 2017; karunarathna et al. 2017; akata et al. 2019). some species of the genus have also been reported from north and south america and africa in temperate habitats (adeniyi et al. 2018; silva-filho et al. 2019; teke et al. 2019). to date, 1293 macrofungal species belonging to 411 genera, 115 families, and 24 orders have been reported from pakistan (aman et al. 2022). out of these, 1117 species, 338 genera, 83 families, and 16 orders belong to basidiomycota, and 176 species, 73 genera, 32 families, and eight orders belong to ascomycota (ahmad et al. 1997: aman et al. 2022). so far, five species of panaeolus, i.e., p. fimicola (pers.) gillet (1878: 621), p. papilionaceus (bull.) quél. (1872: 152), p. sphinctrinus (fr.) quél. (1872: 151), p. semiovatus (sowerby) s.lundell & nannf. (1938: 537) and p. rickenii hora (1960: 454) have been reported from pakistan (ahmad et al. 1997; razaq et al. 2012). during the past two decades, both morphological and phylogenetic analyses have been used in mycological research for the identification of new species of the genus panaeolus (drehmel et al. 1999; zhang et al. 2004; zhao et al. 2011; razaq et al. 2012; jayasiri et al. 2015; wang & tzean 2015; li et al. 2016; zhao et al. 2016). but, only a few species of panaeolus have been reported based on both morphological and phylogenetic analyses (ma 2014; ediriweera et al. 2015; wang & tzean 2015; undan 2016) and most panaeolus species have been described based only on morphology (gerhardt 1996; amandeep et al. 2014; halama et al. 2014; silva-filho et al. 2019). in the present study, some interesting collections of panaeolus were made from three different locations in punjab, pakistan. all the localities lie in the semi-arid region with a maximum average temperature of 45°c and a long rainy season, i.e., july to september (ahmad et al. 2019). both morphological characteristics and phylogenetic analyses of its and 28s sequence data were used to determine the taxonomic position of the new panaeolus species which is subsequently described here in detail. material and methods type locality the type specimen was collected from haroonabad, bahawalnagar district, punjab, pakistan (29°60′81″ n, 73°14′68″ e, 163 m a.s.l.) during the monsoon rainy season of august 2019. the asif m. et al., new species of panaeolus from punjab, pakistan 79 temperature of the collection site varies from a minimum of 11°c to a maximum of 50°c and the average annual rainfall is 99 mm (ahmed et al. 2014a, 2014b). the main vegetation of the area includes dalbergia sissoo roxb., vachellia nilotica (l.) p.j.h.hurter & mabb., eucalyptus camaldulensis dehnh., azadirachta indica a.juss. and albizia lebbeck (l.) benth. (ahmed et al. 2014b). the region falls under a hot semi-arid climate (bsh) following the climate map and classification (peel et al. 2007; belda et al. 2014). the second collection site is kasur, punjab, pakistan (31°12′79″ n, 74°44′08″ e, 218 m a.s.l.). climatic conditions are tremendously variable and described as scorching hot summers and cold winters. monsoons occur towards the end of june with the rainy season lasting 2–3 months. the common woody flora of the district includes capparis decidua edgew. (forssk.), dalbergia sissoo roxb., prosopis cineraria (l.) druce, senegalia modesta (wall.) p.j.h.hurter, and vachellia nilotica (l.) p.j.h.hurter & mabb. (nasir et al. 1995; zabihullah et al. 2006; lateef et al. 2008; durrani & shakoori 2009; anwar et al. 2012; waheed et al. 2020). the third collection site is lahore, the capital of punjab, pakistan (32°52′04″ n, 74°35′87″ e, 217 m a.s.l.). it has a hot semi-arid climate (köppen climate classification bsh) with long, wet, and exceptionally hot summers, dry and warm winters, annual monsoons and dust storms. the monthly mean temperature ranges between 10 and 38°c during the year in lahore (https://rmcpunjab.pmd.gov.pk/). it has a long rainy season (from the end of june to mid-september), with annual mean rainfall of 838 mm, which increases the humidity of the area (siddiqui et al. 2020; tanveer et al. 2020). morphoanatomical study basidiomata were photographed in the field, and morphological features such as size, shape, and color of basidiomata were recorded at the time of collection. munsell’s soil color chart (1975) was used for color notations and for morphological terminologies, vellinga (2001) was followed. specimens were air-dried, kept in zipper bags, and deposited in the herbarium of the institute of botany, university of the punjab, lahore (lah). for the microscopic study, slides were prepared (lamellae, pileus, stipe) using free-hand sections of the dried materials rehydrated in 5% aqueous koh (percentage weight/volume (w/v)) and stained with congo red (2%) and melzer’s reagent following the microscopic procedures of liang et al. (2011, 2018) and cai et al. (2018) and observed under the light microscope (cxrii, labomed labo america inc., fremont, ca, usa) equipped with a camera to examine the following microscopic characteristics under 400× and oil immersion 1000× magnification: size and shape of basidiospores, basidia, cheilocystidia, pileipellis, and stipitipellis. data for morphoanatomical characteristics was based on at least 25 measurements each of basidia, cheilocystidia, and basidiospores. length and width ratios of basidiospores is designated as ‘q’, while the average length and average width ratios of all the basidiospores measured is given as ‘avq’. the notation ‘n/b/p’ is given, where ‘n’ is the number of basidiospores measured, from ‘b’ basidiomata and from ‘p’ collections (bas 1969; yu et al. 2020). dna extraction, pcr amplification, and sequencing genomic dna was extracted using the ctab method (porebski et al. 1997). we amplified the its and 28s regions of nuclear ribosomal dna, using the primer combination its1f/its4 and lr0r/lr5 for its and 28s, respectively (white et al. 1990; vilgalys & hester 1990). polymerase chain reaction (pcr) was performed in a 25 μl reaction volume following the protocol given by warnke (2020). the (pcr) products were sequenced with the same primers in both directions at ©macrogen inc. (238, teheran-ro, gangnam-gu, seoul, republic of korea). the newly generated sequences of p. punjabensis m. asif, q. firdous, a. izhar, niazi & khalid sp. nov. were submitted to genbank: ky636363, mz265142, mz823627, op681142 (its) and on116490, on116491, on116492 (28s). https://rmcpunjab.pmd.gov.pk/ european journal of taxonomy 888: 77–96 (2023) 80 phylogenetic analyses for phylogenetic analysis, its and 28s sequences of nrdna, generated from the pakistani collections were compared with sequences in genbank using the blast tool, priority was given to those sequences which showed high bootstrap value in phylogenetic analyses (altschul et al. 1990). the datasets were created by adding newly generated sequences of p. punjabensis sp. nov. plus the highest-scored blast hits were chosen from genbank and the other sequences of the genus from previous studies (malysheva et al. 2019; hu et al. 2020; voto & angelina 2021). four its and three 28s sequences were newly generated during this study, and 39 its sequences of family galeropsidaceae including psathyrella vesterholti örstadius & e.larss. (örstadius et al. 2015: 29) (kc992938) as an outgroup and 30 lsu sequences of family galeropsidaceae including psathyrella vesterholti as an outgroup, were used for phylogenetic analyses. the sequence alignment of both datasets was done separately using muscle ver. 3.8 (edgar 2004), and initial phylogenetic analyses were performed using mega-x software (tamura et al. 2011) using the maximum likelihood (ml) method. to calculate the appropriate model of nucleotide evolution, the nrits dataset was segmented into three partitions, its1, 5.8s, and its2. the best fit model of nucleotides substitution based on the lowest bic (bayesian information criterion) values for each partition and for 28s based dataset was chosen with jmodeltest2 on xsede via cipres science gateway (darriba et al. 2012). the final phylogenetic analyses of the its and 28s datasets were carried out separately using raxmlhpc2 ver. 8.1.11 under the cipres science gateway (miller et al. 2010). in the ml analysis, 1000 bootstrap repetitions were obtained as statistical support with rapid bootstrapping for both datasets. phylogenetic trees generated by bayesian inference (bi) analyses were performed with a markov chain monte carlo (mcmc) coalescent approach implemented in beast ver. 1.8.2 (drummond & rambaut 2007). both analyses resulted in a similar topology. significant support was considered to be ≥ 80%. in the resulting trees, bootstrap values obtained from maximum likelihood analyses and values of bayesian posterior probabilities > 0.7 were reported. figtree ver. 1.4.3 (rambaut 2014) was used for displaying the phylogenetic trees, and both trees were annotated using adobe illustrator 2020 ver. 24.1.2.408. results phylogenetic analysis of its dataset the fragment size of the target region was 758 bp long. from blast results, the its sequences of panaeolus punjabensis sp. nov. show 99% similarity with p. papilionaceus. in the its phylogenetic tree, the four sequences of the new species formed a separate lineage with strong statistical support. it formed a sister clade with these species of panaeolus: p. papilionaceus, p. campanulatus (bull.) quél., and p. sphinctrinus. in this analysis, p. guttulatus bres. (ku725993) is also closely related to the new species (fig. 1; table 1). phylogenetic analysis of 28s dataset the 28s alignment contained 892 total characters of which 815 were conserved and 73 were variable. in the 28s phylogenetic tree, the three sequences of the new species also formed a separate lineaged with good bootstrap support. it also formed a sister clade with these species of panaeolus: p. papilionaceus, p. campanulatus, and p. sphinctrinus. in this analysis, p. semiovatus (mh868191) is also closely related to the new species (fig. 2; table 1). asif m. et al., new species of panaeolus from punjab, pakistan 81 fig. 1. molecular phylogenetic placement of panaeolus punjabensis m. asif, q. firdous, a. izhar, niazi & khalid sp. nov. based on maximum likelihood (ml) method of its sequences. newly generated sequences are in bold. panaeolus punjabensis m. asif, q. firdous, a. izhar, niazi & khalid sp. nov. (lah36793, t = type specimen) is referring to the holotype. bootstrap values > 80% and bayesian posterior probabilities > 0.7 are shown above the branches. 0.04 ab158633 panaeolus cambodginiensis hm035081 panaeolus sphinctrinus jf908514 panaeolus fimicola hm035084 panaeolus cyanescens jf908517 panaeolus semiovatus mh592651 panaeolus guttulatus jf908518 panaeolus acuminatus mf497586 panaeolus antillarum op681142 panaeolus punjabensis lah37417 op549249 panaeolina foenisecii kf830093 panaeolus papilionaceus ky559329 panaeolus rickenii mh285992 panaeolus olivaceus ky636363 panaeolus punjabensis lah36792 mh593015 panaeolus olivaceus mz265143 panaeolus punjabensis lah36793 t mk397578 panaeolus plantaginiformis jf908515 panaeolus antillarum on561653 panaeolina foenisecii kc992938 psathyrella vesterholti mf497585 panaeolus antillarum fj755227 panaeolus sphinctrinus mz856314 panaeolus mexicanus jf908522 panaeolus campanulatus mg966283 panaeolus bisporus jf961376 panaeolus campanulatus dq182503 panaeolus sphinctrinus km982723 panaeolus alcis mk397577 panaeolus plantaginiformis ku725993 panaeolus guttulatus mn482689 panaeolus axfordii eu834287 panaeolus cyanescens jf961377 copelandia tropicalis mg250381 copelandia sp. jf961370 panaeolus subbalteatus jf908521 panaeolus retirugis mz823627 panaeolus punjabensis lah36794 jf908516 panaeolus rickenii fj478119 panaeolus retirugis 82/0.75 88/1 100/1 100/1 98/1 93/1 98/0.87 100/1 90/1 80/0.75 99/1 87/0.91 98/0.85 97/0.92 98/0.81 91/0.87 100/1 95/1 86/1 82/1 98/1 100/1 family galeropsidaceae european journal of taxonomy 888: 77–96 (2023) 82 0.01 dq071696 panaeolina foenisecii mk278436 panaeolus sphinctrinus mh867057 panaeolus subbalteatus on116492 panaeolus punjabensis lah36794 dq470817 panaeolus sphinctrinus on116490 panaeolus punjabensis lah36793 t on116491 panaeolus punjabensis lah36792 kf830082 panaeolus papilionaceus mk397600 panaeolus plantaginiformis mh867059 panaeolus subbalteatus mk278435 panaeolus papilionaceus mh867785 panaeolus retirugis dq071694 panaeolus semiovatus mk278427 panaeolopsis nirimbii mh867781 panaeolus acuminatus ay207263 panaeolus acuminatus mk397599 panaeolus plantaginiformis mk278429 panaeolus cyanescens mh867058 panaeolus subbalteatus mh867786 panaeolus retirugis mh867787 panaeolus retirugis u11924 panaeolina foenisecii mh868191 panaeolus semiovatus ab104646 panaeolus sphinctrinus mh867784 panaeolus retirugis mh867056 panaeolus subbalteatus kc992938 psathyrella vesterholti ay207265 panaeolus papilionaceus mk278431 panaeolus fimicola mh867557 panaeolina foenisecii 97/1 83/1 82/1 81/0.99 91/0.87 97/1 93/1 92/1 80/1 81/0.92 95/1 85/1 86/0.87 85/0.79 97/1 84/1 family galeropsidaceae fig. 2. molecular phylogenetic placement of panaeolus punjabensis m. asif, q. firdous, a. izhar, niazi & khalid sp. nov. based on maximum likelihood (ml) method of 28s sequences. newly generated sequences are in bold. panaeolus punjabensis m. asif, q. firdous, a. izhar, niazi & khalid sp. nov. (lah36793, t = type specimen) is referring to the holotype. bootstrap values > 80% and bayesian posterior probabilities > 0.7 are shown above the branches. asif m. et al., new species of panaeolus from punjab, pakistan 83 copelandia sp. 294130 mg250381 no data usa unpublished c. tropicalus – jf961377 no data china unpublished panaeolina foenisecii fo 46609 no data dq071696 – garnica et al. 2007 p. foenisecii cbs 142.40 no data mh867557 – vu et al. 2018 p. foenisecii – no data u11924 chapela et al. 1994 p. foenisecii pul00038201 on561653 no data usa unpublished p. foenisecii – op549249 no data usa unpublished panaeolopsis nirimbii perth 7680368 no data mk278427 australia varga et al. 2019 panaeolus acuminatus 4084 jf908518 no data italy osmundson et al. 2013 p. acuminatus glm 45986 no data ay207263 germany walther et al. 2005 p. acuminatus cbs 268 no data mh867781 hungary varga et al. 2019 p. alcis 88085 km982723 no data canada moser 1984 p. antilarum 748 jf908515 no data – – p. antilarum cort:013830 mf497586 no data dominican desjardin & perry 2017 republic p. antilarum sfsu:ded 7874 mf497585 no data thailand desjardin & perry 2017 p. axfordii mflu 19-2367 mn482689 no data china hu et al. 2020 p. bisporus 188954 mg966283 no data usa ediriweera et al. 2015 p. cambodginiensis nbrc30222 ab158633 no data – maruyama et al. 2006 p. campanulatus 10141 jf908522 no data italy osmundson et al. 2013 p. campanulatus no data jf961376 no data china ma 2014 p. cyanescens no data hm035084 no data – broussal & dumesny 2015 p. cyanescens 6576 aqui eu834287 no data italy han et al. 2010 p. cyanescens nl-0429 no data mk278429 hungary varga et al. 2019 p. fimicola  474 jf908514 no data italy wang & tzean 2015 p. fimicola  nl-0232 no data mk278431 hungary varga et al. 2019 p. guttulatus 137 mh592651 no data – unpublished p. guttulatus amb n. 18101 ku725993 no data – unpublished p. mexicanus ange1557 mz856314 no data dominican voto & angelini 2021 republic p. olivaceus 89608 mh285992 no data usa unpublished p. olivaceus – mh593015 no data iran unpublished p. plantaginiformis le 2864 mk397578 mk397600 uzbekistan malysheva et al. 2019 table 1. (continued on next page) taxa information and genbank accession numbers of nrits and lsu sequences of panaeolus (fr.) quél. used in the molecular phylogenetic analyses. sequences generated for this study are shown in bold. species voucher no. genbank accession no. origin reference its lsu european journal of taxonomy 888: 77–96 (2023) 84 p. plantaginiformis le 2862 mk397577 mk397599 russia malysheva et al. 2019 p. papilionaceus dna1940 kf830093 kf830082 usa ediriweera et al. 2015 p. papilionaceus dna1940 kf830093 kf830082 usa ediriweera et al. 2015 p. papilionaceus db 4552 no data mk278435 austria varga et al. 2019 p. papilionaceus glm 45989 no data ay207265 germany walther et al. 2005 p. punjabensis lah36792 ky636363 on116491 pakistan this study p. punjabensis lah36794 mz823627 on116492 pakistan this study p. punjabensis t lah36793 mz265143 on116490 pakistan this study p. punjabensis lah37417 op681142 no data pakistan this study p. reckenii tenn:054965 ky559329 no data argentina unpublished p. retirugis 7070 jf908521 no data italy osmundson et al. 2013 p. retirugis xsd08077 fj478119 no data china undan 2016 p. retirugis cbs 271 no data mh867784 france vu et al. 2019 p. retirugis cbs 273 no data mh867786 france vu et al. 2019 p. retirugis cbs 274 no data mh867787 france vu et al. 2019 p. retirugis cbs 272 no data mh867785 france vu et al. 2019 p. rickenii 749 jf908516 no data italy osmundson et al. 2013 p. semiovatus 4083 jf908517 no data italy osmundson et al. 2013 p. semiovatus glm 51235 no data dq071694 – garnica et al. 2007 p. semiovatus cbs 388 no data mh868191 france vu et al. 2019 p. sphinctrinus cbs 582 hm035081 no data pakistan razaq et al. 2012 p. sphinctrinus pbm 2009 dq182503 dq470817 pakistan razaq et al. 2012 p. sphinctrinus cz519-3 fj755227 no data pakistan razaq et al. 2012 p. sphinctrinus ky7130 no data ab104646 japan maruyama et al. 2003 p. sphinctrinus nl-3955 no data mk278436 slovakia varga et al. 2019 p. subbalteatus no data jf961370 no data china sette et al. 2010 p. subbalteatus cbs 331 no data mh867059 france vu et al. 2019 p. subbalteatus cbs 329 no data mh867058 france vu et al. 2019 p. subbalteatus cbs 327 no data mh867056 france vu et al. 2019 p. subbalteatus cbs 328 no data mh867057 france vu et al. 2019 outgroup psathyrella vesterholtii jhp10.086 kc992938 kc992938 denmark örstadius et al. 2015 species voucher no. genbank accession no. origin reference its lsu table 1. (continued). asif m. et al., new species of panaeolus from punjab, pakistan 85 taxonomy phylum basidiomycota r.t.moore class agaricomycetes doweld order agaricales underw. family galeropsidaceae singer genus panaeolus (fr.) quél. panaeolus punjabensis m. asif, q. firdous, a. izhar, niazi & khalid sp. nov. mycobank mb 840898 figs 3–4 diagnosis the new species turns bluish on handling so it is hallucinogenic and can be distinguished by its broadly fusiform basidiospores, claviform cheilocystidia with rounded tips, and clavate caulocystidia. etymology specific epithet ‘punjabensis’ refers to the type locality, punjab province, pakistan. type material holotype pakistan • punjab province, haroonabad city, district bahawalnagar; 29°60′81″ n, 73°14′67″ e; alt. 163 m a.s.l.; on nutrient-rich loamy soil; 4 aug. 2019; muhammad asif, bwn-45; genbank nos mz265143 (nrits); on116490 (28s); lah[36793]. additional material examined pakistan • punjab province, lahore, 32°52′04″ n, 74°35′87″ e; alt. 217 m a.s.l.; on loamy soil; 10 jul. 2015; qudsia firdous, brb s-01; genbank nos ky636363 (nrits); on116491 (lsu); lah[36792] • same collection data as for preceding; 28 jul. 2016; qudsia firdous, brb s-22; genbank nos mz823627 (nrits); on116492 (lsu); lah[36794] • punjab province, kasur district, 31°12′79″ n, 74°44′08″ e; alt. 218 m a.s.l.; on fallen plant debris; 6 sep. 2020; aiman izhar, ks-0018; genbank no. op681142 (nrits); lah[37417]. description basidiomata 4.4–5.8 cm tall. pileus 1–1.5 cm diam, conic to parabolic when young, becoming convex with maturity, dry; surface light brown at the center (7.5yr8/4), light grayish green (7.5gy8/1) toward margins, smooth when young becoming rugulose at maturity; margin straight in young stage, striate at maturity (fig. 3a, d). lamellae free, olive black (5gy2/1), even margins, distantly placed, two tiers of regularly arranged lamellulae (fig. 3b–c). stipe 3.8–5.3 × 0.3–0.6 cm, surface light grayish-green (10gy8/1), central, equal, surface smooth and glabrous, dry, slightly bulbous base, bruising blue on handling (fig. 3e). annulus and volva absent. odor is indistinct. basidiospores [75/3/3] (13.2–)13.4–16.4(–16.7) × (7.5–)8.2–9.6(–11.4) µm, on average 15 × 9.5 µm, q = 1.4–1.6, qav = 1.5, broadly fusiform, smooth, apiculus absent, thick-walled, hyaline in koh, no colour change in melzer‘s reagent, non-guttulate, germ pore obvious (fig. 4b). basidia (24.8–)24.9–27.9 (–28.1) × (14.3–)14.4–15.6(–15.9) µm, on average 26.2 × 15.1 µm, broadly clavate, mostly bi-spored, rarely trior tetra-spored, thick-walled, hyaline in koh, non-guttulate (fig. 4a). cheilocystidia (30.5–) 32.1–41.4(–44.3) × (6–)6.1–9.1(–9.5) µm, on average 37.4 × 7.8 µm, claviform with flexuous neck and rounded apices, thin-walled, hyaline in koh, non-guttulate (fig. 4c). pileocystidia (18–)19.2–32 https://www.mycobank.org/page/name%20details%20page/840898 european journal of taxonomy 888: 77–96 (2023) 86 fig. 3. morphology of panaeolus punjabensis m. asif, q. firdous, a. izhar, niazi & khalid sp. nov. (lah36793, a–b holotype). scale bars: a–d = 1 cm, e = 10 cm. photos by muhammad asif. asif m. et al., new species of panaeolus from punjab, pakistan 87 fig. 4. microscopic characters of panaeolus punjabensis m. asif, q. firdous, a. izhar, niazi & khalid sp. nov. (lah36793, holotype). a. basidia. b. basidiospores. c. cheilocystidia. d. caulocystidia. e. pileocystidia. scale bars: a–d = 10 µm, e = 20 µm. drawings by aiman izhar. european journal of taxonomy 888: 77–96 (2023) 88 p . p un ja be ns is 10 –1 5 pa ra bo lic to lig ht b ro w n ol iv e bl ac k 38 –5 3 × lig ht 13 .2 –1 6. 7 × br oa dl y 30 .5 –4 4. 3 × cl av if or m w ith t hi s st ud y sp . n ov . co nv ex at th e ce nt er , 3– 6 gr ay is h 7. 5– 11 .4 fu si fo rm 6– 9. 5 fle xu ou s ne ck lig ht g ra yi sh gr ee n an d ro un de d gr ee n to w ar d ap ic es m ar gi ns p. a cu m in at us 17 –2 0 br oa dl y co ni c da rk re dd is h gr ay is h bl ac k 62 –7 5 × 2 re dd is h 11 .4 –1 5 × le m on 15 .5 –2 5. 5 × fu so id -v en tr ic os e k au r e t a l. to b ro ad ly br ow n to br ow n 7. 8– 11 sh ap ed 5. 5– 8. 5 to ir re gu la rl y 20 14 ; o s be llsh ap ed gr ay is h cy lin dr ic m un ds on et a l. 20 13 br ow n p. a lc is 4– 10 co m pa nu la te pa le g ra y da rk g ra y 20 –9 0 × pe x pa le 18 –2 1 × el lip so id 25 –3 5 × 4– 6 ve rs if or m m os er 1 98 4 to c on ic al 0. 5– 1. 5 oc hr ac eo us 9. 5– 12 gr ay , b as e br ow ni sh p. a xf or di i 16 –2 1 he m i re dd is h m ot tle d 42 –5 1 × gr ay is h 8. 8– 11 .4 × lim on i 24 .6 –4 2. 7 × na rr ow ly u tr i h u et a l. sp he ri ca l t o br ow n to da rk g ra y 1. 5– 2. 5 or an ge to 6. 3– 9 fo rm to 5. 9– 10 .7 fo rm 20 20 ca m pa nu la te gr ay is h lig ht b ro w n el lip so id w hi te p. c am bo dg in ie ns is 12 –2 5 co nv ex to ch oc ol at e pa lli d, th en 55 –9 5 × w hi tis h to 10 .5 –1 2 × le m on 26 –3 9 × po ly m or ph ic st am et s 19 96 ; br oa dl y co nv ex br ow n be gr ay is h bl ac k 3. 5– 5 cr ea m , 6. 5– 9 sh ap ed 10 –1 2. 2 w ee ks e t a l. co m in g ye l to b la ck br ow n ne ar 19 79 lo w is h br ow n th e ba se p. c ya ne sc en s 10 –1 5 he m is ph er ic to pu re w hi te da rk g ra yi sh 50 × 2 –3 un if or m ly 11 –1 8 × lim on i ab se nt ab se nt w ar tc ho w e t co nv ex br ow n w hi te 9– 13 .7 fo rm to al . 2 01 0 el lip so id p.  fi m ic ol a  10 –2 0 ca m pa nu la te di ng y gr ay m ot tle d gr ay 60 –1 00 × di ng y pa le 11 –1 4 × su bam yg 22 .7 –2 7 × cy lin dr ic to st am et s 19 96 ; to c on ve x to b la ck , 1– 2 to w hi tis h 7– 9. 5 da lif or m 5. 7– 10 su bl ag en if or m w an g & sl ig ht re dd is h t ze an 2 01 5 ta bl e 2. ( co nt in ue d on n ex t p ag e) c om pa ri so n of m ac ro an d m ic ro -c ha ra ct er is tic s of p an ae ol us p un ja be ns is m . a si f, q . f ir do us , a . i zh ar , n ia zi & k ha lid s p. n ov ., an d its c lo se ly re la te d ta xa . a bb re vi at io ns : l = l en gt h; w = w id th . sp ec ie s na m e p ile us p ile us p ile us l am el la e st ip e st ip e sp or e sp or e c he ilo c he ilo r ef er en ce di am et er sh ap e co lo r co lo r l × w co lo r l × w sh ap e cy st id ia cy st id ia (m m ) (m m ) (µ m ) l × w (µ m ) sh ap e asif m. et al., new species of panaeolus from punjab, pakistan 89 p. g ut tu la tu s 10 co nv ex da rk o liv e bl ac k 40 × 2 sn uff 7. 5– 10 × el lip tic al 25 × 5 cy lin dr ic al se id m oh am br ow n br ow ni sh 4– 6 m ad i e t a l. 20 19 ; s ul ia m an 2 01 9 p . p ap ili on ac eu s 10 –5 0 co ni ca l t o br ow ni sh to gr ay to 60 –1 20 × w hi tis h to 12 –1 4 × ci tr if or m 35 –5 3 × ve rm if or m , u rn a br ah am ca m pu la na te gr ay is h br ow n en tir el y bl ac k 2– 4 sn uff b ro w n 7– 8 7– 12 .5 sh ap ed o r t ub ul ar 20 07 ; k uo 20 07 ; a m an de ep e t a l. 20 14 ; e di ri w ee ra e t a l. 20 15 p. r ic ke ni i 18 –2 0 ca m pa nu la te to da rk b ro w n m ot tle d gr ay 15 –1 40 × bu ff , l at e 9– 12 × el lip so id 19 .5 –3 7 × cl av at e to m a 20 14 co nv ex is h br ow n 1– 2 da rk b ro w n 7– 9. 5 to b ro ad ly 7– 13 la ge ni fo rm w ith w hi tis h to re dd is h el lip so id m ar gi n br ow n p. s em io va tu s 30 –6 0 eg gsh ap ed to ci nn am on b uff pa le b ro w n, 10 0– 16 0 × w hi tis h to 18 .5 –2 1 × el lip tic al ab se nt ab se nt st am et s 19 96 ; co ni co -c on ve x to p in ki sh b uff la te r m ot tle d 6– 10 pa lli d bu ff 10 –1 1. 5 o sm un ds on an d fa di ng to bl ac ki sh et a l. 20 13 w hi tis h p. s ph in ct ri nu s 10 –2 0 be ll sh ap ed to br ow ni sh w he n gr ay a nd 25 –9 0 × sn uff b ro w n 0. 4– 11 .3 × ci tr if or m 11 .0 –1 6. 5 × po ly m or ph ic e di ri w ee ra um bo na te im m at ur e, m ot tle d bl ac k, 1– 2 6. 6– 7. 5 to le m on 5. 5– 9. 2 et a l. 20 15 gr ay is h br ow n l at er e nt ir e sh ap ed w he n m at ur e bl ac k ta bl e 2. (c on tin ue d) . sp ec ie s na m e p ile us p ile us p ile us l am el la e st ip e st ip e sp or e sp or e c he ilo c he ilo r ef er en ce di am et er sh ap e co lo r co lo r l × w co lo r l × w sh ap e cy st id ia cy st id ia (m m ) (m m ) (µ m ) l × w (µ m ) sh ap e european journal of taxonomy 888: 77–96 (2023) 90 (–34.4) × (10.4–)11.9–13.9(–17.4) µm, on average 25.5 × 13.4 µm, clavate to vesiculose, thin-walled, hyaline (fig. 4e). caulocystidia (25.3–)26.7–36.9(–37.5) × (6–)8.1–9.7(–10.5) µm, on average 31.4 × 8.7 µm), clavate, thick-walled (fig. 4d). clamp connections are absent in all tissues. habitat solitary or in small groups on loamy soil containing herbivore (cattle) dung. known distribution known only from three localities, bahawalnagar, kasur, and lahore, punjab, pakistan. discussion the genus panaeolus is quite similar in appearance to panaeolina in the field (kalichman et al. 2020). the two genera can be differentiated on the basis of basidiospores morphology and lamellae color. lamellae in panaeolus are grayish-black and basidiospores are smooth, in panaeolina spores are ornamented and lamellae are dark brown (kaur et al. 2014). all previously reported species of panaeolus from pakistan are not hallucinogenic, but our new species turns bluish on handling which indicates that it is a hallucinogenic species. several different tests can be performed to test the hallucinogenic various mushrooms including panaeolus, such as amplified fragment length polymorphism (aflp) and high-resolution melting essays (hrm) (lee et al. 2000; zhang et al. 2022). a detailed comparison of macroand micro-characteristics of all the closely related species of panaeolus is given in table 2, and from the molecular phylogenetic analyses of its and 28s and morphoanatomical comparison given in table 2, we conclude that panaeolus punjabensis sp. nov. is a new species. in the its-based phylogenetic analysis, p. papilionaceus (kf830093), p. retirugis (fr.) gillet (jf908521), p. sphinctrinus (hm03581, dq182503), p. companulatus (l.) quél. (jf908522), and p. alcis m.m.moser (km982723) lie in the same clade and are closly related to the newly described species, while in the 28s-based phylogenetic analysis, p. semiovatus (mh868191, dq071694), p. acuminatus (p.kumm.) quél. (mh867781, ay207263), p. sphinctrinus (mk278436, ab104646), p. papilionaceus (kf830082, mk278435, ay207265), and p. retirugis (mh867784, mh867785, mh867786, mh867787) are close relatives of our new species. in both phylogenetic analyses, different sequences of the same species appear on different positions such as p. retirugis (gillet 1878: 621) (current name, p. papilionaceus), p. sphinctrinus, and p. campanulatus (quélet 1872: 151), and this could result to misidentification of species owing to close similarities among the species in the genus panaeolus (razaq et al. 2012). acknowledgements the authors are grateful to dr. shah hussain (sultan qaboos university, muscat, oman) and dr. thatsanee luangharn (mae fah luang university, chiang rai, thailand) for their critical review, valuable comments, and suggestions on an earlier version of the manuscript which helped us a lot to improve the article. we are thankful to dr. francis q. brearley (manchester metropolitan university, united kingdom) for the linguistic review of the manuscript. we are also highly obliged to all the anonymous reviewers for their corrections and suggestions to improve this paper. references abraham w.r. 2007. bioactive sesquiterpenes produced from fungi: possibilities and limitations. in: rai m. 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meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; leibniz institute for the analysis of biodiversity change, bonn – hamburg, germany; national museum of the czech republic, prague, czech republic. https://doi.org/10.1016/b978-0-12-372180-8.50042-1 https://doi.org/10.11646/phytotaxa.450.1.2 https://doi.org/10.1080/20961790.2021.1875580 https://doi.org/10.1007/s13225-011-0136-7 https://doi.org/10.1007/s13225-016-0357-x a new mesoserphid wasp from the middle jurassic of northeastern china (hymenoptera, proctotrupoidea) yan zheng 1,* & jun chen 2 1,2 institute of geology and paleontology, linyi university, shuangling rd., linyi 276000, china. 2 state key laboratory of palaeobiology and stratigraphy, nanjing institute of geology and palaeontology, east beijing road, nanjing 210008, china. * corresponding author: zhengyan536@163.com 2 email: rubiscada@sina.com 1 urn:lsid:zoobank.org:author:28eb8d72-5909-4435-b0f2-0a48a5174cf9 2 urn:lsid:zoobank.org:author:f15c888b-77fc-4f68-9272-e549823de4cf abstract. a new genus and species of mesoserphidae (hymenoptera), juraserphus modicus gen. et sp. nov., is described based on a well-preserved fossil specimen from the middle jurassic jiulongshan formation of northeastern china. it is characterized by the following forewing features: the forking of rs+m located approximately one-third of the distance between 1m-cu and 2r-rs, both 1cu-a and 2cu-a antefurcal; 1-m more than twice as long as 1m-cu and hind wing with cells r and rm closed. in addition, it has a short ovipositor, only extending slightly beyond the metasomal apex. its new morphological characters broaden the diversity of mesoserphidae in the mesozoic and provide new insights into the evolution and relationships of mesoserphidae. keywords. mesoserphidae, new taxon, taxonomy, daohugou, jiulongshan formation. zheng y. & chen j. 2017. a new mesoserphid wasp from the middle jurassic of northeastern china (hymenoptera, proctotrupoidea). european journal of taxonomy 379: 1–8. https://doi.org/10.5852/ejt.2017.379 introduction the superfamily proctotrupoidea, a basal group of prototrupomorpha within apocrita, includes 11 extant families and one extinct family (grimaldi & engel 2005). mesoserphidae was erected by kozlov (1970), representing the only extinct family of proctotrupoidea. fossil records of mesoserphids can be traced back to the middle jurassic according to the earliest fossil from the upper middle jurassic jiulongshan formation of northeastern china (shih et al. 2011). this family was once considered to comprise two subfamilies: mesoserphinae kozlov, 1970 and karataoserphinae rasnitsyn, 1994 (grimaldi & engel 2005). however, the latest research based on a phylogenetic analysis indicated that these two subfamilies are paraphyletic and should be abandoned (li et al. 2016). so far, 22 fossil genera with 53 species have been described worldwide (including their locality, stratigraphic level, external morphological characteristics and body measurements), mostly distributed in european journal of taxonomy 379: 1–8 issn 2118-9773 https://doi.org/10.5852/ejt.2017.379 www.europeanjournaloftaxonomy.eu 2017 · zheng y. & chen j. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:46363f91-1864-4aeb-b8f8-9b185d285578 1 mailto:zhengyan536%40163.com?subject= mailto:rubiscada%40sina.com?subject= http://zoobank.org/urn:lsid:zoobank.org:author:28eb8d72-5909-4435-b0f2-0a48a5174cf9 http://zoobank.org/urn:lsid:zoobank.org:author:f15c888b-77fc-4f68-9272-e549823de4cf https://doi.org/10.5852/ejt.2017.379 https://doi.org/10.5852/ejt.2017.379 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:46363f91-1864-4aeb-b8f8-9b185d285578 kazakhstan, transbaikalia and china (kozlov 1968; rasnitsyn 1983, 1986, 1990, 1994; zhang & zhang 2000; shih et al. 2011; shi et al. 2013; zhang et al. 2013; li et al. 2016). among these mesoserphids, 19 species were recorded from the middle jurassic, 23 from the late jurassic and 11 from the early cretaceous (li et al. 2016). studies have demonstrated the mesoserphidae to be highly morphologically divergent, such as in body length (from 1.7 mm to 12.8 mm), forewing length (from 1.5 mm to 10.4 mm), number of antennomeres, wing venation, morphology of the external ovipositor, etc. (zhang & zhang 2000; shih et al. 2011; shi et al. 2013; li et al. 2016). the new, well preserved specimen was collected from the jiulongshan formation (daohugou, inner mongolia, china), famous for its rich fossil insect fauna, including hymenoptera (shih et al. 2011; wang et al. 2014), coleoptera (pan et al. 2011; liu et al. 2015), diptera (chen et al. 2014; shi et al. 2015), odonata (zhang et al. 2008; nel & huang 2015) and others (ren et al. 2009; chen et al. 2015, 2016). the geological age of this formation is the latest middle jurassic (late callovian, ca 165–164 ma), measured by 40k/40ar, ar-ar and shrimp u-pb (he et al. 2004; liu et al. 2006; li et al. 2013). material and methods the fossil specimen studied herein was collected from the upper middle jurassic jiulongshan formation at daohugou village, wuhua town, ningcheng city, inner mongolia, china and is housed in the shandong tianyu museum of nature in pingyi county, shandong province, china. the fossil was examined and photographed, both dry and in ethanol, using a vhx 5000 digital microscope platform. line drawings were prepared with coreldraw 14.0 and adobe photoshop cs5. the specimen was quantitatively measured using the nih imagej software (http://rsb.info.nih.gov/ij/). the morphological terminology used herein follows huber & sharkey (1993) and rasnitsyn & zhang (2004). venation symbols: main longitudinal veins are c (costa), r (radius), rs (radial sector), m (media), cu (cubitus) and a (anal vein); crossveins are r-rs (radial crossvein), m-cu (mediocubita crossvein) and cu-a (anal crossvein); cells are r, mcu and cua. results order hymenoptera linnaeus, 1758 suborder apocrita gerstaecker, 1867 superfamily proctotrupoidea latreille, 1802 family mesoserphidae kozlov, 1970 juraserphus gen. nov. urn:lsid:zoobank.org:act:38cda49b-f168-47b0-9cfd-97ee52f82733 type species juraserphus modicus gen. et sp. nov., designated herein. diagnosis forewing 1-rs as long as 1-m; 1-m more than twice as long as 1m-cu; 2r-rs arising from middle of pterostigma, its width more than twice as long as width of pterostigma; 1cu-a antefurcal, 2cu-a antefurcal; forking of rs+m located approximately one-third of distance bewteen 1m-cu and 2r-rs, closer to 1m-cu; cell 1mcu complete trapezoid and less than half of cua. hind wing with cells r and rm closed. metasoma spindle-shaped, with elongated segments. short ovipositor, only extending slightly beyond metasomal apex. european journal of taxonomy 379: 1–8 (2017) 2 http://rsb.info.nih.gov/ij/ http://zoobank.org/urn:lsid:zoobank.org:act:38cda49b-f168-47b0-9cfd-97ee52f82733 etymology the generic name is composed of the prefix ‘jura’ from the jurassic period and the suffix of the genus name ‘serphus’. the gender is masculine. species included type species only. juraserphus modicus gen. et sp. nov. urn:lsid:zoobank.org:act:34160863-3f15-422f-81b7-70971337541c figs 1–2 diagnosis as for genus. etymology the specific epithet is an adjective derived from the latin word ‘modicus’, which means ‘moderate, average’, indicating that both the body and forewing lengths are medium. fig. 1. holotype of juraserphus modicus gen. et sp. nov. a. part. b. counterpart. c. left forewing and left hind wing of part. d. head and mesosoma of part (in ethanol). e. metasoma of part. scale bars = 2 mm. zheng y. & chen j., juraserphus modicus gen. et sp. nov. from the middle jurassic of china 3 http://zoobank.org/urn:lsid:zoobank.org:act:34160863-3f15-422f-81b7-70971337541c material examined holotype china: part and counterpart, female in dorsal view (tynm-48-1244 a–b). locality and age china: daohugou village, ningcheng county, inner mongolia; callovian (latest middle jurassic), jiulongshan formation. description body length 8.66 mm. head transversely ovoid, length 1.31 mm and width 0.63 mm. eye large, oviod and protruding. antenna filiform, thin, with nine antennomeres preserved, the first flagellomere long, subsequent flagellomeres gradually decreasing in length from base to apex. mesosoma about as wide as head; pronotum trapezoid, extremely short and obviously narrower than head and about 3.6 times as wide as long (length 0.96 mm; width 0.27 mm); mesoscutum subquadrate, 0.86 mm long and 0.37 mm wide; mesoprescutum length 0.52 mm and width 0.43 mm; notauli present and reaching transverse mesonotal suture; mesoscutellum triangular; mesopostnotum rectangular, 0.5 times as wide as long; metascutellum preserved and almost as long as wide. metasoma spindle-shaped, fig. 2. line drawings of juraserphus modicus gen. et sp. nov. a. habitus. b. left forewing. c. left hind wing. abbreviations: n1 = pronotum; n2 = mesonotum; n3 = metanotum; nv = notaulus; tms = transverse mesonotal suture; sc = mesoscutellum; pr = propodeum; vf1 = valvifer 1; vf2 = valvifer 2; vf3 = valvula 3. scale bar = 2 mm. european journal of taxonomy 379: 1–8 (2017) 4 with seven segments, first observed segment very short, subtrapezoidal; second to fifth segments similar to first in shape and length, but gradually increasing in width; sixth segment reversely trapezoidal and about twice as long as fourth segment; seventh segment about as long as sixth segment; valvifer 1, valvifer 2 and valvula 3 clearly discernible; ovipositor short, slightly stretched out of metasomal apex. left foreleg with only femur preserved, small. left midleg well preserved, with femur wider than tibia; tibia slender; tarsus thin, with basitarsus longest and about twice as long as 2nd tarsomere, 3rd tarsomere obviously shorter than 2nd, 4th shorter and thinner than 3rd, 5th twice as long as 4th and possessing two short claws. left hindleg with femur narrow basally and widened apically, with a spindle-like shape; tibia thicker than midtibia; tarsi similar to midtarsi. forewing length 7.05 mm, width 2.73 mm. pterostigma long and slightly oblique apicad, six times as long as wide. 1-rs and 1-m equal in length; 1-m more than twice as long as 1m-cu; 2r-rs arising from middle of pterostigma, oblique, about 2.5 times as long as pterostigmal width. cell 1+2r five-sided and surrounded by r, 1-rs, 1-rs+m, (2-rs+m) + (2-rs) and 2r-rs; m+cu straight and distinct, m and cu straight basally, slightly curved apically; 1-rs origin at a distance from pterostigma (about twice as long as 1-rs), and 2r-rs arising from middle of pterostigma; forking of rs+m located approximately onethird of distance between 1m-cu and 2r-rs; both 1cu-a and 2cu-a antefurcal; cell 1mcu subtrapezoidal, 1.8 times as long as wide basally; 1cu-a slightly longer than 1-m and two-thirds of 2cu-a in length. hindwing with long, oblique r-m meeting rs near its base; cell r long and closed; m+cu and cu partly tubular; m tubular. discussion various morphological characters of mesoserphidae have been recorded from the latest middle jurassic to early cretaceous (zhang & zhang 2000; shih et al. 2011; shi et al. 2013; li et al. 2016). compared with other mesoserphids, juraserphus gen. nov. has the rare combination of both 1cu-a and 2cu-a being antefurcal in the forewing, and cells rm and r closed in the hindwing. additionally, juraserphus gen. nov. is unique among mesoserphidae in having a female metasoma with five basal terga very short and jointly occupying only half of metosoma’s length, whilst the other half is formed by terga 6 and 7 only. juraserphus gen. nov. resembles beipiaoserphus zhang & zhang, 2000 and basiserphus li et al., 2016 with both 1cu-a antefurcal and 2cu-a antefurcal in the forewing, but differs from them in having the forking of rs+m located at a distance of one-third between 1m-cu and 2r-rs, and 1-m about twice as long as 1m-cu (vs one half and 1-m much shorter than 1m-cu in beipiaoserphus; one-sixth or onequarter and 1-m slightly longer than 1m-cu in basiserphus). juraserphus gen. nov has long crossvein 2r-rs, which is more than twice as long as the width of the pterostigma. the length of 2r-rs, however, is less than 1.5 times the width of the pterostigma in codoserphus shi et al., 2013 and more than five times as long as the width of the pterostigma in turgoserphus rasnitsyn, 1990 (rasnitsyn 1990; shi et al. 2013). 1cu-a and 2cu-a of the forewing offer significant characteristics. juraserphus gen. nov has both 1cu-a and 2cu-a antefurcal, as in campturoserphus rasnitsyn, 1986, scoliuroserphus rasnitsyn, 1986 and basiserphus li et al., 2016, but is different from the latter two and from lordoserphus rasnitsyn, 1994 and sinoserphus shih et al., 2011 in the 1cu-a and 2cu-a being interstitial, from apiciserphus li et al., 2016 in 1cu-a and 2cu-a postfurcal, and from amboserphus li et al., 2016 and yanliaoserphus shih et al., 2011 in 1cu-a postfurcal and 2cu-a antefurcal (li et al. 2016). furthermore, the position of the rs+m forking in the forewing could be taken as a key feature. the forking of rs+m is located one-third of the distance between 1m-cu and 2r-rs in juraserphus gen. nov, whereas it is about one-fifth of this distance in mesoserphus kozlov, 1968, sinoserphus shih et al., 2011 and basiserphus li et al., 2016, and approximately one half in amboserphus li et al., 2016 and beipiaoserphus zhang & zhang, 2000. zheng y. & chen j., juraserphus modicus gen. et sp. nov. from the middle jurassic of china 5 the ovipositor of mesoserphidae demonstrates significant variations, especially in its length, among the fossils described. the ovipositor in juraserphus gen. nov. is similar to the one in novserphus li et al., 2016 and ozososerphus li et al., 2016 in being short, extending beyond the metasomal apex. in contrast, the ovipositor in some other genera (for example, apiciserphus li et al., 2016, amboserphus li et al., 2016, udaserphus rasnitsyn, 1983 and choriserphus li et al., 2016) does not extend beyond the metasomal apex. on the other hand, sinoserphus shih, feng & ren, 2011 (especially s. grossus shih, feng & ren, 2011; 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botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain. european journal of taxonomy 379: 1–8 (2017) 8 https://doi.org/10.1016/j.palwor.2013.02.001 european journal of taxonomy 377: 1–7 issn 2118-9773 https://doi.org/10.5852/ejt.2017.377 www.europeanjournaloftaxonomy.eu 2017 · mavrodiev e.v. & yurtseva o.v. this work is licensed under a creative commons attribution 3.0 license. o p i n i o n p a p e r urn:lsid:zoobank.org:pub:172c34f7-e132-4635-9972-46cab5f6dad8 1 “a character does not make a genus, but the genus makes the character”: three-taxon statement analysis and intuitive taxonomy evgeny v. mavrodiev 1,* & olga v. yurtseva 2 1 university of florida, florida museum of natural history, museum road and newell drive, dickinson hall, 301, gainesville, fl, 32611, usa 2 m.v. lomonosov moscow state university, department of higher plants, 1–12, leninskie gory, 119234, moscow,russia * corresponding author: evgeny@ufl.edu 2 email: olgayurtseva@yandex.ru 1 urn:lsid:zoobank.org:author:ea16f665-354d-4d52-845b-341deb506d1a 2 urn:lsid:zoobank.org:author:b532893b-945a-4342-ab52-1a2da307efcd abstract. three-taxon statement analysis (3ta) is a method that may help to formalize the taxonomical intuition of the synapomorphy of the clade as a combination of its diagnostic traits, even if each trait, if taken separately, may be found in one or many other taxa of the same relationship. using example based on the real morphological data, we are showing that 3ta can recognize clade in case of the complete lack of it synapomorphies, as optimized under the criterion of standard parsimony. keywords. three-taxon statement analysis, intuitive taxonomy, synapomorphy, diagnostic traits. mavrodiev e.v. & yurtseva o.v. 2017. “a character does not make a genus, but the genus makes the character”: three-taxon statement analysis and intuitive taxonomy. european journal of taxonomy 377: 1–7. https://doi.org/10.5852/ejt.2017.377 this should be particularly clear to taxonomists. for anyone who has dealt with large and peculiarly diverse varieties of living forms, it is impossible to disregard that each group of organisms is characterized with its own distinct image. this image is not subject to verbal description. the only method of objective characterization of the group lies in the listing of its diagnostic features, i.e. those features that directly distinguish this group from others. however, such features are scarce… this “general image” is none but the type of group. b.s. kuzin (1992). the decadence of systematics, i. (the first author has translated the passage; bold and italic formatting added). as summarized in williams & ebach (2016), when commenting on scotland & steel (2015), character compatibility analysis in systematics captures the “intuitive taxonomic practice of recognizing taxa based on conserved non-homoplastic characters” (scotland & steel 2015: 493). according to williams & http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:172c34f7-e132-4635-9972-46cab5f6dad8 mailto:evgeny%40ufl.edu?subject= mailto:olgayurtseva%40yandex.ru?subject= http://zoobank.org/urn:lsid:zoobank.org:author:ea16f665-354d-4d52-845b-341deb506d1a http://zoobank.org/urn:lsid:zoobank.org:author:b532893b-945a-4342-ab52-1a2da307efcd https://doi.org/10.5852/ejt.2017.376 european journal of taxonomy 377: 1–7 (2017) 2 ebach (2016), who presented evidence of compatibility analysis within many different contexts (e.g., williams & ebach 2008, 2016), the definition of scotland & steel (2015) is not relevant to actual intuitive taxonomic practice. as stated in williams & ebach (2016), the taxonomic practice is based on the different procedure – on the finding of the taxa using the combinations of the characters, which are not necessarily compatible or unique. in support of this view, williams & ebach (2016) citing nelson (1996, 2004: 137), who in turn was summarizing antoine laurent de jussieu (1748–1836), who clearly stated that the combination of the characters actually “constitutes the essential and invariable character [=synapomorphy]” (italics is ours), even if “each character, taken separately”, may be found in one or many other taxa of the same relationship. as a recent relevant example of this, we note similar passages in mavrodiev et al. (2014), crespo et al. (2015) and yurtseva et al. (2016), who all found or summarized that unique combinations of partially overlapping morphological characters can successfully be used as a diagnostic for the taxonomic recognition of the intuitively clear genera of the iridaceae (previously circumscribed with the “broadly” defined iris (incl. belamcanda), aloe (aloeaceae), asparagaceae subfam. scilloideae tribe ornithogaleae (= hyacinthaceae subfam. ornithogaloideae), atraphaxis (polygonaceae), centaurium (gentianaceae), chenopodium (amaranthaceae), coreopsis (asteraceae), nothofagus (nothofagaceae), typha (typhaceae), among others. however, is there an exact method that may help researchers to invoke taxonomic intuition of “actual synapomorphy” as a combination of traits? as mentioned by williams & ebach (2016), three-taxon statement analysis (3ta), one of the two approaches to hennigian cladistics (e.g., williams & siebert 2000), originally proposed by nelson & platnick (1991), is just such a method. as was stressed by williams & ebach (2008: 210), the 3ta “… sees systematic data as reducible to the simplest relational form, the three-taxa statement or minimal relations: hence, the statement a(bc), where b and c share a relationship to the exclusion of a…”. therefore, 3ta based on the original way dealing with information: on the representation and analysis of the relationships directly (williams & ebach 2008). 3ta can recognize clades, for which the standard (conventional) maximum parsimony (mp) analysis provides no unequivocal synapomorphies, as was clearly demonstrated by nelson (1996) (see also williams & ebach 2005, 2008, 2016), using the hypothetical matrices of the binary characters, or even no synapomorphies, as optimized under the criterion of conventional parsimony (e.g., nelson & platnick 1991, see also mavrodiev 2016). to illustrate this analytical power of the 3ta, we would like to provide an example not from hypothetical data, but from an actual comprehensive taxonomic study based on recent treatment of the genus atraphaxis l. (polygonaceae) and the related taxonomical entities (yurtseva et al. 2016). a few words are still necessary to describe the proper context. recent molecular phylogenetic analyses of tavakkoli et al. (2015) and yurtseva et al. (2016) discovered the sister position of polygonum subsect. spinescentia boiss. (included by tavakkoli et al. (2015) in atraphaxis as atraphaxis sect. polygonoides s. tavakkoli, kaz. osaloo & mozaff.) to atraphaxis l. s. str., as well as the sisterhood of the clade (polygonum subsect. spinescentia plus atraphaxis) and genus bactria yurtseva & mavrodiev (a. ovczinnikovii (czukav.) yurtseva). however, due to the vague and extremely complicated morphology, the morphological cladistic treatment of atraphaxis and its closely related taxonomic entities has never been performed until the recent past (yurtseva & mavrodiev 2017; yurtseva et al. 2017). marodiev e.v. & yurtseva o.v., three-taxon statement analysis and intuitive taxonomy 3 recently, polygonum subsect. spinescentia has been accepted at the generic rank as persepolium o.v. yurtseva & e.v. mavrodiev (yurtseva & mavrodiev 2017; yurtseva et al. 2017). this genus comprises several narrow endemics of west and south iran (reviewed in tavakkoli et al. 2015 and yurtseva et al. 2016, 2017). by applying standard mp analysis and 3ta to the comprehensive 27 characters’ morphological dataset (table s1 (available as an electronic supplement and taken from yurtseva et al. 2017: 194–194, see their appendices 2 and 3)) of the widely distributed genus atraphaxis (incl. a. section ovczinnikovia), we have confirmed that the monophyletic persepolium is sister to the narrowly defined atraphaxis and the position of this taxon is strongly supported (fig. 1). observing table s1, it is easy to see that the characters 1, 2, 4, 8, 12, 16, 18, 19 and 26 provide evidence for the clade (persepolium) and/or are optimized as synapomorphies of this clade under the standard mp criterion (yurtseva et al. 2017: 179–181). but can we recognize the clade (persepolium), if all the supported conventional characters 1, 2, 4, 8, 12, 16, 18, 19 and 26 (table s1) are excluded from future analyses? in such cases the only intuitively clear, but verbally inexpressible “general image” (eidos) of persepolium, as well as the set of the combinations of non-unique diagnostic features, nested “around” this “image”, may be considered by somebody as a source of taxonomic evidence for this section. the conventional mp analysis (as well as a maximum likelihood and bayesian methods) is unable to recognize the clade (persepolium) if standard characters 1, 2, 4, 8, 12, 16, 18, 19 and 26 (table s1) were excluded from the analysis (fig. 1c). due to the lack of the conventional synapomorphic characters none of the six trees based on the reduced matrix contains the clade (persepolium). however under the same conditions, 3ta still found the clade (persepolium) with 100% confidence (fig. 1d). we would like to stress that neither of the three-taxon statements that support the clade (persepolium) are derived from the “evident” conventional characters (one, two, four, eight, 12, 16, 18, 19, and 26 in the table s1), because all these were excluded from the matrix before the three-taxon permutation had been performed. therefore, within the framework of the 3ta, neither of the “unique” characters is actually necessary for the recognition of the clade (persepolium). this clade exists, but no conventional characters (table s1) need define it. contrary to richter (2016) and others (summarized in richter 2016), this result is clearly arguing in favor of nelson’s concepts of either taxon or homology as a relationship (summarized in williams & ebach 2008 and nelson 2011): “if synapomorphies are understood as relationships rather than homologues, then homologies, synapomorphies, taxa, and relationships become equivalent invisibilities” (nelson 2011: 139). but what do “invisibilities” mean in this context? why not the impossibility of formalizing the vision verbally? the impossibility to “translate” the clear intuition of the general image of the plant into a list of standard characters? if the last interpretation of nelson’s view is correct, then the pure aesthetic as well as husserlian contexts (husserl 1913; seamon 1998; see also schuetz 1959 and uehlein 1992 among others) of 3ta are possible. numerous philosophical contexts, however, are still relevant to discuss the 3ta (e.g., mavrodiev 2016). in this manner, the characters do not actually make the genus, but rather the genus gives the characters, which follow the distinct image (eidos) of the genus, not subject to verbal description (kuzin 1992). http://www.europeanjournaloftaxonomy.eu/index.php/ejt/rt/suppfiles/502/0 european journal of taxonomy 377: 1–7 (2017) 4 marodiev e.v. & yurtseva o.v., three-taxon statement analysis and intuitive taxonomy 5 fig. 1. a. strict consensus of 61 most parsimonious phylogenetic trees; tree length = 90 steps; ci = 0.6000; ri = 0.8302, recovered from a standard cladistic analysis (mp) (fitch parsimony) of the complete conventional morphological matrix of atraphaxis s. l. (table s1). all 27 unordered (non-additive) characters are parsimony informative. b. strict consensus of two nested most parsimonious hierarchies of patterns; length = 8328 steps; ci = 0.8521; ri = 0.8264), recovered from a mp analysis of the three-taxon statement representation of the complete conventional 27 characters’ morphological matrix of atraphaxis s. l. (table s1). the number of 3tss (characters) is equal to 7096, all are parsimony-informative. c. strict consensus of six most parsimonious phylogenetic trees; tree length = 56 steps; ci = 0.6071; ri = 0.8370, recovered from a mp analysis of the reduced conventional 18 characters’ morphological matrix of atraphaxis s. l. (table s1) with the characters one, two, four, eight, 12, 16, 18, 19, and 26 excluded. d. strict consensus of two nested, most parsimonious hierarchies of patterns; length = 5844 steps; ci = 0.8665; ri = 0.8460), recovered from a mp analysis of the three-taxon statement representation of the reduced conventional 18 characters’ morphological matrix of atraphaxis s. l. (table s1) with characters 1, 2, 4, 8, 12, 16, 18, 19 and 26 excluded. the number of 3tss (characters) is equal to 5064, all are parsimony-informative. all mp analyses as in paup* 4.0a150 (swofford 2002) were conducted using either conventional matrices or taxodium’s output 3ts nexus files with a heuristic search of 1000 random addition replicates (saving no more than 100 trees per replicate), and the tbr branch swapping/multrees option into effect. branches with a minimum length of zero were collapsed. the three-taxon statement analysis (3ta) of the unordered morphological matrix was established after it three-taxon (3ts) williams-siebert (ws) representation (williams & siebert 2000) using taxodium v. 1.2 (mavrodiev & madorsky 2012). the 3ts permutations were performed with the following command: taxodium input_file_name. csv –ium –ob –og –nex the value of the operational outgroup was fixed as a value of bactria lazkovii. all 3tss were weighted uniformly and treated as ‘‘ordered’’ (wagner parsimony). the bootstrap resampling of both conventional and 3ts matrices have been performed as described in mavrodiev & madorsky (2012). the diagnostic traits are optimized using mesquite (maddison & maddison 2011). acknowledgments dr. david m. williams (the natural history museum, london, uk) and prof. malte c. ebach (university of new south wales & the sydney’s australian museum, au) are greatly acknowledged for their helpful comments and related discussion. the morphological study (ovy) was carried out in accordance to government order for the lomonosov moscow state university (project no. аааа-а16-116021660045-2). references crespo m.b., martinez-azorin m. & mavrodiev e.v. 2015. can a rainbow consist of a single colour? a new comprehensive generic arrangement of the ‘iris sensu latissimo’ clade (iridaceae), congruent with morphology and molecular data. phytotaxa 232: 1–78. https://doi.org/10.11646/phytotaxa.232.1.1 husserl e. 1913. ideas pertaining to a pure phenomenology and to a phenomenological philosophy – first book: general introduction to a pure phenomenology. martinus nijhoff publishers, boston. 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(eds) conference on taxonomy and evolutionary plant morphology dedicated to the 85th anniversary of v.n.tikhomirov, january 31–february 3, 2017. m.v. lomonossov moscow state university, moscow, russia. yurtseva o.v., kuznetsova o.i., mavrodieva m.e. & mavrodiev e.v. 2016. what is atraphaxis l. (polygonaceae, polygoneae): cryptic taxa and resolved taxonomic complexity instead of the formal lumping and the lack of morphological synapomorphies. peerj 4. e1977 https://doi.org/10.7717/peerj.1977 yurtseva o.v., severova e.e. & mavrodiev e.v. 2017. persepolium (polygoneae): a new genus in polygonaceae based on conventional maximum parsimony and three-taxon statement analyses of a comprehensive morphological dataset. phytotaxa 314 (2): 151–194. https://doi.org/10.11646/phytotaxa.314.2.1 manuscript received: 24 february 2017 manuscript accepted: 16 may 2017 published on: 11 december 2017 topic editor: koen martens desk editor: jeroen venderickx printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain. https://doi.org/10.7717/peerj.1977 https://doi.org/10.11646/phytotaxa.314.2.1 european journal of taxonomy 272: 1–13 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2017.272 www.europeanjournaloftaxonomy.eu 2017 · ramage t. et al. this work is licensed under a creative commons attribution 3.0 license. d n a l i b r a r y o f l i f e , r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:8ec5f9fb-5a94-4fa6-8274-86326e8404b3 1 a dna barcode-based survey of terrestrial arthropods in the society islands of french polynesia: host diversity within the symbiocode project thibault ramage 1, patricia martins-simoes 2, gladys mialdea 3, roland allemand 4,†, anne duplouy 5, pascal rousse 6, neil davies 7, george k. roderick 8 & sylvain charlat 9,* 1 9 quartier de la glacière, 29900 concarneau, france. 2,3,4,9 laboratoire de biométrie & biologie evolutive, cnrs – université lyon 1, bat. mendel, 43 boulevard du 11 november, 69622 villeurbanne, france. 2 ciri, international center for infectiology research, lyon, france. 5 university of helsinki, metapopulation research centre, department of biosciences, p.o. box 65, viikinkaari 1, 00014 helsinki, finland. 6 38 rue des primevère, 35160 le verger, france. 7,8 richard b. gump south pacific research station, university of california berkeley, bp 244 maharepa, 98728 moorea, french polynesia. 8 environmental science, policy and management, university of california, berkeley, california 94720, usa * corresponding author: sylvain.charlat@univ-lyon1.fr 1 email: thibault.ramage@hotmail.fr 2 email: patmsimoes@gmail.com 3 email: gmialdea@gmail.com 5 email: duplouyanne@yahoo.fr 6 email: rousse.pascal@wanadoo.fr 7 email: neiltahiti@gmail.com 8 email: roderick@berkeley.edu † deceased 1 urn:lsid:zoobank.org:author:8de31f66-13bf-4516-a205-60f2ea39e3dd 2 urn:lsid:zoobank.org:author:bb4451a0-44ac-46fd-a60d-f019aa87d62e 3 urn:lsid:zoobank.org:author:e18e3969-c359-4474-aeac-0d53027672c7 4 urn:lsid:zoobank.org:author:e1e1055d-d791-4882-ab67-ef29f1392f6a 5 urn:lsid:zoobank.org:author:af956717-d0e6-44a3-8f4b-8e312993edee 6 urn:lsid:zoobank.org:author:b06c2640-700a-429b-aa2f-1be09251c845 7 urn:lsid:zoobank.org:author:fb092614-b8dc-40f6-bad5-454a0c880799 8 urn:lsid:zoobank.org:author:16ffe533-cecc-44be-ae1a-3e4b543bf48a 9 urn:lsid:zoobank.org:author:a9ae69c2-039d-47fd-9dd2-b34c4363cb71 http://dx.doi.org/10.5852/ejt.2017.272 www.europeanjournaloftaxonomy.eu https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:8ec5f9fb-5a94-4fa6-8274-86326e8404b3 mailto:sylvain.charlat%40univ-lyon1.fr?subject= mailto:thibault.ramage%40hotmail.fr?subject= mailto:patmsimoes%40gmail.com?subject= mailto:gmialdea%40gmail.com?subject= mailto:duplouyanne%40yahoo.fr?subject= mailto:rousse.pascal%40wanadoo.fr?subject= mailto:neiltahiti%40gmail.com?subject= mailto:roderick%40berkeley.edu?subject= http://zoobank.org/urn:lsid:zoobank.org:author:8de31f66-13bf-4516-a205-60f2ea39e3dd http://zoobank.org/urn:lsid:zoobank.org:author:bb4451a0-44ac-46fd-a60d-f019aa87d62e http://zoobank.org/urn:lsid:zoobank.org:author:e18e3969-c359-4474-aeac-0d53027672c7 http://zoobank.org/urn:lsid:zoobank.org:author:e1e1055d-d791-4882-ab67-ef29f1392f6a http://zoobank.org/urn:lsid:zoobank.org:author:af956717-d0e6-44a3-8f4b-8e312993edee http://zoobank.org/urn:lsid:zoobank.org:author:b06c2640-700a-429b-aa2f-1be09251c845 http://zoobank.org/urn:lsid:zoobank.org:author:fb092614-b8dc-40f6-bad5-454a0c880799 http://zoobank.org/urn:lsid:zoobank.org:author:16ffe533-cecc-44be-ae1a-3e4b543bf48a http://zoobank.org/urn:lsid:zoobank.org:author:a9ae69c2-039d-47fd-9dd2-b34c4363cb71 european journal of taxonomy 272: 1–13 (2017) 2 abstract. we report here on the taxonomic and molecular diversity of 10 929 terrestrial arthropod specimens, collected on four islands of the society archipelago, french polynesia. the survey was part of the ‘symbiocode project’ that aims to establish the society islands as a natural laboratory in which to investigate the flux of bacterial symbionts (e.g., wolbachia) and other genetic material among branches of the arthropod tree. the sample includes an estimated 1127 species, of which 1098 included at least one dna-barcoded specimen and 29 were identified to species level using morphological traits only. species counts based on molecular data emphasize that some groups have been understudied in this region and deserve more focused taxonomic effort, notably diptera, lepidoptera and hymenoptera. some taxa that were also subjected to morphological scrutiny reveal a consistent match between dna and morphology-based species boundaries in 90% of the cases, with a larger than expected genetic diversity in the remaining 10%. many species from this sample are new to this region or are undescribed. some are under description, but many await inspection by motivated experts, who can use the online images or request access to ethanol-stored specimens. keywords. arthropods, dna barcoding, french polynesia, moorea biocode, symbiocode. ramage t., martins-simoes p., mialdea g., allemand r., duplouy a., rousse p., davies n., roderick g.k. & charlat s. 2017. a dna barcode-based survey of terrestrial arthropods in the society islands of french polynesia: host diversity within the symbiocode project. european journal of taxonomy 272: 1–13. http://dx.doi.org/10.5852/ ejt.2017.272 introduction in this paper, we report on the diversity of a large and non-taxonomically focused sample of terrestrial arthropods, collected on four islands of the society archipelago (french polynesia) from 2005 to 2007. this sample was obtained as part of the symbiocode initiative (coordinated by sc) in collaboration with the moorea biocode project (http://biocode.berkeley.edu) (check 2006). although primarily focused on symbiotic bacteria and their flux across host species (bailly-bechet et al. in press), this dataset also offers an opportunity to complement biodiversity records from this region, which is the subject of the present paper. this study focused on four of the five main islands of the society, situated along a 200 km northwest/ southeast axis, corresponding to the movement of the pacific plate over a unique hot spot during the last three million years (guillou et al. 2005). typical of young, small and isolated volcanic islands, their fauna is generally characterised by a low diversity of species and a high level of endemism, with an increasing proportion of introduced and invasive species (whittaker & fernández-palacios 2009). much of the south pacific’s terrestrial biota originates from the west (australasia), colonizing the oceanic islands via stepping-stone dispersal (miller 1996; gillespie & roderick 2002; gressitt 1956). fairmaire (1849, 1850) was the first to focus on the insect fauna of french polynesia. this fauna was again intensively studied between 1926 and 1940 as a result of the collections of saint-george and the pacific entomological survey (reviewed in ramage in press). among the 2972 valid arthropod species names reported from french polynesia, 61% are considered endemic, but the level of introduced species is also high, representing 10% of the overall species count (ramage in press). in the present study, we report on the occurrence of 1127 arthropod species (supplementary file, sheet s1): 228 of them were assigned to previously described species, and of the rest, 105 to genus, 567 to family and 227 to order. dna barcodes were the main source of taxonomic information, but some taxa (listed in table 1) were also thoroughly characterised based on morphology. for groups that were analysed by taxonomic experts, combined genetic and morphological evidence suggests the occurrence http://dx.doi.org/10.5852/ejt.2017.272 http://dx.doi.org/10.5852/ejt.2017.272 http://biocode.berkeley.edu http://www.europeanjournaloftaxonomy.eu/index.php/ejt/rt/suppfilemetadata/396/0/40 ramage t. et al., symbiocode: arthropod diversity in the society islands 3 of many species new to this region or new to science. some of these species are in the process of formal description by various colleagues (dierkens & ramage 2016; rousse et al. in prep.), but many more await inspection by motivated experts, on the basis of online photos or ethanol-stored material, available upon request. material and methods sampling arthropods were collected in the society islands of tahiti, moorea, huahine and raiatea, with most effort, for logistical reasons, on the island of moorea and to a lesser extent tahiti. collecting took place from 2005 to 2007 using sweep nets, malaise traps (only in moorea and tahiti) and light traps, or by hand, without focusing on any particular habitat or taxonomic group. details of the 317 collecting events, defined as a unique combination of location and date, are given in the supplementary file, sheet s2. raw material from the field was stored at ambient temperature in 95% ethanol in 50 ml centrifuge tubes for further processing. photographs and morpho-species assignments specimens showing no apparent morphological differences to a non-expert eye were attributed the same morpho-species numbers. up to ten specimens per morpho-species were individually photographed (under binocular microscope or with a reflex camera depending on their size) and stored in 95% ethanol at -20°c before further processing. in total, 10 929 specimens were processed in this way (listed in the supplementary file, sheet s3). all photographs can be accessed through the bold database (http:// www.boldsystems.org) (syc project, ds-symc dataset, http://dx.doi.org/10.5883/ds-symc) and the table 1. taxa that have been identified on the basis of morphology. for these groups, names of previously described species were used and new species are being described. taxon expert araneae michaël dierkens blattodea, phasmida, psocodea, scolopendromorpha, mantodea thibault ramage coleoptera (anthribidae, buprestidae, cerambycidae, chrysomelidae, coccinellidae, curculionidae, dryophthoridae, elateridae, endomychidae, lucanidae, monotomidae, mycetophagidae, nitidulidae, oedemeridae, scarabaeidae, silvanidae) thibault ramage coleoptera (carabidae) alexander anichtchenko diptera (neriidae, platystomatidae, stratiomyidae, syrphidae, tephritidae) thibault ramage hemiptera (aphrophoridae, aradidae, cicadellidae, coreidae, geocoridae, gerridae, miridae, oxycarenidae, pentatomidae, plataspidae, scutelleridae, tingidae) thibault ramage hemiptera (cixiidae) hannelore hoch hemiptera (delphacidae) manfred asche hymenoptera (apidae, chrysididae, crabronidae, evaniidae, formicidae, sphecidae, vespidae) thibault ramage hymenoptera (ichneumonidae) pascal rousse lepidoptera (lycaenidae, nymphalidae, sphingidae) thibault ramage odonata daniel grand orthoptera (gryllidae, mogoplistidae) thibault ramage http://www.europeanjournaloftaxonomy.eu/index.php/ejt/rt/suppfilemetadata/396/0/40 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/rt/suppfilemetadata/396/0/40 http://biocode.berkeley.edu http://biocode.berkeley.edu http://dx.doi.org/10.5883/ds-symc european journal of taxonomy 272: 1–13 (2017) 4 moorea biocode database (http://biocode.berkeley.edu). additional specimens from overly represented morpho-species were mass stored in 95% ethanol at -20°c for potential future analysis (additional specimens, see the supplementary file, sheet s1). dna extraction in order to maximize the phylogenetic and geographic diversity of the study, we selected specimens for dna extraction and further molecular analysis using the following criteria when possible: up to three specimens per morpho-species per island and from different locations. we thus selected 4837 specimens, from which dna was extracted using nucleospin 96 tissue kits (macherey-nagel gmbh & co. kg, düren, germany), according to the manufacturer’s instructions, with the following modifications: (a) the pre-lysis step consisted of incubation with proteinase k overnight and (b) the elution step included two sub-steps (each with 50 μl of water). if specimens were smaller than 5 mm long, the whole body was used for dna extraction; otherwise, we used tissue from the mid-lower abdomen, including the gonads but leaving the genitalia intact for potential subsequent taxonomic work. we stored 20 µl of each genomic extract at -20°c for further analysis; the remaining 80 µl were placed at -80°c for long-term storage. dna barcoding a 643–664 bp (most commonly 658 bp) region of the mitochondrial gene cytochrome oxidase 1 (co1) was amplified with standard lco1490 and hco2198 primers (folmer et al. 1994). dna amplification (pcr) was performed in a total volume of 30 µl with 1.5 mm of mgcl2, 2 mm of all four dntps, 0.2 µm of each primer, 0.02 units/µl of eurotaq r dna polymerase (eurobio, les ulis, france) and 2 µl of template using the following temperature profile: initial denaturation at 95°c for 120 seconds (s); 35 cycles of 94°c for 30 s, 47°c for 30 s and 72°c for 90 s; and a final extension at 72°c for 600 s. all reactions took place in a tetrad r thermocycler (bio-rad, hercules, ca, usa). pcr products were purified and sanger-sequenced on an abi 3730 using both the forward and reverse pcr primers. trace files were imported in geneious v. 5.4.0 (biomatters) (kearse et al. 2012). the first 45 bp of the 5’ end of each read were trimmed, as well as further low-quality end regions with error probability larger than 0.1%. heteroplasmic positions were identified based on peak similarity (more than 50% overlap between two conflicting peaks at a given position in one read); ambiguous base calls were assigned to heteroplasmic positions using the iupac code. forward and reverse reads were then assembled and a consensus sequence integrating phred quality scores was produced. in the consensus sequence, the phred score used for a given position was the phred score from a single read if only one read covered this position, or the sum of the forward and reverse scores in case of agreement between the forward and reverse reads. in case of conflict between the reads at a given position, the base used and the associated phred score in the consensus were those of the highest quality read at the position. an ambiguous base call (n) was assigned to poorly supported positions (phred scores below 20) in the consensus. high quality reads (i.e., those carrying more than 80% nucleotides with phred scores larger than 40), but not assembled to their reverse complement because of failure of one reaction, were also recovered. consensus sequences and recovered single reads were then all aligned to an arbitrarily chosen reference co1 sequence (genbank kf226475, from the butterfly hypolimnas bolina) in order to unambiguously identify and trim primer regions. reads longer than 200 bp were then selected and the longest open reading frame was determined for all reads, yielding only 12 sequences with stop codons, which were discarded. in addition, 18 sequences were identified as experimental contaminants (precisely matching another sequence in the dataset) and thus were excluded. all such contaminations occurred locally in dna extraction or pcr plates: the http://dx.doi.org/10.5883/ds-symc http://www.europeanjournaloftaxonomy.eu/index.php/ejt/rt/suppfilemetadata/396/0/40 ramage t. et al., symbiocode: arthropod diversity in the society islands 5 contaminant well was never more than two wells away from the contaminated well. notably, specimens were randomly distributed across 54 plates, so that specimens from the same morpho-species were not clustered, allowing cross contaminations to be detected when it occurred. we also excluded five natural contaminants, harbouring an insect parasitoid instead of host barcode. sequences were aligned in geneious using the predicted protein sequences. clustering of sequences within otus of 3627 co1 sequences, 3180 passed the bold database quality filters and were clustered using the refined single linkage (resl) algorithm into 1026 species-level groups or otus (operational taxonomic units), each identified with a unique barcode index number (bin) (ratnasingham & hebert 2013). in brief, resl employs a single linkage clustering procedure (producing clusters including all sequences connected by a genetic distance below a certain threshold) followed by a refinement step in which clusters including at least three sequences can be split if they show significant discontinuity in genetic distances. of the 447 sequences that did not pass the bold quality filters, 314 were assigned to one of the bins identified by resl. the remaining 133 specimens were clustered in groups of sequences diverging by no more than 3% substitutions per site, resulting in an additional 84 otus also included as they represent significant information, although they should be considered more cautiously. the procedure used for each specimen is indicated in the supplementary file, sheet s3. taxonomic assignments with the exception of spiders, which were classified to the lowest possible taxonomic level based on morphology prior to dna extraction (dierkens & charlat 2009), all specimens were only assigned the taxonomic rank of order or class upon morpho-species assignments. for barcoded specimens, the consistency of this assignment with the 1st blast hit was verified (using blastn with default options against the ncbi nr database). closer inspections of photographs or specimens allowed correction of any remaining inconsistencies. such errors were either due to contaminations (as noted above) or, more often, to an incorrect initial assignment. a more accurate dna barcode-based taxonomic assignment was then proposed based on the quality of the best blast hit, with the following categories: (1) species level assignment for at least 98% sequence identity over at least 80% of the barcode; (2) genus level for 95% sequence identity and 80% overlap; (3) family level for 90% sequence identity and 80% overlap. in combination with blast, we developed a phylogenetic approach to propose a taxonomic assignment based on dna sequence data. we developed r scripts to extract one representative co1 sequence from each arthropod genus from genbank (resulting in 20 350 sequences). for each arthropod order, we added these sequences to our own alignment using profile alignment in mafft (katoh & standley 2013) and constructed a tree for each order using fasttree (price et al. 2010). poorly supported nodes (less than 80% alrt support) (anisimova & gascuel 2006) were collapsed in archaeopteryx (han & zmasek 2009). we then inspected these trees in dendroscope (huson & scornavacca 2012), with colour coding of tips based on family names, to refine and verify blast-based assignments. an improved classification of our specimen was then proposed using the “strict” criteria defined by wilson et al. (2011), which results in a very low level (2%) of false positives. in brief, a taxon name is assigned to a specimen if (1) its position within a clade is strongly supported (at least 80% alrt support) and (2) the clade where it is placed is taxonomically homogeneous. for groups listed in table 1, we found no inconsistencies between morphology and barcode-based assignments. for the other groups, most taxonomic assignments relied on barcodes only, as detailed in the supplementary file, sheet s1. some morpho-species that had not produced a good quality barcode were also identified to the species level when this was feasible without ambiguity. http://www.europeanjournaloftaxonomy.eu/index.php/ejt/rt/suppfilemetadata/396/0/40 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/rt/suppfilemetadata/396/0/40 european journal of taxonomy 272: 1–13 (2017) 6 results characteristics of the sample in total, the symbiocode sample includes 10 929 arthropod specimens that were assigned to morphospecies, individually photographed, and deposited in the bold database (http://www.boldsystems. org) (syc project, ds-symc dataset, http://dx.doi.org/10.5883/ds-symc) and the moorea biocode databases (http://biocode.berkeley.edu). the supplementary file (sheet s3) provides a complete list of the specimens with a unique url for each specimen. more than half (52%) of the specimens were collected at low elevations or near sea level (below 100 m elevation, fig. 1) in potentially disturbed habitats, which are more likely to harbour non-native species than locations at higher elevations. malaise traps were used only on the islands of moorea and tahiti, and mostly in remote locations at high altitude. fig. 1. distributions of the elevations at which specimens were collected on each island. figure'1.'! tahiti altitude (m) f re qu en cy 0 500 1000 1500 0 50 0 15 00 25 00 moorea altitude (m) f re qu en cy 0 500 1000 1500 0 50 0 15 00 25 00 huahine altitude (m) f re qu en cy 0 500 1000 1500 0 50 0 15 00 25 00 raiatea altitude (m) f re qu en cy 0 500 1000 1500 0 50 0 15 00 25 00 n .'o f's pe ci m en s' n .'o f's pe ci m en s' n .'o f's pe ci m en s' n .'o f's pe ci m en s' http://www.boldsystems.org http://www.boldsystems.org http://dx.doi.org/10.5883/ds-symc http://biocode.berkeley.edu http://www.europeanjournaloftaxonomy.eu/index.php/ejt/rt/suppfilemetadata/396/0/40 ramage t. et al., symbiocode: arthropod diversity in the society islands 7 hence, island, collecting method, and elevation are not independent (fig. 1). considering the four islands together, 30% of the specimens from low elevation were collected with malaise traps, compared to 90% of those from high altitude. this sampling effort should be kept in mind when quantifying diversity, because malaise traps tend to collect relatively more diptera and hymenoptera compared to other orders, especially coleoptera (gressitt & gressitt 1962; lamarre et al. 2012). among the 317 collecting events, 80% took place at low elevation and contained one or a few samples (thanks, notably, to the contribution of middle-school students from the paopao school of moorea). in contrast, high elevation collecting events were generally more intense and contributed more specimens per event. barcoding success rate dna was extracted from 4837 specimens. as detailed in table 2 (by order), the supplementary file, sheet s2 (by species) and the supplementary file, sheet s3 (by specimen), amplification and sequencing of the co1 locus was successful in 3627 specimens, that is, 75% of the genomic extracts, but the success rate varied widely among arthropod orders (table 2). phylogeny explained a large part of this variation, with some clades being clearly less efficiently sequenced than others, possibly because of divergence in the primer target regions. most of the barcode data were of high quality (92% of the co1 sequences were longer than 400 bp and included fewer than 1% ambiguous positions); 8% were of lower quality (between 200 and 400 bp long, including up to 20 ambiguous positions) but carried enough information to be maintained in the analysis. sequences are available in bold, and also in genbank (bankit1909431: kx051578–kx055204). table 2. rate of dna barcoding success in the main taxa under study. groups with fewer than 10 specimens tested are not included. the barcoding success rate is defined as the proportion of specimens tested that yielded a quality dna barcode (see material and methods for quality criteria). taxa barcoding success rate n specimens n species amphipoda 61.50% 13 4 araneae 59.50% 412 50 blattodea 68.20% 44 11 coleoptera 62.70% 413 119 collembola 26.70% 60 8 dermaptera 5.90% 17 1 diptera 90.80% 1109 305 hemiptera 65.70% 696 133 hymenoptera 74.60% 689 172 isopoda 5.00% 20 1 isoptera 30.80% 13 1 ixodida 13.00% 23 1 lepidoptera 91.50% 885 223 neuroptera 100.00% 18 5 odonata 63.60% 44 8 orthoptera 65.10% 172 16 polydesmida 9.10% 11 1 psocodea 70.20% 104 24 thysanoptera 86.40% 22 5 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/rt/suppfilemetadata/396/0/40 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/rt/suppfilemetadata/396/0/40 european journal of taxonomy 272: 1–13 (2017) 8 biodiversity the co1 sequences clustered into 1693 different haplotypes and into 1110 species-like groups or otus defined on the sole basis of molecular data. of the 1110 otus, 1026 (92%) were defined using the resl algorithm implemented in bold and thus received a unique identifier in this database (bin); 642 of the bins were new to the bold database. the remaining 84 otus comprised sequences that did not pass the bold quality filters, but were clustered in groups of sequences diverging by no more than 3% substitutions per site. some specimens, from which dna was not extracted, or for which co1 sequencing failed, were nevertheless assigned on the basis of morphology (when possible to do so with confidence) to otus from this study (n = 4295 specimens) or to a named species not represented in the other otus (n = 411 specimens). in summary, the biodiversity reported here is based on 8248 specimens, representing an estimated 1127 species designated as follows: (i) 199 named species (2924 specimens) including at least one sequenced specimen, (ii) 899 otus (5177 specimens), including at least one sequenced specimen, not named at the species level but taken as a proxy for species identity, and (iii) 29 named species (147 specimens) without dna sequence data. each species used in these counts was assigned a species id, as listed in the supplementary file, sheet s1. the 1127 species are distributed among major arthropod groups as shown in table 3. most of the biodiversity collected during this survey comprises the five major insect orders (hemiptera, coleoptera, diptera, hymenoptera, lepidoptera). importantly, the species richness in the three orders most frequently represented in the sample (diptera, lepidoptera and hymenoptera) is higher than documented in all earlier reports combined for the society islands (ramage in press). for hymenoptera, the diversity reported here is greater than known previously for the entirety of french polynesia, including all of its five archipelagos. considering groups that have been well characterised in our study (listed in table 1), we observe that only 60% of the species found in our sample have previously been reported. our sample includes 107 species from which we can assess the level of correlation between otus and morphology-based species identity, because they were distinguished on the basis of morphology and include at least two barcoded specimens (supplementary file, sheet s1). no specimens from the same otu received different species names based on morphology, suggesting the dna sequence-based approach was at least as efficient as morphology in distinguishing species. in contrast, 11 out of 107 proposed species included more than one bin, resulting in a 10% error rate, which is in the lower range of what has been found in other studies (meier et al. 2006; ratnasingham & hebert 2013) (see the supplementary file, sheet s4 for a list of these species and summary statistics on their mitochondrial diversity). new records and other notable occurrences in this section we consider the 227 species that were named and highlight notable occurrences. although coleoptera has been the most intensively collected order in this region, we note new records (supplementary file, sheet s5). for hymenoptera, the present sample generally shows equal or higher diversity than all previous reports combined. our sample also contributes two new family records for french polynesia, the sphecidae and the chrysididae, as reported previously (ramage et al. 2015; ramage & kimsey 2015). among species from the aculeata infra-order identified to genus or species, new records for french polynesia include two species of apidae (genera xylocopa latreille, 1802 and braunsapis michener, 1969), three species of crabronidae (genera liris fabricius, 1804 and polemistus saussure, 1892) and two species of vespidae (genera pachodynerus saussure, 1875 and vespula thomson, 1869). these collections will be the subject of upcoming, more specific publications (ramage in press). the examination of the family ichneumonidae led to a similar conclusion: among the 19 species belonging to this family, 14 were identified to genus or species level and five were already known in french polynesia, but four are new records for the country and five represent new species (rousse et al. in prep.). http://www.europeanjournaloftaxonomy.eu/index.php/ejt/rt/suppfilemetadata/396/0/40 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/rt/suppfilemetadata/396/0/40 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/rt/suppfilemetadata/396/0/40 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/rt/suppfilemetadata/396/0/40 ramage t. et al., symbiocode: arthropod diversity in the society islands 9 the other insect orders, which to date have been less thoroughly studied taxonomically in this project, are also likely to harbour many species previously undocumeted in french polynesia. for example, the family stratiomyidae (diptera) was only known for two species in french polynesia, hermetia illucens (linnaeus, 1758) and chromatopoda annulipes (walker, 1849), while our data show that four other species are likely present. within the order hemiptera, new records for french polynesia are likely in the families plataspidae (genus coptosoma laporte de castelnau, 1832), scutelleridae (genus calliphara germar, 1839; ramage & gourvès 2016), tingidae (genus corythucha stål, 1873), oxycarenidae (genus oxycarenus fieber, 1837) and membracidae (genus spissistilus caldwell, 1949). among the 227 named species in our sample, 88 are indigenous or with unknown status, 122 are introduced (including new records), and 17 are endemic (supplementary file, sheet s1). discussion the symbiocode collection, described in the present study, provides an overview of the terrestrial arthropod diversity of the society archipelago. approximately one fifth of the species present in this sample have been identified at the species level, extending the known range of some species to french polynesia and enriching knowledge of other groups of taxa. table 3. species counts in this study and in previous records from the society archipelago (based on ramage submitted). taxon n species earlier surveys (society) n species earlier surveys (french polynesia) n species symbiocode named sp. symbiocode: previous record / new record / new sp. blattodea 12 28 14 5 / 0 / 0 chelicerata 238 337 77 24 / 11 / 0 coleoptera 558 772 119 42 / 3 / 0 collembola 10 33 8 0 / 0 / 0 crustacea 37 78 6 1 / 0 / 0 dermaptera 8 9 1 0 / 0 / 0 diptera 201 328 307 20 / 8 / 0 embioptera 1 1 0 0 / 0 / 0 hemiptera 224 401 135 17 / 0 / 0 hymenoptera 126 172 178 32 / 5 / 5 lepidoptera 151 494 224 30 / 4 / 0 mantodea 1 1 1 1 / 0 / 0 myriapoda 15 19 5 2 / 0 / 0 neuroptera 11 14 5 0 / 0 / 0 odonata 12 19 8 7 / 0 / 0 orthoptera 17 32 16 2 / 0 / 0 phasmida 2 2 1 1 / 0 / 0 psocodea 42 56 25 4 / 0 / 0 siphonaptera 2 3 1 1 / 0 / 0 thysanoptera 43 59 5 1 / 0 / 0 zygentoma 1 5 0 0 / 0 / 0 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/rt/suppfilemetadata/396/0/40 european journal of taxonomy 272: 1–13 (2017) 10 although not exhaustive, and constrained by the collecting and taxonomic methods used, the study reveals that many arthropod groups are far more diverse than suggested by previous reports. focusing on groups that have been morphologically well characterised in our study, we observe that only 60% of the species collected have previously been reported. in other words, many of the collected species are new to this region or to science (dierkens & ramage 2016; rousse et al. in prep.), emphasising the scale of the challenge to characterize tropical arthropod biodiversity, even for small and isolated islands. the correlation between molecular-based otus and morphology-based species identity was assessed in 107 species that were distinguished on the basis of morphology and included at least two barcoded specimens. no specimens from the same otu received different species names based on morphology, but 10% of the proposed species based on morphology included more than one otu. three explanations can be proposed to explain such high sequence diversity within these groups. some species may have very large effective population sizes and/or high natural levels of genetic diversity (for example, invasive species introduced from multiple distant sources). alternatively, some proposed species names may correspond to more than one true species that may or may not be distinguishable morphologically following deeper scrutiny. finally, some species may have been subject to mitochondrial introgression, possibly fuelled by cytoplasmic symbionts (hurst & jiggins 2005) and thus carry mitochondrial dna lineages from distinct species. further work is needed, including more thorough morphological inspection and sequencing of nuclear dna markers, to determine which explanations hold. while this study confirms that dna barcoding offers a powerful means to quantify the diversity of complex samples, it also illustrates that molecular data are more valuable when associated with that from morphological characters. we thus hope this report will stimulate further synergistic projects to characterize the biodiversity of this region and encourage experts to complement our analysis, using the on-line photographs and metadata, or the many remaining specimens, available upon request. acknowledgments we are grateful to the cnrs for the atip grant “symbiocode” to sc, the genoscope for dna sequencing through “bibliothèque du vivant” and “speed id” projects, and the moorea biocode team and the gordon and betty moore foundation for financial and logistic support, including access to malaise traps. we also thank julien varaldi for handy r scripts; alexander anichtchenko, daniel grand, hannelore hoch and manfred asche for invaluable taxonomic expertise; and finally marjorie saillan and the “entomology club” of the paopao school in moorea for their unique contribution to the collecting effort. this article is dedicated to roland allemand for his early encouragement, and more generally for his major contributions to the field of entomology. references anisimova m. & gascuel o. 2006. approximate likelihood-ratio test for branches: a fast, accurate, and powerful alternative. systematic biology 55: 539–552. bailly-bechet m., simoes p., szöllősi g., mialdea g., sagot m.-f. & charlat s. in press. how long does wolbachia remain on board? molecular biology and evolution. check e. 2006. treasure island: pinning down a model ecosystem. nature 439: 378–379. http://dx.doi. org/10.1038/439378a dierkens m. & charlat s. 2009. contribution à la connaissance des araignées des îles de la société (polynésie française). revue arachnologique 17: 63–81. dierkens m. & ramage t. 2016. deuxième contribution à la connaissance des araignées de polynésie française. bulletin mensuel de la société linnéenne de lyon 85: 134–172. http://dx.doi.org/10.1038/439378a http://dx.doi.org/10.1038/439378a ramage t. et al., symbiocode: arthropod diversity in the society islands 11 fairmaire l.m.h. 1849. essai sur les coléoptères de la polynésie. revue et magasin de zoologie 2: 277–291, 352–365, 410–422, 445–460, 504–516, 550–559. fairmaire l.m.h. 1850. essai sur les coléoptères de la polynésie. revue et magasin de zoologie 2: 50–64, 115–122, 181–185. folmer o., black m., hoeh w., lutz r. & vrijenhoek r. 1994. dna primers for amplification of mitochondrial cytochrome c oxidase subunit i from diverse metazoan invertebrates. molecular marine biology and biotechnology 3: 294–299. gillespie r.g. & roderick g.k. 2002. arthropods on islands: colonization, speciation, and conservation. annual review of entomology 47: 595–632. gressitt j.l. 1956. some distribtion patterns of pacific island faunae. systematic zoology 5: 11–47. gressitt j.l. & gressitt m.k. 1962. an improved malaise trap. pacific insects 4: 87–90. han m.v. & zmasek c.m. 2009. phyloxml: xml for evolutionary biology and comparative genomics. bmc bioinformatics 10: e356. http://dx.doi.org/10.1186/1471-2105-10-356 hurst g.d. & jiggins f.m. 2005. problems with mitochondrial dna as a marker in population, phylogeographic and phylogenetic studies: the effects of inherited symbionts. proceedings of the royal society b 272: 1525–1534. huson d.h. & scornavacca c. 2012. dendroscope 3: an interactive tool for rooted phylogenetic trees and networks. systematic biology 61: 1061–1067. http://dx.doi.org/10.1093/sysbio/sys062 katoh k. & standley d.m. 2013. mafft multiple sequence alignment software version 7: improvements in performance and usability. molecular biology and evolution 30: 772–780. http://dx.doi.org/10.1093/ molbev/mst010 kearse m., moir r., wilson a., stones-havas s., cheung m., sturrock s., buxton s., cooper a., markowitz s., duran c., thierer t., ashton b., meintjes p. & drummond a. 2012. geneious basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. bioinformatics 28: 1647–1649. http://dx.doi.org/10.1093/bioinformatics/bts199 lamarre g.p.a., molto q., fine p.v.a. & baraloto c. 2012. a comparison of two common flight interception traps to survey tropical arthropods. zookeys 216: 43–55. http://dx.doi.org/10.3897/ zookeys.216.3332 meier r., shiyang k., vaidya g. & ng p.k.l. 2006. dna barcoding and taxonomy in diptera: a tale of high intraspecific variability and low identification success. systematic biology 55: 715–728. http:// dx.doi.org/10.1080/10635150600969864 miller s. 1996. biogeography of pacific insects and other terrestrial invertebrates: a status report. in: keast a. & miller s. (eds) the origin and evolution of pacific island biotas, new guinea to eastern polynesia: patterns and processes: 463–475. academic publishers, amsterdam. price m.n., dehal p.s. & arkin a.p. 2010. fasttree 2 approximately maximum-likelihood trees for large alignments. plos one 5 (3): e9490. http://dx.doi.org/10.1371/journal.pone.0009490 ramage t. in press. checklist of the terrestrial and freshwater arthropods of french polynesia (chelicerata; myriapoda; crustacea; hexapoda). zoosystema. ramage t. & gourvès j. 2016. un nouveau scutelleridae pour la polynésie française, calliphara bifasciata white, 1839 (hemiptera). bulletin de la société entomologique de france, in press. ramage t. & kimsey l.s. 2015. the aculeata of french polynesia. iv. first record of chrysis angolensis (hymenoptera, chrysididae). bulletin de la société entomologique de france 120: 209–211. http://dx.doi.org/10.1186/1471-2105-10-356 http://dx.doi.org/10.1093/sysbio/sys062 http://dx.doi.org/10.1093/molbev/mst010 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evolution, and conservation. second edition. oxford university press, oxford. wilson j.j., rougerie r., schonfeld j., janzen d.h., hallwachs w., hajibabaei m., kitching i.j., haxaire j. & hebert p.d. 2011. when species matches are unavailable are dna barcodes correctly assigned to higher taxa? an assessment using sphingid moths. bmc ecology 11: e18. http://dx.doi. org/10.1186/1472-6785-11-18 manuscript received: 16 february 2016 manuscript accepted: 22 july 2016 published on: 7 february 2017 guest editors: line le gall, frédéric delsuc, stéphane hourdez, guillaume lecointre and jean-yves rasplus desk editor: danny eibye-jacobsen printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands. http://dx.doi.org/10.1371/journal.pone.0066213 http://dx.doi.org/10.1186/1472-6785-11-18 http://dx.doi.org/10.1186/1472-6785-11-18 ramage t. et al., symbiocode: arthropod diversity in the society islands 13 information on supplementary file sheet s1. species list. n_otu: number of otus, relevant and only provided for named species. nap: not applicable. md: missing data. n_specimens: number of specimens stored in individual tubes. additional_ specimens: indicates whether additional specimens, mass stored in ethanol, are available in addition to those indicated in the “n_specimens” column. dna_tissue: body part used for dna extraction. n_dna_ extract: number of specimens from which dna was extracted and co1 sequencing was attempted. biogeo_status: biogeographical status (p: previously reported taxa, either indigenous or of unknown status; e: endemic taxa, endemic to a single or several islands, or to a single or several archipelagos of french polynesia; i: introduced taxa). bin: list of barcode index numbers included in this species. otus: list of operational taxonomic units (using our internal otu index) included in this species. clustering_method: method used to create the otu(s) making a species, when relevant (resl for most species, 3%_threshold for a few species that did not pass the bold quality filter). morpho-species: a morpho-species id, given only for specimens that did not receive a species id (that is, specimens that were not barcoded, not assigned to a barcoded species based on morphological characters and not named at the species level). sheet s2. collecting events. md: missing data. sheet s3. specimens. md: missing data. symbiocode_id: unique identifiers for specimens, internal to this study (matches with bold and biocode ids are also given, as well as links to specimen pages, including photographs and all metadata). col_event: collecting event (see sheet s2 for details). specimen_ available: indicates whether the specimen is available upon request for further study. species_id: unique identifier for species, as listed in sheet s1. included_in_biodiversity_survey: indicates whether the specimen was included in the species count, which is the case for specimens belonging to barcoded species or named at the species level on the basis of morphological characters. bin: barcode index number as defined in the bold database. otu: our internal operational taxonomic unit number, corresponding to the bin when a sequence was assigned to a bin. clustering_method: “resl” if the sequence was directly assigned a bin number in bold, “resl_indirect” for lower quality sequences that were assigned to a bin or “3%_threshold” for lower quality sequences that were not assigned to a bin, but clustered in groups at a maximum of 3% divergence. sheet s4. mitochondrial diversity in 11 species containing more than one otu based on the resl algorithm. pi: mean of the raw genetic distances calculated among all individuals. max_dist: maximum raw genetic distance within this species. bins: list of barcode index numbers included in each species. additional relevant information for each species can be found in sheet s1. sheet s5. new species records for french polynesia. http://www.europeanjournaloftaxonomy.eu/index.php/ejt/rt/suppfilemetadata/396/0/40 european journal of taxonomy 271: 1–19 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2017.271 www.europeanjournaloftaxonomy.eu 2017 · ohler a. & nicolas v. this work is licensed under a creative commons attribution 3.0 license. d n a l i b r a r y o f l i f e , r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:20e56a57-baca-44f3-b310-5d5bc91de040 1 which frog’s legs do froggies eat? the use of dna barcoding for identification of deep frozen frog legs (dicroglossidae, amphibia) commercialized in france annemarie ohler 1,* & violaine nicolas 2 1,2 institut de systématique, évolution, biodiversité, isyeb_umr7205_cnrs, mnhn, upmc, ephe, muséum national d’histoire naturelle, sorbonne universités, paris, france. * corresponding author: ohler@mnhn.fr 2 email: vnicolas@mnhn.fr 1 urn:lsid:zoobank.org:author:fc72f206-744c-4ba9-a609-1c604ac1af6a 2 urn:lsid:zoobank.org:author:fa1b1fa8-cb99-4ce8-b87a-f9b329e5df07 abstract. several millions frogs captured in the wild in indonesia are sold for food yearly in french supermarkets, as deep frozen frog legs. they are commercialized as rana macrodon, but up to 15 lookalike species might also be concerned by this trade. from december 2012 to may 2013, we bought 209 specimens of deep frozen frog legs, and identified them through a barcoding approach based on the 16s gene. our results show that 206 out of the 209 specimens belong to fejervarya cancrivora, two to limnonectes macrodon and one to f. moodiei. thus only 0.96 % of the frogs were correctly identified. unless misclassification was intentional, it seems that indonesian frog leg exporters are not able to discriminate between the species. the quasi absence of l. macrodon in our samples might be an indication of its rarity, confirming that its natural populations are declining rapidly, in agreement with its “vulnerable” status according to the iucn red list. our results show that the genetic and morphological diversity of the frogs in trade is much higher than the genetic and morphological diversity measured so far by scientific studies. these results underline the need for large scale studies to assess the status of wild populations. key words. fejervarya cancrivora, limnonectes macrodon, frog leg trade, barcoding, identification, conservation, red list. ohler a. & nicolas v. 2017. what frog’s leg do froggies eat? the use of dna barcoding for identification of deep frozen frog legs (dicroglossidae, amphibia) commercialized in france. european journal of taxonomy 271: 1–19. http://dx.doi.org/10.5852/ejt.2017.271 introduction the international traffic of wild animals is considered an important threat to many animal species and is subject to international regulations. from 1998 to 2007 (10 years), 35 million animals and plants registered on the cites lists have been exported from south-east asian countries (nijman 2010). for http://dx.doi.org/10.5852/ejt.2017.271 www.europeanjournaloftaxonomy.eu https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:20e56a57-baca-44f3-b310-5d5bc91de040 mailto:ohler%40mnhn.fr?subject= mailto:vnicolas%40mnhn.fr?subject= http://zoobank.org/urn:lsid:zoobank.org:author:fc72f206-744c-4ba9-a609-1c604ac1af6a http://zoobank.org/urn:lsid:zoobank.org:author:fa1b1fa8-cb99-4ce8-b87a-f9b329e5df07 http://dx.doi.org/10.5852/ejt.2017.271 european journal of taxonomy 271: 1–19 (2017) 2 the same period, between 180 million and 1 billion specimens of frogs were collected every year in the wild in indonesia, and one eighth of these frogs were then exported to europe (kusrini 2005; kusrini & alfold 2006). for the ten-year period from 2000 to 2009, 46 400 tonnes of frog legs were imported into europe, which corresponds to approximately 928 million to 2.3 billion frogs (altherr et al. 2011). the main european countries that import frog legs from indonesia are france and belgium (kusrini & alfold 2006; altherr et al. 2011). the consumption of frogs is a tradition in french cuisine, causing the french to be called “froggies” by their neighbours, and in former days private people and restaurants would collect local frog specimens at certain periods of the year. however, in the 1970s deep freezing technology was developed, which allowed long term storage and large scale transport of frog legs, and large numbers of frogs were collected by commercial enterprises (dubois 1983; neveu 2004). this modified the conditions of traditional local sustainable collection and drove some local frog populations to extinction. a wildlife protection law was voted in 1979 to protect the french frog species (le serrec 1988). the market turned to tropical countries, mainly bangladesh, india and indonesia, but india and bangladesh quickly stopped frog exports: the main species concerned (hoplobatrachus tigerinus (daudin, 1802)) was put on cites lists and very protective laws for the local anuran species were enacted. thus, in 1987 indonesia became the main exporter of frogs and more than 80% of european frog leg imports now come from this country (altherr et al. 2011). from 1973 to 1987, 830 to 2659 tonnes of frog legs were imported to france from indonesia (le serrec 1988). this import was 5600 tonnes in 1992 (kusrini & alford 2006), 4600 tonnes per year in 2000– 2009 (altherr et al. 2011), and 2906 to 3275 tonnes per year in 2009–2011 (french customs 2012). frog legs from indonesia are sold in supermarkets, particularly in those specialised in deep frozen food, or in asian food markets. kusirini & alford (2006) showed that three species are predominantly collected in indonesia for consumption: the giant javan frog limnonectes macrodon (duméril & bibron, 1841), the crab-eating frog fejervarya cancrivora (gravenhorst, 1829) and the common grass frog fejervarya limnocharis (gravenhorst, 1829), the latter probably representing a mixture of f. limnocharis and f. iskandari veith, kosuch, ohler & dubois, 2001 (djong 2007). about fifteen look-alike species of large size might also be involved in collection for international trade (altherr et al. 2011; ohler unpubl. report), and they all belong to the genera fejervarya bolkay, 1915 and limnonectes fitzinger, 1843. some of these species are common in the wild (e.g., f. cancrivora) while others are uncommon and listed as vulnerable in the iucn red list (e.g., l. macrodon, see iskandar et al. 2004). nonetheless, the european regulation no. 2065/2001 (anonymous 2001) asks that the labels of fishery products show the common and scientific names on pre-packed, non-transformed fresh products in order to assure traceability of specimens. in france, deep frozen frog legs are usually sold in the supermarkets in plastic bags of 500 g or 1 kg and bear labels with a latin species name and their country of origin. monitoring and management of harvested populations require accurate species identification; yet, high error rates are likely (warkentin et al. 2009). frog species can often be distinguished on various morphological characters such as body proportions, foot morphology and coloration pattern. because exported frogs are skinned and the bodies cut off before packing, the characters allowing a morphological identification cannot be observed on deep frozen legs and thus identification is difficult (warkentin et al. 2009). in this paper we explore the use of dna barcoding as an identification tool to identify deep frozen frog legs commercialized in france. dna barcoding is a technique that uses a short dna sequence from a standard locus as a species identification tool (hebert et al. 2003). from december 2012 to may 2013, we bought 209 frogs in different supermarkets in france that were labeled as rana macrodon (a variant of the valid name limnonectes macrodon). these frogs were sold under three different brands but all come from the same importer. the samples were identified through a barcoding approach based on the 16s gene. the 16s gene was previously recognized as one of the ohler a. & nicolas v., dna barcoding for identification of frog legs 3 most effective genes for molecular amphibian species identification (vences et al. 2005; grosjean et al. 2015). moreover, 16s reference sequences were available in the genbank database for the 15 species potentially collected in indonesia for commercialization. a diminution in size of adult frogs due to overharvesting has been suggested (iskandar, cited in anonymous 2007). as snout-vent length and tibia length are highly correlated in frogs (emerson 1978; vidal-garcía et al. 2014), we used tibia length to obtain an estimate for the body size of commercialized frogs. material and methods biological samples, dna extractions, pcr conditions, dna sequencing in this study, a total of 209 biological samples of commercialized frogs were bought in different french supermarkets from december 2012 to may 2013 (table 1, appendix). all these samples were sold as frozen frog legs in bags of 500 or 1000 g. they came from indonesia and were sold as rana macrodon. all these samples are stored in the collections of the national museum of natural history, paris, france (appendix). dna extracts were prepared from muscle tissue using the nucleospin tissue core kit (macherey nagel). 16s amplification and sequencing were obtained using the primer pair 16sa-l (5’-cgcctgtttatcaaaaacat-3’) and 16sb-h (5’-ccggtctgaactcagatcacgt-3’) (palumbi et al. 1991). the thermal profile consisted of 38 cycles at 94°c for 45 s, 55°c for 45 s and 72°c for 1 min. the amplicons obtained were about 550 bp long. pcr products were purified and sequenced using abi technology at the genoscope (evry, france). sequences were checked by eye in codoncode v. 5.1. the sequences obtained were deposited in the genbank database under accession numbers kx055940−kx055957. molecular species identification of samples to identify all our samples we first determined the number of haplotypes in our dataset using tcs (clement et al. 2000). each haplotype was then assigned to a species through a nucleotide blast approach conducted in genbank (http://blast.ncbi.nlm.nih.gov) and through a phylogenetic method (bayesian tree). blast species identification was accomplished following three different criteria (meier et al. 2006): best match (bm), best close match (bcm) and all species barcode (asb). the bm criterion assigns table 1. species identification recovered from a blast in genbank (16s gene) of the 209 specimens of frogs bought in french supermarkets from december 2012 to may 2013 and identified on the label as “rana macrodon from indonesia”. month fejervarya cancrivora fejervarya moodiei limnonectes macrodon total december 2012 22 – – 22 january 2013 48 – – 48 february 2013 26 1 – 27 march 2013 26 – – 26 april 2013 46 – 1 47 may 2013 38 – 1 39 total 206 1 2 209 http://blast.ncbi.nlm.nih.gov european journal of taxonomy 271: 1–19 (2017) 4 identifications to the closest match regardless of genetic distance. the bcm criterion is similar to bm, but the query is identified by the closest match with a distance below a defined threshold. finally, the asb criterion is similar to the bcm by applying a threshold but it returns all the sequences within it. a query is identified when all the matching sequences below the threshold are conspecific. for bcm and asb, a query may provide ambiguous results if sequence divergences of different species are below the threshold (asb) or sequences from different species are the closest match below the threshold (bcm). a tree-based approach of species delimitation was also used. the tree-based criterion of reciprocal monophyly was used to define species boundaries. in our analysis we included: 1) all the haplotypes recovered in our deep frozen frog legs, 2) all species of the genera fejervarya and limnonectes known from indonesia (except limnonectes dammermani (mertens, 1929), l. kenepaiensis (inger, 1966), l. rhacodus (inger, boedi & taufik, 1996), l. sinuatodorsalis matsui, 2015, l. timorensis (smith, 1927) and fejervarya schlueteri (werner, 1893) for which no 16s gene sequences are currently available in genbank), and 3) one sequence of hoplobatrachus rugulosus (wiegmann, 1834) and one sequence of occidozyga laevis (günther, 1858) included as outgroups. depending on 16s gene sequence availability in genbank, we included one to four individuals per species in our analysis. sequences were aligned with clustalw (thompson et al. 1994). in order to minimize the number of missing data we kept 553 sites in our final analyses. evolutionary relationships among sequences were estimated by conducting bayesian markov chain monte carlo phylogenetic analyses (mcmc) with mrbayes v. 3.1.2 (huelsenbeck & ronquist 2001). mrmodeltest v. 3.04 (nylander 2004) was used to evaluate the fit of 24 nested models of nucleotide substitution to the data. according to the akaike information criterion, mrmodeltest recommended the gtr + i + g model. this model was used in the bayesian analysis. three heated chains and one single cold chain were employed, and runs were initiated with random trees. two independent mcmc runs were conducted with five million generations per run; trees (and parameters) were sampled every 100 generations. stationarity was assessed by examining the average standard deviation of split frequencies and the potential scale reduction factor (ronquist et al. 2005). for each run, the first 25% of sampled trees were discarded as burn-in. geographical origin of commercialized frogs for the species fejervarya cancrivora, 16s sequences of specimens from different geographical regions are available in genbank (table 2). we conducted a phylogeographic analysis to see if there is any phylogeographic signal within this species and whether the geographical origin of the commercialized frogs could thus be determined based on 16s gene sequences. to this aim, 16s gene sequences of f. cancrivora for which the geographical origin was known were retrieved from genbank. to minimize the number of missing data, 257 sequences of 494 bp were retained in final analyses (206 sequences of commercialized frogs + 51 specimens from genbank). relationships among haplotypes were inferred by constructing a network using the median-joining method available in network v. 5.0.0.0 (bandelt et al. 1999). morphometry snout vent length (svl) and tibia length (tl) of fejervarya cancrivora were obtained from boulenger (1920) and from measurements of adult specimens from borneo, java and sumatra stored at the field museum of natural history, chicago (fmnh 256671, 256688, 256690, 256692–97, 256709–14). tibia length of frog legs was measured with a slide caliper (appendix). the pearson correlation coefficient showed a highly significant correlation between snout vent length and tibia length (r = 0.973; p < 0.001***). for all specimens measured we calculated the ratio r = tl/svl. we used r = mean of all r to estimate body size depending on tl. we obtained the value r = 0.48177; n = 28; standard deviation = 0.197. thus, we could use the following formula to calculate an estimate for snout vent length for the frog legs: svl = tl / 0.48177. ohler a. & nicolas v., dna barcoding for identification of frog legs 5 table 2. list of specimens of fejervarya cancrivora used in our phylogeographic analysis. genbank no. source country island locality no. of individuals kx055940 this study indonesia – – 1 kx055941 this study indonesia – – 54 kx055942 this study indonesia – – 7 kx055943 this study indonesia – – 66 kx055944 this study indonesia – – 1 kx055945 this study indonesia – – 6 kx055946 this study indonesia – – 1 kx055947 this study indonesia – – 5 kx055948 this study indonesia – – 1 kx055949 this study indonesia – – 8 kx055950 this study indonesia – – 49 kx055951 this study indonesia – – 1 kx055952 this study indonesia – – 1 kx055953 this study indonesia – – 1 kx055954 this study indonesia – – 1 kx055955 this study indonesia – – 3 ab570273 kurniawan et al. 2014 indonesia bali denpasar 6 ab570274 kurniawan et al. 2014 indonesia bali denpasar 1 ab570275 kurniawan et al. 2014 indonesia bali denpasar 1 ab570276 kurniawan et al. 2014 indonesia bali denpasar 1 ab570277 kurniawan et al. 2014 indonesia bali denpasar 1 ab444684 kurniawan et al. 2010 indonesia bangka tempilang 1 ab570273 kurniawan et al. 2014 indonesia java banyumas 1 ab570273 kurniawan et al. 2014 indonesia java banyuwangi 1 ab444684 kurniawan et al. 2010 indonesia java bogor 1 ab570273 kurniawan et al. 2014 indonesia java cilacap 9 ab570273 kurniawan et al. 2014 indonesia java kediri 2 ab570273 kurniawan et al. 2014 indonesia java malang 2 ab444684 kurniawan et al. 2010 indonesia sumatra padang 1 ab570273 kurniawan et al. 2010 indonesia java pelabuhan ratu 1 ab570273 kurniawan et al. 2014 indonesia kalimantan pontianak 6 ab570273 kurniawan et al. 2014 indonesia sumatra jamb 1 ab444684 kurniawan et al. 2014 indonesia sumatra jamb 1 ab570273 kurniawan et al. 2014 indonesia sumatra lampung 1 ab444684 kurniawan et al. 2010 indonesia sumatra langkat 3 ab444685 kurniawan et al. 2010 indonesia sumatra padang 1 ab570273 kurniawan et al. 2014 indonesia sumatra palembang 1 ab444685 kurniawan et al. 2010 indonesia sumatra panti 1 ab444685 kurniawan et al. 2010 indonesia sumatra payakumbuh 1 ab444684 kurniawan et al. 2010 malaysia selangor selangor 1 eu435279 kung et al. unpubl. taiwan donggang donggang 1 eu435280 kung et al. unpubl. taiwan fangliao fangliao 1 eu365387 hsu et al. unpubl. taiwan jiadong jiadong 3 eu365389 hsu et al. unpubl. taiwan sinpi sinpi 1 european journal of taxonomy 271: 1–19 (2017) 6 results species identification after alignment, our dataset contained 585 sites. eighteen haplotypes were recovered (appendix), and they clustered in three groups: (1) haplotypes h01 to h16 (206 individuals) which differ from one another by less than 6 mutations (i.e., 1% sequence divergence), (2) haplotype h17 (one individual) and (3) haplotype h18 (2 individuals). haplotype h17 differs from haplotypes h01 to h16 by 50 to 56 mutations (8.5 to 9.5% of absolute sequence divergence) and from haplotype h18 by 104 mutations (17.8% of absolute sequence divergence). haplotype h18 differs from haplotypes h01 to h16 by 108 to 111 mutations (18.7 to 19.0% of absolute sequence divergence). our nucleotide blast analysis showed that haplotypes h01 to h16 correspond to the species fejervarya cancrivora (best close match: 99 to 100% of sequence identity; table 1). three distinct species have been recognized in “fejervarya cancrivora” (designated as large, mangrove and sulawesi types; hasan et al. 2012). the large type of f. cancrivora was designated as the nominal f. cancrivora (kotaki et al. 2010), while the mangrove and sulawesi types were designated as f. moodiei (taylor, 1920) and an undescribed species, respectively (kurniawan et al. 2011). our haplotypes h01 to h16 cluster with the large f. cancrivora (99% sequence identity). the percentage of sequence identity with the mangrove type (represented in genbank by 10 sequences: eu652694, dq458252, af206473, ab543602, ab070738, ab530508, ab372018, ay841754, ab444692, ab444691) is 91%, and it is 93% with the sulawesi type (represented in genbank by 13 sequences: eu979849, ab444693, ab570278, ab570283, ab570287, ab570286, ab570290, ab570288, ab570289, ab570284, ab570282, ab570280, ab570281). based on the percentage of sequence identity, the other closest species are several species of fejervarya (88–89% of identity with f. vittigera (wiegmann, 1834), f. triora stuart, chuaynker, chan-ard & inger, 2006, f. iskandari, f. multistriata (hallowell, 1861), f. limnocharis, f. sakishimensis matsui, toda & ota, 2008) and hoplobatachus rugulosus. a threshold of 5% is often considered to correspond to distinct species of fejervarya (kurabayashi et al. 2005). based on this threshold, the results of the bm, bcm and asb methods are congruent and show that haplotypes h01 to h16 can be attributed to the species f. cancrivora. haplotype h17 is highly similar (99–100% identity) to the mangrove type (f. moodiei). this haplotype is also close (90–92%) to the large type and the sulawesi type of f. cancrivora. based on the percentage of sequence identity, the other closest species are several species of fejervarya (88–90% of identity with f. vittigera, f. triora, f. iskandari, f. multistriata, f. limnocharis and f. sakishimensis) and hoplobatachus rugulosus (88–89%). haplotype h18 corresponds to the species limnonectes macrodon (99% of sequence identity). based on the percentage of sequence identity, the other closest species are limnonectes shompenorum das, 1996 (94%), l. leporinus anderson, 1923 (91%), l. leytensis (boettger, 1893) (91%) and l magnus (stejneger, 1910) (90%). in our phylogenetic tree (fig. 1), all indonesian species of the genera fejervarya and limnonectes are reciprocally monophyletic and bayesian posterior probabilities are high (pp > 0.98) for all species. this tree clearly shows that haplotypes h01 to h16 correspond to the species fejervarya cancrivora, haplotype 17 corresponds to the species f. moodiei and haplotype h18 corresponds to the species limnonectes macrodon. phylogeography of fejervarya cancrivora the results of our network analysis show that there is some genetic variability within f. cancrivora (fig. 2): 21 haplotypes differing by 1 to 5 mutations were found. thirteen out of the 16 haplotypes ohler a. & nicolas v., dna barcoding for identification of frog legs 7 identified in the commercialized frogs (representing 95 individuals) have not been recorded in previous analyses. haplotype h02, found in 54 commercialized frogs, was recovered in taiwan and several indonesian islands (kalimantan, sumatra, bali and java). haplotype h11 (49 commercialized frogs) was recovered in malaysia and several indonesian islands (bangka, sumatra, java). haplotype h10 fig. 1. phylogeny of indonesian species of fejervarya and limnonectes recovered by the bayesian analysis (gtr + i + g model). hoplobatrachus rugulosus (wiegmann, 1834) and occidozyga laevis (günther, 1858) were used as outgroups. numbers on nodes represent bayesian posterior probabilities, * indicates a value higher than 0.98. only values higher than 0.75 are represented. h01 to h18 indicate the 18 haplotypes from frozen frog legs recovered in this study. european journal of taxonomy 271: 1–19 (2017) 8 (8 commercialized frogs) is present in taiwan. due to the lack of phylogeographic structure within previously sequenced f. cancrivora specimens and to the high number of new haplotypes detected in commercialized frogs, it is not possible to infer the geographical origin of the frogs sold in france. estimation of snout-vent length of frog legs in trade in france for 192 frog legs, the estimated snout-vent length ranged from 59.4 to 111.7 mm (mean value: 79.0; standard deviation: 11.37). the histogram (fig. 3, appendix) of these measurements shows a bimodal distribution which does not correspond to the size distribution of fejervarya cancrivora specimens collected in natural populations without sampling bias. this is also reflected by the relatively low standard deviation. measurements of specimens from borneo, java and sumatra published in boulenger (1920) vary from 54 to 88 mm (mean value: 72.8; standard deviation: 13.34), whereas specimens collected in the early 1990s range from 49.7 to 101.6 mm (mean value: 78.9; standard deviation: 14.98). discussion misidentification of frogs commercialized in france results based on the phylogentic tree and the blast approach (bm, bcm and asb) are congruent and show (threshold of 5%; kurabayashi et al. 2005) that 206 out of the 209 analyzed specimens belong fig. 2. minimum spanning network depicting relationships among 16s haplotypes of fejervarya cancrivora (gravenhorst, 1829). the size of each circle is proportional to the haplotype frequency and the lengths of the connecting lines are proportional to the number of mutations. colors refer to distinct regions (indonesia: java, sumatra, bali, kalimantan, bangka; malaysia; taiwan) and commercialized frogs of unknown origin are in black. ohler a. & nicolas v., dna barcoding for identification of frog legs 9 to the species fejervarya cancrivora, two to the species limnonectes macrodon and one to the species f. moodiei. thus only 0.96% of the frogs were correctly identified. in a previous study on frog legs imported to belgium as f. cancrivora, l. macrodon, l. limnocharis and rana catesbeiana shaw, 1802, all samples proved to be fejervarya cancrivora. in this case, 34.5% of the identifications were correct (veith et al. 2000). in that study 36.8% of the frog legs were sold as limnonectes macrodon. although all our samples were labelled as rana macrodon, only 2 frog legs (0.96%) could be identified as this species. as proposed by several authors (kusrini 2005; kusrini & alford 2006; veith et al. 2000), it seems likely that misclassification is not intentional but due to the fact that frog leg exporters are not able to discriminate between the species in trade. managers of export companies stipulate that they should be supplied only with frog legs of l. macrodon (kusrini & alford 2006). on the other hand, in the local markets l. macrodon and f. cancrivora are correctly identified and sold at different prices, because the meat of l. macrodon is considered of better taste (kusrini & alford 2006; a. ohler, pers. obs.). sustainability and conservation the species concerned by international trade are of similar, large size but show differences in breeding biology and habitat, although relevant data are scarce. fejervarya cancrivora inhabits marshes and paddy fields, not avoiding habitat modified by man and thus a large area of potential habitat is available (inger 1966; iskandar 1998; yuan et al. 2004). virtually nothing is known of the habitat and ecology of f. moodiei (ohler 2004). fejervarya cancrivora lays a relatively large number (up to more than 2500) of small-sized eggs in successive clutches (inger 1956; alcala 1962), which develop in lotic habitats. limnonectes macrodon is present in riparian secondary forests (inger 1966). it can be observed in clearings and secondary growth, and it is very rare in primary forests. a single clutch of about 1000 eggs is laid in side pools of rivers (iskandar 1998). overharvesting should thus have a higher impact on l. macrodon, as breeding capacity is smaller in l. macrodon than in f. cancrivora and riverside habitats are scarcer than ponds and paddy fields. the conservation status of f. cancrivora was evaluated as “least concern” “in view of its wide distribution, tolerance of a broad range of habitats, presumed large fig. 3. histograms. a. snout vent length (in mm) in adult fejervarya cancrivora (gravenhorst, 1829) from samples collected for scientific purposes (boulenger 1920) and collection specimens as mentioned in material and methods. b. snout vent length estimated from tibia length of genetically identified frog legs from french supermarkets (specimen list, see appendix). european journal of taxonomy 271: 1–19 (2017) 10 population, and because it is unlikely to be declining to qualify for listing in a more threatened category” (yuan et al. 2004). fejervarya moodiei is listed as “data deficient in view of continuing doubts as to its extent of occurrence, status and ecological requirements” (ohler 2004). limnonectes macrodon is listed as “vulnerable because it depends on streams in lowland forest, and so its area of occupancy is probably less than 2000 km2, its distribution is severely fragmented, and there is continuing decline in the extent and quality of its habitat, the number of locations, and the number of mature individuals” (iskandar et al. 2004). these evaluations are not based on population studies that might measure the impact of the heavy harvesting ongoing for decades. however, the quasi absence of l. macrodon in our samples might be an indication of its rarity in the field and the fact that its natural populations are declining rapidly. in the 1990s, researchers already considered that large-sized frogs had completely disappeared in many parts of java and sumatra (manthey & grossmann 1997; inger & stuebing 1997). in 2006, 18.8% of the frogs captured by harvesters in west java and east java belonged to the species l. macrodon (kuzrini & alford 2006). in 2012–2013, only 0.96% of the samples from trade sold in france belonged to this species. a diminution in size of adult frogs due to overharvesting has been suggested (iskandar, cited in anonymous 2007). the estimated size of the frogs from deep frozen specimens of f. cancrivora is not smaller than specimens collected 20 or a hundred years ago. the frogs measured in boulenger (1920) are slightly smaller in size than the frogs collected in the early 1990s and in 2012–2013. however, the body size of the frogs in trade is clearly biased. after collection, frogs are separated into size categories and only large specimens are chosen for export (kusrini & alford 2006). in samples studied by church (1960) and kusrini & alford (2006) that had been collected for trade, very large-sized frogs were present (maximum size of 132 mm for 1325 frogs, and of 162 mm for 555 frogs, respectively). in the samples bought in the supermarkets in france imported by the same company, two size classes of frogs are present, but very large frogs have not been observed in our sample. the absence of small-sized frogs is due to the sorting out of larger specimens for export (kusrini & alford 2006). nevertheless, the presence of numerous large frogs means that such large specimens can be found in the field. based on our genetic results, we could not allocate our samples to precise geographic origins. the absence of an effect of overharvesting on body size can only be studied when the precise geographic origin of specimens is known, because body size varies geographically (kurniawan et al. 2011). the specimens studied in the present work may well come from newly harvested populations. in fact, the presence of f. moodiei in our samples, a species which has a known range outside java and sumatra, a range that includes the philippines, the northern coasts of the gulf of thailand, bangladesh and orissa (kurniawan et al. 2010), indicates that the specimens studied here may have come from other islands than java and south-eastern sumatra. “at present, nothing is known about the impact of extensive frog harvesting on indonesian frog populations and agricultures.” this phrase of veith et al. (2000) can be cited as such, as there has been no change in the last 15 years. there are still no studies on the effects of frog harvesting on natural populations. our results show that the genetic and morphological diversity of the frogs in trade is much higher than the genetic and morphological diversity measured so far by scientific studies (kurniawan et al. 2010, 2011). these results underline the need for large-scale studies on the taxonomy, population structure, reproductive data and ecology of the species concerned in international trade. in 2000, veith et al. concluded that the development of a “quick and cheap test for management authorities” was necessary to monitor the international trade in frog legs. such a test is already available through dna sequencing, and the price is decreasing rapidly with the development of next generation sequencing techniques. the lack of any progress in the development of identification tools and assessing the status of wild populations is not due to an absence of convenient methodology but it reflects the fact that conservation is focused on flagship species, such as tigers and pandas, or species that qualify for rarity. ohler a. & nicolas v., dna barcoding for identification of frog legs 11 the edible frogs of indonesia, although billions of individuals are killed annually, are the scarcely visible part of the bulk of species that are disappearing. acknowledgements this project was supported by the network ‘bibliothèque du vivant’ funded by the cnrs, the muséum national d’histoire naturelle (mnhn), the institut national de recherche agronomique (inra) and the commissariat à l'energie atomique (cea, genoscope). this work was also supported by the ‘action transversale du muséum: taxonomie moléculaire, dna barcode & gestion durable des collections’ and the ‘service de systématique moléculaire’ of the mnhn (ums 2700, paris, france). we are grateful to tesceline tremaud and dario zuccon for their help in molecular lab work. the 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editor: danny eibye-jacobsen printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands. http://dx.doi.org/10.1007/s10531-009-9758-4 http://cs.mcgill.ca/~birch/doc/mrbayes/mb3.1_manual.pdf http://cs.mcgill.ca/~birch/doc/mrbayes/mb3.1_manual.pdf http://dx.doi.org/10.1093/nar/22.22.4673 http://dx.doi.org/10.1023/a:1008906918236 http://dx.doi.org/10.1186/1742-9994-2-5 http://dx.doi.org/10.1186/1742-9994-2-5 http://dx.doi.org/10.1111/jeb.12292 http://dx.doi.org/10.1111/j.1523-1739.2008.01165.x http://dx.doi.org/10.2305/iucn.uk.2004.rlts.t58269a11759436.en http://dx.doi.org/10.2305/iucn.uk.2004.rlts.t58269a11759436.en european journal of taxonomy 271: 1–19 (2017) 14 appendix list of samples of deep frozen frog legs with purchase date, collection number, haplotype number, taxonomic identification, tibia length (tl) and estimated snout vent length (svl). date museum number haplotype number species identification (blast) tl svl (estimated) april 2013 2013.0317 h01 fejervarya cancrivora 34.2 71.0 april 2013 2013.0318 h02 fejervarya cancrivora 34.9 72.4 april 2013 2013.0319 h03 fejervarya cancrivora 32.5 67.5 april 2013 2013.0320 h04 fejervarya cancrivora 32.2 66.8 april 2013 2013.0321 h03 fejervarya cancrivora 36.7 76.2 april 2013 2013.0322 h02 fejervarya cancrivora 32.9 68.3 april 2013 2013.0323 h02 fejervarya cancrivora 36.8 76.4 april 2013 2013.0324 h02 fejervarya cancrivora 37.0 76.8 april 2013 2013.0325 h02 fejervarya cancrivora 33.9 70.4 april 2013 2013.0326 h04 fejervarya cancrivora 33.8 70.2 april 2013 2013.0327 h02 fejervarya cancrivora 38.5 79.9 april 2013 2013.0328 h02 fejervarya cancrivora 34.1 70.8 april 2013 2013.0329 h02 fejervarya cancrivora 32.0 66.4 april 2013 2013.0330 h02 fejervarya cancrivora 36.8 76.4 april 2013 2013.0331 h02 fejervarya cancrivora 36.3 75.3 april 2013 2013.0332 h04 fejervarya cancrivora 33.6 69.7 april 2013 2013.0333 h02 fejervarya cancrivora 32.9 68.3 april 2013 2013.0334 h05 fejervarya cancrivora 35.5 73.7 april 2013 2013.0335 h04 fejervarya cancrivora 38.4 79.7 april 2013 2013.0336 h06 fejervarya cancrivora 32.8 68.1 april 2013 2013.0337 h04 fejervarya cancrivora 35.1 72.9 april 2013 2013.0338 h04 fejervarya cancrivora 33.3 69.1 april 2013 2013.0339 h04 fejervarya cancrivora 35.4 73.5 april 2013 2013.0340 h07 fejervarya cancrivora 34.5 71.6 april 2013 2013.0341 h02 fejervarya cancrivora 33.2 68.9 march 2013 2013.0342 h02 fejervarya cancrivora 33.3 69.1 march 2013 2013.0343 h02 fejervarya cancrivora 37.2 77.2 march 2013 2013.0344 h02 fejervarya cancrivora 36.2 75.1 march 2013 2013.0345 h02 fejervarya cancrivora 39.7 82.4 march 2013 2013.0346 h02 fejervarya cancrivora 39.2 81.4 march 2013 2013.0347 h04 fejervarya cancrivora 34.6 71.8 march 2013 2013.0348 h02 fejervarya cancrivora 34.0 70.6 march 2013 2013.0349 h08 fejervarya cancrivora 36.6 76.0 march 2013 2013.0350 h08 fejervarya cancrivora 33.1 68.7 march 2013 2013.0351 h02 fejervarya cancrivora 34.7 72.0 march 2013 2013.0352 h09 fejervarya cancrivora 37.8 78.5 march 2013 2013.0353 h10 fejervarya cancrivora 35.6 73.9 march 2013 2013.0354 h02 fejervarya cancrivora 33.3 69.1 ohler a. & nicolas v., dna barcoding for identification of frog legs 15 march 2013 2013.0355 h04 fejervarya cancrivora 33.5 69.5 march 2013 2013.0356 h02 fejervarya cancrivora 36.0 74.7 march 2013 2013.0357 h04 fejervarya cancrivora 35.7 74.1 march 2013 2013.0358 h04 fejervarya cancrivora 36.5 75.8 march 2013 2013.0359 h02 fejervarya cancrivora 35.3 73.3 march 2013 2013.0360 h02 fejervarya cancrivora 33.8 70.2 march 2013 2013.0361 h08 fejervarya cancrivora 36.6 76.0 march 2013 2013.0363 h02 fejervarya cancrivora 33.8 70.2 march 2013 2013.0364 h04 fejervarya cancrivora 32.9 68.3 march 2013 2013.0365 h04 fejervarya cancrivora 31.1 64.6 march 2013 2013.0366 h04 fejervarya cancrivora 38.9 80.7 march 2013 2013.0367 h02 fejervarya cancrivora 34.9 72.4 march 2013 2013.0368 h02 fejervarya cancrivora 37.3 77.4 february 2013 2013.0369 h11 fejervarya cancrivora 33.3 69.1 february 2013 2013.0371 h11 fejervarya cancrivora 34.4 71.4 february 2013 2013.0372 h11 fejervarya cancrivora 32.6 67.7 february 2013 2013.0373 h11 fejervarya cancrivora 31.3 65.0 february 2013 2013.0374 h11 fejervarya cancrivora 33.3 69.1 february 2013 2013.0375 h11 fejervarya cancrivora 36.2 75.1 february 2013 2013.0376 h11 fejervarya cancrivora 35.3 73.3 february 2013 2013.0377 h11 fejervarya cancrivora 32.4 67.3 february 2013 2013.0378 h11 fejervarya cancrivora 28.6 59.4 february 2013 2013.0379 h11 fejervarya cancrivora 31.0 64.3 february 2013 2013.0380 h11 fejervarya cancrivora 32.1 66.6 february 2013 2013.0381 h11 fejervarya cancrivora 32.7 67.9 february 2013 2013.0382 h11 fejervarya cancrivora 31.8 66.0 february 2013 2013.0383 h11 fejervarya cancrivora 31.5 65.4 february 2013 2013.0384 h11 fejervarya cancrivora 33.6 69.7 february 2013 2013.0385 h11 fejervarya cancrivora 32.0 66.4 february 2013 2013.0386 h17 fejervarya moodiei 31.2 – february 2013 2013.0387 h11 fejervarya cancrivora 33.1 68.7 february 2013 2013.0388 h11 fejervarya cancrivora 32.7 67.9 february 2013 2013.0389 h11 fejervarya cancrivora 33.5 69.5 february 2013 2013.0390 h11 fejervarya cancrivora 30.4 63.1 february 2013 2013.0391 h11 fejervarya cancrivora 31.2 64.8 february 2013 2013.0392 h11 fejervarya cancrivora 30.9 64.1 february 2013 2013.0393 h11 fejervarya cancrivora 34.6 71.8 february 2013 2013.0394 h11 fejervarya cancrivora 31.6 65.6 february 2013 2013.0395 h11 fejervarya cancrivora 32.6 67.7 february 2013 2013.0396 h11 fejervarya cancrivora 35.4 73.5 appendix (cont.) date museum number haplotype number species identification (blast) tl svl (estimated) european journal of taxonomy 271: 1–19 (2017) 16 january 2013 2013.0397 h02 fejervarya cancrivora 42.6 88.4 january 2013 2013.0398 h02 fejervarya cancrivora 41.9 87.0 january 2013 2013.0399 h04 fejervarya cancrivora 43.4 90.1 january 2013 2013.0400 h04 fejervarya cancrivora 45.8 95.1 january 2013 2013.0401 h02 fejervarya cancrivora 43.5 90.3 january 2013 2013.0402 h04 fejervarya cancrivora 46.6 96.7 january 2013 2013.0403 h02 fejervarya cancrivora 42.3 87.8 january 2013 2013.0404 h08 fejervarya cancrivora 44.7 92.8 january 2013 2013.0405 h04 fejervarya cancrivora 43.4 90.1 january 2013 2013.0406 h10 fejervarya cancrivora 47.4 98.4 january 2013 2013.0407 h06 fejervarya cancrivora 40.0 83.0 january 2013 2013.0408 h04 fejervarya cancrivora 43.0 89.3 january 2013 2013.0409 h02 fejervarya cancrivora 44.3 92.0 january 2013 2013.0410 h02 fejervarya cancrivora 46.0 95.5 january 2013 2013.0411 h04 fejervarya cancrivora 43.5 90.3 january 2013 2013.0412 h04 fejervarya cancrivora 39.9 82.8 january 2013 2013.0413 h06 fejervarya cancrivora 43.5 90.3 january 2013 2013.0414 h02 fejervarya cancrivora 50.2 104.2 january 2013 2013.0415 h02 fejervarya cancrivora 47.1 97.8 january 2013 2013.0416 h02 fejervarya cancrivora 41.0 85.1 january 2013 2013.0417 h04 fejervarya cancrivora 48.0 99.6 january 2013 2013.0418 h02 fejervarya cancrivora 46.7 96.9 april 2013 2013.0419 h04 fejervarya cancrivora 45.4 94.2 april 2013 2013.0420 h04 fejervarya cancrivora 43.4 90.1 april 2013 2013.0421 h04 fejervarya cancrivora 45.2 93.8 april 2013 2013.0422 h04 fejervarya cancrivora 53.8 111.7 april 2013 2013.0423 h04 fejervarya cancrivora 44.7 92.8 april 2013 2013.0424 h04 fejervarya cancrivora 45.2 93.8 april 2013 2013.0425 h04 fejervarya cancrivora 42.5 88.2 april 2013 2013.0426 h04 fejervarya cancrivora 44.1 91.5 april 2013 2013.0427 h04 fejervarya cancrivora 45.9 95.3 april 2013 2013.0428 h04 fejervarya cancrivora 46.7 96.9 april 2013 2013.0429 h04 fejervarya cancrivora 45.2 93.8 april 2013 2013.0430 h18 limnonectes macrodon 43.3 – april 2013 2013.0431 h04 fejervarya cancrivora 51.8 107.5 april 2013 2013.0432 h04 fejervarya cancrivora 44.2 91.7 april 2013 2013.0433 h10 fejervarya cancrivora 45.8 95.1 april 2013 2013.0434 h04 fejervarya cancrivora 44.7 92.8 april 2013 2013.0435 h10 fejervarya cancrivora 46.4 96.3 april 2013 2013.0436 h04 fejervarya cancrivora 42.0 87.2 appendix (cont.) date museum number haplotype number species identification (blast) tl svl (estimated) ohler a. & nicolas v., dna barcoding for identification of frog legs 17 april 2013 2013.0437 h04 fejervarya cancrivora 45.3 94.0 april 2013 2013.0438 h04 fejervarya cancrivora 45.7 94.9 april 2013 2013.0439 h04 fejervarya cancrivora 47.4 98.4 april 2013 2013.0440 h04 fejervarya cancrivora 48.5 100.7 january 2013 2013.0441 h04 fejervarya cancrivora 35.4 73.5 january 2013 2013.0442 h12 fejervarya cancrivora 35.2 73.1 january 2013 2013.0443 h02 fejervarya cancrivora 34.1 70.8 january 2013 2013.0444 h11 fejervarya cancrivora 37.3 77.4 january 2013 2013.0445 h04 fejervarya cancrivora 33.5 69.5 january 2013 2013.0446 h03 fejervarya cancrivora 32.3 67.0 january 2013 2013.0447 h13 fejervarya cancrivora 32.0 66.4 january 2013 2013.0448 h04 fejervarya cancrivora 37.1 77.0 january 2013 2013.0449 h03 fejervarya cancrivora 34.9 72.4 january 2013 2013.0450 h02 fejervarya cancrivora 31.8 66.0 january 2013 2013.0451 h03 fejervarya cancrivora 37.9 78.7 january 2013 2013.0452 h14 fejervarya cancrivora 34.4 71.4 january 2013 2013.0453 h02 fejervarya cancrivora 34.1 70.8 january 2013 2013.0454 h04 fejervarya cancrivora 36.0 74.7 january 2013 2013.0455 h04 fejervarya cancrivora 36.5 75.8 january 2013 2013.0456 h04 fejervarya cancrivora 34.9 72.4 january 2013 2013.0457 h11 fejervarya cancrivora 33.5 69.5 january 2013 2013.0458 h03 fejervarya cancrivora 35.8 74.3 january 2013 2013.0459 h02 fejervarya cancrivora 36.4 75.6 january 2013 2013.0460 h02 fejervarya cancrivora 36.5 75.8 january 2013 2013.0461 h02 fejervarya cancrivora 34.2 71.0 january 2013 2013.0462 h04 fejervarya cancrivora 35.7 74.1 january 2013 2013.0463 h04 fejervarya cancrivora 37.3 77.4 january 2013 2013.0464 h04 fejervarya cancrivora 33.0 68.5 january 2013 2013.0465 h11 fejervarya cancrivora 39.2 81.4 january 2013 2013.0466 h03 fejervarya cancrivora 32.7 67.9 december 2012 2013.0467 h02 fejervarya cancrivora 41.0 85.1 december 2012 2013.0468 h02 fejervarya cancrivora 48.3 100.3 december 2012 2013.0469 h11 fejervarya cancrivora 44.8 93.0 december 2012 2013.0470 h11 fejervarya cancrivora 43.6 90.5 december 2012 2013.0471 h11 fejervarya cancrivora 43.2 89.7 december 2012 2013.0472 h11 fejervarya cancrivora 43.5 90.3 december 2012 2013.0473 h11 fejervarya cancrivora 46.8 97.1 december 2012 2013.0474 h11 fejervarya cancrivora 45.2 93.8 december 2012 2013.0475 h11 fejervarya cancrivora 45.5 94.4 december 2012 2013.0476 h11 fejervarya cancrivora 47.8 99.2 appendix (cont.) date museum number haplotype number species identification (blast) tl svl (estimated) european journal of taxonomy 271: 1–19 (2017) 18 december 2012 2013.0477 h11 fejervarya cancrivora 47.7 99.0 december 2012 2013.0478 h11 fejervarya cancrivora 42.5 88.2 december 2012 2013.0479 h11 fejervarya cancrivora 42.0 87.2 december 2012 2013.0480 h11 fejervarya cancrivora 41.5 86.1 december 2012 2013.0481 h11 fejervarya cancrivora 46.4 96.3 december 2012 2013.0482 h11 fejervarya cancrivora 46.1 95.7 december 2012 2013.0483 h11 fejervarya cancrivora 46.2 95.9 december 2012 2013.0484 h11 fejervarya cancrivora 45.6 94.7 december 2012 2013.0485 h11 fejervarya cancrivora 48.8 101.3 december 2012 2013.0486 h11 fejervarya cancrivora 41.0 85.1 december 2012 2013.0487 h11 fejervarya cancrivora 44.6 92.6 december 2012 2013.0488 h11 fejervarya cancrivora 43.8 90.9 may 2013 2013.0489 h04 fejervarya cancrivora 37.6 78.0 may 2013 2013.0490 h04 fejervarya cancrivora 34.2 71.0 may 2013 2013.0491 h06 fejervarya cancrivora 35.3 73.3 may 2013 2013.0492 h02 fejervarya cancrivora 34.1 70.8 may 2013 2013.0493 h04 fejervarya cancrivora 32.8 68.1 may 2013 2013.0494 h10 fejervarya cancrivora 35.2 73.1 may 2013 2013.0495 h02 fejervarya cancrivora 34.1 70.8 may 2013 2013.0496 h02 fejervarya cancrivora 35.0 72.6 may 2013 2013.0497 h10 fejervarya cancrivora 31.7 65.8 may 2013 2013.0498 h10 fejervarya cancrivora 35.4 73.5 may 2013 2013.0499 h04 fejervarya cancrivora 36.0 74.7 may 2013 2013.0500 h15 fejervarya cancrivora 35.6 73.9 may 2013 2013.0501 h10 fejervarya cancrivora 36.2 75.1 may 2013 2013.0502 h04 fejervarya cancrivora 31.4 65.2 may 2013 2013.0503 h04 fejervarya cancrivora 36.3 75.3 may 2013 2013.0504 h06 fejervarya cancrivora 36.5 75.8 may 2013 2013.0505 h02 fejervarya cancrivora 33.8 70.2 may 2013 2013.0506 h02 fejervarya cancrivora 34.0 70.6 may 2013 2013.0507 h02 fejervarya cancrivora 34.4 71.4 may 2013 2013.0508 h04 fejervarya cancrivora 36.9 76.6 may 2013 2013.0509 h02 fejervarya cancrivora 36.9 76.6 may 2013 2013.0510 h06 fejervarya cancrivora 34.3 71.2 may 2013 2013.0511 h02 fejervarya cancrivora 32.7 67.9 may 2013 2013.0512 h04 fejervarya cancrivora 33.0 68.5 may 2013 2013.0515 h02 fejervarya cancrivora – – may 2013 2013.0516 h16 fejervarya cancrivora – – may 2013 2013.0517 h08 fejervarya cancrivora – – may 2013 2013.0518 h04 fejervarya cancrivora – – appendix (cont.) date museum number haplotype number species identification (blast) tl svl (estimated) ohler a. & nicolas v., dna barcoding for identification of frog legs 19 appendix (cont.) may 2013 2013.0519 h04 fejervarya cancrivora – – may 2013 2013.0520 h04 fejervarya cancrivora – – may 2013 2013.0521 h16 fejervarya cancrivora – – may 2013 2013.0522 h16 fejervarya cancrivora – – may 2013 2013.0523 h18 limnonectes macrodon – – may 2013 2013.0524 h04 fejervarya cancrivora – – may 2013 2013.0525 h04 fejervarya cancrivora – – may 2013 2013.0526 h04 fejervarya cancrivora – – may 2013 2013.0527 h04 fejervarya cancrivora – – may 2013 2013.0528 h02 fejervarya cancrivora – – may 2013 2013.0529 h04 fejervarya cancrivora – – date museum number haplotype number species identification (blast) tl svl (estimated) new members of the genera neanura macgillivray, 1893 and deutonura cassagnau, 1979 (collembola: neanuridae) from the middle east adrian smolis 1,*, masoumeh shayanmehr 2 & elham yoosefi-lafooraki 3 1 institute of environmental biology, department of invertebrate biology, evolution and conservation, university of wrocław, przybyszewskiego 65, 51-148 wrocław, poland. 2,3 department of plant protection, sari agricultural science and nature resources university, sari, mazandaran 578, iran. * corresponding author: adrian.smolis@uwr.edu.pl 2 email: shayanm30@yahoo.com 3 email: eyoosefi@ymail.com 1 urn:lsid:zoobank.org:author:ff99fab6-0735-4aa5-bee0-8012e90516f4 2 urn:lsid:zoobank.org:author:34df6ad6-9748-4963-9c68-3553b5ecdc72 3 urn:lsid:zoobank.org:author:f7a47f39-1dea-4e12-bed8-23d8b315c7d0 abstract. new species of neanura macgillivray, 1893 and deutonura cassagnau, 1979 are described from northern iran. both taxa are characterized by unusual features that place them in isolated positions within the genera. neanura deharvengi sp. nov. differs from congeners by the extreme reduction of head chaetotaxy and fused lateral tubercles on the head. these characteristics of the new species broaden the existing diagnosis of the genus neanura. an updated diagnosis is provided herein. deutonura persica sp. nov. is most similar to d. plena (stach, 1951), known from the carpathians. the new species can be distinguished by the strong reduction of its head, labial, and labral chaetotaxy as well as the relative length of chaetae de2 and de3 on abdominal segments i–iii. the iranian records of d. decolorata (gama & gisin, 1964) are questioned. brief remarks on the importance of the newly described species for the knowledge of both genera are also provided. keywords. springtails, neanurinae, western palaearctic, iran. smolis a., shayanmehr m. & yoosefi-lafooraki e. 2018. new members of the genera neanura macgillivray, 1893 and deutonura cassagnau, 1979 (collembola: neanuridae) from the middle east. european journal of taxonomy 406: 1–16. https://doi.org/10.5852/ejt.2018.406 introduction in recent years, there have been a series of publications about iranian species in the subfamily neanurinae (shayanmehr et al. 2013; mayvan et al. 2015; smolis et al. 2012, 2016a, 2016b, 2017). to date, the local fauna of this subfamily encompasses 13 species classified into 8 genera. due to the size and diversity of european journal of taxonomy 406: 1–16 issn 2118-9773 https://doi.org/10.5852/ejt.2018.406 www.europeanjournaloftaxonomy.eu 2018 · smolis a. et al. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:8e7cde52-df77-4f87-8efc-14a41a674bf7 1 mailto:adrian.smolis%40uwr.edu.pl?subject= mailto:shayanm30%40yahoo.com?subject= mailto:eyoosefi%40ymail.com?subject= http://zoobank.org/urn:lsid:zoobank.org:author:ff99fab6-0735-4aa5-bee0-8012e90516f4 http://zoobank.org/urn:lsid:zoobank.org:author:34df6ad6-9748-4963-9c68-3553b5ecdc72 http://zoobank.org/urn:lsid:zoobank.org:author:f7a47f39-1dea-4e12-bed8-23d8b315c7d0 https://doi.org/10.5852/ejt.2018.406 https://doi.org/10.5852/ejt.2018.406 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:8e7cde52-df77-4f87-8efc-14a41a674bf7 iran these numbers are certainly an underestimate. confirming this, we supply the descriptions of two new iranian species of this group. the genus neanura macgillivray, 1893 is the oldest taxonomic unit in the subfamily. cassagnau (1979) redefined the genus and split it into four subgenera: neanura sensu stricto, cryptonura, deutonura and endonura. shortly after, this was expanded by deharveng (1982b). as currently understood, neanura includes only 7 species (deharveng 1982b; buşmachiu & deharveng 2008; smolis & deharveng 2017). the distribution of most of them, excluding the cosmopolitan n. muscorum (templeton, 1835), is restricted to europe, especially to the western and central parts of the continent. taxonomically, neanura is probably most closely related to the siberian genus kalanura smolis, 2007, from which it differs in the arrangement of lateral tubercles and the relative length of chaetae a and b on the head (smolis 2007). in contrast to the genus neanura, deutonura cassagnau, 1979 is nowadays one of the largest genera of neanurinae, comprising 57 valid species (e.g., deharveng 1982a; deharveng & weiner 1984; porco et al. 2010; deharveng et al. 2015). the highest diversity of the genus is documented for western europe; for example, 19 species have been recorded in france (deharveng et al. 2015). due to the fusion of tubercles di and de on the head and tubercles di on the penultimate abdominal segment, deutonura strongly resembles the monotypic genus albanura deharveng, 1982, although it differs from the latter by having separate tubercles di on abdominal tergite iv (fused in albanura) and by the number of chaetae di on abdomen v (3+3 or very rarely 2+2 chaetae in deutonura, 2+2 in albanura) (deharveng 1982b). in the present contribution, we describe neanura deharvengi sp. nov. and deutonura persica sp. nov. from northern iran. the former species is especially interesting since it possesses two unusual characters: extreme reduction of cephalic chaetotaxy and fused lateral tubercles on the head. nevertheless, this new species is also characterised by several features (i.e., 3+3 eyes, the cross-type arrangement of chaetae on the head, separateness of cephalic chaetae dl and presence of male ventral organ) which are characteristic of the genus neanura. it convinced us that a new genus shouldn’t be erected for it at the moment and that the diagnosis of neanura can be modified to accommodate the new species. we additionally include short remarks on the published record of d. decolorata (gama & gisin, 1964) from iran and on the global distribution of the genera neanura and deutonura. material and methods specimens were collected from soil and litter samples. the samples were extracted using a berlesetullgren apparatus. specimens were cleared in nesbitt’s fluid, subsequently mounted on slides in swan’s medium and studied using a nikon eclipse e600 phase contrast microscope. figures were drawn with a camera lucida and prepared for publication using adobe photoshop cs3. the material is deposited in the department of invertebrate biology, evolution and conservation, institute of environmental biology, university of wrocław, poland (dibec). terminology and abbreviations the terminology for the descriptions follows those of deharveng & weiner (1984), smolis & deharveng (2006) and smolis (2008). the following abbreviations are used in the text, figures and tables: general morphology: abd. = abdomen ant. = antenna european journal of taxonomy 406: 1–16 (2018) 2 aoiii = sensory organ of antennal segment iii cx = coxa fe = femur scx2 = subcoxa 2 t = tibiotarsus th. = thorax tr = trochanter vt = ventral tube groups of chaetae: ag = antegenital an = chaetae of anal lobes ap = apical ca = centroapical cm = centromedial cp = centroposterior d = dorsal fu = furcal ve or ve = ventroexternal vea = ventroexternoanterior vec = ventroexternocentral vei = ventroexternointernal vel = ventroexternolateral vem = ventroexternomedial vep = ventroexternoposterior vc = ventrocentral vi or vi = ventrointernal vl = ventrolateral tubercles: af = antenna-frontal cl = clypeal de = dorsoexternal de, ee = elementary tubercles on head di = dorsointernal dl = dorsolateral l = lateral oc = ocular so = subocular types of chaetae: b4, b5 = ordinary chaetae on tibiotarsi brs = border s-chaeta on ant. iv bs = s-chaeta on ant. iv i = ordinary chaeta on ant. iv iv = ordinary chaetae on ventral ant. iv l’ = ordinary lateral chaeta on abd. v mc = short macrochaeta mcc = very short macrochaeta smolis a. et al., new species of neanura and deutonura from the middle east 3 me = mesochaeta mi = microchaeta mia = microchaetae on ant. iv ml = long macrochaeta mou = cylindrical s-chaetae on ant. iv (‘soies mousses’) ms = s-microchaeta oca, ocm, ocp = ocular chaetae on head or = subapical pit of ant. iv s or s = chaeta s sgd = dorsal s-chaeta on ant. iii sgv = ventral s-chaeta on ant. iii x = labial papilla x results class collembola lubbock, 1873 order poduromorpha börner, 1913 family neanuridae börner, 1901 subfamily neanurinae börner, 1901 tribe neanurini salmon, 1951 genus neanura macgillivray, 1893 type species achorutes muscorum templeton, 1835. since the new species described below is characterized by a number of characters unknown in other members of the genus, e.g., the absence of cephalic chaetae a and the fusion of lateral tubercles on the head into one mass, the currently accepted diagnosis of the genus as proposed by deharveng (1982) is emended here. moreover, three recently erected genera (kalanura, xylanura smolis, 2011 and persanura mayvan et al., 2015; see smolis 2007, 2011 and mayvan et al. 2015) morphologically strongly resemble the genus neanura. as a result, its diagnosis requires extension based on new morphological characters (e.g., relative length of cephalic chaetae a and b, presence of tubercles di on abd., arrangement of tubercles on abd. v, presence of male ventral organ) diagnostic and useful for generic characterization within the tribe neanurini. emended diagnosis body colour blue to grey or whitish. eyes with 3+3 dark pigmented ocelli. dorsal tubercles and reticulations present, well developed. abdominal segments iv and v subequal in length. chaetotaxy of body and appendages not polychaetotic. ant. iv with 8 subequal s–chaetae. labral chaetotaxy 4/2, 4 or 2/2,4. mouthparts reduced, maxilla styliform, mandible tridentate. tubercles af and cl on head separate. chaetae a on head present or rarely absent. if present, chaetae a and b of same length. chaetae ocp on head present or absent. head with tubercles (dl + l) and so or rarely they are fused into single mass. sometimes tubercle (l+dl) on head consists of two parts (l+1/2dl) and dl’. arrangement of chaetae di and de on head of cross-type, line of chaetae di2–de2 crosses line di1–de1. tubercles di and de on head separate. tubercles di on th. i present or absent. th. ii–iii with 3 chaetae di. tubercles di on abd. i–iv present. tubercles di never fused on abdominal segment iv, but fused, partially fused or separate on abd. v. each half of abd. v with 3 or 2 chaetae di. cryptopygy absent or slightly developed. abd. vi bilobed. furcal remnant without microchaetae. chaeta l’ present or absent. adult males usually with european journal of taxonomy 406: 1–16 (2018) 4 modified chaetae on ventral side of abdomen (male ventral organ). tibiotarsi with 19, 19, 18 chaetae, clavate chaetae absent. claw without inner tooth. neanura deharvengi sp. nov. urn:lsid:zoobank.org:act:1ef90002-5e99-4451-9855-815658f5a9b0 figs 1–2; table 1 diagnosis habitus typical of genus. dorsal tubercles present and well developed. body colour light bluish grey. buccal cone short, labrum without ogival sclerifications. dorsal cephalic chaetotaxy strongly reduced: chaetae a, c, o, e, ocp, dl5, dl3, so2, so6 and l2–3 absent. tubercles di on th. i absent. abd. i–iii without chaetae de3. tubercle (di+di) of abd. v with 2+2 chaetae, chaetae di2 absent. cryptopygy absent. male ventral organ present. etymology the new species is dedicated to our colleague and friend louis deharveng for his magnificent contribution to our knowledge of collembola. material examined holotype iran: adult ♀, on slide, mazandarn province, behshahr region, abbas-abad forest, 36°40′ n, 53°32′ e, leaf litter and soil, 28 mar. 2013, e. yoosefi-lafooraki leg. (dibec). paratypes iran: adult ♂ and subadult ♂, on slides, same collection data as for holotype (dibec). other material iran: juvenile, on slide, mazandarn province, noor region, kadirsar village, 36°26′ n, 51°49′ e, leaf litter and soil, 1 mar. 2013, e. yoosefi-lafooraki leg. (dibec). description body. length (without antennae): 0.58 to 0.80 mm (holotype: 0.8 mm). body convex. colour of body light bluish grey. 3+3 black eyes of medium size, in typical arrangement of genus (two anterior and one posterior). chaetal morphology. dorsal ordinary chaetae of three types: long macrochaetae (ml), mesochaetae and microchaetae. long macrochaetae relatively thick, slightly arc-like or straight, narrowly sheathed, feebly serrated, mostly apically rounded or rarely, mainly in lateral position, pointed (figs 1a–b, 2c). mesochaetae similar to ventral chaetae, thin, smooth and pointed. microchaetae similar to mesochaetae but shorter. s–chaetae of tergites thin, smooth and short, notably shorter than nearby macrochaetae (figs 1a–b, 2c, e–f). antennae. typical of genus. ant. i–ii with 7 or 11 ordinary chaetae, respectively. ant. iii with 5 s-chaetae (aoiii) and 18 ordinary chaetae: 5 d, 4 vi, 4 vc and 5 ve (fig. 1f). ant. iv dorsally with complete set of chaetae: 8 s, or, i, 12 mou and 3 brs (fig. 1e). s–chaetae of ant. iv of medium length, subequal and moderately thickened. apical vesicle distinct, trilobed (fig. 1c–d). ant. iv ventrally with complete set of chaetae (3 brs, 2 iv; ap: 8 bs and 5 mia; ca: 2 bs and 3 mia; cm: 3 bs and 1 mia; cp: 8 mia and 1 brs). smolis a. et al., new species of neanura and deutonura from the middle east 5 http://zoobank.org/urn:lsid:zoobank.org:act:1ef90002-5e99-4451-9855-815658f5a9b0 mouthparts. buccal cone relatively short, with labral sclerifications non-ogival. labrum chaetotaxy: 2/2, 4 (fig. 2a). labium with four basal, three distal and four lateral chaetae, papillae x absent. maxilla styliform, mandible thin and tridentate. fig. 1. neanura deharvengi sp. nov. a. dorsal chaetotaxy of head, th. and abd. i. b. lateral chaetotaxy of head. c. apical bulb, ventral view. d. apical bulb, dorsal view. e. dorsal chaetotaxy of ant. iii–iv. f. ventral chaetotaxy of ant. iii. scale bars: a–b, e–f = 0.1 mm; c–d = 0.01 mm. european journal of taxonomy 406: 1–16 (2018) 6 fig. 2. neanura deharvengi sp. nov. a. labrum. b. ventral chaetotaxy of abd. ii–vi, adult male. c. ml di1 of abd. v. d. modified chaetae of male ventral organ. e. sensillum of abd. v. f. dorsal chaetotaxy of abd. iv–vi. scale bars: a = 0.01 mm; b–f = 0.1 mm. smolis a. et al., new species of neanura and deutonura from the middle east 7 dorsal chaetotaxy and tubercles. head with fused tubercles so and (l+1/2dl) (fig. 1a–b). cephalic chaetae dl2, dl6 and l4 free. th. ii–iii with free chaetae di2, di3 and de2. on th. ii–iii, chaetae de3 and dl3 absent. on abd. i–iii, line of chaetae de1–chaeta s nearly parallel to dorsomedian line. on abd. i–iii chaetae de2 free (fig. 1a). abd. v with 3 tubercles: (di+di) and 2 (de+dl+l). on abd. v chaetae di2 absent. no cryptopygy, abd. vi visible from above (fig. 2f). ventral chaetotaxy. on head, groups vea, vem and vep with 3 chaetae each; group vi with 6 chaetae. on abd. v, 2+2 chaetae ag, chaetae vl and l’ present. male with thick and forked chaetae (“male ventral organ”) in groups vei and vec (abd. iv), and in fu (abd. iii; fig. 2b, d). legs. claw without internal tooth. on tibiotarsi, chaeta m present and chaetae b4 and b5 relatively short and pointed. remarks neanura deharvengi sp. nov. is most similar to n. pallida deharveng, 1979 and n. minuta gisin, 1963, both of which are also characterized by the presence of chaetae l’ on abd. v, the absence of chaetae ocp on the head and tubercles di on th. i. besides the unique characters of the new species, a reduction tubercle number of chaetae types of chaetae names of chaetae cl 4 ml me f g af 4 ml mi b d oc 2 ml mi ocm oca di 2 ml mi di1 di2 de 2 ml mi de1 de2 (dl+ l+so) 8 ml me mi dl5, l1, so1 dl4, l4 so3–5 table 1a. chaetotaxy of neanura deharvengi sp. nov.: cephalic chaetotaxy–dorsal side. tergites legs di de dl l scx2 cx tr fe t th.i 1 2 1 0 3 6 13 19 th.ii 3 2+s 2+s+ms 3 2 7 6 12 18 th.iii 3 2+s 2+s 3 2 8 6 11 18 sternites abd.i 2 2+s 2 2 vt: 4 abd.ii 2 2+s 2 2 ve: 5–6 ve1 present abd.iii 2 2+s 2 3 ve: 4 fu: 5 me, 0 mi abd.iv 2 1+s 3 4 vei: 2 vec:2 vel: 4 vl: 4 abd.v (2+2) 5+s ag: 2 vl: 1 l’:1 abd.vi 7 ve: 12–13 an: 2 mi table 1b. chaetotaxy of neanura deharvengi sp. nov.: postcephalic chaetotaxy. european journal of taxonomy 406: 1–16 (2018) 8 of cephalic chaetotaxy and a fusion of lateral tubercles on the head, these species can additionally be distinguished by the following features: coloration (whitish to pale bluish in n. deharvengi sp. nov. and n. pallida; dark bluish in n. minuta), chaetotaxy of head (chaetae a, c, e, dl1, dl3, so2, so6 and l2–3 absent in deharvengi sp. nov.; present in pallida and minuta), number of ordinary chaetae de on th. ii and iii (2, 2, respectively, in deharvengi sp. nov.; 2, 3 in pallida; 3, 4 in minuta), number of ordinary chaetae de on abd. i–iii (2 in deharvengi sp. nov.; 3 in pallida and minuta), number of chaetae di on abd. v (2+2 in deharvengi sp. nov.; 3+3 in pallida and minuta) and number of tubercles on penultimate abdominal segment (3 in deharvengi sp. nov.; 4 in pallida and minuta). the only other member of the genus besides n. deharvengi sp. nov. recorded to date from iran is n. muscorum (cox 1982; yahyapour 2012). these species are significantly dissimilar and differ in a number of characters including, e.g., cephalic chaetae ocp (absent in deharvengi sp. nov.; present in muscorum), tubercle di on th. i (absent in deharvengi sp. nov.; present in muscorum), the number of chaetae de on th. ii and iii (2+s, 2+s, respectively, in deharvengi sp. nov.; 3+s, 4+s in muscorum), male ventral organ (present in deharvengi sp. nov.; absent in muscorum), chaetae l’ on abd. iv (present in deharvengi sp. nov.; absent in muscorum), and the number of tubercles on abd. v (3 in deharvengi sp. nov.; 4 in muscorum). genus deutonura cassagnau, 1979 type species achorutes phlegraeus caroli, 1912. deutonura persica sp. nov. urn:lsid:zoobank.org:act:412b7b74-6689-4d39-a09a-871119ed0754 figs 3–4; table 2 diagnosis habitus typical of genus. dorsal tubercles present and well developed. body white, with 2+2 dark pigmented eyes. buccal cone short, labrum without ogival sclerifications. cephalic tubercles af and cl generally not fused. no granular area between chaetae a and b. head without chaetae o, c, e, dl3, l2 and l3. th. iii with 3+s chaetae de. tubercles di on abd. v not bilobed. cryptopygy present and strongly developed. male ventral organ present. etymology the specific epithet ‘persica’ refers to the historical name of iran, the terra typica of the new species. material examined holotype iran: adult ♀, on slide, mazandarn province, neka region, hezarjarib forest, 36°37′ n, 53°21′ e, dead wood, 28 mar. 2013, e. yoosefi lafooraki leg. (dibec). paratypes iran: 4 adult ♀♀, 2 adult ♂♂, 1 subadult ♂, 1 juvenile, on slides, collected from moss on tree, leaf litter, dead wood in a hole of persian ironwood (parotia persica), same collection data as for holotype (dibec). smolis a. et al., new species of neanura and deutonura from the middle east 9 http://zoobank.org/urn:lsid:zoobank.org:act:412b7b74-6689-4d39-a09a-871119ed0754 description body. length (without antennae): 0.49 to 1.50 mm (holotype: 0.85 mm). body slightly flattened. body colour white. 2+2 medium black eyes, in typical arrangement for genus. chaetal morphology. dorsal ordinary chaetae of five types: long macrochaetae (ml), short macrochaetae (mc), very short macrochaetae (mcc), mesochaetae and microchaetae. long macrochaetae thickened, subcylindrical, slightly arc-like or straight, narrowly sheathed, serrated, apically rounded or rarely, in lateral part of body, pointed (figs 3b, f–g, 4b). macrochaetae mc and mcc thickened, slightly fusiform, straight and apically rounded. mesochaetae similar to ventral chaetae, thin, smooth and pointed. microchaetae similar to mesochaetae, but distinctly shorter. s–chaetae of tergites thin, smooth and short, distinctly shorter than nearby macrochaetae (figs 3b, f–g, 4b–c). antennae. typical of genus. ant. i–ii with 7 or 11 ordinary chaetae, respectively. ant. iii with 5 s-chaetae (aoiii) and 17 ordinary chaetae: 4 d, 4 vi, 4 vc and 5 ve (fig. 3a, i). sensillum sgv long and thin. ant. iv dorsally with the complete set of chaetae: 8 s, or, i, 12 mou and 3 brs. s–chaetae of ant. iv of medium length, subequal and moderately thickened. apical vesicle distinct, trilobed (fig. 3c–d). ant. iv ventrally with complete set of chaetae (3 brs, 2 iv; ap: 8 bs and 5 mia; ca: 2 bs and 3 mia; cm: 3 bs and 1 mia; cp: 8 mia and 1 brs). dorsal chaeta d5 absent on ant. iii. mouthparts. buccal cone relatively short, with labral sclerifications non-ogival. labrum chaetotaxy: 2/2, 4 (fig. 3h). labium with four basal, three distal and three lateral chaetae, papillae x absent (fig. 3e). maxilla styliform, mandible thin and tridentate. dorsal chaetotaxy and tubercles. head without granular area between chaetae a and b. chaetotaxy of head strongly reduced (fig. 3b, f). elementary tubercles de and ee on head absent. cephalic chaetae a distinctly shorter than b. chaetae ocm and ocp of nearly equal length. chaetae de2 on head usually as mi, rarely as mcc. chaetae dl2 on head as mi or mcc (fig. 3b, f). chaetae di 3 on th. ii-iii free (fig. 3b). on th. iii, chaetae de2 longer than de3. on abd. i–iii chaetae de2 shorter than de3 (fig. 3g). chaeta di2 on abd. v as microchaeta, di3 as mi or mcc. cryptopygy strong, abd. vi hardly visible from above. ventral chaetotaxy. on head, groups vea, vem and vep with 4, 3, 4 chaetae respectively (fig. 3f); group vi with 6 chaetae. on abd. iv, furcal rudiment with 4 microchaetae (fig. 4a). tubercle l on abd. iv with 5–6 chaetae. male with thick and forked chaetae (“male ventral organ”) on even anal valves (abd. vi), in groups vei, vec and vel (abd. iv), and fu (abd. iii; fig. 4a). legs. claw without internal tooth. on tibiotarsi, chaeta m present and chaetae b4 and b5 relatively short and pointed. variability a complete fusion of the cephalic tubercles af and cl has been observed in a single specimen. nevertheless, based on the available material it is hard to assess definitively whether it is just an aberrant form or an example of morphological variation within the species. remarks using the recent published key of the genus (deharveng et al. 2015), deutonura persica sp. nov. seems to be most similar to d. plena (stach, 1951), a species described and only known from the western and eastern carpathians. these two taxa, however, can easily be distinguished using a number of characters: chaetae c and e on the head (absent in persica; present in plena), elementary tubercles de and ee on the head (absent in persica; present in plena), number of lateral chaetae (l+so) on the head (8 in european journal of taxonomy 406: 1–16 (2018) 10 fig. 3. deutonura persica sp. nov. a. dorsal chaetotaxy of ant. iii–iv. b. dorsal chaetotaxy of head and th. c. apical bulb, ventral view. d. apical bulb, dorsal view. e. chaetotaxy of labium. f. ventral and lateral chaetotaxy of head. g. dorsal chaetotaxy of abd. iii–vi. h. labrum. i. ventral chaetotaxy of ant. iii. scale bars: a, c–e, h–i = 0.01 mm; b, f–g = 0.1 mm. smolis a. et al., new species of neanura and deutonura from the middle east 11 persica; 10 in plena), number of prelabral chaetae (2 in persica; 4 in plena), number of lateral chaetae on the labium (3 in persica; 4 in plena), microchaetae on fu (present in persica; absent in plena), male ventral organ (present in persica; absent in plena) and relative length of chaetae de2 and de3 on abd. i–iii (de3 longer than de2 in persica; de3 shorter than de2 in plena). fig. 4. deutonura persica sp. nov. a. ventral chaetotaxy of abd. iii–vi, adult male. b. ml di1 of abd. v. c. sensillum of abd. v. scale bars: a–c = 0.1 mm. table 2a. chaetotaxy of deutonura persica sp. nov.: cephalic chaetotaxy–dorsal side. tubercle number of chaetae types of chaetae names of chaetae cl 4 mc me f g af 6 ml mc mi b a d oc 3 ml mi ocm, ocp oca (di+de) 4 ml mc or mcc mcc or mi di, de1 di2 de2 dl 5 ml mcc mcc or mi mi dl1, dl5 dl4 dl2 dl6 (l+so) 8 ml me mi l1, l4, so1 so3–6 so2 european journal of taxonomy 406: 1–16 (2018) 12 only one species of deutonura, d. decolorata (gama & gisin, 1964), has previously been recorded (cox 1982; shayanmehr et al. 2013) from iran. this species was described and is otherwise known only from the french alps (gisin 1964; porco et al. 2010). because of the fused cephalic tubercles af and cl, d. decolorata belongs to the small conjuncta species group, containing only 4 european species (deharveng 1979). we have detected a similar fusion of cephalic tubercles in a single individual of d. persica sp. nov. (see variability). in light of the known distribution of d. decolorata and the above observation, the record of d. decolorata from iran should probably be treated as highly uncertain. discussion the two new taxa described in this paper broaden and enrich our knowledge of the genera neanura and deutonura, especially with regard to their morphology and biogeography. for example, as mentioned in the introduction, the genus neanura, excluding the cosmopolitan n. muscorum, is primarily restricted to western and central europe. the discovery of n. deharvengi sp. nov. has shown that other species of the genus can be expected outside europe. in addition, the new species is characterized by a peculiar set of characters, including the absence of chaetae a on the head. although many genera and species of neanurinae are characterized by a more or less advanced degree of reduction of cephalic chaetotaxy, these processes usually do not include the mentioned chaetae. for instance, within the western palaearctic a similar reduction of chaetae a is observed exclusively in the genus bilobella caroli, 1912, belonging to paleonurini, another tribe of neanurinae (e.g., deharveng 1981; smolis & kaprus’ 2008). in contrast to the previous genus, the range of deutonura is notably broader as its members are distributed both in the western palaearctic (western, southwestern, and central europe, northern africa; 46 species) and in the eastern palaearctic (korea, japan and russian far east; 11 species). two species are exceptions to this general distribution pattern: d. frigida (yosii, 1969), which ranges from central siberia (the basin of the yenisei river) to north-eastern america (deharveng & weiner 1984; babenko & fjellberg 2006) and d. gibbosa porco, bedos & deharveng, 2010, which was probably introduced by humans to south africa (deharveng et al. 2015). in spite of the fact that more than 80% of the known members of the genus occur in the western palaearctic, the range of deutonura in the region is highly concentrated and localized. for example, there are no species of deutonura in such areas as the british isles, scandinavia, the balkan peninsula, crimea, asia minor, or the islands of the east mediterranean basin (e.g., rhodes, crete, cyprus). the absence of deutonura in northern europe is easy to understand, as the neanurinae fauna of these areas is generally impoverished. the rest of the present picture can probably be explained by incompleteness of our knowledge. nevertheless, some areas like greece (including crete and rhodes), albania, bulgaria, turkey, or the caucasus have been more or less well investigated for neanurinae diversity and many species belonging to the subfamily have been recorded table 2b. chaetotaxy of deutonura persica sp. nov.: postcephalic chaetotaxy. tergites legs di de dl l scx2 cx tr fe t th.i 1 2 1 0 3 6 13 19 th.ii 3 2+s 3+s+ms 3 2 7 6 12 18 th.iii 3 3+s 3+s 3 2 8 6 11 18 sternites abd.i 2 3+s 2 3 vt: 4 abd.ii 2 3+s 2 3 ve: 5 ve1 present abd.iii 2 3+s 2 3 ve: 4 fu: 4–5 me, 4 mi abd.iv 2 2+s 3 5–6 vei: 2 vec:2 vel: 4 vl: 4 abd.v (3+3) 4+s ag: 2 vl: 1 l’:1 abd.vi 7 ve: 11–12 an: 2 mi smolis a. et al., new species of neanura and deutonura from the middle east 13 there (e.g., cassagnau & péja 1979; ellis 1976; deharveng 1982a, 1982b; smolis & kuznetsova 2016). the observed gap in the known distribution of deutonura is striking, and the answer to this question can undoubtedly be pivotal to understanding its history and evolution. acknowledgments we are very grateful to two anonymous reviewers for their valuable suggestions and corrections to the manuscript. the work was financially supported by the institute of environmental biology, faculty of biological science, university of wrocław, poland (project no. 1076/ś/ibś/2017). references babenko a. & fjellberg a. 2006. collembola septentrionale. a catalogue of springtails of the arctic regions. kmk scientific press ltd, moscow. buşmachiu g. & deharveng l. 2008. neanurinae and 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[in persian with english abstract.] manuscript received: 23 march 2017 manuscript accepted: 1 august 2017 smolis a. et al., new species of neanura and deutonura from the middle east 15 https://doi.org/10.3897/zookeys.335.5491 https://doi.org/10.5962/bhl.part.80349 https://doi.org/10.5252/z2017n1a5 https://doi.org/10.5962/bhl.part.80440 https://doi.org/10.11646/zootaxa.4200.1.2 https://doi.org/10.11646/zootaxa.4092.2.11 https://doi.org/10.3897/zookeys.553.6009 https://doi.org/10.3897/zookeys.673.12084 published on: 22 february 2018 topic editor: gavin broad desk editor: kristiaan hoedemakers printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain. european journal of taxonomy 406: 1–16 (2018) 16 european journal of taxonomy 6: 1-10 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2012.6 www.europeanjournaloftaxonomy.eu 2012 • kjell a. johanson & hong-thai pham this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e 1 three new species of helicopsyche (trichoptera, helicopsychidae) from northern vietnam, with a key to helicopsyche species of vietnam johanson k.a. & pham h.-t. 2011. three new species of helicopsyche (trichoptera, helicopsychidae) from northern vietnam, with a key to helicopsyche species of vietnam. european journal of taxonomy 6: 1-10. http:// dx.doi.org/10.5852/ejt.2012.6 kjell a. johanson1 & hong-thai pham2 1swedish museum of natural history, entomology department, box 50007, se-104 05 stockholm. e-mail: kjell.arne.johanson@nrm.se (corresponding author) 2department of insect systematics, institute of ecology and biological resources, 18 hoang quoc viet, cau giay, ha noi, vietnam. e-mail: phamthai1976@yahoo.com abstract. three new species of helicopsyche siebold, 1856 are described from vietnam: helicopsyche melina sp. nov., helicopsyche meander sp. nov., and helicopsyche lamnata sp. nov. all species were described from melinh station for biodiversity in the me linh district of vinh phuc province. the species were collected mainly in malaise traps situated across a small stream surrounded by lowland forest. some individuals were also collected on light in traps situated at the stream bank. key words. vietnam, helicopsychidae, taxonomy, vinh phuc, stream. introduction with 270 described species the snail-case caddisfly genus helicopsyche siebold, 1856, ranks as the tenth largest genus in the order trichoptera. the genus has been recorded from all major faunal regions, except antarctica (johanson 1998) and has the highest species diversity southern latitude interval 5-30° and southern latitude interval 15-45° (johanson 1997). with 30 described helicopsyche species (johanson 1999), the 17,000 km2 large new caledonian grande terre has the highest density of species in the world. the larva of all species are reported to build dextrally coiled larval cases made from sand grains and glued together by silk. they live in permanent and preferably running water bodies, often in well-shaded forest streams and rivers. the 71 previously described helicopsyche species of the oriental region (morse 2011) are classified in two subgenera, galeopsyche johanson, 1998 and helicopsyche siebold, 1856 (johanson 1998). the first record of helicopsyche from vietnam was that of helicopsyche azwudschgal malicky, 1995, described from tam dao national park at 800-1,100 m altitude. schefter & johanson (2001) described three more species from the country: helicopsyche khemoiensis schefter & johanson, 2001 (from nghe an district), helicopsyche azunensis schefter & johanson, 2001, and helicopsyche dacklestensis schefter & johanson, 2001 (the two latter from gia lai district). malicky (2010) listed the following two additional species from vietnam: helicopsyche boniata malicky & chantaramongkol, 1992, helicopsyche admata malicky & chantaramongkol, 1992, both originally described from thailand. with the addition of the three new species recorded and described below, the http://creativecommons.org/licenses/by/3.0/ www.europeanjournaloftaxonomy.eu european journal of taxonomy 6: 1-10 (2012) 2 known diversity of helicopsyche in vietnam is nine species, only slightly lower than found in thailand (fourteen spp.). this research was executed to increase our knowledge about the taxonomic species diversity and morphological variation in the family helicopsychidae, as well as to increase our understanding of the biogeography of individual species in the group. material and methods the material was collected in light traps between 4-5th april and malaise traps between 4-12th april 2011 in the vinh phuc province, me linh district, at a stream near ngoc thanh village, melinh station for biodiversity. it comprises 357 male and 81 female helicopsyche; the males divided into three species while the females are not associated or determined. the specimens were identified to morphospecies without initial treatment of the genitalia. before illustrated the abdomen of the species were macerated in proteinasek for extraction of dna, and for a short period in hot 8% koh to remove remaining tissue. each abdomen was temporarily mounted in euparal on a microscope slide and illustrated using a drawing tube mounted on a leitz ortholux ii light microscope. the individual illustrations were scanned in 600 dpi gray scale and used as a digital background layer in adobe® photoshop® cs v.8.0. the final illustration was drawn using a wacom® sketchpad and digital pen. after illustration, upper layer was saved into final illustrations. the abdomen with genitalia was thereafter transferred into 80% ethanol in a micro-vial, together with the rest of the specimen. terminology for morphological characters mainly follows that of johanson (1998) and oláh & johanson (2008). the material is deposited in the following institutions: iebr = institute of ecology and biological resources, hanoi, vietnam nhrs = swedish museum of natural history, stockholm, sweden results helicopsyche melina sp. nov. fig 1 diagnosis helicopsyche melina sp. nov. is most similar to h. admata malicky & chantaramongkol, 1992, h. dacklestenensis schefter & johanson, 2001 and h. nastia malicky & melnitsky (melnitsky & malicky 2008) particularly due to the presence of four-segmented maxillary palps in combination with the nearly similar shape of the gonopods and tergum x in lateral view. the new species is easily distinguished from these by the presence of a well-developed transverse plate of segment ix located at mid-height of posterior part of the segment; the primary branch of each gonopod is produced anterad; and the secondary branch of each gonopod is long, straight, and oriented in right angle to the primary branch of the gonopods. etymology melina, named after the melinh station for biodiversity, near the type locality. johanson k.a. & pham h.-t., new helicopsyche from vietnam 3 material examined holotype ♂ vietnam: vinh phuc prov, me linh district, stream near ngoc thanh village, melinh station for biodiversity, 21°23'28.2"n, 105°42'46.7"e, 233 m, malaise trap 4-12 apr. 2011, loc#vn001, leg. k.a. johanson, p.h. thai & t.t. du (nhrs, alcohol) [dna voucher ip7]. fig. 1. helicopsyche melina sp. nov., ♂ holotype. a. abdominal sternum vi, lateral view. b. abdominal sternal process vi, ventral view. c. genitalia, lateral view. d. genitalia, dorsal view. e. genitalia, ventral view. f. phallus, lateral view. g. phallus, ventral view. european journal of taxonomy 6: 1-10 (2012) 4 paratypes ♂ vietnam: 8 ♂♂, vinh phuc prov, me linh distr, stream near ngoc thanh village, 21°23'40.1"n, 105°42'54.9"e, light trap 5 apr. 2011, loc#vn008, leg. k.a. johanson, p.h. thai & t.t. du (iebr, alcohol); 241 ♂♂, vinh phuc prov, me linh district, stream near ngoc thanh village, 21°23'48.4"n, 105°42'56.1"e, 186 m, malaise trap 5-12 apr. 2011, loc#vn006, leg. k.a. johanson, p.h. thai & t.t. du (nhrs, alcohol); 20 ♂♂, vinh phuc prov, me linh district, stream near ngoc thanh village, 21°23'48.4"n, 105°42'56.1"e, 186 m, malaise trap 5-12 apr. 2011, loc#vn006, leg. k.a. johanson, p.h. thai & t.t. du (iebr, alcohol); 34 ♂♂, vinh phuc prov, me linh distr, stream near ngoc thanh village, 21°23'38.9"n, 105°42' 50.7"e, light trap 4 apr. 2011, loc#vn002, leg. k.a. johanson, p.h. thai & t.t. du (nhrs, alcohol); 2 ♂♂, vinh phuc prov, me linh distr, stream near ngoc thanh village, 21°23'58.7"n, 105°42'49.8"e, light trap 5 apr. 2011, loc#vn007, leg. k.a. johanson, p.h. thai & t.t. du (nhrs, alcohol). description maxillary palps. four-segmented, each segment gradually shorter distally. pair of interantennal setal warts bean shaped, each about as long as width of individual scapus. each scapus about as long as each basal segment of maxillary palps. spur formula 1,2,4; each foreleg spur slightly shorter than each first tarsomere. length. forewing 2.8-3.6 mm, hind wing 2.3-2.9 mm (n = 10). male abdomen and genitalia. (fig. 1) sternal process vi about one-fifth as long as sternum vi, straight, oriented posteroventrad; slightly tapering in lateral view (fig. 1a); with slightly club-shaped apex in ventral view (fig. 1b). segment ix about as long as high in lateral view (fig. 1c); each anterior lobe of segment ix (fig. 1c) narrowly and symmetrically ellipsoid in lateral view, oriented anterad; anterodorsal and anteroventral margins almost straight (fig. 1c); in dorsal view, inner margin forming narrowly ellipsoid cavity (fig. 1d); in ventral view, without central posterior process (fig. 1e); inner margin forming nearly triangular cavity; lateral apodemes nearly horizontal (fig. 1c), widening anteriorly, meeting anterior margin, sub-marginal line absent; tergal transverse apodemes well-developed; sternal transverse apodeme absent. transverse plate of segment ix located at mid-height of posterior margin of segment ix; in lateral view almost club-shaped, curving ventrad (fig. 1c); in ventral view forming broadly heart-shaped plate above basal part of gonopods (fig. 1e). tergum x, in lateral view, oriented posterad (fig. 1c), almost straight, tapering along its length; apex strongly pointed (fig. 1d); in dorsal view (fig. 1d), deeply divided into pair of tapering, divergent branches, with about 3 pairs of equally long megasetae in apical group, starting opposite to apex of gonopods in lateral view (fig. 1c). superior appendages tubular (fig. 1d), slightly curving dorsally. primary branch of gonopods, in lateral view (fig. 1c), oriented dorsally, broadened distad to mid-height; anterior margin smooth, strongly concave; dorsal and posterior margins edged, undulating, posteroventral margin almost straight (fig. 1c); apices produced mesad into rounded lobes above basolateral part of tergum x. secondary branch of gonopods nearly as long as secondary branch of gonopods; slender, almost straight in lateral view, except apex slightly curving dorsad; with few setae; curving mediad in ventral view (fig. 1e). basomesal lobes absent. basal plate, in lateral view (fig. 1c), nearly straight, narrowing along its length, slightly produced beyond anteroventral margin of segment ix in lateral view (fig. 1c); narrow in ventral view (fig. 1e). phallic apparatus, lateral view, tube-shaped along its length (fig. 1f), narrowest shortly after basis, apex about as broad as base; nearly straight after mid-length; in ventral view, with basis narrow (fig. 1g); widest immediately after mid-length; endotheca weakly produced, posteroventral part not sclerotized, except pair of weakly sclerotized pair of triangular processes distally of phallic sclerite. johanson k.a. & pham h.-t., new helicopsyche from vietnam 5 helicopsyche meander sp. nov. fig. 2 diagnosis helicopsyche meander sp. nov. is most similar to h. lamnata sp. nov., h. anaktangga malicky, 1995, h. anaksaku malicky, 1995, h. minyas malicky & nawvong (malicky et al. 2004) and h. cymodoce schmid, 1993, particularly due to the presence of two-segmented maxillary palps in combination with the nearly similar shape of the gonopods in lateral view. the new species is distinguished from h. anaktangga, h. anaksaku, h. minyas and h. cymodoce by the pointed apex of tergum x in lateral view; and from h. lamnata sp. nov., h. anaksaku and h. minyas by the more strongly s-shaped gonopods. it is furthermore separated from h. lamnata sp. nov. by the absence of a central posterior process of segment ix. etymology meander, named after the shape of the gonopods, resembling a meandering river. material examined holotype ♂ vietnam: vinh phuc prov, me linh district, stream near ngoc thanh village, melinh station for biodiversity, 21°23'28.2"n, 105°42'46.7"e, 233 m, malaise trap 4-12 apr. 2011, loc#vn001, leg. k.a. johanson, p.h. thai & t.t. du (nhrs, alcohol) [dna voucher ip7]. description maxillary palps. two-segmented, with distal segment slightly longer than basal segment. pair of interantennal setal warts semi-spherical, elevated. each scapus about as long as each basal segment of maxillary palps. spur formula 1,2,4; each foreleg spur slightly shorter than each first tarsomere. lenth. forewing 3.7 mm, hind wing 2.8 mm (n = 1). male abdomen and genitalia. (fig. 2) sternal process vi about one-eighth as long as sternum vi, straight, oriented posteroventrad; slightly tapering in lateral view (fig. 2a); pointed in ventral view (fig. 2b). segment ix about 1.5 x taller than long in lateral view (fig. 2c); each anterior lobe of segment ix (fig. 2c) irregularly narrowing anteriorly in lateral view, asymmetrical, oriented anterad; anterodorsal margin almost straight; anteroventral margin concave (fig. 2c); in dorsal view, inner margin forming narrowly ellipsoid cavity (fig. 2d); in ventral view, without central posterior process (fig. 2e); inner margin forming wide, almost rectangular cavity; lateral apodemes nearly horizontal (fig. 2c), each parallel-sided along their length, meeting anterior margin; sub-marginal line present above lateral apodemes; tergal transverse apodemes present, fading immediately above superior appendages; sternal transverse apodeme present along posterior margin of segment ix (fig. 2c). transverse plate absent. tergum x, in lateral view, oriented posteroventrad (fig. 2c), slightly sigmoid, tapering towards mid-length, apex strongly club-shaped, slightly curving dorsally (fig. 2c); in dorsal view (fig. 2d), deeply and widely divided into pair of parallel-sided and parallel-running branches, with 3 pairs of equally long megasetae in apical group, starting well before apex of gonopods in lateral view (fig. 2c). superior appendages club-shaped (fig. 2c), oriented ventrad. primary branch of gonopods, in lateral view (fig. 2c), with basal part oriented posterad before strongly bending dorsad, angling posterad at half-length, apical part produced mesad above tergum x; almost parallel-sided along their length; anterior margin smooth, strongly concave; dorsal and posterior margins undulating. secondary branch of gonopods absent. basomesal lobes long, tuboid, each with pair of apical setae; sigmoid in lateral view (fig. 2d), parallel-sided with diverging apex in ventral view (fig. 2e). basal plate, in lateral view (fig. 2c), curving anteroventrad, narrowing along its length from posterior to anterior european journal of taxonomy 6: 1-10 (2012) 6 end, slightly produced beyond anteroventral margin of segment ix in lateral view (fig. 2c); widely triangular in ventral view (fig. 2e). phallic apparatus, lateral view, irregularly tube-shaped along its length (fig. 2f), apex about as broad as base; nearly straight after mid-length; in ventral view, with basis narrow (fig. 2g); widest after mid-length; endotheca weakly produced, posteroventral part not sclerotized, except single, weakly sclerotized triangular processes distally of phallic sclerite. fig. 2. helicopsyche meander sp. nov., ♂ holotype. a. abdominal sternum vi, lateral view. b. abdominal sternal process vi, ventral view. c. genitalia, lateral view. d. genitalia, dorsal view. e. genitalia, ventral view. f. phallus, lateral view. g. phallus, ventral view. johanson k.a. & pham h.-t., new helicopsyche from vietnam 7 helicopsyche lamnata sp. nov. fig. 3 diagnosis helicopsyche lamnata sp. nov. is most similar to h. vongsombathi johanson & malm, 2007, from which it is easily distinguished by the presence of a central posterior process of segment ix and the distal part of the gonopods are more strongly produced posteriorly. the species also resembles h. meander sp. nov., particularly by the nearly club-shaped apex of tergum x in lateral view, but in h. lamnata sp. nov. the vith sternal process is much longer and the gonopods are less sharply sigmoid. etymology lamnata, plate-like in latin, refers to the process on the posterior margin of sternite ix. material examined holotype ♂ vietnam: vinh phuc prov, me linh district, stream near ngoc thanh village, melinh station for biodiversity, 21°23'28.2"n, 105°42'46.7"e, 233 m, malaise trap 4-12 apr. 2011, loc#vn001, leg. k.a. johanson, p.h. thai & t.t. du (nhrs, alcohol) [dna voucher ip9]. paratype same data as holotype, except (iebr, alcohol). description maxillary palps. two-segmented, all segments sub-equal in length. pair of interantennal setal warts very large, spherical, elevated, densely covered by long setae. each scapus slightly longer than individual segments of maxillary palps. spur formula 1,2,4; each foreleg spur nearly as long as each first tarsomere. length. forewing 4.0 mm, hind wing 3.1 mm (n = 1). male abdomen and genitalia. (fig. 3) sternal process vi about two-fifths as long as sternum vi, straight, oriented posteroventrad; parallel-sided in lateral view (fig. 3a); almost parallel-sided in ventral view (fig. 3b). segment ix slightly taller than long in lateral view (fig. 3c); each anterior lobe of segment ix (fig. 3c) smoothly narrowing anteriorly in lateral view, asymmetrical, oriented anterodorsad; anterodorsal margin almost straight; anteroventral margin concave (fig. 3c); in dorsal view, inner margin forming narrowly ellipsoid cavity (fig. 3d); large central posterior process located below basis of gonopods, sharply triangular in lateral view (fig. 3c); in ventral view widely trapezoid, with slightly concave posterior margin. lateral apodemes nearly horizontal (fig. 3c), parallel-sided along their length, located at mid-height of segment ix, ending in sub-marginal line; tergal transverse apodemes absent; sternal transverse apodeme oriented in right angle to lateral apodeme, located along posterior margin of segment ix (fig. 3c). transverse plate absent. tergum x, in lateral view, oriented posteroventrad (fig. 3c), strongly tapering towards mid-length, with single, dorsolateral, stout seta at mid-length; distal half almost parallel-sided, apex slightly club-shaped, orienting posteroventrally (fig. 3c); in dorsal view (fig. 3d), deeply and narrowly divided into pair of parallel-sided and parallelrunning branches; each with 4 pairs equally long megasetae in apical group, and two pairs stout, posteradoriented spines (fig. 3c). superior appendages club-shaped (fig. 3c), oriented posteroventrad. primary branch of gonopods, in lateral view (fig. 3c), with basal part oriented posterad before strongly bending posterodorsad, curving posterad after mid-length, apical part rounded in dorsal view; produced mesad into pointed plates above tergum x in dorsal view (fig. 3d); slightly widening along their length; basal part of anterior margin smooth; dorsal and posterior margins undulating. secondary branch of gonopods european journal of taxonomy 6: 1-10 (2012) 8 absent. basomesal lobes long, slightly broadening before apex, each with pair of apical setae; straight in lateral view (fig. 3c), diverging in ventral view (fig. 3e). basal plate, in lateral view (fig. 3c), straight, strongly narrowing anteriorly towards mid-length, anterior half very slender, not produced beyond anteroventral margin of segment ix in lateral view (fig. 3c); almost parallel-sided in ventral view (fig. 3e), with rounded anterior apex. phallic apparatus, lateral view, irregularly tube-shaped along its length (fig. 3f), apex about as broad as base; nearly straight after mid-length; irregular in ventral view (fig. 3g); widest after mid-length; endotheca weakly produced, posteroventral part not sclerotized, except long, single, weakly sclerotized triangular processes distally of phallic sclerite. fig. 3. helicopsyche lamnata sp. nov., ♂ holotype. a. abdominal sternum vi, lateral view. b. abdominal sternal process vi, ventral view. c. genitalia, lateral view. d. genitalia, dorsal view. e. genitalia, ventral view. f. phallus, lateral view. g. phallus, ventral view. johanson k.a. & pham h.-t., new helicopsyche from vietnam 9 key to male helicopsyche from vietnam 1. maxillary palps each with two segments; gonopods without secondary branch...................................2 – maxillary palps each with four segments; gonopods with secondary branch.......................................6 2.(1) in genitalia, basomesal lobe absent (as in fig. 1c)....h. boniata malicky & chantaramongkol, 1992 – in genitalia, basomesal lobe present (fig. 2c)......................................................................................3 3.(2) in genitalia, basomesal lobe originated from ventral part of gonopod basis, resembling secondary branch of gonopods (as in schefter & johanson 2001, figs 5, 9)........................................................4 – in genitalia, basomesal lobe originated from mid-height of gonopod basis (fig. 2c)........................5 4.(3) in genitalia, segment ix about as long as high; tergum x slender along its length in lateral view; in ventral view basomesal lobes fused mesally into common basal plate; in dorsal view, tergum x shallowly divided apically...............................................h. khemoiensis schefter & johanson, 2001 – in genitalia, segment ix much higher than long; tergum x wide and posteriorly tapering in lateral view; in ventral view basomesal lobes separated; in dorsal view, tergum x deeply divided apically...... ...............................................................................................................h. azwudschgal malicky, 1995 5.(3) sternite vi with sternal process about two-fifths as long as segment vi (fig. 3a); posteroventral margin of segment ix produced posterad into large plate (figs 3c, e); gonopods slightly sigmoid (fig. 3c)...................................................................................................................h. lamnata sp. nov. – sternite vi with sternal process about one-seventh as long as segment vi (fig. 2a); posteroventral margin of segment ix not produced posterad (fig. 2e); gonopods strongly sigmoid, bending about 90° (fig. 2c)................................................................................................................h. meander sp. nov. 6.(1) in genitalia, dorsal branch of gonopods almost parallel-sided in lateral view.................................... .....................................................................................h. admata malicky & chantaramongkol, 1992 – in genitalia, primary branch of gonopods widening distally in lateral view (fig. 1c)......................7 7.(6) in genitalia, primary branch of gonopods produced anterad (fig. 1c)..............h. melina sp. nov. – in genitalia, primary branch of gonopods produced posterad.............................................................8 8.(7) in genitalia, primary branch of gonopods shorter than secondary branch of gonopods; each lateral branch of tergum x divided apically...............................................................h. angusta ulmer, 1951 – in genitalia, primary branch of gonopods longer than secondary branch of gonopods; each lateral branch of tergum x undivided apically..................................h. dacklestenensis schefter & johanson, 2001 acknowledgements this study was a collaboration between institute of ecology and biodiversity resources (iebs), hanoi, and the swedish museum of natural history, stockholm. we are particularly thankful to the director of iebr, prof. dr. le xuan canh, and vice director of iebr prof. dr. ta huy thinh for giving us the opportunity to perform this research. the present study was broadly supported by the basis project of the department of insect systematics, iebr, particularly by mr. thieu du tran who assisted us in the field. references johanson k.a. 1997. zoogeography and diversity of the snail case caddisflies (trichoptera: helicopsychidae). in: holzenthal r.w. & flint o.s. jr. (eds) proceedings of the 8th international symposium on trichoptera: 205-212. ohio biological survey, columbus, ohio. european journal of taxonomy 6: 1-10 (2012) 10 johanson k.a. 1998. phylogenetic and biogeographic analysis of the family helicopsychidae (insecta: trichoptera). entomologica scandinavica, supplement 53: 1-172. johanson k.a. 1999. seventeen new species of helicopsyche from new caledonia (trichoptera: helicopsychidae). tijdschrift voor entomologie 142: 37-64. johanson k.a. & malm t. 2007. three new helicopsyche from laos (trichoptera: helicopsychidae). zootaxa 1407: 13-22. malicky h. 1995a. neue köcherfliegen (trichoptera, insecta) aus vietnam. linzer biologische beiträge 27: 851-885. malicky h. 1995b. zwei neue helicopsyche (helicopsychidae) aus perak, malaysia. braueria 22: 4. malicky h. 2010. atlas of southeast asian trichoptera. faculty of science printing unit, chiang mai university, thailand. malicky h. & chantaramongkol p. 1992. neue köcherfliegen (trichoptera) aus thailand und angrenzenden landern. braueria 19: 13-23. malicky h., chantaramongkol p., bunlue p., changthong n., nawvong j., nuntakwang a., prommi t., thamsenanupap p. & thapanya d. 2004. 27 neue köcherfliegen aus thailand (insecta, trichoptera). linzer biologische beiträge 36: 287-304. melnitsky s. & malicky h. 2008. trichoptera from chang island, southeastern thailand, with the description of three new species. braueria 35: 25-27. morse j.c. (ed.) 2011. trichoptera world checklist. http://entweb.clemson.edu/database/trichopt/index. htm [accessed 19 jul. 2011.] oláh j. & johanson k.a. 2008. generic review of hydropsychinae, with description of schmidopsyche, new genus, 3 new genus clusters, 8 new species groups, 4 new species clades, 12 new species clusters and 62 new species from the oriental and afrotropical regions (trichoptera: hydropsychidae). zootaxa 1802: 1-248. schefter p.w. & johanson k.a. 2001. three new species of helicopsyche from vietnam (trichoptera: helicopsychidae). pan-pacific entomologist 77: 9-18. schmid f. 1993. considérations sur les helicopsychides (trichoptera, integripalpia). beaufortia 43: 65100. siebold c.t.e. von 1856. wahre parthenogenesis bei schmetterlingen und bienen. wilhelm engelmann, leipzig. ulmer g. 1951. köcherfliegen (trichopteren) von den sunda-inseln. teil i. archiv für hydrobiologie, supplement 19: 1-528. manuscript received on: 27 july 2011 manuscript accepted on: 16 november 2011 published on: 2 february 2012 topic editor: malcom scoble in compliance with article 8.6 of the iczn, printed versions of all papers are deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d'histoire naturelle, paris, france; national botanic garden of belgium, meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark. http://entweb.clemson.edu/database/trichopt/index.htm http://entweb.clemson.edu/database/trichopt/index.htm european journal of taxonomy 363: 1–20 issn 2118-9773 https://doi.org/10.5852/ejt.2017.363 www.europeanjournaloftaxonomy.eu 2017 · schneider c. & deharveng l. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:5720cf48-37a8-4814-93f3-192493488435 1 first record of the genus spinaethorax papáč & palacios-vargas, 2016 (collembola, neelipleona, neelidae) in asia, with a new species from a vietnamese cave clément schneider 1,* & louis deharveng 2 1 mécanismes adaptatifs & evolution, mecadev umr 7179 cnrs, mnhn, dpt systematics & evolution, muséum national d'histoire naturelle, cp50 entomology, 45 rue buffon, 75005 paris, france. 2 institut de systématique, evolution, biodiversité, isyeb – umr 7205 – cnrs, mnhn, upmc, ephe, muséum national d’histoire naturelle, sorbonne universités, 57 rue cuvier, cp 50, 75005 paris, france. *corresponding author: clement.schneider@mnhn.fr 2 email: dehar.louis@wanadoo.fr 1 urn:lsid:zoobank.org:author:c0bec337-0235-4e4b-8efb-134b3eed1b90 2 urn:lsid:zoobank.org:author:d8f5c679-c30c-442c-8621-d3b8edb17ef7 abstract. a new species of the genus spinaethorax papáč & palacios-vargas, 2016, recently erected for  two cave species of mexico, is described from a vietnamese cave. it differs from the mexican species most noticeably by the dorsal chaetotaxy of the head (number and morphology of chaetae), the shape of s-chaetae on the third antennomere, the dorsal chaetotaxy of the abdomen and the chaetotaxy of the dens. the pattern of special τ-chaetae is described for the first time in the genus. the affinities between  spinaethorax and the other genera of neelipleona are discussed. spinaethorax is probably closely related to neelus folsom, 1896. a table of the differential characters is provided for the three known species of spinaethorax. spinaethorax appears to be restricted to caves, but its presence in vietnam indicates that this genus has a much larger distribution than previously recognized. keywords. taxonomy, chaetotaxy, vietnam, cave fauna, springtail. schneider c. & deharveng l. 2017. first record of the genus spinaethorax papáč & palacios-vargas, 2016  (collembola, neelipleona, neelidae) in asia, with a new species from a vietnamese cave. european journal of taxonomy 363: 1–20. https://doi.org/10.582/ejt.2017.363 introduction neelipleona is one of the four orders of collembola (massoud 1971, 1976) and contains a single family, neelidae. the group remained little studied since the discovery of its first representative, neelus murinus folsom, 1896. a recent series of morphological and/or molecular contributions has greatly improved the systematics of neelipleona (kováč & papáč 2010; schneider et al. 2011, 2016; papáč &  kováč 2013; schneider & d’haese 2013; papáč et al. 2016), especially the genera neelus folsom, 1896 and megalothorax willem, 1900. schneider (2017) redescribed the generotypes of neelus, neelides caroli, 1912 and acanthoneelidus bretfeld & griegel, 2006 and gave an account of the morphological https://doi.org/10.5852/ejt.2017.363 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:5720cf48-37a8-4814-93f3-192493488435 mailto:clement.schneider@mnhn.fr mailto:dehar.louis@wanadoo.fr http://zoobank.org/urn:lsid:zoobank.org:author:c0bec337-0235-4e4b-8efb-134b3eed1b90 http://zoobank.org/urn:lsid:zoobank.org:author:d8f5c679-c30c-442c-8621-d3b8edb17ef7 https://doi.org/10.582/ejt.2017.363 european journal of taxonomy 363: 1–20 (2017) 2 knowledge on the entire neelipleona. this latter work, however, did not include first-hand observations of  zelandothorax delamare deboutteville & massoud, 1963 or of the recently erected genus spinaethorax papáč & palacios-vargas, 2016. spinaethorax was created by papáč & palacios-vargas (2016) to accommodate two cave species of mexico, megalothorax spinotricosus palacios-vargas & sánchez, 1999 and m. tonoius palacios-vargas & sánchez, 1999. papáč & palacios-vargas (2016) defined  spinaethorax with the following character set: presence of sensory fields (shared with all genera, except neelides), sword-like macrochaeta on oral fold (unique), 6 + 6 chaetae on basomedian fields of labium (unique), fusion of antennomeres iii/ iv (shared with megalothorax and zelandothorax), 3 + 3 teeth on tenaculum (shared with neelus and some megalothorax), 1 + 1 neosminthuroid chaetae on sternum of abdomen iv (shared with neelus and acanthoneelidus), two chaetae on proximal part of dens (shared with adults of most neelus species), absence of chaeta e3 on distal part of dens (in adult; shared with megalothorax, zelandothorax and acanthoneelidus) and three chaetae close to the abdominal sensory field (unique). collecting carried out during a long-term biodiversity assessment of the hon chong karst in south vietnam led to the discovery of a new species, spinaethorax adamantis sp. nov. the present paper includes a thorough morphological description of spinaethorax adamantis sp. nov. and a review of the definition of the genus spinaethorax. the pattern of τ-chaetae (schneider et al. 2016) is for the first time  described in a species of spinaethorax. additional information is given on the morphological affinities  of spinaethorax with the other genera of neelipleona. homologies between the genera are suggested for certain morphological characters. a table of the differential characters between the three known species of spinaethorax is provided. material and methods environmental context the new species was collected in a small cave of a karstic hill of the hon chong karst system in southern vietnam, close to the border with cambodia. this karst, an exceptionally rich hot-spot of endemism, is currently under critical threat due to limestone exploitation (deharveng et al. 2009). the cave hang kim cuong itself is small, 67 m long (laumanns 2011), developed at the foot of a rocky hill close to the seaside. the species was obtained by berlese extraction of thin debris spread irregularly on the floor,  but was not found in the small patches of guano present in the cave. this dark, moderately humid cave hosts a remarkable fauna when visited in 2006, including numerous collembola, some of which are apparently undescribed: acrocyrtus sp., ceratophysella sp., folsomides sp. near f. americanus denis, 1931, lepidonella lecongkieti deharveng & bedos, 1995, megalothorax laevis denis, 1948, a blind rambutsinella sp. and several new species of pronura and willemia. sampling and observations the specimens were extracted from the sample with a simple berlese funnel and stored in 96 % ethanol. they were cleared in lactic acid and mounted on microscope slides in marc andré ii medium. they were examined with a leica dmlb differential phase contrast microscope with magnifications of ×250  to ×1000. drawings were made with a drawing tube and vectorized with inkscape. characters of s. spinotricosus and s. tonoius are based on the descriptions of palacios-vargas & sánchez (1999) and papáč & palacios-vargas (2016). measurements were performed with a graduated slide. values given  in µm were measured on a single specimen chosen among the biggest, ratios and proportions are average measures from several relevant specimens. schneider c. & deharveng l., first record of spinaethorax in asia 3 terminology and conventions the terminology follows schneider & d’haese (2013), schneider et al. (2016) and schneider (2017). chaetal categories used in the description are defined in schneider (2017). when referenced, neelus murinus is always sensu massoud & vannier (1967). the genus names neelus and neelides will not be abbreviated for their respective species in order to avoid ambiguity. abbreviations and symbols in text antenna ant. i–iv = antennomere i to iv s1–s5 = s-chaetae of ant. iii or = ant. iv organite s, sx, sy = s-chaetae of ant. iv head and trunk abd. i–vi = abdominal segment i to vi av = chaetae of anal valves s1, s2, s3, s3’ = s-chaetae on trunk sf1–6 =  sensory fields 1 to 6 and associated wax rod secretory element th. i–iii = thoracic segment i to iii τ  =  τ-chaetae wrc1–wrc8 = free wax rod secretory elements 1 to 8 claw la, lp, bp = auxiliary lamellae of unguis institutional abbreviations hcmu = ho chi minh city national university, department of biology, university of natural sciences, ho chi minh city, vietnam mnhn = muséum national d’histoire naturelle, paris, france results class collembola lubbock, 1873 order neelipleona massoud, 1971 family neelidae folsom, 1896 genus spinaethorax papáč & palacios-vargas, 2016 figs 1–8, 9a, d; table 1 revised diagnosis eyes and post-antennal organ absent. secondary granulation made of fine grains; dermastra missing;  integumentary channels absent. anterior process of labrum absent or very weak. ant. iii and iv fused; s-chaeta s1 short and wide; s-chaeta s5 present; two apical sensory rod-like chaetae (a, sa). basomedian fields of labium with 4 + 4 chaetae; basolateral fields of labium with 3 + 3 chaetae. oral fold with  enlarged lateral chaeta. sensory fields well-developed with swollen inner chaetae; sf6 with three guard chaetae. trunk with 4 + 4 s-chaetae (s1, s2, s3, s3’). abd. iv sternum with 1 + 1 neosminthuroid chaetae. distal articular process of manubrium with convex tip. dens without median articular process. unguis with basal narrowing; with well developed lamella bp; lamella la missing on unguis ii. tenaculum with 3 + 3 teeth. european journal of taxonomy 363: 1–20 (2017) 4 keys or tables for determination of the genera of neelidae are available in papáč & palacios-vargas  (2016) and schneider (2017). spinaethorax adamantis sp. nov. urn:lsid:zoobank.org:act:b0f131ab-279a-4890-a33b-ed70d437828d figs 1–8, 9a, d; table 1 diagnosis postero-dorsal area of head with 12 + 12 chaetae including 5 + 5 lanceolate chaetae. antero-dorsal area of head with two unpaired chaetae. ant. iii with 14 chaetae; s-chaetae s2, s3, s4 long and tubular. dorsal area of th. ii with 4 + 4 thickened or lanceolate chaetae. abd. i–v terga without spine-like microchaetae. abd. vi tergum with thickened chaetae. subcoxa 2 iii and coxa iii with thickened chaetae. dens proximal part with one strong chaeta. differential diagnosis spinaethorax adamantis sp. nov. differs from s. spinotricosus and s. tonoius in several characters, summarized in table 1. most notably, s. adamantis sp. nov. has only one proximal chaeta on dens (instead of two in the other species), long and tubular s-chaetae s2, s3 on ant. iii (vs short in the other species) and a long and tubular s-chaetae s4 (vs short and globular in the other species). s. adamantis sp. nov. and s. tonoius differ most strikingly from s. spinotricosus by the absence of the numerous spinelike microchaetae (reduced number of ordinary chaetae instead). fig. 1. spinaethorax adamantis sp. nov., habitus, with enlargement of the protuberance bearing wax rod secretory element of sf6. http://zoobank.org/urn:lsid:zoobank.org:act:b0f131ab-279a-4890-a33b-ed70d437828d schneider c. & deharveng l., first record of spinaethorax in asia 5 etymology the name is derived from the latin adamas (diamond) and is inspired by the local name for the cave hang kim cuong, “diamond cave”, in which the new species was collected. the name emphasizes that one of its richnesses lies in the biodiversity instead of actual diamonds. material examined holotype vietnam: ♂, on slide (mnhn-ea040281), kien giang province, kien luong, hon chong karst, nui hon chong hill (also called nui chua hang), hang kim cuong (diamond cave), 28 nov. 2006, x=104.639784 e, y=10.137582 n, elevation about 15 m, extracted with berlese funnel from soil in the dark zone of the cave, l. deharveng and a. bedos leg. (sample vn06-088) (mnhn). paratypes vietnam: 3 ♀♀, 1 ♂, 1 specimen of unknown sex on 3 slides, same data as the holotype, three paratypes deposited in the collection of the mnhn (france), two paratypes in the collection of the hcmu (vietnam). description size. length from labrum to anus of largest specimen ~550 µm. pigmentation. whitish in alcohol. general shape and body segmentation (fig. 1). orthognathous. antennae directed laterally with a slight curve toward the anterior direction, slender. ant. iii and iv fused. total length of antenna ~ 148 µm; max width (ant. iii) ~ 18 µm; ratio width : length ~ 0.12; relation of length of ant. i : ant. ii : ant. iii : ant. iv 0.15 : 0.5 : 1 : 1. point of insertion of head on th. i aligned with antero-posterior axis of the head. diameter of th. i slightly reduced without steep curve between th. i–head and th. i–ii. th. i in one part, neck missing. th. i–head articulary folds not observed. terga from th. ii to abd. v fused. posterior granule t of the dorso-median line missing, replaced by a small unpaired patch devoid of secondary granules (t, fig. 4a). wax rod secretory elements of sf3 (dorsal area of th. ii) and of sf6 (dorsal area of abdomen) standing on strong, protruding processes (fig. 1). precoxal area of th. ii and iii without clear anterior lobe. each coxa long and bent. on leg i and ii length of coxa > tibiotarsus > femur > trochanter. on leg iii length of coxa > tibiotarsus ~ femur > trochanter. abd. vi sternum clearly separated posteriorly from abd. vi tergum by large anal aperture, separated anteriorly from abd. v sternum by a steep curve, abd. vi sternum distinct laterally from the great abdomen terga by the absence of secondary granulation. abd. iv sternum enlarged, creating a large ventro-posterior side bearing the furca. abd. i–iii sterna reduced with tenaculum close to ventral tube. manubrium/dens articular processes: manubrium with weak process bearing hardened peg with convex tip that connects to concave tip of strong, projecting process of dens (fig. 6c). dens proximal part wider than the distal part with a steep curve between the two. no suture line or articular process between proximal and distal part of dens (fig. 6d). relation of length of manubrium : dens proximal part : dens distal part : mucro ~ 0.90 : 0.32 : 1 : 0.82. integument. with a simple secondary granulation made of fine granules. secondary granulation present  on posterior part of th. ii to abd. v terga (fig. 4a), absent on head, antennae, abd. vi tergum, furca, legs, thoracic and abdominal sterna. dermastra missing. integumentary channels missing. chaetae. seven morphological variations of chaetae on head and trunk: (i) ordinary chaetae; (ii) s-chaetae, short and swollen chaetae (s1–s3, s3’) found on trunk; (iii) τ-chaetae, long and flexible chaetae shaped as  trichobothria with basal ring implanted in a small depression of the integument and weakly contrasted in light microscopy, found on trunk (~30–35 µm, fig. 4b); (iv) wax rod secretory crypts associated with european journal of taxonomy 363: 1–20 (2017) 6 ta bl e 1. d if fe re nt ia l c ha ra ct er s fo r th e th re e sp ec ie s of th e ge nu s sp in ae th or ax . s . t on oi us a ft er p al ac io sv ar ga s & s an ch ez ( 19 99 ) an d pa pá č  &  p al ac io sv ar ga s  (2 01 6) , s . s pi no tr ic os us a ft er  p ap áč  &  p al ac io sv ar ga s  (2 01 6) . s. a da m an tis s p. n ov . s. to no iu s s. s pi no tr ic os us h ea d n um be r o f c ha et ae : p os te ro 12 +1 2 (5 +5 la nc eo la te ) / 3 +3 12 +1 2 (1 +1 la nc eo la te ) / 2 +2 10 +1 0 (4 +4 la nc eo la te ) / 2 +2 do rs al a re a / p os te ro -l at er al a re a n um be r o f c ha et ae : a nt er odo rs al 10 +1 0 + 2 un pa ir ed / 5+ 5 ? / ? 9+ 9 + 1 un pa ir ed / 2+ 2 ar ea / an te ro -l at er al a re a l an ce ol at e ch ae ta e re la tiv e si ze il 1 > il 2 ~ il 4 ~ il 5 > il 3 n /a il 1 > il 2 > il 4 > il 5 a nt en na a nt . i ii 14 c ha et ae (6 b el ow s 4) , 12 c ha et ae (5 b el ow s 4) , 15 c ha et ae (7 b el ow s 4) , s2 , s 3, s 4 lo ng a nd tu bu la r s2 a nd s 3 sh or t, s4 g lo bu la r s2 a nd s 3 sh or t, s4 g lo bu la r a nt . i v 14 c ha et ae (i nc lu di ng 5 w ith s lig ht 11 c ha et ae , 9 s -c ha et a s 13 c ha et ae , 1 3 sch ae ta e s   m od ifi ca tio n) , 1 2  sch ae ta e  s tr un k t h. ii d or sa l a re a— nu m be r o f 4+ 4 ?( 3+ 3– 4+ 4) 6+ 6 th ic ke ne d or la nc eo la te c ha et ae a bd . i –v te rg a ch ae to ta xy ra ng in g fr om 2 7+ 27 to 3 1+ 32 c ha et ae ?( >2 5 + 25 c ha et ae ) ?( 3 + 3 ch ae ta e) + >8 0 sp in elik e m ic ro ch ae ta e a bd . v i t er gu m — ch ae ta l m or ph ol og y th ic ke ne d ? th in le gs l eg i (s ub co xa 1 –t ib io ta rs us ) 1, 1 , 1 , 4 , 9 , 1 3 1, 1 , ? , 3 , 8 , 1 2 1, 1 , 1 , 4 , 8 , 1 3 l eg ii (s ub co xa 1 –t ib io ta rs us ) 1, 1 , 1 , 3 , 8 , 1 4 1, 1 , ? , 2 , 7 , 1 2 1, 1 , 1 , 3 , 8 , 1 4 l eg ii i ( su bc ox a 1– tib io ta rs us ) 2, 1 , 2 , 4 , 8 , 1 3 1, 1 , ? , 3 , 7 , 8 3, 1 , 2 , 3 , 8 , 1 3 su bc ox a 2 ii i a nd c ox a ii i th ic ke ne d ? or di na ry ch ae ta e m or ph ol og y u ng ui s ii i la a bs en t ?l a ab se nt la p re se nt f ur ca pr ox im al  p ar t o f d en s,   1  2  (a ft er  p ap áč  &   2 nu m be r o f c ha et ae (a du lt) pa la ci os -v ar ga s 20 16 ) schneider c. & deharveng l., first record of spinaethorax in asia 7 fig. 2. spinaethorax adamantis sp. nov. a–b. chaetotaxy of the head. a. dorsal side. b. ventral side, anterior part. c. labrum dorsal side. d–f. labium. d. focus on papillate chaetae of the palp. e. focus on chaetae in proximal field. f. focus on papilla h. g–h. maxilla, different views. i. left mandible. j–k. maxilla outer lobe. j. lateral side. k. ventral side. abbreviations: blf = basolateral field of labium;  bmf = basomedian field of labium; mol = maxilla outer lobe; of = oral fold. labial palp lettering after fjellberg (1999). european journal of taxonomy 363: 1–20 (2017) 8 a sensory field; (v) free wax rod secretory crypts (wrc1–8); (vi) inner swollen chaetae of sensory fields;  (vii) neosminthuroid chaetae. ordinary chaetae size ranging from microchaetae [1–6 µm], mesochaetae [7–15 µm] to macrochaetae [16 µm–42 µm]. ordinary chaetae morphology ranging from simple to lanceolate; including spectacular lanceolate macrochaetae on head and thorax (figs 2a, 4a). sensory fields and wax rods. six pairs of sensory fields (sf1–6) on head and trunk, each field associated  with a wax rod secretory crypt (figs 2a, 4a). on head: sf1 preantennal without special inner chaeta (fig. 2a); sf2 postantennal without special inner chaeta (fig. 2a). on trunk: sf3 dorsally on th. ii with three short candlelight-shaped inner chaetae (fig. 4a); sf4 in precoxal area of th. ii with two short candlelight-shaped inner chaetae (fig. 4a); sf5 in precoxal area of th. iii and sf6 dorsally on posterior part of abdomen, each with a short candlelight-shaped inner chaeta (fig. 4a). sf6 well developed (fig. 4a). eight pairs of free wax rod secretory crypts (seven in th. iii tergum area, eight in abdominal terga area) (fig. 4a). edges of sf5 and wrc5 separate, wrc8 in antero-ventral position and close to sf6 (figs 4a, 9d). tertiary wax rod secretory elements apparently missing. labrum. labrum without clearly differentiated anterior process, a-row of chaetae very close to labral ridge (fig. 2c). a-row consisting of two pairs of smooth, slightly thickened and strongly curved mesochaetae (a1, a2). m- and p-rows of chaetae each with five mesochaetae (fig. 2c). labral ridge without apical hairs or spines (fig. 2c). labium.  basomedian  fields  with  4  +  4  mesochaetae  (fig. 2b,  d–e).  basolateral  fields  with  3  +  3  mesochaetae (lateral chaetae on weak tubercles) and with small antero-lateral integumentary processes (fig. 2b, d–e). proximal field with four mesochaetae (fig. 2e). palp with six papillate chaetae and six guard hairs as (h, h1, h2), (a), (b), (b2, b4), (c), (d, d2), (e, e) (lettering sensu fjellberg 1999). hypostomal papilla (h) with a strong, apically enlarged chaeta, other papillae with acuminate, straight or slightly curved chaetae (fig. 2d, f). other mouthparts. oral fold with two chaetae and a microspine, ventral chaeta enlarged and strongly curved near the base, lateral chaeta of ordinary morphology (fig. 2a–b, j). maxilla outer lobe with sub-basal mesochaeta and apical papilla bearing a mesochaeta (fig. 2j–k), apical papilla without annex hair (fig. 2j–k), sublobal plate with strong hair (fig. 2j–k). left and right mandibles each with four apical, rounded teeth (fig. 2i). maxilla with at least four cilliated lamellae, capitulum with two hooked teeth (fig. 2g–h). head chaetotaxy. postero-dorsal area with 12 + 12 chaetae: 5 + 5 macrochaetae, 6 + 6 mesochaetae in f.p–pa.a-rows, 1 + 1 mesochaetae on the border of sf2 (figs 2a, 7a). postero-lateral area with 3 + 3 mesochaetae (figs 2a, 7a). antero-dorsal area with 10 + 10 mesochaetae and two unpaired mesochaetae (figs 2a, 7a). antero-lateral area with 1 + 1 macrochaetae and 4 + 4 mesochaetae (figs 2a, 7a). ventral area with 3 + 3 post-labial mesochaetae (figs 2b, 7b). antennal chaetotaxy. ant. i with three dorsal chaetae (fig. 3a). ant. ii with six chaetae in two whorls, one in basal whorl, five in apical whorl (fig. 3a–b). ant. iii with 14 chaetae in 4 rough whorls distributed as 1, 2, 4, 7 (fig. 3a–b), of varied morphology, ranging from thin ordinary chaetae to “s-chaetae s-like”; five s-chaetae (s1–s5). s1 as a plump, stalked chaeta, possibly with four folds (fig. 3a, d), s2–s4 long and tubular with blunt apex, s5 short with pointed apex (fig. 3a–b). ant. iv with 14 chaetae, 14 s-chaetae (12 s, sx, sy), one organite (or) and two apical and subapical sensory rods (a, sa) (fig. 3a–c). five chaetae slightly departing from the ordinary morphology in a large and contrasted basal ring, a subtly more tubular aspect (vs conical aspect) and outward curving (vs inward curving) (fig. 3a–b). s-chaetae s scarcely differentiated from ordinary macrochaetae, with apex finely  rounded. sy thick and tubular with round apex, sx as long as sy but slender with round apex (fig. 3a, c). schneider c. & deharveng l., first record of spinaethorax in asia 9 organite apparently bifid with two thin arms apically converging, one perceptibly larger than the other  (fig. 3a, c). ornamentation of s-chaetae not seen with light microscopy. fig. 3. spinaethorax adamantis sp. nov. a–d. chaetotaxy of the antenna. a. dorsal side. b. ventral side. c. focus on s-chaeta sx and organite or with enlargement of the latter. d. s-chaeta s1, different aspects. asterisks indicate modified chaetae. chaetae implanted on the other side represented with gray stroke.  european journal of taxonomy 363: 1–20 (2017) 10 th. ii–abd. v terga chaetotaxy. th. ii–abd. v terga with a minimum of 52 + 52 chaetae, 12 + 12 τ-chaetae, 4 + 4 s-chaetae (s1, s2, s3, s3’) and 8 + 8 wrc (figs 4a, 7c). dorso-lateral areas of th. ii with 5 + 5 macrochaetae, 4 + 4 mesochaetae and 2 + 2 τ-chaetae (figs 4a, 7c), lengths of the 5 macroand 4 mesochaetae 13–40 µm: a4 (13 µm)<a1, p1, p4<a2<p3<a5, p2<a3 (40 µm); a2, a3, a5 and p2 thickened. precoxal areas of th. ii with 5 + 5 macrochaetae, 1 + 1 s-chaetae s1 and 1 + 1 τ-chaetae (figs 4a, 7c); lengths of chaetae a8, a9 (22 µm) <p7<p8<a7 (42 µm); a7, p7 and p8 thickened. dorso-lateral areas of th. iii–anterior abdomen with 16 + 16 chaetae, 2 + 2 s-chaetae (s3, s3’) 5 + 5 wrc (wrc1–4, wrc7) and 7 + 7 τ-chaetae (figs 4a, 7c); chaetae consisting of 1 + 1 macrochaetae (p7, 35 µm), the others as mesochaetae; p2 and p3 slightly thickened, p7 strongly thickened; wrc3 thickened. precoxal areas of th. iii with 3 + 3 chaetae, 2 + 2 wrc (wrc5–6) and 1 + 1 τ-chaetae (figs 4a, 7c); chaetae including 2 + 2 thickened macrochaetae (a8–45 µm, p8–22 µm); wrc5 thickened. posterior area of the abdomen (until abd. v included) with chaetae in variable numbers and tendency to asymmetrical distribution (range seen 19 + 19 to 23 + 24 chaetae), 1 + 1 s-chaetae (s2) and 1 + 1 wrc (wrc8) (figs 4a, 7c); chaetae including 8 + 8 macrochaetae (17–34 µm) and 13 + 13 mesochaetae. s1 tubular, s2 bean-shaped (fig. 4c), s3, s3’ light bulb-shaped (fig. 4a). leg chaetotaxy. leg i (fig. 5a) subcoxa 1, subcoxa 2 and coxa each with one chaeta; trochanter with four chaetae (2 proximal, 2 distal); femur with nine chaetae; tibiotarsus with 13 chaetae in five rough  whorls distributed as 2, 2, 1, 3, 5; one of the apical anterior chaetae slightly different (weak contrast, thin and with subtly blunt apex). leg ii (fig. 5b) subcoxa 1, subcoxa 2 and coxa each with one chaeta; trochanter with three chaetae; femur with eight chaetae; tibiotarsus with 14 chaetae in four rough whorls distributed as 3, 4, 3, 4. leg iii (fig. 5c) subcoxa 1 with two chaetae; subcoxa 2 with one chaeta; coxa with two chaetae in proximal position; trochanter with four chaetae; femur with 8 chaetae; tibiotarsus with 13 chaetae in four rough whorls distributed as 2, 4, 3, 4. each pretarsus with two microchaetae, one fig. 4. spinaethorax adamantis sp. nov. a. chaetotaxy of the trunk. b. focus on the τ-chaetae (asterisks  link τ-chaetae in a and b). c. focus on s-chaeta s2. d. focus on the neosminthuroid chaeta. secondary granulation of th.ii–abd.v terga partially shown (anterior, ventral and posterior limits, also limits near sensory fields and wrc1–8). schneider c. & deharveng l., first record of spinaethorax in asia 11 fig. 5. spinaethorax adamantis sp. nov. a–c. chaetotaxy of legs. a. leg i with focus on apical chaetae of tibiotarsus, asterisk indicates a special chaeta. b. leg ii (also with sf4 on th. ii precoxal area). c. leg iii (also with sf5 on th. iii precoxal area). d–g. morphology of claws. d. claw i posterior side. e. claw i anterior side. f. claw ii posterior side. g. claw iii posterior side. european journal of taxonomy 363: 1–20 (2017) 12 on anterior side and one on posterior side, the anterior chaeta larger than the posterior chaeta (fig. 5d– g). claws. ratio unguis length: pretarsus width on leg i–iii, respectively ~3.5, 3.3, 2.8. ratio of unguiculus: unguis for claw i, ii, iii ~0.5, 0.55, 0.45 (fig. 5d–g). unguis with basal narrowing (fig. 5d–g). on claw i lamella la moved in anterior position, shaped as a thin spine (fig. 5d); on claw ii and iii lamella la missing (fig. 5f, g). on each claw lamellae lp and bp well developed (fig. 5d–g). fig. 6. spinaethorax adamantis sp. nov. a–b. male sexual apparatus. a. male genital plate (asterisk indicates a chaeta not seen on other side, possibly unpaired). b. immature male genital plate. c. manubrium/ dens anterior articular process. d. chaetotaxy of manubrium and dens posterior side. e. distal part of dens and mucro anterior side. f. mucro posterior side. g–h. tenaculum. h. focused on apical teeth and basal process. i. ventral tube lateral side. schneider c. & deharveng l., first record of spinaethorax in asia 13 on unguiculus basal tubercle reduced to very small posterior integumentary process, lamella of the unguiculus with slender, conical aspect, without perceptible crests and with smooth edges reduced to apical part (fig. 5d–g). ratio of unguis length : tibiotarsus length on leg i–iii ~0.38, 0.4, 0.4. abd. vi and genital chaetotaxy. abd. vi tergum with 3 + 3 macrochaetae, two unpaired mesochaetae (a0, av) and 1 + 1 microchaetae (µ.av) (figs 4a, 7c). abd. vi sternum with 5 + 5 chaetae (including av), 2 + 2 microchaetae µ.av and 1 + 1 small basal rings without visible chaetal element (figs 4a, 7c). no clear delimitation between anal valves and rest of abd. vi tergum and sternum. male genital plate either with 6 + 6 circumgenital chaetae or 5 + 5 and one unpaired circumgenital chaetae and 7 + 7 eugenital chaetae (fig. 6a). immature male observed with 8 + 8 or 9 + 9 chaetae or microspines (fig. 6b). female genital plate not observed. abd. iv sternum and furca. abd. iv sternum with 4 + 4 chaetae and 1 + 1 neosminthuroid chaetae (fig. 4a). neosminthuroid chaetae flame-shaped, with very minute teeth (fig. 4a, d). manubrium with 4 + 4 chaetae (fig. 6d). posterior side of dens with a strong proximal chaeta with minute external teeth, two subdistal conical spines with basal ring, a distal chaeta and two lobe-like distal spines without basal ring; anterior side distally with small acute spine and two lobes (blunt spines) without basal ring (fig. 6d–e). mucro with thin lamellae, posterior lamellae serrate (15–17 teeth), apex rounded (fig. 6f–g). tenaculum and ventral tube. tenaculum with 3 + 3 large teeth, basal tubercle produced as 1 + 1 well-developed processes (fig. 6g–h). ventral tube with 2 + 2 apical chaetae, without clear posterior lobe (fig. 6i). ecology and distribution the presence of a species of spinaethorax in vietnam indicates that this genus has a much larger distribution than thought when it was described. the species is likely to be cave-restricted, in spite of its weak troglomorphic characters (the claws appear thinned in their distal part, but this may be phyletic rather than adaptive), as we never encountered it in the numerous surface samples we examined in this small region of southern vietnam. spinaethorax may have been overlooked because of its cave habitat, but also because it could be a rare species inside caves. the presence of the genus was confirmed in  two other caves of the hon chong karst (one cave yielding a species new to science, the other yielding juveniles of an unidentified species). many caves sampled in southeast asia contained unidentified  megalothorax, but spinaethorax remains so far restricted to the hon chong karst. however, thorough re-examination of cave neelidae from the region as well as new sampling may change the figure. general discussion threat to the biodiversity of hang kim cuong a visit to this cave in october 2014 did not allow to retrieve the undescribed species of the genera ceratophysella, folsomides, pronura and willemia, probably in relation with complete trampling of the whole floor of the cave by visitors, and disappearance of guano patches. s. adamantis sp. nov. was however recovered. morphology integument the genus spinaethorax differs from neelides, neelus, acanthoneelidus and megalothorax in its integumentary characters. the complete absence of integumentary channels is probably unique in neelipleona (this character was, however, not studied in zelandothorax) and the absence of dermastra separates it from the genera neelus and neelides. the functions of those two characters are unknown. european journal of taxonomy 363: 1–20 (2017) 14 their absence in spinaethorax (only cave-dwelling species known) and presence in neelus (both in troglobitic and edaphic species) suggest the subterranean environment does not put a high selective pressure on those characters. fig. 7. spinaethorax adamantis sp. nov. a–b. diagram of the chaetotaxy of the head: a dorsal side, b ventral side. c. diagram of the chaetotaxy of the trunk. enlarged or lanceolate chaetae with thicker line. nomenclature of chaetotaxy after schneider & d’haese (2013), schneider et al. (2016) and schneider (2017), partial equivalence with head nomenclature of s. tonoius (palacios-vargas & sánchez 1999) is given in a (il1–5 chaetae). schneider c. & deharveng l., first record of spinaethorax in asia 15 labrum one of the most remarkable characters of the genus neelus and megalothorax is the highly modified  labrum with an anterior process bearing the mand a-rows of chaetae (schneider 2017). this seems also to be the case of the genus zelandothorax (delamare deboutteville & massoud 1963; schneider 2017) while the genera neelides and acanthoneelidus have an ordinary labrum. the labrum of spinaethorax seems to have a somewhat intermediate morphology. the anterior process seems to exist, but is rudimentary (fig. 9a), with only weak marks of separation from the base of the labrum. the labral ridge is not well contrasted in light microscopy, but appears to be slightly in anterior position relative to the a-row of chaetae (figs 2c, 9a). the labrum of spinaethorax is probably related to the labrum of neelus or megalothorax, either being an ancestral state or a vestigial state. labium the labium of spinaethorax is very similar to the that of of neelus, with the same chaetotaxic pattern (palp and basal fields). from our observations on s. adamantis sp. nov. we conclude that the chaetotaxy of the basal fields consists of 4 + 4 basomedian chaetae and 3 + 3 basolateral chaetae. while the 2 + 2  most median chaetae of the basolateral fields are much smaller than the 1 + 1 lateral chaetae in neelus murinus sensu massoud & vannier (1967), the 3 + 3 chaetae are subequal in s. adamantis sp. nov. and s. spinotricosus. oral fold the presence of a third chaetotaxic element on the oral fold of s. adamantis sp. nov. is an unusual character among collembola. it was not reported for s. tonoius and s. spinotricosus. it is present in at least one other undescribed species of vietnam and could be an additional characteristic of the genus. the enlargement of the lateral chaeta of the oral fold is present in all material of spinaethorax examined so far. chaetotaxy of the head the chaetotaxic diagram in fig. 7a–b supports a hypothesis of chaeta position homology for s. adamantis sp. nov. and n. folsomi, neelus murinus, acanthoneelidus pratensis and several megalothorax species (discussion in schneider 2017). the general pattern seems rather similar to that of neelus murinus. palacios-vargas & sánchez (1999) introduced a nomenclature for some chaetae of the dorsal, posterior part of the head of s. tonoius, which was used by papáč & palacios-vargas (2016) for s. spinotricosus. the new species s. adamantis sp. nov. has 5 il-chaetae (fig. 7a), whereas the mexican species have fewer. chaetae il3 and il4 of s. spinotricosus (in papáč & palacios-vargas 2016) are homologized to il4 and il5 of s. adamantis sp. nov. and s. tonoius lacks il3. the chaetotaxic pattern of s. spinotricosus is very similar to that of s. adamantis sp. nov., but with a few interesting differences: one pair of chaetae missing in the inter-antennal region, likely in the ia.-row, consistent with neelus murinus. spinaethorax spinotricosus also has only one axial unpaired chaeta in the anterior, dorsal area, which is unique in neelipleona both as an absolute character and as an intra-generic variation (megalothorax, neelides and neelus with two axial unpaired chaetae, acanthoneelidus with three). antennae the chaetotaxic diagram in fig. 8 represents a partial hypothesis of chaetal position homology between s. adamantis sp. nov. and neelides folsomi, neelus murinus, a. pratensis and several species of megalothorax (discussion in schneider 2017). the antenna of spinaethorax is similar to that of neelus in terms of chaetotaxic pattern. it is distinctive mostly due to the morphology of the s-chaetae of ant iii (s1–s4, fig. 3a–c), the presence of five modified chaetae on ant. iv (so far confirmed  only in s. adamantis sp. nov., fig. 3a–b) and the fusion of ant. iii and iv. the morphology of the ant. iv organite (fig. 3c) is similar to that observed in the genus neelus. this peculiar morphology was first reported by massoud & vannier (1967) in  neelus murinus and neelus labralisetosus. the european journal of taxonomy 363: 1–20 (2017) 16 drawings in massoud & vannier (1967, figs 1a, 2a) seem to indicate the presence of a thin membrane between the filaments. such a membrane does not seem to exist in neelus koseli kováč & papáč, 2010 (a larger species) (kováč & papáč 2010) and we cannot confirm a membrane in neelus murinus or s. adamantis sp. nov. sensory fields the  pattern  of  sensory  fields  of  spinaethorax is the same as that in neelus, megalothorax and acanthoneelidus. in spinaethorax, sf1 and sf2 are both devoid of a swollen inner chaeta. instead, an ordinary chaeta is present on the border of the field. this condition is similar to what is observed in  neelus murinus, for which the sf2 chaeta is hypothesized to be homologous with the sf2 special inner fig. 8. spinaethorax adamantis sp. nov., partial diagram of the chaetotaxy of the antenna (ant. iii/ant. iv). schneider c. & deharveng l., first record of spinaethorax in asia 17 chaeta of megalothorax and acanthoneelidus (schneider 2017). spinaethorax has only one special inner chaeta in sf5, the same state is observed in adults of a. pratensis and juveniles of neelus murinus and megalothorax (schneider 2017). the position of wrc8 and s2 relative to sf6 may be useful at the genus level. in neelus murinus, wrc8 is in dorsal position and relatively far from sf6, wrc8 seems directed anterodorsally while sf6 wax secretory element is directed ventrally and s2 is in ventral position (fig. 9b). in megalothorax species (except m. granulosus schneider & d’haese, 2013), wrc8 is in anterior position to sf6, wrc8 seems directed antero-dorsally, sf6 wax secretory element is directed antero-ventrally and s2 is in dorsal position (fig. 9c). in spinaethorax, wrc8 is in antero-ventral position to sf6, both wrc8 and sf6 wax secretory elements are, respectively, directed antero-ventrally and ventrally, and s2 is in ventral position (fig. 9d). in a. pratensis and m. granulosus, wrc8 is in anterior position to sf6, both wrc8 and sf6 wax secretory elements seem directed anteriorly or antero-dorsally and s2 is in dorsal position (fig. 9e–f). pseudopore-like elements spinaethorax adamantis sp. nov. lacks the head and trunk pseudopore-like elements reported for megalothorax (schneider et al. 2016). however, a pair of similar elements (basal ring without visible chaeta) was observed on the abd. vi sternum of s. adamantis sp. nov. (absent in the other genera). fig. 9. a. schematic interpretation of the labrum of spinaethorax adamantis sp. nov. b–f. chaetotaxic organization of sf6. b. neelus murinus. c. megalothorax minimus. d. spinaethorax adamantis sp. nov. e. megalothorax granulosus. f. acanthoneelidus pratensis. g–l. outline of claw iii, normalized in length, of representatives of the six genera of neelipleona. g. spinaethorax adamantis sp. nov. h. neelus murinus. i. megalothorax minimus. j. zelandothorax novaezealandiae. k. neelides folsomi. l. acanthoneelidus pratensis. h, i, k, l after schneider (2017), j after bonet (1947). european journal of taxonomy 363: 1–20 (2017) 18 wax rod secretory elements while live observations of wax rods growth have not yet been reported for spinaethorax, the genus displays the associated wax rod secretory element of sensory fields (sf1–sf6) and the free wax rod secretory elements (wrc1–8) that are involved with wax rods growth in other neelipleona. spinaethorax and neelus have similar wrc patterns, with wrc2 and wrc7 moved anteriorly from what is observed in megalothorax and acanthoneelidus. in s. adamantis sp. nov., wrc3 and wrc5 have a specially thickened chaetal element. a similar thickening of wrc3 only is observed in neelus. an interesting observation is the numerous microspines on abdominal terga of s. spinotricosus. similar microspines are present in neelides folsomi (adult) and several representatives of the genus neelus. it seems to be associated with numerous wax rods growth in live animals and are designated as “tertiary wax rod elements” (schneider 2017). the microspines of s. spinotricosus might be tertiary wax rod elements. more intriguingly, those microspines are absent in s. adamantis sp. nov. and s. tonoius. the intrageneric variation (presence/absence) of this character is also observed in neelus (e.g., neelus murinus and neelus koseli with microspines, neelus lackovici papáč et al., 2016 and neelus klisurensis kováč & papáč, 2010 without) (kováč & papáč 2010; papáč et al. 2016; schneider 2017). in neelides folsomi, the microspines are completely absent from early instars and are replaced with a reduced number of ordinary chaetae (schneider 2017). those observations hint that such a spectacular character, whose function is as yet uncertain, was selected (or dismissed) several times during the evolution of neelipleona, with underlying heterochronic evolutionary processes. τ-chaetae the  pattern  of  τ-chaetae  differs  between  neelides (20 + 20), neelus and spinaethorax (12 + 12), megalothorax and acanthoneelidus (9 + 9). it is an interesting character for neelipleona systematics at the genus level. the intrageneric stability of the pattern is supported by observations on three species of neelus (papáč et al. 2016; schneider 2017) and eight species of megalothorax (schneider & d’haese 2013; schneider et al. 2016; schneider 2017). the τ-chaetae of s. adamantis sp. nov. are long, similarly to those of neelus and megalothorax. trunk chaetotaxy the chaetotaxic diagram in fig. 7c represents a partial hypothesis of chaeta position homology between s. adamantis sp. nov. and neelides folsomi, neelus murinus, a. pratensis and several megalothorax species (discussion in schneider 2017). the number of ordinary chaetae in the posterior region of the great abdomen (abd. iv, v terga) is greater in s. adamantis sp. nov., s. tonoius, neelus lackovici and neelus klisurensis than in other neelipleona. in s. adamantis sp. nov. the chaetotaxic pattern in this part is variable and prone to asymmetry: on four specimens we observed 6 + 6, 6 + 6, 8 + 9 and 10 + 11 chaetae in the area indicated in fig. 7c. we also observed a much greater polychaetosis in an additional specimen of spinaethorax (undescribed new species, pers. obs.). however, we did not study the region exhaustively and do not propose a chaetal nomenclature for it. the abd. vi sternum of the available mature male of s. adamantis sp. nov. does not display the special swollen chaetae (sm) that are a sexual character of male megalothorax (schneider & d’haese 2013, schneider et al. 2016) and apparently of male neelus (schneider 2017). claws the morphology of the claws of the species of spinaethorax is added to the definition of the genus as  it is very peculiar. it differs from all other neelipleona in the following features: lamella la in anterior position (instead of antero-lateral) on claw i; la missing on claw ii (instead of present); lamella of the unguiculus apparently conical and with smooth edges reduced, limited to the apex (instead of flattened  lamella with smooth edges and possibly bearing anterior and posterior crests). lamella la can be present schneider c. & deharveng l., first record of spinaethorax in asia 19 on claw iii (s. spinotricosus) or missing (s. adamantis sp. nov., maybe s. tonoius). palacios-vargas & sánchez (1999) reported a small tooth on each unguiculus of s. spinotricosus while papáč & palaciosvargas (2016) indicated a small tooth on unguiculus i and none on unguiculus ii and iii. all unguiculi of s. adamantis sp. nov. are toothless. the general shape of the claw, with basal narrowing, stands out from the other genera (fig. 9g–l). it remains, nevertheless, more similar to neelus with a slender unguis and unguiculus. the presence of a well-developed lamella bp is a character shared by spinaethorax, neelus, megalothorax and zelandothorax. furca and tenaculum the various characters of the furca and the morphology of the tenaculum of spinaethorax are similar to what is observed within the genus neelus. the most noticeable variation is the absence of chaeta e3 on the distal part of dens, also absent in megalothorax, zelandothorax, acanthoneelidus and juveniles of neelus and neelides (present in adult). conclusion the morphological observations indicate a close relationship between neelus and spinaethorax. a few characters could, however, raise the alternative hypothesis of a relationship with megalothorax and zelandothorax, essentially the fusion of ant. iii/iv, the absence of e3 on dens, the presence of a fine  secondary granulation, the absence of dermastra and perhaps the organization of sf6. the comparative morphology of genera of neelipleona suggests a complex evolutionary history that will need to be investigated in a larger phylogenetic framework than those yet developed (schneider et al. 2011; schneider & d’haese 2013). acknowledgments the field trip to southern vietnam was facilitated by truong quang tam (institute of tropical biology,  ho chi minh city) and prof. le cong kiet (ho chi minh city national university). we deeply appreciated the thorough anonymous reviews, that greatly improved the quality of the manuscript, and we express our thanks to their authors. references bonet f. 1947. monografía de la familia neelidae (collembola). revista de la sociedad mexicana de historia natural 8: 131–192. deharveng l., bedos a., le c.k., le c.m. & truong q.t. 2009. endemic arthropods of the hon chong  hills (kien giang), an unrivaled biodiversity heritage in southeast asia. in: le c.k., truong q.t. & ly  n.s. 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(both from namibia) and h. leroyi sp. nov. (from south africa) – and the male of h. dolichocephala (lawrence, 1928) is described for the first time. the distribution of the species is also revised and a new updated key to the species is compiled. keywords. africa, six-eyed crab spiders, haplogynae. lotz l.n. 2018. an update on the spider genus hexophthalma (araneae: sicariidae) in the afrotropical region, with descriptions of new species. european journal of taxonomy 424: 1–18. https://doi.org/10.5852/ejt.2018.424 introduction in a very recent paper on sicariidae, magalhaes et al. (2017) revalidated the genus name hexophthalma karsch, 1879 for six species occurring in africa (world spider catalog 2017). the genus was firstly described as hexomma (karsch 1878) for the species h. hahni, but this name was preoccupied and replaced with hexophthalma (karsch 1879). simon (1893) synonymized hexophthalma with sicarius walckenaer, 1847. all subsequent authors published new afrotropical species as sicarius (pocock 1900; purcell 1908; lawrence 1928). the distribution of hexophthalma is restricted to the southern part of the afrotropical region. the majority of the species occur in the drier western half of southern africa, extending eastward to the north. as very little collecting has been done in angola, it is possible that the distribution may extend further north into this country. in a recent paper on the afrotropical species of hexophthalma (lotz 2012), a short review was given of the described species. this was based on the type specimens or specimens collected close to the type locality. since then an intensive study of all available specimens from nine collections was undertaken. this led to the discovery of a lot of variation in the female genitalia and of unexpected prominent differences between the palps of the different species that can be used to differentiate between them. https://doi.org/10.5852/ejt.2018.424 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:2a452a63-0b25-4110-b9e1-61a69cd0e7fb mailto:leonlotz57%40gmail.com?subject= http://zoobank.org/urn:lsid:zoobank.org:author:d55d7b33-baf0-48ac-b826-d3ff6787eea7 https://doi.org/10.5852/ejt.2018.424 european journal of taxonomy 424: 1–18 (2018) 2 the dorsal setae on the leg femora were found to be too variable to be used as a diagnostic character for species, except in the case of h. spatulata (pocock, 1900), where the setae are scoop shaped and situated on a slight mound (fig. 23). in all other species the setae vary from thin to broad, with some having both thin and broad setae. in this publication the female genitalia and male palps are drawn from photos of actual specimens. in some cases the female genitalia of several specimens of the same species are depicted to show the intraspecific variation as well as the variation between the left and right side of the same specimen. during the study three new species and the male of h. dolichocephala (lawrence, 1928) were discovered and are described here. the variation in female genitalia and the differences found in the male palps also led to the realization that h. testacea (purcell, 1908) and h. hahni (karsch, 1878) is one and the same species and h. testacea is therefore here synonymized with h. hahni. the distribution of the different species is also updated here with maps showing the localities of the studied specimens only, because not all specimens mentioned in published records (newlands 1986) could be found to corroborate their identification. material and methods all the specimens that could be obtained on loan, from the collections mentioned below, were studied and are discussed here (responsible curator(s) mentioned in parentheses). illustrations of the male palps and female genitalia were made from photos of the actual genitalia studied. male palps were taken from the right side and studied and drawn from the prolateral side. illustrations were made with coreldraw x4. photos of the genitalia were taken with a paxcam 5 digital camera, mounted on a novel dissection microscope. photos are not included, as they are not of publishable quality. all female genitalia and male palps were photographed and compared. coordinates given in square brackets are not included on the specimen labels, but were looked up on physical maps and added. abbreviations morphological terms ame-le = distance between anterior median and anterior lateral eyes cl = carapace length cll = clypeus length cw = carapace width tl = total length institutions amsa = albany museum, grahamstown, south africa (j. mitchley). the code “am” is used on the specimen labels ppri = plant protection research institute, pretoria, south africa (a. dippenaar-schoeman, p. marais). the code “arc (nca)” is used on the specimen labels bmnh = british museum of natural history, london, england (j. beccaloni) cas = california academy of sciences, usa (c. griswold) dmsa = durban museum, durban, south africa mnhn = muséum national d’histoire naturelle, paris, france (c. rollard) nmba = national museum, bloemfontein, south africa (araneae collection) (l. lotz) nmnw = national museum of namibia, windhoek (t. bird). the code “smn” is used on the specimen labels nmzb = national museum of zimbabwe, arachnology department bulawayo, zimbabwe (m. fitzpatrick). the code “nmz/a” is used on the specimen labels samc = iziko museum, cape town, south africa (d. larson) lotz l.n., spider genus hexophthalma in the afrotropical region 3 results class arachnida cuvier, 1812 order araneae clerck, 1757 family sicariidae keyserling, 1880 genus hexophthalma karsch, 1879 key to the species of hexophthalma 1. females ............................................................................................................................................. 2 – males ................................................................................................................................................. 9 2. enlarged dorsal setae on femora raised on a slight mound and scoop-shaped (fig. 23); female spermathecae consist of numerous tubes, each ending in one or more spherical vesicles (fig. 16) .. ................................................................................................................h. spatulata (pocock, 1900) – enlarged dorsal setae on femora not raised on a slight mound and not scoop-shaped; spermathecae consist of one or more finger-like tubes (figs 1–15) ........................................................................ 3 3. inner spermathecae consist of one main finger-like tube on each side (fig. 10) ................................ ..................................................................................................h. dolichocephala (lawrence, 1928) – inner spermathecae consist of more than one finger-like tube on each side ..................................... 4 4. spermathecae without outer spermathecae near uterus externus and without reduced outer spermathecae (fig. 9) ..................................................................... h. damarensis (lawrence, 1928) – spermathecae with outer spermathecae near uterus externus or with reduced outer spermathecae ... ........................................................................................................................................................... 5 5. spermathecae with outer spermathecae near uterus externus ........................................................... 6 – spermathecae with reduced outer spermathecae .............................................................................. 7 6. inner spermathecae with long finger-like tubes (figs 6–8) ...............h. albospinosa (purcell, 1908) – inner spermathecae with shorter, more bulbous, finger-like tubes (figs 11–15) ................................ ...................................................................................................................... s. hahni (karsch, 1878) 7. inner spermathecae hammer-like, bent slightly laterally (fig. 1) ..................... h. binfordae sp. nov. – inner spermathecae not hammer-like, bent more or less medially ................................................... 8 8. inner spermathecae with two finger-like tubes, outer spermathecae relatively round (fig. 5) ........... ..................................................................................................................................h. leroyi sp. nov. – inner spermathecae with three or more finger-like tubes, outer spermathecae long (figs 2–4) ......... ...............................................................................................................h. goanikontesensis sp. nov. 9. embolus with a broad, blunt apex (fig. 22) ..........................................h. spatulata (pocock, 1900) – embolus with a narrow apex (figs 17–21) ..................................................................................... 10 10. embolus with a short, flattened apex (fig. 20) ........................h. dolichocephala (lawrence, 1928) – embolus with a thin, sharp apex ......................................................................................................11 11. broader, basal part of embolus straight-edged (figs 18, 21) .......................................................... 12 – broader, basal part of embolus with a hump along one edge (figs 17, 19) .................................... 13 european journal of taxonomy 424: 1–18 (2018) 4 12. thin apical part of embolus almost as long as thicker basal part (fig. 17) ........................................ ............................................................................................................h. albospinosa (purcell, 1908) – thin apical part of embolus much shorter than thicker basal part (fig. 21) ....................................... ......................................................................................................................h. hahni (karsch, 1878) 13. thin apical part of embolus without embolus keel at base (fig. 19) .................................................. ........................................................................................................ h. damarensis (lawrence, 1928) – thin apical part of embolus with embolus keel at base (fig. 18) .........h. goanikontesensis sp. nov. hexophthalma binfordae sp. nov. urn:lsid:zoobank.org:act:dd563de2-a5bd-4ab0-8319-021e01b366da figs 1, 24 diagnosis female inner spermathecae consist of one lateral extending sac and a divided round sac medially, on each side, with a small sclerotized reduced outer spermatheca (fig. 1) which is unlike that of any of the other species. male unknown. etymology the species name is a matronym for greta binford, collector of the type specimen. material examined holotype namibia: ♀, khorxas, twyfelfontein, 18°30′53.64″ s, 14°22′21.84″ e, 19 aug. 2005, g. binford leg. (nmnw 45842). description female (n = 1) tl = 11.9; cl = 5.5; cw = 5.5; cll = 0.7. ame-le = 0.6; eye diameter 0.2. carapace reddish-brown, cephalic area slightly darker; abdomen and legs yellow-brown. body and legs with lancet-like curved macrosetae, more strongly curved on body than on legs. spermathecae consist of one lateral extending sac and a divided round sac medially, on each side, with a small sclerotized reduced outer spermatheca basally (fig. 1). leg measurements: i ii iii iv palp femur 7.0 7.2 7.2 7.0 1.6 patella 2.6 2.6 2.5 2.3 1.0 tibia 6.9 7.0 6.3 6.0 1.0 metatarsus 4.8 4.7 4.6 4.6 – tarsus 2.9 3.0 2.8 3.0 2.1 total 24.2 24.5 23.5 22.9 5.7 natural history hexophthalma binfordae sp. nov. was collected from a savanna area. distribution hexophthalma binfordae sp. nov. is only known from the type locality in namibia (fig. 24). http://zoobank.org/urn:lsid:zoobank.org:act:dd563de2-a5bd-4ab0-8319-021e01b366da lotz l.n., spider genus hexophthalma in the afrotropical region 5 hexophthalma goanikontesensis sp. nov. urn:lsid:zoobank.org:act:ea012b80-652f-43df-b071-33618a497072 figs 2–4, 18, 24 diagnosis female inner spermathecae consist of sac-like fingers on each side, with a sclerotized reduced outer spermatheca; the number of sac-like spermathecae varies from three to seven between specimens and from side to side (figs 2–4); may at first glance resemble h. leroyi sp. nov., but with more than two spermathecae sacs, the other sacs hidden behind two top sacs. male embolus with a thin apical part with small keel at the base and a thicker basal part with a slight hump on one side, similar to that of h. damarensis, but differing by the presence of the small keel and the side hump being apical of median, on the thicker part of the embolus (fig. 18). etymology the specific name is an adjective derived from the type locality. material examined holotype namibia: ♂, swakpomund, namib-naukluft park, goanikontes, 22.68565° s, 14.84964° e, 7 feb.– 28 mar. 2009, j. irish & r. scholtz leg. (nmnw 47282). paratypes namibia: 2 ♂♂, 2 ♀♀, same data as for holotype (ex nmnw 47282); 1 ♀, cape cross [21°46′ s, 13°59′ e], 8 jul. 1963, a. els leg. (nmnw 35066); 2 ♀♀, damaraland, klein graias, 2014cc, 5 aug. 1985, e. griffin leg. (nmnw 41688); 1 ♂, 1 imm., kaokoland, 17°37′ s, 12°12′ e, 14 oct. 1988, h. kleinhans leg. (nmnw 40797); 1 ♂, kaokoland, 15 miles w of orupembe at park border, 1812ac, r.e.g. leg. (nmnw 35964); 2 ♀♀, 3 imm., koichab river, sceleton coast, 20 miles inland [ca 20°21′ s, 13°33′ e], 8 nov. 1968, r.f. lawrence leg. (amsa); 1 ♀, rössing mine, arandis water, 2214bd, 7 jun. 1984, e. griffin leg. (nmnw 38930); 1 ♂, rössing mine, crusher dust area, 2215ca, 19 mar.–9 apr. 1984, e. griffin leg. (nmnw 37021); 1 ♂, rössing mine, panner gorge, 22°29′ s, 15°01′ e, 9 apr.– 8 may 1984, e. griffin leg. (nmnw 37086); 1 ♀, same data as for preceding but 11 mar.–9 apr. 1985, irish and rust leg. (nmnw 38737); 1 ♂, rössing mine, upper ostrich gorge, 22°29′ s, 14°59′ e, 9 apr.–8 may 1984, e. griffin leg. (nmnw 37057); 1 ♂, same data as for preceding but 5 apr.–6 may 1985, irish and rust leg. (nmnw 38794); 1 ♂, swakopmund, 22.67° s, 14.52° e, 15 mar. 2010, g. ackerman leg. (ppri 2010/2540); 3 ♂♂, 1 ♀, swakpomund, namib-naukluft park, goanikontes, 22°43′11.9″ s, 14°49′23.2″ e, 7–28 feb. 2009, j. irish & r. scholtz leg. (nmnw 47221, 47233); 1 ♀, same data as for preceding but 5 feb. 1929, r.d. bradfield leg. (amsa 6003); 1 ♀, ugab river [21°10′ s, 13°39′ e], 25 apr. 1987 (nmnw 40495); 1 ♂, vogelfederberg, 23°06′ s, 14°55′ e, 5 oct. 1984, e. griffin leg. (nmnw 42837). note hexophthalma goanikontesensis sp. nov. appears to be the same species as h. sp. 2 in magalhaes et al. (2017), judging from the figures in their publication (figs 20c–d, 21c–d). description female (n = 7) tl = 10.26 (8.8–12.4); cl = 4.80 (3.7–6.0); cw = 4.76 (3.8–5.6); cll = 0.59 (0.5–0.8). ame-le = 0.6; eye diameter = 0.2. carapace reddish-brown, cephalic area slightly darker; abdomen and legs yellowbrown. body and legs with lancet-like curved macrosetae, more strongly curved on body than on legs. inner spermathecae consist of sac-like fingers on each side, with a sclerotized reduced outer spermatheca; http://zoobank.org/urn:lsid:zoobank.org:act:ea012b80-652f-43df-b071-33618a497072 european journal of taxonomy 424: 1–18 (2018) 6 the number of sac-like inner spermathecae varies from three to seven sacs among specimens and from side to side (figs 2–4). leg measurements: i ii iii iv palp femur 6.0 6.3 6.0 6.2 1.8 patella 2.3 2.4 2.3 2.3 1.0 tibia 5.5 5.6 5.1 5.1 1.0 metatarsus 3.9 4.0 3.5 3.8 – tarsus 2.8 2.6 2.5 2.8 1.9 total 20.5 20.9 19.4 20.2 5.7 male (n = 10) tl = 9.01 (8.0–9.8); cl = 4.65 (4.1–5.2); cw = 4.62 (4.2–5.1); cll = 0.61 (0.60–0.65). ame-le = 0.55; eye diameter = 0.2. colour and spination as in female. embolus with a thin apical part with small keel at the base and a thicker basal part with a slight hump on one side, apical of median on the thicker part of the embolus (fig. 17). leg measurements: i ii iii iv palp femur 6.3 7.0 6.4 6.4 1.6 patella 2.2 2.2 2.2 2.0 1.0 tibia 5.8 5.9 5.5 5.4 1.2 metatarsus 4.1 4.3 4.0 3.8 – tarsus 2.8 2.8 2.6 2.6 0.6 total 21.2 22.2 20.7 20.2 4.4 natural history hexophthalma goanikontesensis sp. nov. have been collected in desert areas, by pit traps and by hand from under rocks. distribution hexophthalma goanikontesensis sp. nov. is known from the northern coastal areas in namibia (fig. 24). hexophthalma leroyi sp. nov. urn:lsid:zoobank.org:act:3eaaf34d-d7db-4c5f-84be-231233e064b4 figs 5, 24 diagnosis female inner spermathecae consist of two sac-like fingers on each side, with a round sclerotized reduced outer spermatheca (fig. 5), to a certain extent similar to h. goanikontesensis sp. nov., but with only two inner spermathecae sacs. male unknown. etymology the specific name is a patronym for john leroy, collector of the type specimen. http://zoobank.org/urn:lsid:zoobank.org:act:3eaaf34d-d7db-4c5f-84be-231233e064b4 lotz l.n., spider genus hexophthalma in the afrotropical region 7 material examined holotype south africa: ♀, northern cape province, augrabies national park, 28.66° s, 20.42° e, 24 feb. 1983, j. leroy leg., lr270 (ppri 89/786). description female (n = 1) tl = 12.9; cl = 5.9; cw = 6.0; cll = 0.9. ame-le = 0.65; eye diameter = 0.2. carapace reddish-brown, cephalic area slightly darker; abdomen and legs yellow-brown. body and legs with lancet-like curved macrosetae; macrosetae more curved on body than on legs. inner spermathecae consist of two sac-like fingers on each side, with a round, sclerotized, reduced outer spermatheca (fig. 5). leg measurements: i ii iii iv palp femur 7.2 7.4 6.8 7.0 1.6 patella 2.6 2.8 2.7 2.7 1.0 tibia 7.1 7.1 6.8 6.2 1.1 metatarsus 4.9 4.8 4.3 4.6 – tarsus 2.8 2.8 2.6 3.2 2.0 total 24.6 24.9 23.2 23.7 5.7 natural history hexophthalma leroyi sp. nov. was collected in a grassy area in the nama karoo biome. distribution hexophthalma leroyi sp. nov. is known only from the type locality in south africa (fig. 24). hexophthalma albospinosa (purcell, 1908) figs 6–8, 17, 25 sicarius albospinosus purcell, 1908: 224. sicarius albospinosus – lawrence 1938: 215. — newlands 1986: 45. — lotz 2012: 3. hexophthalma albospinosa – magalhaes et al. 2017: 851, figs 20a–b, 21a–b, 62a–c. revised diagnosis female inner spermathecae consist of long finger-like sacs that branch close to the copulatory tube end, similar to in h. hahni except that the finger-like tubes are longer and thinner (figs 6–8). male embolus consisting of a thin pointed apical part and straight-sided thicker basal section (fig. 17), similar to in h. hahni except for the longer thin apical part in relation to the thicker basal part. material examined lectotype namibia: ♀, lüderitz bay (angra pequena, great namaqualand [26°39′ s, 15°09′ e], jan. 1904, l. schultze leg. (samc x150521). european journal of taxonomy 424: 1–18 (2018) 8 paralectotypes namibia: 2 juv., same data as for lectotype (samc x150521). other material namibia: 1 ♂, agate beach, 2615ca, 9 oct. 1984, m. griffin leg. (nmnw 42838); 1 ♀, same data as for preceding (nmnw 42927); 1 ♂, atlas bay, diamond area i, 26°49′ s, 15°08′ e, 13–26 nov. 1993, e. marais leg. (nmnw 43252); 1 ♂, between chameis head and south rock, 27°51′ s, 15°39′ e, 6 aug. 1997, e. griffin leg. (nmnw 43810); 1♀, 1 imm., griffiths bay, diamond area 1, 26°39′ s, 15°07′ e, 16 jun. 1993, m. griffin leg. (nmnw 42894); 1 ♂, jammerbucht, 27°10′ s, 15°16′ e, 20 jun. 1993, m. griffin leg. (nmnw 42912); 1 ♂, 3 ♀♀, 4 imm., klinghardts mountain foothills, diamond area 1, figs 1–8. female internal genitalia. 1. hexophthalma binfordae sp. nov. 2–4. h. goanikontesensis sp. nov. 5. h. leroyi sp. nov. 6–8. h. albospinosa (purcell, 1908). scale bar = 1 mm. lotz l.n., spider genus hexophthalma in the afrotropical region 9 27°25′ s, 15°36′ e, 17–18 jun. 1993, m. griffin and e. griffin leg. (nmnw 42905, 42907); 2 ♂♂, 2 ♀♀, 2 imm., lüderitz, diamond area 2, kowis mountain, 26°38′25.7″ s, 15°20′46.7″ e, 22 aug. 2006, eduventures 9 leg. (nmnw 47257, 47258); 1 ♂, lüderitz, lüderitz peninsula, 26°39′58.5″ s, 15°07′19.9″ e, 14 dec. 2008–3 feb. 2009, i. wiesel, t.l. bird and c. bird leg. (nmnw 47339); 1 ♂, lüderitz, spergebiet national park, base of tsaukhaib mountain, 26°43′01.1″ s, 15°40′05.2″ e, 20 may 2009, eduventures leg. (nmnw 47367); 1 ♂, namdeb southern mine, old articulated dumptruck, 30 jan.–12 feb. 1997, v. ward and j. daneel leg. (nmnw 43565); 2 ♀♀, oranjemund, 28.55° s, 16.43° e, jan. 1991, n. larson leg. (ppri 91/1530); 1 ♀, oranjemund site 3, namdeb mine dumps, 2816cb, 23–26 jan. 1998, v. ward leg. (nmnw 43844); 1 ♂, oranjemund mining area, control site 5, near orange river, 20 sept.–2 oct. 1996, e. griffin leg. (nmnw 43554); 1 ♀, orange river mouth, coastal dunes, 2826cb, 10 nov. 1986, e. griffin leg. (nmnw 42957). natural history hexophthalma albospinosa have been collected in the succulent karoo biome, by pitfall traps and by hand from under rocks and old roofing tin. distribution hexophthalma albospinosa is distributed along the southern coastal areas of namibia (fig. 25). hexophthalma damarensis (lawrence, 1928) figs 9, 19, 25 sicarius damarensis lawrence, 1928: 221. sicarius damarensis – newlands 1986: 47. — lotz 2012: 4. hexophthalma damarensis – magalhaes et al. 2017: 852. diagnosis female inner spermathecae unique, with short, bulbous, finger-like sacs that branch off the copulatory tube part, without reduced outer spermathecae (fig. 9). male embolus ending in a thin, pointed apex; thicker basal part of embolus with a hump on one side, slightly basal from median (fig. 19); this is similar to in h. goanikontesensis sp. nov. except that the thinner apical part lacks a basal embolus keel and the hump on the side of the thicker part is more basal. material examined holotype namibia: ♀, outjo [20°07′ s, 16°09′ e] (samc 7057). other material namibia: 1 ♂, helio, 19°03′ s, 16°29′ e, 14 feb.–23 mar. 1987, e. griffin leg. (nmnw 40160); 1 ♀, 1 imm., same data as for preceding but 8 aug. 1987–27 mar. 1988 (nmnw 40986); 1 ♂, same data as for preceding but 27 mar.–4 may 1988 (nmnw 41135). natural history hexophthalma damarensis have been collected in savanna biomes using pitfall traps. distribution hexophthalma damarensis is distributed in the northern central part of namibia (fig. 25). european journal of taxonomy 424: 1–18 (2018) 10 hexophthalma dolichocephala (lawrence, 1928) figs 10, 20, 25 sicarius dolichocephalus lawrence, 1928: 222. sicarius dolichocephalus – newlands 1986: 49. — lotz 2012: 8. hexophthalma dolichocephala – magalhaes et al. 2017: 850. diagnosis female inner spermathecae consist of one finger-like sac on each side, with a very short outer spermatheca (fig. 10). male embolus apical part thin, blunt and slightly twisted and flattened, unlike the other species where the apical part is thin and pointed (fig. 20). figs 9–16. female internal genitalia. 9. hexophthalma damarensis (lawrence, 1928). 10. h. dolichocephala (lawrence, 1928). 11–15. h. hahni (karsch, 1878). 16. h. spatulata (pocock, 1900). scale bar = 1mm. lotz l.n., spider genus hexophthalma in the afrotropical region 11 material examined lectotype namibia: ♀, kaoko otavi [18°18′ s, 13°40′ e], apr. 1926 (samc 6937). other material namibia: 1 ♂, 1 ♀, 1 imm., kaokoland, opuwo, 17°24′20.34″ s, 14°13′01.20″ e, 18 aug. 2005, g. binford leg. (nmnw 45843); 1 ♀, kaokoland, opuwo, baynes mountain, epembe, 17°04′32.9″ s, 13°02′08.5″ e, 20 apr. 2006, eduventures 8 leg. (nmnw 47266); 1 ♂, 1 ♀, munutum river [18°17′ s, 12°04′ e], 19 oct. 1988, h. kleinhans leg. (nmnw 40863); 1 ♂, okahama water point, 17°10′34.1″ s, 12°59′54.7″ e, 25 apr. 2006, eduventures 8 leg. (nmnw 47271); 1 ♂, möwe bay, 19°20′ s, 12°40′ e, 1990, d. and j. bartlett leg. (nmnw 42684); 1 ♂, ruacana, 17°25′ s, 14°19′ e, 25–27 aug. 1973, state museum leg. (nmnw 35801). description male (n = 5) tl = 19.5 (11.6–28.7); cl = 4.98 (4.7–5.6); cw = 4.94 (4.7–5.6); cll = 0.59 (0.50–0.65). amele = 0.5; eye diameter = 0.2. carapace reddish-brown, cephalic area slightly darker; abdomen and legs yellow-brown. body and legs with lancet-like curved macrosetae, more strongly curved on body than on legs. embolus thick over most of the length, tapered apically, point blunt and slightly twisted at base of thinner part (fig. 20). leg measurements: i ii iii iv palp femur 7.4 8.6 7.4 7.7 1.5 patella 2.3 2.4 2.2 2.2 0.9 tibia 8.4 8.8 7.6 7.3 1.2 metatarsus 6.2 7.5 6.5 6.5 – tarsus 2.9 3.3 2.9 3.3 0.7 total 27.2 30.6 26.6 27.0 4.3 natural history hexophthalma dolichocephala have been collected in desert, nama karoo and savanna biomes, by pitfall traps and by hand from under rocks. distribution hexophthalma dolichocephala is distributed in the northwestern corner of namibia (fig. 25). hexophthalma hahni (karsch, 1878) figs 11–15, 21, 26 hexomma hahnii karsch, 1878: 325. sicarius testaceus purcell, 1908: 224. syn. nov. hexophthalma hahni – karsch, 1879: 109. — magalhaes et al. 2017: 850, figs 13b, 14b–c, 16g, 18g, 19c, 23a–c, 24a, 61a–d. sicarius hahni – simon, 1893: 269. — purcell 1908: 225. — newlands 1986: 51. — lotz 2012: 8. european journal of taxonomy 424: 1–18 (2018) 12 sicarius testaceus – newlands 1986: 57. — lotz 2012: 12. hexophthalma testacea – magalhaes et al. 2017: 852. note: sicarius oweni newlands, 1986: 53 is an unpublished synonym. diagnosis female inner spermathecae consist of finger-like sacs that branch close to the copulatory tube end, similar to in h. albospinosa except that the finger-like tubes are shorter and more bulbous (figs 11–15). male embolus consisting of a thin pointed apical part and a straight-sided thicker basal section (fig. 21), similar to in h. albospinosa except that the thin apical part is shorter in relation to the thicker basal part. figs 17–23. 17–22. male palps (right side prolateral). 17. hexophthalma albospinosa (purcell, 1908). 18. h. goanikontesensis sp. nov. 19. h. damarensis (lawrence, 1928). 20. h. dolichocephala (lawrence, 1928). 21. h. hahni (karsch, 1878). 22. h. spatulata (pocock, 1900). 23. h. spatulata, enlarged femoral seta (not to scale). scale bars = 1 mm. lotz l.n., spider genus hexophthalma in the afrotropical region 13 material examined country unknown: 2 ♂♂, 2 ♀♀, 4 imm. (samc b7732, b8870). namibia: 1 ♂ [no locality or date], g. brown leg. (samc b5704); 1 ♂, ganab, 23°04′ s, 15°10′ e, 13– 16 sep. 1971, p.g. olivier leg. (nmnw 35311); 1 ♀, same data as for preceding (nmnw 35313); 1 ♀, grootfontein, nosib cave, 1917bd, 8 feb. 1995, segl leg. (nmnw 43454); 1 ♂, 1 ♀, keetmanshoop, khabus 146, 2618ac, 7 dec. 1987–17 jan. 1988, mar. 1992, n. and g. olivier leg. (nmnw 40752); 2 ♂♂, same data as for preceding but 23 jul.–1 dec. 1988 (nmnw 42120, 42788); 1 ♀, naukluft park, gifgat 2, 2215da, 11 jul. 1995, e. marais leg. (nmnw 43673); 1 ♀, 1 imm., naukluft park, gobabeb, 23°33′28.2″ s, 15°02′14.32″ e, 25 oct. 1968, r.f. lawrence leg. (amsa); 1 ♂, 2 ♀♀, same data as for preceding but 21 aug. 2005, g. binford leg. (nmnw 45835); 1 ♂, 2 ♀♀, naukluft park, kuiseb river, near gobabeb [23°34′ s, 15°03′ e], 18 feb.–20 mar. 1983, kuiseb survey leg. (ppri 83/341); 1 ♂, same data as for preceding but 27 aug. 1985, c.s. crawford leg. (nmnw 41116); 1 ♂, 1 ♀, otjiwarongo, waterberg plateau park, 20°30′17.94″ s, 17°14′32.16″ e, 14 aug. 2005, g. binford leg. (nmnw 45838); 1 ♂, sossusvlei, 24.7° s, 15.42° e, dec. 1988, b. grobbelaar leg. (ppri 92/54); 1 ♂, valencia, 96 km sw of windhoek, 2316ab, 23 jun. 1969, a. kemp leg. (amsa); 1 ♂, 1 imm., windhoek, daan viljoen game reserve, 22°32′16.14″ s, 16°54′44.16″ e, 12 aug. 2005, g. binford leg. (nmnw 45845); 1 ♂, windhoek, verloren cave, 23°09′ s, 16°20′ e, 28 jun. 1993, j. irish leg. (nmba 7361); 1 ♂, 1 imm., messum crater, 21°20.430′ s, 14°13.218′ e, 4–10 apr. 2000, e. griffin leg. (nmnw 44724); 1 ♀, 10 miles w of okombahe [ca 21°24′ s, 15°14′ e], 10 may 1958, e.s. ross and r.e. leech leg. (cas). south africa – limpopo province: 1 ♂, kruger national park, pafuri area, 22.46° s, 31.3° e, 14 oct. 1995, a. leroy leg. (ppri 98/405); 1 ♀, limpopo valley national park, 22.22° s, 29.13° e, 29 apr. 1997, j. leroy leg. (ppri 97/816); 4 ♂♂, makgabeng area, w of senwabawana, 23.24° s, 28.85° e, 22 dec. 2006, m. burger and r. hawkins leg. (ppri 2008/3788, 2008/3807); 1 ♂, rochdale, near waterpoort, 22.54° s, 29.41° e, 4 apr. 1993, a. leroy leg. (ppri 2007/1626); 1 ♂, vhembi biosphere, gondeni, 22.914° s, 30.050° e, 4 dec. 2012, c. schoeman leg. (ppri 2015/632). – mpumalanga: 1 ♀, 2 imm., bango poort, cave poort, 24.58° s, 31.1° e, mar. 1984, l. braak leg. (ppri 85/174). – northern cape province: 1 ♂, blackridge, e of langberge, e-ne of groblershoop, 28°49′ s, 22°32′ e, 22 dec. 2005, m. burger leg. (ppri 2009/3835); 1 ♂, bo-seekoebaard, se of groblershoop [29°05′ s, 22°17′ e], 13 dec. 2005, m. burger leg. (ppri 2009/3834); 1 ♂, bushmanland, krapohl island [28°50′30″ s, 18°43′30″ e], 1909 (samc 14708); 2 ♀♀, 4 imm., calvinia, klein arendskraal, 31°24′ s, 19°03′ e, 19 feb. 2004, l.n. lotz leg. (nmba 9259, 9269); 1 ♂, douglas [29°04′ s, 23°47′ e], r. broom leg. (amsa); 3 ♂♂, eselsfontein, south of grootdrink, 28.62° s, 21.68° e, 13 nov. 2005, m. burger, p. braad and a. hill leg. (ppri 2009/3830–32); 1 ♂, same data as for precedings but 22 nov. 2005, m. burger leg. (ppri 2009/3833); 1♂, 1 imm., little namaqualand, kamies [kamieskroon?] [ca 30°13′ s, 17°56′ e] (samc b7442); 1♂, 1 imm., namaqualand, kap kap 382, 30°16′ s, 18°22′ e, 20 oct. 1990, l.n. lotz leg. (nmba 5122); 1 ♂, 1 ♀, nieuwoudtville, oorlogskloof nature reserve, 31.45° s, 19.1° e, 27 apr. 2000, a. leroy leg. (ppri 2010/187); 1 ♀, 2 imm., nw gordonia, c.a. anderson leg. (amsa 1967); 1 ♂, port nolloth, 2916bd, 13 mar. 1973, m.-l. penrith leg. (nmnw 35722); 2 ♂♂, 1 ♀, quaggafontein 476, 15 km s-sw of soebatsfontein, 30°11′14.6″ s, 17°32′58.6″ e, 29 sept. 2002, biota leg. (ppri 2009/5561, 2009/5562). – north west province: 1 ♀, groot marico, 25.6° s, 26.43° e, 21 jan. 1991, m. filmer leg. (ppri 91/1323). – western cape province: 1 ♀, 10 km n of bitterfontein [ca 30°59′ s, 18°14′ e], 15 sept. 1983, j.v. leg. (samc c1981); 1 ♀, clanwilliam district (samc 3548); 1 ♀, clanwilliam, sanddrif (dwarsrivier 330), 32.16° s, 18.89° e, feb. 1993, g. müller leg. (ppri 93/261); 1 ♀, donkins bay, 3118cd, 14 mar. 1973, m.j. and m.-l. penrith leg. (nmnw 35729); 1 ♀, doringbaai [31°49′ s, 18°14′ e], nov. 1956, samc expedition leg. (samc c1235); 3 ♀♀, swartberg nature reserve, gamkaskloof, 33.36° s, 21.69° e, dec. 1995, m. de jager leg. (ppri 96/399); 1 ♀, 12 imm., same data as for preceding but 1 jul. 2000, z. van der walt leg. european journal of taxonomy 424: 1–18 (2018) 14 (ppri 2002/44); 1 ♀, hans strydom dam nature reserve, mogol, 33.95° s, 22.46° e, 19 jan. 1983, m. mansell leg. (ppri 83/198); 1 ♂, the island, s of herbertsdale, 34.08° s, 21.77° e, 3 jan. 2006, m. burger leg. (ppri 2009/3828); 1 ♂, same data as for preceding but 30 jan. 2006 (ppri 2009/3829); 1 ♀, 19 km se of klawer [ca 31°54′ s, 18°39′ e], 1 may 1958 (cas); 1 ♀, lutzville, vlêrmuisklip, near vredendal, 31.39° s, 18.32° e, 2 oct. 1986, m.w. mansell and j. hoffman leg. (ppri 86/650); 1 ♀, papendorp, near olifants river estuary [ca 31°42′ s, 18°13′ e], 12 nov. 1949, b. malkin leg. (cas); 2 ♂♂, 1 ♀, 1 imm., prince albert, swartberg nature reserve, gamkaskloof, die hel, 33°20′ s, 21°40′ e, sep. 1998, z. van der walt leg. (nmba 8695, 8860); 1 ♀, 3 imm., vanrhynsdorp, van rhyn’s pass, 31.6° s, 18.75° e, 3 oct. 1986, m.w. mansell and j. hoffman leg. (ppri 86/648). zimbabwe: 1 ♀, pande mine, 2230ad, 9 feb. 1994, f. nyathi leg. (nmzb a/11518); 1 ♀, 1 imm., sentinel ranch, hunters camp, 2229ab, 16 apr. 1990, falcon collage-nhmz leg. (nmzb a/9940). natural history hexophthalma hahni have been collected in the fynbos, grassland, karoo, desert and savanna biomes, by pitfall traps and by hand, in caves and from under rocks and logs. distribution hexophthalma hahni is distributed in namibia, south africa and southern zimbabwe (fig. 26). hexophthalma spatulata (pocock, 1900) figs 16, 22–23, 25 sicarius spatulatus pocock, 1900: 321. sicarius spatulatus – newlands 1986: 56. — lotz 2012: 10. hexophthalma spatulata – magalhaes et al. 2017: 853, figs 6a–d, 12c, 14d, 18h, 19d, 20e–f, 22a–e, 23d–f, 31h. diagnosis leg femora in both sexes with scoop-shaped enlarged dorsal setae (fig. 23), differing from the setae of all other species. female inner spermathecae consist of a broad chamber, extending anteriorly in numerous tubes, each ending in one or more spherical vesicles (fig. 16), differing from all other species ,where the inner spermathecae are finger-like sacs. male embolus ending in a broad, blunt triangular apex (fig. 22), whereas the embolus of all the other species have thin apices. material examined south africa – eastern cape province: 5 ♂♂, 1 ♀, 2 imm., alicedale [33°18′ s, 26°05′ e], dec. 1914, f. cruden leg. (amsa); 1 ♀, same data as for preceding (amsa 2525). – western cape province: 1 ♂, borrelfontein, 8 km w of gouritz river mouth, 34°20′ s, 21°51′ e, 4 mar. 2007, h. leibel leg. (ppri 2008/745); 1 ♂, same data as for preceding but nov. 2007–may 2008 (ppri 2009/5601); 1 ♂, bredasdorp [34°32′ s, 20°03′ e] (samc b7435); 4 ♀♀, calidon [34°14′ s, 19°26′ e] (samc 8348); 1 ♂, 1 ♀, 2 imm., same data as preceding but w.f. purcell leg. (samc b363); 1 ♂, 2 ♀♀, 1 imm., hermanus [34°25′ s, 19°13′ e], (samc 11632, b8482); 2 ♂♂, 2 ♀♀, 5 imm., houwhoek, 3419aa (samc 8327); 1 ♂, kammanasie nature reserve, 33°39′ s, 22°44′ e, 31 jul. 2003, reserve staff leg. (ppri 2010/2403); 1 ♀, piquetberg, 3218dd (samc b8490); 1 ♀, swellendam, tradows pass [33°58′ s, 20°42′ e] (samc b6767). lotz l.n., spider genus hexophthalma in the afrotropical region 15 natural history hexophthalma spatulata have been collected in a fynbos biome. distribution hexophthalma spatulata is distributed along the southern coastal areas of south africa (fig. 25). figs 24–25. distribution maps for species of hexophthalma karsch, 1879. 24. h. goanikontesensis sp. nov., h. binfordae sp. nov. and h. leroyi sp. nov. 25. h. albospinosa (purcell, 1908, h. damarensis (lawrence, 1928), h. dolichocephala (lawrence, 1928) and h. spatulata (pocock, 1900). european journal of taxonomy 424: 1–18 (2018) 16 discussion species of hexophthalma, the former african sicarius, have raised interest because of their supposed medical importance (newlands & atkinson 1988; van aswegen et al. 1997). the venom of some of the species has even been considered as one of the strongest spider venoms (newlands 1986), but this has never been corroborated. studies on this topic, mainly on the interspecific variation of the venom, which can be very large (müller et al. 2012), are particularly difficult, because the identification of the species is virtually impossible. the study of zobel-thropp et al. (2010) is symptomatic: it was hindered by the lack of systematic knowledge and had to use approximate identifications like sicarius cf. hahni and s. cf. damarensis. the present study has made a step forward, but it is clear that more efforts will be needed to facilitate studies on this genus, be it systematic, ecological or medical. in several species the males are still unknown, more species are to be expected and the monophyly of the genus is questionable because of the morphologically aberrant h. spatulata. future studies will therefore have to be carried out along several axes. in the first place, further collecting will be needed to fill the gaps outlined above. molecular studies in combination with detailed morphological assessments will be necessary to outline the interspecific relationships and the phylogenetic placement of the group, mainly in connection with its possible polyphyly. acknowledgments i am very grateful to the institutions mentioned in the text for the loan of their specimens. the council and director of the national museum bloemfontein are also acknowledged for supporting this project. fig. 26. distribution map for hexophthalma hahni (karsch, 1878). lotz l.n., spider genus hexophthalma in the afrotropical region 17 references karsch f. 1878. exotisch-araneologisches. zeitschrift für die gesammten naturwissenschaften 51: 323– 333. karsch f. 1879. sieben neue arachniden von st martha. entomologische zeitung herausgegeben von dem entomologische vereine zu stettin 40: 106–109. available from https://biodiversitylibrary.org/page/9188954 [accessed 23 mar. 2018]. lawrence r.f. 1928. contributions to a knowledge of the fauna of south-west africa vii. arachnida (part 2). annals of the south african museum 25: 217–312. available from https://biodiversitylibrary.org/page/40656433 [accessed 23 mar. 2018]. lawrence r.f. 1938. transvaal museum expedition to south-west africa and little namaqualand, may to august 1937. spiders. annals of the transvaal museum 19: 215–226. lotz l.n. 2012. present status of sicariidae (arachnida: araneae) in the afrotropical region. zootaxa 3522: 1–41. magalhaes i.l.f., brescovit a.d. & santos a.j. 2017. phylogeny of sicariidae spiders (araneae: haplogynae), with a monograph on neotropical sicarius. zoological journal of the linnean society 179 (4): 767–864. müller g.j., wium c.a., marks c.j., du plessis c.e. & veale d.j.h. 2012. spider bite in south africa: diagnosis and management. continuing medical education 30: 382–392. available from http://www.cmej.org.za/index.php/cmej/article/view/2547/2582 [accessed 23 mar. 2018]. newlands g. 1986. necrotic arachnidism in southern africa. unpublished ph.d. thesis. university of the witwatersrand, johannesburg. newlands g. & atkinson p. 1988. review of southern african spiders of medical importance, with notes on the signs and symptoms of envenomation. south african medical journal 73: 235–239. pocock r.i. 1900. some new arachnida from cape colony. annals and magazine of natural history, series 7 6 (31): 316–333. https://doi.org/10.1080/00222930008678382 purcell w.f. 1908. araneae (1). in: schultze l. forschungsreise im westlichen und zentralen südafrika ausgeführt in den jahren 1903–1905. denkschrift der medicinisch-naturwissenschaftlichen gesellschaft, jena 13: 203–246. available from https://biodiversitylibrary.org/page/37102545 [accessed 23 mar. 2018]. simon e. 1893. famille filistatidae. histoire naturelle des araignées 1: 255–488. https://biodiversitylibrary.org/page/38670557 van aswegen g., van rooyen j.m., van der nest d.g., veldman f.j., de villiers t.h. & oberholzer g. 1997. venom of a six-eyed crab spider, sicarius testaceus (purcell, 1908), causes necrotic and haemorrhagic lesions in the rabbit. toxicon 35: 1149–1152. https://doi.org/10.1016/s0041-0101(96)00203-6 world spider catalog 2017. world spider catalog. version 18.0. natural history museum bern. available from https://wsc.nmbe.ch/ [accessed on 3 april 2017]. zobel-thropp p.a., bodner m.r. & binford g.j. 2010. comparative analyses of venoms from american and african sicarius spiders that differ in sphingomyelinase d activity. toxicon 55: 1274–1282. https://doi.org/10.1016/j.toxicon.2010.01.019 manuscript received: 18 april 2017 manuscript accepted: 11 september 2017 https://biodiversitylibrary.org/page/9188954 https://biodiversitylibrary.org/page/40656433 http://www.cmej.org.za/index.php/cmej/article/view/2547/2582 https://doi.org/10.1080/00222930008678382 https://biodiversitylibrary.org/page/37102545 https://biodiversitylibrary.org/page/38670557 https://doi.org/10.1016/s0041-0101%2896%2900203-6 https://wsc.nmbe.ch/ https://doi.org/10.1016/j.toxicon.2010.01.019 european journal of taxonomy 424: 1–18 (2018) 18 published on: 10 april 2018 topic editor: rudy jocqué desk editor: chloë chester printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain. european journal of taxonomy 509: 1–14 issn 2118-9773 https://doi.org/10.5852/ejt.2019.509 www.europeanjournaloftaxonomy.eu 2019 · rogers d.c. et al.. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:ea21763a-c185-4650-a075-5b9750640f50 1 a review of dendrocephalus (dendrocephalinus) (crustacea: anostraca) with the first records of male-male anostracan aggressive competition d. christopher rogers 1, *, ann dunn 2 & w. wayne price 3 1 kansas biological survey, kansas university, higuchi hall, 2101 constant avenue, lawrence, ks 66047-3759, usa. 2 14391 campbell farm lane, lovettsville, va 20180, usa. 3 department of biology, university of tampa, tampa, florida, usa. * corresponding author: branchiopod@gmail.com 2 email: ard224@cornell.edu 3 email: wprice@ut.edu 1 urn:lsid:zoobank.org:author:77d95892-8588-4016-88a9-73a2a1e1815f 2 urn:lsid:zoobank.org:author:3f2619c0-cdf7-40d5-aaec-61190c976889 3 urn:lsid:zoobank.org:author: abstract. we present a review of dendrocephalus (dendrocephalinus) with an updated diagnosis for the subgenus and a key to all known species. we provide new records of dendrocephalus alachua, which was previously supposed to be extinct, and we describe a new species, dendrocephalus proeliator sp. nov., which is separated from all other species based on the form of the male frontal appendage. dendrocephalus proeliator sp. nov. appears to be morphologically intermediate between d. alachua and d. lithaca. in addition, we provide conservation assessments for all four species in the subgenus, according to iucn red list standards. we also report for two species the first known examples of direct male-male agonistic behaviour and competition for access to areas frequented by receptive females. keywords. fairy shrimp, florida, hill topping, sexual selection, new species. rogers d.c., dunn a. & price w.w. 2019. a review of dendrocephalus (dendrocephalinus) (crustacea: anostraca) with the first records of male-male anostracan aggressive competition. european journal of taxonomy 509: 1–14. https://doi.org/10.5852/ejt.2019.509 introduction dendrocephalus daday, 1908 is a new world genus in the thamnocephalidae, with two subgenera (rogers 2006). dendrocephalus s. str. ranges from argentina, north through to costa rica, the galápagos islands, and aruba, with 17 described species (brendonck et al. 1990; belk et al. 2002; rogers 2013; rogers et al. 2012; rogers & volcan 2016; rabet et al. 2018). rogers & volcan (2016) provided keys to the species of this subgenus, and improved keys are in development (rogers et al., in press). https://doi.org/10.5852/ejt.2019.509 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ http://zooank.org/urn:lsid:zoobank.org:pub:ea21763a-c185-4650-a075-5b9750640f50 mailto:branchiopod%40gmail.com?subject= mailto:ard224%40cornell.edu?subject= mailto:wprice%40ut.edu?subject= http://zoobank.org/urn:lsid:zoobank.org:author:77d95892-8588-4016-88a9-73a2a1e1815f http://zoobank.org/urn:lsid:zoobank.org:author:3f2619c0-cdf7-40d5-aaec-61190c976889 https://doi.org/10.5852/ejt.2019.509 european journal of taxonomy 509: 1–14 (2019) 2 the subgenus dendrocephalinus is limited to the extreme southern united states of america (georgia, florida, and texas), and has three described species (rogers 2006, 2013). we review the subgenus, provide an updated key to the species, and discuss the conservation status of each. we present a new species of dendrocephalinus from florida and describe its remarkable male-male territorial aggressive behaviour. material and methods material was examined in museum collections or collected live from the wild, and cultured using the wild collections. material of the new species was examined from the florida museum of natural history, sent to the first author for identification. all preserved material was examined under a wild m8 stereo dissection microscope. all drawings are freehand. live adult specimens were collected using a dip net from the water column. the specimens are preserved in 70% ethyl alcohol, which was changed after 24 hours. substrate from one of the localities was collected together with live eggs of the new species. frontal appendage terminology follows pereira (1983). conservation assessments follow the guidelines produced by the iucn (2001). cultures were started using techniques that simulate natural seasonal temperature and hydrologic cycles of the original collection habitat. a 1 l portion of the substrate was gently deposited in a plastic culture container (58 cm × 41 cm × 16 cm), and lightly sprayed with deionized water to prevent dust or debris and eggs from being displaced into the air during inundation. the substrate was initially inundated by adding 16 l of deionized water that had been warmed to 25°c. the culture was then gently stirred to break up substrate clumps, and gentle aeration via an air stone was added at one corner of the container. the culture was incubated at 25°c for 12 hours, then let come to room temperature. after 24 h, a second 16 l of 25°c deionized water was added to the culture container. the culture was maintained at room temperature under t12 fluorescent grow lights on a timed 12:12-h light:dark cycle. cultures were fed 5 ml to 10 ml of a diet high calcium plant flake food (worldwide aquatics, inc.) ground to a fine powder. in addition, 48 h after the second addition of water, 20 ml of a free swimming green alga culture (selenastrum sp.) was added for additional food. repositories dcr = author’s own collection flmnh = florida museum of natural history, florida, usa usnm = national museum of natural history, washington, d.c., usa results subphylum crustacea brünnich, 1772 class branchiopoda latreille, 1817 order anostraca sars, 1867 family thamnocephalidae packard, 1883 (sensu rogers 2006) genus dendrocephalus daday, 1908 (sensu rogers 2006) subgenus dendrocephalinus rogers, 2006 dendrocephalinus rogers, 2006: 7, 12, fig. 1d. chirocephalus – creaser 1940: 435, figs 1–6. rogers d.c. et al. review of dendrocephalinus 3 branchinella – dexter 1953: 756, 761, figs 24–28. — moore 1967: 219, figs 1–3. — belk & sissom 1992: 312, figs 1–3. — belk & brtek 1995: 323, 325. — brendonck 1997: 451–454. — brtek & mura 2000: 1077, figs12–17. — brtek 1997: 21–22. — brtek 2002: 54. type species chirocephalus lithacus creaser, 1940 (by designation herein) diagnosis (updated from rogers 2006) fully extended gonopods extending to third or base of fourth abdominal segment. gonopod basal portion chitinised, rigid, subcylindrical, bearing a single ventromedial tubercle. gonopod distal eversible portion soft, distal third expanded, comb plate lacking, distal half with four longitudinal rows of short spines, each row evenly spaced around gonopod. brood pouch expanded proximally, distal portion either narrowing abruptly to posteriorly directed subcylindrical tube, extending to base of fourth to eighth abdominal segment, or absent. male abdomen first two or three segments with ventrolateral spiniform projections. male first thoracopods unmodified. nearctic (usa, mexico (?)). all species occur within the coastal plain anostracan biogeographic region (rogers, 2014a). remarks we suspect that this taxon may actually represent an independent genus, however molecular analyses will be required to demonstrate this. at this time we choose to conservatively maintain the taxon as a subgenus. dendrocephalus (dendrocephalinus) acacioidea (belk & sissom, 1992) fig. 1a, d branchinella acacioidea belk & sissom, 1992: 312, figs. 1–3. branchinella acacioidea – belk & brtek 1995: 323. — brendonck 1997: 451, 454. — brtek 1997: 21; 2002: 54. dendrocephalus (dendrocephalinus) acacioidea – rogers 2006: 7, 12, fig. 1d. material examined usa • 15 ♀♀, 11 ♂♂; texas, hidalgo county, roadside pool, east side of texas 281, 12.9 km north of edenberg; 26°25΄ n, 98°30΄ w; 21 aug. 1980; s.l. sissom leg.; gift from d. belk; dcr #321. remarks this species is unique in that the 2a sub-branch (the apical branch) is absent, and sub-branch 2v and 1d are identical. the female shares the elongated brood pouch form with d. proeliator sp. nov. the only member of the subgenus occurring outside the extreme southeastern usa: all records are from southern, coastal texas. all known locality substrates have 10–15% calcium carbonate and 2–8 ms/cm salinity (rogers 2014b). so far, this species is known from three nearby locations, all seem to be on either private land or in public right of ways (belk & sissom 1992). due to the limited distribution of d. acacioidea, this species meets the iucn red list cr b1a,b species criteria (iucn 2000), which means the species is critically endangered, with an extent of occupancy less than 100 km2, and a projected decline in extent of occurrence. european journal of taxonomy 509: 1–14 (2019) 4 fig. 1. dendrocephalus spec. a. dendrocephalus acacioidea, male head, left side, anterior view. b. dendrocephalus alachua, male head, left side, anterior view. c. dendrocephalus lithacus, male head, left side, anterior view. d. dendrocephalus acacioidea, brood pouch, right, lateral view. e. dendrocephalus alachua, brood pouch, right, lateral view. f. dendrocephalus lithacus, brood pouch, right, lateral view. g. dendrocephalus lithacus, male, labrum, ventral view. h. dendrocephalus lithacus, gonopods, right, lateral view. scale bar: a–c = 1 mm; d–f, h = 4 mm; g = 0.25 mm. rogers d.c. et al. review of dendrocephalinus 5 dendrocephalus (dendrocephalinus) alachua (dexter, 1953) fig. 1b, e branchinella alachua dexter, 1953: 761, figs 24–28. branchinella alachua – moore 1967: 219–220. — belk & sissom, 1992: 314. — belk & brtek 1995: 323. — brendonck 1997: 451–454. — brtek 1997: 21. — brtek 2002: 54. dendrocephalus alachua – rogers 2006: 7, 12. material examined holotype usa • ♂; florida, alachua county; 6 jul. 1947; r.w. dexter leg.; usnm 93538. paratypes usa • 3 ♂♂; same data as for holotype; usnm 93539. other material usa • 5 ♀♀, 7 ♂♂, plus 26 ♀♀, 33 ♂♂ reared in culture; hillsborough county, tampa, cemetery pond, corner of martin luther king boulevard and east lake avenue; 27°58.859΄ n, 82°24.256΄ w; 13 sep. 2000; substrate collected for culture 6 jul. 2000; w.w. price leg.; dcr #772 • 41 ♀♀, 6 ♂♂; david e. west park, 2212 east 97th street, corner of martin luther king boulevard and east lake avenue; 28°02.270΄ n, 82°26.063΄ w; 8 jan. 2008; w.w. price leg.; dcr #774. remarks the original description of the female has the brood pouch elongate, almost fusiform. in many, but not all, female specimens i have cultured, the brood pouch is abbreviated, lacking the free distal portion (fig. 3e). this species is known from five unspecified collections in alachua county, florida, near gainesville (dexter 1953; moore 1967; two unpublished records at usnm), and was suggested as possibly extinct, pending further surveys, even after many surveys were performed (rogers, 2006). however, collecting efforts by w.w. price yielded two new localities in the tampa region. the tampa locality is at least 180 km south of the unspecified type locality (a “barrow pit” (dexter 1953) which has probably been destroyed (rogers 2006)). for both new locations for this species the substrate has no measurable calcium carbonate and salinity 0 to ~2 ms/cm, as both locations are heavily vegetated, and have tree cover, with the substrate covered in vegetative debris (rogers 2014b; this study). similarly, both locations are less than a meter above the water table. dendrocephalus alachua co-occurred at both new locations with the smooth clam shrimp lynceus gracilicornis packard, 1871, and the spiny clam shrimp eulimnadia cylindrova belk, 1989. both dexter (1953) and moore (1967) reported d. alachua co-occurring with the anostracan streptocephalus sealii ryder, 1879. however, this species was not present at either of the two new localities. rogers (2006) conservatively designated d. alachua as cr a1 species (iucn 2000) pending further surveys. but with these new collections this species meets the iucn red list cr b1a,b species criteria (iucn 2000), that the species is ‘critically endangered’, with an extent of occupancy less than 100 km2, and a projected decline in extent of occurrence. dendrocephalus (dendrocephalinus) lithacus (creaser, 1940) fig. 1c, g–h chirocephalus lithacus creaser, 1940: 435, figs 1–6. european journal of taxonomy 509: 1–14 (2019) 6 chirocephalus lithacus – dexter 1953: 761. branchinella lithaca – belk & sissom 1992: 314. — belk & brtek 1995: 325. — brendonck 1997: 451–454. — brtek 1997: 22. — brtek 2002: 54. dendrocephalus lithica – rogers 2006: 7, 12. material examined holotype usa • ♂; georgia, dekalb county, stone mountain, within 100 yards of airway beacon; 9 jun. 1939; usnm 79294. allotype usa • ♀; same data as for holotype; usnm 79295. paratype usa • ♀; same data as for holotype; usnm 79296. remarks this species is known from a set of rock outcrop pools on stone mountain in georgia and has only been collected twice (dexter 1953). a petition was made to list d. lithacus as endangered under the united states federal endangered species act. however, after extensive survey efforts no evidence of extant populations of d. lithacus was found (federal register may 27, 1998) and the petition was denied (rogers 2006). surveys made by the georgia department of natural resources at stone mountain and other granitic plutons in the area have so far yielded no new populations. this species is most likely extinct. the clam shrimp eulimnadia graniticola rogers et al., 2010 occurs in these pools. dendrocephalus (dendrocephalinus) proeliator sp. nov. urn:lsid:zoobank.org:act:203c0a81-a3cd-4a02-9001-47259ee2ea7c fig. 2 diagnosis males are typical of the subgenus dendrocephalinus, and readily identified in that the frontal appendage primary arms are each triramal, with branch 2a bearing an apical hamulate projection. the antenna-like appendage is extremely small and may be obscured. females are not separable. etymology the specific epithet ‘proeliator’ is latin for ‘warrior’. the gender is masculine. material examined holotype usa • ♂; florida, dade county; 25°34.781΄ n, 80°34.541΄ w; 1.5 m a.s.l., on west side of sw 237th avenue, south of grossman hammock; 30 jul. 2001; e.l. nance leg.; flmnh type 49581. allotype usa • ♀, same data as for holotype; flmnh type 49582. paratypes usa • 4 ♀♀, 1 ♂; same data as for holotype; flmnh type 9041. http://zoobank.org/urn:lsid:zoobank.org:act:203c0a81-a3cd-4a02-9001-47259ee2ea7c rogers d.c. et al. review of dendrocephalinus 7 fig. 2. dendrocephalus proeliator sp. nov. paratypes; florida, dade county; flmnh type 9041. a. male head, anterior view. b. female, right lateral view. c. right limb v, anterior view. d. two examples of the egg. abbreviations: 1v = first ventral branch; 2a = apical branch; 2d = dorsal branch (terminology follows pereira 1983). scale bar: a = 2 mm; b = 7 mm; c = 1 mm; d = 1.5 mm. european journal of taxonomy 509: 1–14 (2019) 8 other material usa • 28 ♀♀, 26 ♂♂; florida, highlands county, archbold biological station, pond # 107.1; 27°11΄36.24˝ n, 81°21΄29.88˝ w; 41 m a.s.l.; 1 jul. 2016; a. dunn; cultured material; dcr-976. type locality the type locality is pond #107.1 at the archbold biological station; a temporary pool in calcareous wet prairie. the substrate is duette sand, the pool fills to a maximum observed depth of 10 cm, and the subcircular basin is approximately 15 m across. these soils are calcareous and have a salinity of ~2 ms/ cm. the basin substrate has 100% vegetative cover, dominated by approximately 70% yellow hatpin (syngonanthus flavidus (michx.) ruhland, 1903) and 30% florida bluestem (andropogon floridanus scribn., 1896). a cluster of scrub oaks (possibly quercus geminata small, 1897) is in the center, providing shade and leaf litter. the surrounding area is scrubby flatwoods; a mixture of saw palmetto and shrubs with scattered pitch pine. description male length. average length of preserved material: 4.5 mm (males) and 8.2 mm (females) from apex of head to apex of telson. head (fig. 2a). male anteriolateral corner of head slightly projecting over eyestalk base. labrum smooth. antenna-like appendage anteriobasal to eyestalk, filiform, obscure, length approximately 0.1× eyestalk peduncle. first antenna approximately 85% the length of eye plus eyestalk. second antenna capable of extending to thoracomere ii or iii. second antennal proximal antennomere subcylindrical, smooth, with length approximately 3.0× distoapical width. proximal antennomere distal end with short, medial, lobiform projection. projection protruding slightly distally, smooth, apically rounded, length approximately 0.1× antennomere. second antennal distal antennomere length slightly longer than proximal antennomere, flattened anterioposteriorly, gently arcuate medially, with apex acute and directed medially. medial edge distal ~55% margined with fine, aciculate denticles, directed subdistally. cephalic appendage prominent, typical of the subgenus. proximal trunk subcylindrical, smooth, lacking spines. primary branches smooth, lacking spines, diverging approximately even with the distal end of the second antenna proximal antennomere. primary branch triramal. distal branches diverging at distal end of primary branch. primary branch length ~3.4 × proximal trunk. branch 2v thick, flattened, with numerous semiannulations. length ~0.7 × primary branch. apex tapering in distal third. distal fourth with four lateral lobes on each margin. lobes and apex tipped with an acute spine. spine length ~2.5 × basal width. branch 2d flagelliform, subequal in length to primary branch, lacking spines or setae. branch 2a roughly triangular, with a chitinised tumid area covering approximately one third of the lateral surface, bearing rows of large chitinised, spiniform projections. spiniform projections smaller distally and medially, length ranging from ~0.5 × to 1.0 × basal width. spiniform projections posteriorly and laterally becoming larger, subcylindrical, length ranging from ~1.0 × to 2.0 × basal width, each bearing an apical ring of spinulae. spinulae number three to five per projection. branch 2a distally chitinised, hamulate, with apex subacute. distal hamulate projection chitinised, length ~0.25 total branch 2a length. maxillae lobiform and transverse, no setae visible. thoracopods (fig. 2c). serially homologous. thoracopod v with endites margined with long, plumose setae. endite vi with two distal spines. endopod subquadrate, with distal margin bearing a submedial emargination. endopod with distomedial margin bearing a row of acute spines, length ~3 × basal width, separated by their basal width. endopod distal margin mesad of emargination with four spines, slightly curved posteriorly, length ~4 × basal width, separated by their basal width. endopod distal margin lateral of emargination bearing a row of cirriform setae, length ~5.0 × to 6.0 × the distal spines. endopod of all thoracopods, except thoracopod i lack setae on basomedial margin. thoracopod i and ii endopods rogers d.c. et al. review of dendrocephalinus 9 as in other limbs, although the basomedial margin bears a few obscure filiform setae. exopod lamellar, sinuate, margined in distal half with long, plumose, recurved setae. epipodite elliptic and without marginal setae. margins crenulate. praepipodite broadly triangular, lamellar, without marginal setae, bearing a light mediolateral cleft. genitalia. genital segments smooth. retracted gonopod as for genus. everted gonopods not observed. abdomen smooth, cercopods ⅔ as long as abdomen, margined with plumose setae. female generally as typical for the genus (fig. 2b). head. rounded, anterolateral corners slightly projecting over eyestalk base. first antenna length 0.25 × second antenna. second antenna broadly elliptic, lamellar. thorax smooth. thoracopods as in male, except endopods lacking emargination. brood pouch. fusiform, extending to proximal margin of postgenital abdominal segment iv. eggs subspherical, diameter approximately 190 μm, with large, broad pentagonal or quadragonal facies, each face diameter approximately 60 μm (fig. 2d). differential diagnosis of the four species of dendrocephalus (dendrocephalinus), males of d. proeliator sp. nov. are morphologically intermediate between d. lithacus (fig. 1c) and d. alachua (fig. 1b). these three species are easily differentiated from d. acacioidea (fig. 1a) which lacks branch 2a in the frontal appendage. the three remaining species bear the lateral, broad surface of branch 2a with its rows of spiniform projections. dendrocephalus proeliator sp. nov. bears three rami, as does d. alachua, and the three rami are superficially similar in overall structure between the two taxa. however, the hamulate distal projection of d. proeliator sp. nov. is absent in d. alachua. this hamulate distal projection is present in d. lithacus. dendrocephalus proeliator sp. nov. is further separated from d. alachua in that branch 2v has eight total lateral lobes verses nine in d. alachua, and branch 2d is long, flagelliform and smooth in d. proeliator sp. nov. instead of vermiform and bearing rings of spines in d. alachua. furthermore, branch 2d is proportionately much longer in d. proeliator sp. nov. than in d. alachua. dendrocephalus alachua also has a distomedial spiniform projection at the apex of the second antennal distal antennomere, which is lacking in d. proeliator sp. nov. dendrocephalus lithacus lacks the branch 2d found in the other two species, and branch 2a is proportionately larger and more robust than the condition found in d. proeliator sp. nov. the hamulate projection of branch 2a in d. proeliator sp. nov. is approximately one fourth the total length of the branch, whereas in d. lithacus it is nearly half the branch length. dendrocephalus lithacus also bears a row of fine spinulae on the second antenna proximal antennomere just basal to the frontal appendage, and a second row on the posteriolateral margin of the frontal appendage main trunk. these are lacking in d. proeliator sp. nov. and d. alachua. the second antennae distal antennomere medial margin bears spaced transverse grooves in d. lithacus versus a row of fine denticles in d. proeliator sp. nov. european journal of taxonomy 509: 1–14 (2019) 10 distribution and habitat so far, this species is only known from two locations in southern peninsular florida. both sites are temporary pools in calcareous wet prairie. both sites are in sandy loam soils, with 50–90% calcium carbonate, and salinity around 2.0 ms/cm. the pool south of grossman hammock is inside everglades national park, in an area of state managed lands in water conservation area 3b (kambly & moorehead 2009). the type locality resides within the land managed by the archbold biological station, established in 1941 by archibold expeditions, a non-profit, independent research organisation, based in florida. due to the limited distribution of d. proeliator sp. nov., this species meets the iucn red list cr b1a,b species criteria (iucn 2000), which means the species is ‘critically endangered’, with an extent of occupancy less than 100 km2, and a projected decline in extent of occurrence. it is our hope that future surveys will find additional populations. behaviour mate searching behaviour was described in detail by rogers (2019). anostracans engage in hill topping behaviour, wherein salient objects are used as congregation points for males and receptive females. we observed this same hill topping behaviour in d. alachua and d. proeliator sp. nov. however, both species engaged in male-male agonistic interactions, wherein males actively attempt to drive each other away from hill topping points. males would patrol salient objects as described by rogers (2019). when another male was encountered (fig. 3a), they would attempt to inspect each other as potential mates (sensu rogers 2019), but in attempting to inspect, each would not let the other beneath them, and they would circle, always facing each other (fig. 3b). this would lead to instances of butting heads and actively feeling each other with the first antennae (fig. 3c), interspersed with more circling. occasionally, one would be able to get past its opponent’s head and push them on the side (fig. 3d). typically, if a male had eyes closer set, or was pushed once or twice on the side, it would flee the other male (fig. 3e–f). presumably, the male with the wider set eyes would be larger and so the smaller one would flee to find a new hill topping site. the activity of the first antennae during these encounters may be entirely tactile, or there may be some sort of chemosensory occurring as is observed in agonistic behaviour in crayfish (bergman & moore 2005). key to the species of dendrocephalinus 1. male frontal appendage with each main branch bearing two sub-branches ..................................... 2 – male frontal appendage with each main branch bearing three sub-branches ................................... 3 2. frontal appendage with sub-branch 1v, lamellar, margined with spiniform papillae; sub-branch 2d absent; sub-branch 2a broad, with chitisied papillae on lateral surface, terminating in a chitinised, hamulate spine (fig. 1c) ..................................................................................................................... ................................ dendrocephalus lithacus (creaser, 1940) (usa: georgia, possibly extinct) – frontal appendage with sub-branch 1v and 2d lamellar, margined with spiniform papillae; subbranch 2a absent (fig. 1a) ..... dendrocephalus acacioidea (belk & sissom, 1992) (usa: texas) 3. frontal appendage with sub-branch 2d lacking spines or papillae; sub-branch 2a terminating in a chitinised, hamulate spine (fig. 2a) .............. dendrocephalus proeliator sp. nov. (usa: florida) rogers d.c. et al. review of dendrocephalinus 11 fig. 3. dendrocephalus proeliator sp. nov, male-male aggressive interactions. a. encounter. b. circling. c. head to head attack. d. broadside attack. e–f. chasing. european journal of taxonomy 509: 1–14 (2019) 12 – frontal appendage with sub-branch 2d with small scattered spines; sub-branch 2a lacking a chitinised, hamulate spine (fig. 1b) ......... dendrocephalus alachua (dexter, 1953) (usa: florida) discussion the subgenus dendrocephalinus was suggested to have split from dendrocephalus sometime in the late paleozoic (around 270 million years ago) when south america and north america were together at the equator, or at some point dendrocephalinus was the result of founder dendrocephalus populations invading north america (rogers 2006). the fact that d. acacioidea is quite different in terms of frontal appendage morphology and is disjunct from the other three species, suggests that it may be basal to the other species. the strong similarity of gonopodal morphology leaves little doubt that it belongs in dendrocephalinus. dendrocephalinus is smaller, more gracile, and the frontal appendage is greatly reduced and less elaborate than in dendrocephalus s. str. sexually mature adult preserved dendrocephalus s. str. material in dcr’s collection range from 10 to 16 mm total body length (from anterior margin of the head to posterior margin of the telson), whereas in dendrocephalinus the range is from 4 to 9 mm. the frontal appendage of dendrocephalus s. str. has two ventral branches (1v, which is biramal, and 2v), a bior triramal dorsal branch (2d), and an apical branch that bears some form of a chitinised hook (2a). in dendrocephalinus only the single ventral branch remains (1v), and it is of a single ramus that strongly resembles the 1v anterior sub-branch of dendrocephalus s. str. the 2d branch, if present, is only one ramus, which either resembles 1v or is flagellar. the 2a, if present, is broad, and covered with chitinised papillae, and may or may not have the apical hook. the elaborate male-male aggressive behaviour over hill topping sites is competition for access to receptive females. anostracans live in seasonally astatic aquatic habitats with highly stochastic hydroperiods. this would suggest that aggressive interactions would be time ill spent; the limited hydroperiod would necessitate that the animals focus on reproduction. however, these species live in the coastal plain anostracan bioregion (rogers 2014a), which frequently has long periods of precipitation, particularly in the summer months, from tropical storms and hurricanes. thus, it is likely that these pools have prolonged hydroperiods providing time to engage in more competitive actions. with the discovery of a new species in the subgenus, the existing key (rogers & hann 2016) is no longer useful. thus, we provide an updated key to the subgenus here. acknowledgments the authors are especially grateful to trisha spears (florida state university, tallahassee), john jensen, thomas floyd, and colleen cunningham (the georgia natural heritage program) for their help collecting in florida and georgia. we are also grateful for the generous and kind reviews provided by luc brendonck and an anonymous reviewer. references belk d. & brtek j. 1995. checklist of the anostraca. hydrobiologia 298: 315–353. https://doi.org/10.1007/bf00033826 belk d. & sissom s.l. 1992. new branchinella (anostraca) from texas, u.s.a., and the problem of antennalike processes. journal of crustacean biology 12: 312–316. https://doi.org/10.2307/1549083 belk d., shug belk m. & reading k.a.l. 2002. survey of large branchiopods on aruba and observations on taxonomic characters in leptestheria (spinicaudata). hydrobiologia 486: 115–118. https://doi.org/10.1007/bf00033826 https://doi.org/10.2307/1549083 rogers d.c. et al. review of dendrocephalinus 13 https://doi.org/10.1023/a:1021386315348 bergman d.a. & moore p.a. 2005. the role of chemical signals in the social behavior of crayfish. chemical senses 30 (supplement 1): 305–306. https://doi.org/10.1093/chemse/bjh236 brendonck l. 1997. the anostracan genus branchinella (crustacea: branchiopoda), in need of a taxonomic revision; 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(eds) thorp & covich’s freshwater invertebrates, 4th edition, volume ii: keys to the nearctic fauna: 437–477. academic press. london. https://doi.org/10.1023/a:1021386315348 https://doi.org/10.1093/chemse/bjh236 https://doi.org/10.1111/j.1096-3642.1997.tb00142.x https://doi.org/10.2307/2485207 https://doi.org/10.2307/3224696 https://doi.org/10.2307/1548264 https://doi.org/10.5852/ejt.2018.478 https://doi.org/10.11646/zootaxa.257.1.1 https://doi.org/10.11646/zootaxa.3838.3.1 https://doi.org/10.11646/zootaxa.3856.1.1 https://doi.org/10.1093/jcbiol/ruy099 european journal of taxonomy 509: 1–14 (2019) 14 rogers d.c. & volcan m.v. 2016. a new dendrocephalus (crustacea, anostraca, thamnocephalidae) from rio grande do sul state, brazil. nauplius 24: 1–9. https://doi.org/10.1590/2358-2936e2016014 rogers d.c., weeks s.c. & hoeh w.r. 2010. a new species of eulimnadia (crustacea; branchiopoda; diplostraca; spinicaudata) from north america. zootaxa 2413: 61–68. rogers d.c., gomes j.p.c. & viera f. 2012. a new species of dendrocephalus (crustacea, anostraca) from serra dos carajás (pará state, brasil). zootaxa 3363: 52–58. rogers d.c., cohen r.g. & hann b.j. in press. class branchiopoda. in: m.c. damborena, d.c. rogers, & j.h. thorp (eds). thorp & covich’s freshwater invertebrates, 4th edition, volume v: keys to the neotropical fauna. academic press. london. manuscript received: 30 october 2018 manuscript accepted: 31 january 2019 published on: 26 march 2019 topic editor: rudy jocqué desk editor: eva-maria levermann printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany. https://doi.org/10.1590/2358-2936e2016014 european journal of taxonomy 125: 1–23 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2015.125 www.europeanjournaloftaxonomy.eu 2015 · minelli a. this work is licensed under a creative commons attribution 3.0 license. o p i n i o n p a p e r urn:lsid:zoobank.org:pub:68f7ac91-1add-4a1e-8c5b-6f1cdcdc9dd4 1 biological systematics in the evo-devo era alessandro minelli university of padova, department of biology, via ugo bassi 58 b, i -35131 padova, italy e-mail: alessandro.minelli@unipd.it urn:lsid:zoobank.org:author:d9fa639f-7538-441c-9572-032fd6c4eede abstract. evolutionary developmental biology (evo-devo) suggests a distinction between modular and systemic variation. in the case of modular change, the conservation of the overall structure helps recognizing affinities, while a single, fast evolving module is likely to produce a bonanza for the taxonomist, while systemic changes produce strongly deviating morphologies that cause problems in tracing homologies. similarly, changes affecting the whole life cycle are more challenging than those limited to one stage. developmental modularity is a precondition for heterochrony. analyzing a matrix of morphological data for paedomorphic taxa requires special care. it is, however, possible to extract phylogenetic signal from heterochronic patterns. the taxonomist should pay attention to the intricacies of the genotype→phenotype map. when using genetic data to infer phylogeny, a comparison of gene sequences is just a first step. to bridge the gap between genes and morphology we should consider the spatial and temporal patterns of gene expression, and their regulation. minor genetic change can have major phenotypic effects, sometimes suggesting saltational evolution. evo-devo is also relevant in respect to speciation: changes in developmental schedules are often implicated in the divergence between sympatric morphs, and a developmental modulation of ‘temporal phenotypes’ appears to be responsible for many cases of speciation. keywords. genotype→phenotype map; heterochrony; saltational evolution; speciation; temporal phenotype minelli a. 2015. biological systematics in the evo-devo era. european journal of taxonomy 125: 1–23. http:// dx.doi.org/10.5852/ejt.2015.125 introduction in the last two centuries, the relationships between biological systematics and developmental biology have widely fluctuated. looking at these relationships from the vantage point of systematics, the basic fact is that the vast majority of taxa have always been diagnosed on adult characters only, especially in the case of species and lower supraspecific taxa. problems with earlier stages are of different nature, among which: objectively limited range of variation, compared with characters only displayed by the adult – think of copulatory organs and secondary sexual characters in animals, or flower and fruit traits in flowering plants inadequate presence of non-adult specimens in collections, except for select groups such as amphibiotic insects http://dx.doi.org/10.5852/ejt.2015.125 http://www.europeanjournaloftaxonomy.eu/index.php/ejt https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:68f7ac91-1add-4a1e-8c5b-6f1cdcdc9dd4 mailto:alessandro.minelli%40unipd.it?subject=alessandro.minelli%40unipd.it http://zoobank.org/urn:lsid:zoobank.org:author:d9fa639f-7538-441c-9572-032fd6c4eede http://dx.doi.org/10.5852/ejt.2015.125 http://dx.doi.org/10.5852/ejt.2015.125 european journal of taxonomy 125: 1–23 (2015) 2 lack of information about the relationships between stages (what is the adult of this larva, or juvenile, and vice versa) lacking or limited interest in a precise identification of non-adult specimens scattered but disquieting evidence that juveniles or larvae may eventually suggest a different pattern of relationships than the corresponding adults (e.g., brassoline nymphalids (penz et al. 2013) and chalcosiine zygaenids (yen et al. 2005) among the lepidoptera; staphyliniformia (beutel & leschen 2005) and melolonthine scarabs (allsopp & lambkin 2006) among the coleoptera; ‘nematocerans’ (oosterbroek & courtney 1995), sabethine mosquitoes (judd 1998) and sepsids (meier 1996) among the diptera; see also a critical discussion in meier & lim (2009)). the problem is worsened by the lack of a sound theoretical background against which to look for a satisfactory resolution of this conflict to be sure, none of these difficulties should be taken as universal and unanswerable. in particular, it is fair to remark: the abundance of useful diagnostic characters, other than those found in the adults, that are exhibited by non-adult specimens in many groups – exemplary are dragonflies and some groups of lepidoptera the taxa of which non-adult specimens are much more accessible than the corresponding (often unknown) adults, and offer adequate diagnostic characters upon which species description and also a generic and suprageneric classification can be based; this is the case of many families of mites (a good example, relative to the erythraeid genus leptus latreille, 1796, is southcott 1992), the species of which are routinarily described on larvae more than at lower taxonomic levels, information about early ontogenetic stages has often been regarded as informative of affinities among major lineages. this has even contributed to the erection of supraphyletic assemblages, the names of which explicitly refer to developmental processes, such as protostomia and deuterostomia. an obvious great success of the strategy of inferring kinship from the shared possession of early developmental traits was the recognizion by kowalewski (1866) of the close affinities of vertebrates with what we now acknowledge as the non-vertebrate chordates. curiously, exactly in the same year haeckel (1866) formulated his biogenetic law, according to which phylogeny recapitulates ontogeny. there is no need to summarize here the history of the fading acceptance of haeckel’s principle in the following decades. the interested reader is referred to gould’s (1977) book for a detailed account. it is, however, fair to mention here that, despite the increasing disrepute into which falls the biogenetic law, an interest for ontogeny survived nevertheless in comparative biology and, specifically, in systematics. an ontogenetic criterion to establish homology between features of two different organisms is present indeed in remane’s (1952) classic (pre-cladistic) approach to comparative anatomy, and was suggested by hennig (1966) as a potential way to identify character polarity, a key step in reconstructing phylogeny. a major conceptual limit of all these approaches was, however, the lack of a comprehensive scheme within which to interpret the relationships between development and evolution. ironically, despite obvious progress in both fields, the problem effectively worsened during the central decades of the past century, because of the lack of dialogue between evolutionary biology and developmental biology (amundson 2005). a renewed interested in the relationships between ontogeny and phylogeny or, more generally, between developmental biology and evolutionary biology, emerged eventually after the publication of gould’s (1977) book, significantly titled ontogeny and phylogeny, and rapidly consolidated, first of all around the rapidly successful growth of comparative developmental genetics – witness raff & kaufman’s (1983) landmark volume on embryos, genes and evolution. soon thereafter, the new discipline of evolutionary developmental biology (more popular under the nickname of evodevo) emerged, its name deriving from the title of hall’s (1992) well-targeted book. minelli a., biological systematics in the evo-devo era 3 today, we can characterize evolutionary developmental biology as a very active frontier of the life sciences. a perception of its scope and research agenda can be obtained by a perusal of books such as hall (1998) and hall & olson (2003) or, perhaps more sizeably, by browsing the volumes of the few dedicated journals launched thus far (evolution & development; journal of experimental zoology (molecular and developmental evolution); evo-devo). it must be acknowledged that evo-devo’s boundaries with respect to the parent disciplines – evolutionary biology and developmental biology – are still far from being well-defined (see, e.g., arthur 2002; müller 2008), partly because evo-devo itself has stimulated a revisitation of the internal structure and research scope of both developmental biology (e.g., minelli & pradeu 2014) and evolutionary biology (e.g., pigliucci & müller 2010). the consequences of the advent of evo-devo for a revisitation of the conceptual foundations and the improvement of practice in taxonomy are multiple and potentially far-reaching, as numerous examples demonstrate, some of which are discussed in this article. however, to date the issue has been explicitly and comprehensively addressed only by a few authors (telford & budd 2003; cracraft 2005; minelli 2007, 2009, in press; minelli et al. 2007; minelli & fusco 2012). before surveying, in the remaining of this article, key areas in which biological systematics can benefit from intellectual exchange with evo-devo, it is fair to mention that evo-devo, in turn, necessitates help from taxonomy. at the most fundamental level, taxonomy is needed to get a correct identification of the species used in experimental research. this is a far from marginal point, not only because many experimental biologists do not care enough for ascertaining the identity of the organisms used in the lab, especially when these belong to taxa very numerous in species and taxonomically difficult, but also because of the widespread occurrence of cryptic species that can be taken apart and eventually given a name only following adequate specialist study. emblematic is the case of the ascidian ciona intestinalis (linnaeus, 1767). only five years after its genome sequence was published (dehal et al. 2002), it was realized that the stocks labeled with this name and held in the labs in different parts of the world actually belonged to two different species, reproductively isolated but hardly distinguishable on morphological traits (iannelli et al. 2007). multiple species have also been treated as one in the case of the leech helobdella triserialis (e. blanchard, 1849), another popular model species (siddall & borda 2003; bely & weisblat 2006; weisblat & kuo 2009; kutschera et al. 2013). the choice of new model species is another problem in relation to which taxonomy – or, more explicitly, phylogenetics – is sometimes called for help by biologists interested in the evolution of development. this choice, however, is generally misguided, being based on misconceptions about phylogenetics and character evolution (discussed in jenner 2006; minelli & baedke 2014). developmental genes and phenotype the so-called developmental genes, i.e., the genes with better documented effects on the control of development, have attracted interest as potential sources of useful phylogenetic information. several studies have focused on the hox genes, an obvious choice in view of their role in controlling major aspects of the overall body architecture such as the orderly sequence of body features along the anteroposterior body axis. for example, hox gene sequences have been used in reconstructing the relationships among bilaterian phyla (de rosa et al. 1999; balavoine et al. 2002; passamaneck & halanych 2004; hueber et al. 2013), or to investigate phylogeny within large phyla (e.g., arthropoda: cook et al. 2001; janssen et al. 2014). the path is long, however, from the gene sequence to a morphological phenotype somehow depending on its expression. european journal of taxonomy 125: 1–23 (2015) 4 the relationships between genotype and phenotype (the so-called genotype→phenotype map) are now largely acknowledged as complex and non-linear (e.g., alberch 1991; altenberg 1995; west-eberhard 2003; pigliucci 2010; wagner & zhang 2011). in other terms, rarely, if ever, does one gene correspond to one phenotypic trait, and vice versa. as a rule, a diversity of phenotypic traits is affected by the expression of one gene (pleiotropy), and different genes, or genetic cascades, may translate into undistinguishable phenotypes (convergence and/or redundancy). this is exactly the place where evo-devo, with its focus on comparative patterns of expression of developmental genes, can play an important role in the service of systematics. next to comparing gene sequences, we can in fact move to compare gene expression patterns, in order to trace homologies, especially in cases where morphological evidence does not seem to be conductive to a definitive assessment. interesting results have been obtained, for example, by hughes & kaufman (2002), copf et al. (2003) and angelini & kaufman (2005) in tracing homologies between body regions of distantly related arthropod taxa by comparing expression patterns of hox genes. the same approach was used by jager et al. (2006) (see also manuel et al. 2006) to align the anterior appendages of pycnogonids with those of other arthropods. an interesting exercise of ‘evo-devo cladistics’ has recently been presented by stach (2014) for the major lineages of the deuterostomes, using a matrix of data such as the number of copies of genes in each hox paralogous group (hox1 to hox13; his characters ## 1, 6, 11, 16, 21, … 61), but also bmpexpression on the prospective oral and aboral sides of the embryo (his characters ## 67 and 68). matrix entries for characters of the latter kind result from a direct translation of the homology hypotheses implicit in evo-devo studies. as said above, the relationships between change in gene sequence and change in phenotype are anything but linear. wray et al. (2003) distinguished the following classes of evolutionary change in gene expression: changes in the spatial extent of gene expression. (e.g., schiff et al. 1992; brunetti et al. 2001; scemama et al. 2002) changes in the timing of gene expression (transcriptional heterochrony), as documented for many taxa (e.g., wray & mcclay 1989; kim et al. 2000; skaer et al. 2002). for example, the bristle pattern on the notum of diptera is regulated by the expression of the gene scute. differences between more distantly related taxa, such as ceratitis capitata (wiedemann, 1824) (tephritidae) (wülbeck & simpson 2000), drosophila melanogaster meigen, 1830 (drosophilidae) (romani et al. 1989) and calliphora vicina robineau-desvoidy, 1830 (calliphoridae) (pistillo et al. 2002) result from different regulations of the spatial expression patterns of scute, but in the presence of identical spatial regulation, differences can emerge, due to differences in the timing of this gene’s expression, as revealed by the differences in the bristle pattern of calliphora vicina compared to another calliphorid, protophormia terranovae (robineau-desvoidy, 1830) (skaer et al. 2002) changes in the level of gene expression. a path-breaking study on the developmental mechanisms underlying differences in beak shape among darwin’s finches (geospiza gould, 1837 spp.) revealed that these are caused by the differential levels of expression of just two genes (bmp4 and calmodulin) (abzhanov et al. 2004, 2006). in drosophila fallén, 1823, marcellini & simpson (2006) found that the development of the dorsocentral bristles – whose number and spatial arrangement are extensively used in species-level taxonomy in this genus and in other diptera – depended on a genetic regulatory element, the dorsocentral enhancer, involved in the activatation of the scute gene in a cluster of cells from which, in species such as d. melanogaster, two bristles on the posterior scutum arise. in a few species, however, e.g., d. quadrilineata meijere, 1911, a derived state has evolved, with anterior as well as posterior dorsocentral bristles. this character state correlates with an anterior expansion of the expression domain of the scute gene. this is in turn an effect of an evolutionary change in minelli a., biological systematics in the evo-devo era 5 the nucleotide sequence of the enhancer. the authors present this case as an example of how gene expression patterns can be modified by a simple “evolutionary tinkering” of pre-existing enhancers, thus contributing to morphological diversification. the scope of variation modular vs. systemic variation in a biology dominated by explanations of the structure of living beings in terms of adaptations, such as was the dominant perspective during much of the xix and xx centuries, it was usual to stress the functional integration among body parts, in animals especially. this was remarkable in cuvier’s (1812) pre-evolutionary law of the correlation among parts, as much as in darwin’s (1859) view of the individual organism as the main unit of selection; the same perspective was also dominant in the evolutionary synthesis of mid xxth century. things have changed in the last few decades, largely under the impact of evolutionary developmental biology. this discipline characteristically focuses on the evolvability of organismal traits (hendrikse et al. 2007), that is, on the arrival of the fittest rather than on the survival of the fittest. focussing on developing systems, rather than on the adaptive value of the eventually ensuing phenotypes, invites acknowledging that a developing organism is a system of local modules dominated by specific dynamics (e.g., wagner 1996; schlosser & wagner 2003; klingenberg 2005). eventually, a body part, which is ‘substantially’ individualized in developmental terms, turns out to represent one of those units the morphologist calls homologues (wagner 1989). in evolutionary terms, modularity is a precondition for the occurrence of heterochrony, as a loose causal and functional coupling between modules allows these to change their dynamic parameters without necessarily disrupting the whole organism’s ontogeny. however, the units of evolutionary change do not necessarily coincide with developmental modules. changes are sometimes modular, affecting individual modules that often emerge as hot points of morphological evolution, but in other instances evolutionary change is systemic, affecting – more or less literally – the whole organism (minelli 2015). this distinction between modular and systemic change is of utmost interest to the taxonomist. in the case of localized, modular change, the conservation of the overall structure helps in recognizing the affinities of the involved species, and the fast evolving module is likely to produce a bonanza for the taxonomist. this is the case with the copulatory structures of a great number of insect groups, with the chewing structures (mastax) of rotifers, with the gonopods of the males of helminthomorph millipedes. the last case deserves a more detailed account. in these millipedes, the number of walking legs varies between 32 and 375 pairs. except for minor peculiarities of the first pair, or the first few pairs, all these walking appendages are morphologically identical in the females and in male juveniles. new pairs of legs are added at each moult. at later stages, however, the eighth pair of legs, often also the ninth, of the male undergo a unique metamorphosis (non-systemic metamorphosis; drago et al. 2008, 2011) and are replaced in the adult by specialized sexual appendages (gonopods) used as claspers or to transfer sperm. in most millipede species, these appendages differ dramatically from walking legs, and show up a fantastic diversity of shapes, often too complex to be adequately described in words. these gonopods are by far the single most important source of taxonomic characters for millipede taxonomy. things are completely different wherever morphological evolution has been systemic, rather than modular. in this case, the whole body architecture is likely to have been so deeply affected as to cause great difficulty in recognizing (on morphology alone) the actual affinities of the derived lineage. two european journal of taxonomy 125: 1–23 (2015) 6 conspicuous examples of systemic morphological evolution are provided by two plant groups, the riverweeds and the duckweeds. the riverweeds (family podostemaceae) are tropical aquatic plants, whose vegetative structures deviate strongly in comparison with all other flowering plants. in the most derived species, none of the conventional vegetative parts of angiosperms are recognizable and the whole plant is more reminiscent of an alga than of a plant with roots, stem, branches and leaves. duckweeds (araceae: lemnoideae) have been found to represent a basally splitting lineage within the arum family (henriquez et al. 2014), but their vegetative structure is the most unusual among the flowering plants: the whole photosynthetic structure is represented here by an irregular disk-shaped body, or by a small cluster of such small, floating bodies, out of which one or more roots sprout out. the ultimate member of this lineage is wolffia arrhiza (l.) horkel ex wimm., a simple subspherical blob of green matter, perhaps one millimeter in diameter. the phenotypic consequences of evolvability are sometimes limited to a single developmental stage, sometimes pervasive over the whole life cycle. think of cirripeds like balanus da costa, 1778 and lepas linnaeus, 1758, which begin the post-embryonic phase of development as typical crustacean larvae, but later change into very unusual adults whose real affinities were discovered only when their metamorphosis was first observed by thompson (1830, 1835). this contrasts with organisms represented along their entire life cycle by oddly shaped stages such as the minuscule cycliophorans that live on the appendages of the norwegian lobster: to describe their developmental stages, zoologists were forced to introduce new terms such as the pandora larva and the prometheus larva, because the terminology available for other animals did not offer adequate labels for these unique forms (obst & funch 2003), whose phylogenetic affinities still remain problematic (sørensen et al. 2000; giribet et al. 2004; neves et al. 2012). ontogenetic patterns of variation most people, a majority of biologists included, tacitly take for granted that development is a sequence of events proceeding from an egg (or a seed, a spore, a bud) towards the adult, with an accompanying increase in complexity, the latter being theoretically difficult to define, but suggested in practice, for example, by the number and diversity of cell types, or differentiated body parts (e.g., bonner 1988; mcshea 2000). this view of development squares well with von baer’s (1828) ‘law’ that traced a parallelism between the ontogenetic progression throughout embryonic development and the emergence of morphological characters diagnostic of increasingly lower taxa. however, this popular view of development is conceptually unsatisfactory (minelli 2011, 2014) and potentially misleading as a guide to extracting information of taxonomic or phylogenetic scope. in part, the problem is caused by heterochrony, a feature of comparative developmental biology of which haeckel was already well aware and to which i will return below. but there is another aspect, only revealed by the recent advances in evo-devo. the range of variation does not regularly increase in time, beginning with egg cleavage. irrespective of our preferences in fixing the starting point and the end of a developmental sequence, the most conservative step is not the egg (which is a very specialized cell; boyden & shelswell 1959; song et al. 2006; minelli 2014), but a multicellular stage called the phylotypic stage (sander 1983; slack et al. 1993) such as the germ-band stage in arthropods and the pharyngula in vertebrates; or, perhaps more sensibly, a phylotypic period (richardson 1995; richardson et al. 1997). if we track the ontogenetic pathways of representatives of a given phylum starting from the zygote, we will see them converging towards the phylotypic stage, and only moving ahead from this stage do they start diverging according to the popular, von baerian model. the latter, eventually, has been replaced by an hourglass model, the narrowest point of which is the phylotypic stage (duboule 1994). minelli a., biological systematics in the evo-devo era 7 this analysis may eventually turn into a guide to searching for phylogenetically more informative stages along an ontogenetic trajectory. the conservative phylotypic stage is arguably a choice target if we are looking for characters useful to diagnose higher-level taxa, whereas for information useful in comparisons between lower taxa, e.g., species ordinarily classified in the same genus, we are advised to look elsewhere. in different groups, different developmental phases may turn out to be more variable and informative. for example, in a comparative study of the developmental sequences of many species of niphargus schiödte, 1847 (crustacea: amphipoda), fišer et al. (2008) found that spine development and shifts in allometric growth do not occur at random times, the highest degree of heterochrony (a proxy for independence between events) being found mainly in early mid-aged instars. similar studies on other taxa may well be rewarding. founder evolvability and character independence systematists should always try to check for the mutual independence of the traits on which a phylogenetic analysis will be performed. however, convincing evidence of independence, or of a lack of it, is not always easy to obtain. as a consequence, every bit of information, even of circumstantial nature, is welcome. useful suggestions can derive, for example, by examining data in the light of flohr et al.’s (2013: 20663) concept of founder evolvability, defined “as the maximal range of derived organisms with different niches that can be accessed from the founding ancestor by mutation and recombination over an interval of evolutionary time.” the authors’ experiments on bacteria have demonstrated how the dynamics of adaptive radiation are constrained by the niche occupied by the founder. the principle, however, applies also to natural radiations, especially if these evolved in exclusive and adaptively demanding environments. in the specific case of evolutionary lineages characterized by a trend towards decreasing body complexity, the principle of founder evolvability turns out to be similar to the economic principle of diminishing returns: the more you have cut away, the less remains to be further deleted in the future. this may provide a simple explanation for the trend observed in the parasitic flatworms by brooks & mclennan (1993). of 1882 character transformations implied by their phylogeny, only 10.8% could be described as a loss. but it could hardly be otherwise: on the principle of diminishing returns, a high prevalence of anatomical simplification in the nodes closest to the root of the tree has arguably determined the scarcity of further losses along the tree branches. discontinuous variation virtual characters a quite unexpected peculiarity of the genotype→phenotype map is the fact that the phenotypic expression of a gene, or gene network, can disappear for a long time, but still remain ready to resurface, following minor genetic change or a targeted external stimulus. for example, simple treatments with hormones have been shown to release the expression of complex phenotypes not known to occur today in nature. in this way, kim et al. (2006) were able to obtain males from all-female populations of cladocerans. the treatment caused the (re)appearance of sex-specific characters that were subsequently used in taxonomic identification and phylogenetic inference. a similar treatment was applied by glenner et al. (2008) (see also scholtz 2008) to representatives of a crustacean clade only known for early larval stages (the so-called y-larvae), thus obtaining more advanced developmental stages from which useful taxonomic characters could be studied. european journal of taxonomy 125: 1–23 (2015) 8 saltational evolution irrespective of the metrics we adopt to evaluate the phenetic distance between two species, it is not always easy to distance oneself from the naïve preconception that the phenetic distance is a prima facie proxy for the age of divergence. but this is not necessarily true: both smaller changes and larger transitions happen in fact along the history of life (orr 1998). in other words, evolution offers examples of discontinuities (cf. frazzetta 2012) that cannot be explained as the end result of a long chain of individually small (possibly adaptive) changes. larger morphological discontinuities have often provided diagnostic traits of higher taxa, such as the gastropods’ torsion of the visceral sac or the dramatic directional asymmetry of the flatfishes. before the advent of evo-devo, the very idea of saltational evolution as an explanation for macroevolutionary transitions (as suggested, for example, by goldschmidt (1940)), was strictly banned as heretical. however, an appreciation of the nonlinear character of the genotype→phenotype map is enough to realize how major phenotypic changes can be accomplished in a leap. for example, a singlegene mutation was probably responsible for the evolution of the bilaterally symmetrical orchid flower from an ancestor with a radially symmetrical one (theißen 2009). similarly, a small genetic change may explain a sudden duplication of the number of leg pairs observed in scolopendromorph centipedes. most of the ca. 700 species of scolopendromorpha described thus far have a fixed number of 21 pairs of legs, although a sizeable minority has 23 (the number of leg-pairs in adult centipedes is always odd). the only scolopendromorph genus from which both numbers (21 and 23) have been recorded is scolopendropsis brandt, 1841 and, more precisely, individuals with either 21 or 23 pairs of legs seem to coexist in s. bahiensis (brandt, 1841). this was the whole story before the discovery of s. duplicata chagas, edgecombe & minelli, 2008, where leg-bearing segments are either 39 or 43 (chagas et al. 2008). this is, indeed, the only major difference between the two species, the split between which is likely to be quite recent. in this case, we can hypothesize that speciation has been accompanied by a duplication of trunk segment number, a phenotypically major leap, but one that was very likely the effect of a minor change in terms of developmental control (minelli et al. 2009). in the flowering plants, dramatic changes in overall morphology, such as a transition from centimeter size and herbaceous habit to meter height and woody tree structure, may happen within a genus, and in such a short time as not to involve the establishment of a complete reproductive barrier. for example, within genera represented on continents by small herbaceous species, huge size and woody stem have recently evolved in species or groups of species colonizing oceanic islands. among the many examples given by carlquist (1974) in his extensive treatment of island floras, there are several representatives of the asteraceae (large woody species of bidens l. in southern polynesia, of senecio l. in new zealand and of centaurea l. in the canary islands), but also conspicuous examples in echium l. (boraginaceae) and euphorbia l. (euphorbiaceae) on madeira, the canary islands and cape verde islands. worthy of special attention are the hawaiian asteraceae usually classified in the genera dubautia gaudich., wilkesia a. gray and argyroxiphion dc. their, probably recent, common ancestor has been inferred to have been similar to some living shrubby dubautia species, but this group has diversified into several distinct life forms, including the thick cushions of densely packed silvery linear leaves known as the silverswords (argyroxiphium spp.). habitus differences among these plants are enormous; nevertheless, they are not completely isolated reproductively, as witnessed by a hybrid between a dubautia species and an argyroxiphium species found in nature (carlquist 1980). woodiness has also evolved in other places, as in the alpine belt of the highest mountains of africa, where some huge senecio and lobelia l. are found, among others. minelli a., biological systematics in the evo-devo era 9 heterochrony and phylogenetics heterochrony is defined (klingenberg 1998) as evolutionary change in rates and timing of developmental processes. patterns of heterochrony may contain phylogenetic signals. for example, guralnick & lindberg (2001) used the timing of cell lineage events to generate a phylogenetic tree of several lophotrochozoan taxa that turned out to agree in many respects with trees based on traditional morphological characters. the problem is how to extract from heterochronic patterns phylogenetically useful data. this problem has repeatedly been addressed since the last decade of the past century, and solutions were proposed following a revisitation of the concept of heterochrony that suggested a perspective other than the traditional one long established by the works of de beer (1930, 1940), gould (1977), alberch et al. (1979), mcnamara (1986, 1995), mckinney (1988) and mckinney & mcnamara (1991). this traditional approach focused on growth heterochrony, i.e., on developmental changes in size and shape relationships. two major classes of heterochronic patterns were thus distinguished: paedomorphosis (an organism reaches sexual maturity while retaining juvenile traits, compared to its close ancestors and/or relatives) and peramorphosis (maturation is delayed and growth period is extended). this approach does not cover the whole range of heterochronic patterns and does not even include those potentially more useful for the taxonomist. this is why smith (1997) and velhagen (1997) suggested a different approach, called sequence heterochrony, in which heterochrony is identified in the changes in the sequence order of developmental events within the ontogenetic sequence. techniques to analyze sequence heterochronies are discussed, e.g., in velhagen (1997), richardson et al. (2001), smith (2001, 2002, 2003), bininda-emonds et al. (2002, 2007), schulmeister & wheeler (2004), jeffery et al. (2005), goswami (2007), blomquist (2008), harrison & larsson (2008), werneburg & sanchez-villagra (2009) and wilson et al. (2010). the taxonomist must be aware of the fact that heterochrony is sometimes noticeable even at the intraspecific level. examples have been provided by de jong et al. (2009) for a lake victoria cichlid and by tills et al. (2011) for the pond snail, radix balthica (linnaeus, 1758). paedomorphosis the misleading effects of paedomorphosis on phylogenetic analysis have been accurately discussed in an important study on salamander phylogeny. wiens et al. (2005) demonstrated that most paedomorphic families had been clustered in a single clade by a previous phylogenetic analysis (gao & shubin 2001) based on morphological data, largely because of the absence, in the paedomorphic lineages, of those synapomorphies that in non-paedomorphic taxa develop at metamorphosis. an additional problem was the parallel retention, in the paedomorphic lineages, of traits associated with the aquatic habitat typical of salamander larvae. heterochrony and flowering plant taxonomy most of the recent progress in evo-devo has involved metazoans, not only at the experimental level, but also in conceptual advances. there is, however, a wonderful opportunity to extend this progress to plants and, indeed, to all organisms beyond metazoans. in the case of flowering plants, in particular, the literature is full of data generated under traditional research schemes that can be fruitfully reinterpreted in the light of evo-devo. this is, specifically, the case of heterochrony. in order to stimulate research in this field, i will devote here a couple of pages to summarizing evidence demonstrating the phylogenetic signal present in heterochronic patterns in flowering plants. european journal of taxonomy 125: 1–23 (2015) 10 1. major phenotypic effects obtained through ‘cheap’ heterochronic changes in a classic paper on the role of single-gene mutations in the evolution of flowering plants, hilu (1983) suggested, that “nonpolygenic macromutations supplemented with micromutation” can be responsible for the large morphological gaps existing between higher taxa. the phenotypic effects of heterochrony are usually small, but can occasionally be so conspicuous as to explain why genus-level distinctions have often been introduced in the past to separate groups of species that differ in one obvious trait from their closest relatives, within whose radiation they are nevertheless deeply nested. box et al. (2008) presented a couple of interesting examples concerning the european orchids. it does not require specialist knowledge to distinguish the species traditionally ascribed to the genus nigritella rich. from those ascribed to gymnadenia r. br., but molecular evidence places the former firmly within the latter genus, suggesting a recent origin for their peculiarities in the shape of flowers and inflorescences. the same is true for the orchid known as coeloglossum viride (l.) hartm., which is nothing but a species of dactylorhiza necker ex nevski, mainly deviating in the relative length of spur vs. labellum. in either instance, the differences formerly taken to deserve acknowledgement at the generic level turn out to be simple heterochronic effects of differences in the growth rate of two parts of the flower. 2. modularity and character independence a traditional problem in retrieving and coding information for phylogenetic analysis is how to ensure the mutual independence of characters. this problem has frequently emerged also in the recent literature on sequence heterochrony (cf. the literature cited above). this difficulty, however, is reduced to a minimum wherever a body part behaves as a developmentally independent module. in plants, this has been shown, for example, in the case of the nectariferous petals of delphinium l. (ranunculaceae) (guerrant 1982). in bauhinia l. (fabaceae) the individual petals and stamens behave as separate homologues; thus, within the genus there are species like b. blakeana dunn, with 5 petals and 3 fertile stamens, alongside species like b. divaricata l., with 2 petals and 1 functional stamen only (wunderlin 1983; chen et al. 2010). 3. peramorphosis and paedomorphosis in campanulaceae in the current circumscription, campanulaceae include a clade (lobelioideae) often segregated in the past as an independent family, lobeliaceae. this clade is extremely diverse, especially in terms of growth form, including very small, centimeter-high herbaceous plants as well as giant woody shrubs, up to 9 m in height. spectacular examples of the latter life form are the species classified in the genus cyanea gaudich., an hawai’ian endemic (givnish et al. 2009). the tallest cyanea species grow through juvenile stages that apparently retain more of the traits of the herbaceous ancestors, thus suggesting a peramorphic trend. however, there are also cyanea species of reduced size and juvenile habitus, which suggest an opposite, paedomorphic trend (lammers 1990). this hypothesis deserves revisitation based on molecular evidence. 4. heterostyly and homoplasy heterostyly is a peculiar kind of polymorphism exhibited by a number of plant species (examples are known for 199 genera in 28 plant families; naiki 2012), where individual plants belong to one or the other of two alternative phenotypes (distyly); quite rare are species with three different morphs (tristyly). in the case of distyly, some plants produce flowers with the stylus much longer than the stamen filaments (pin phenotype), while the others produce flowers with stamen filaments much longer than the stylus (thrum phenotype). pollination is effective only if the pollinated plant belongs to the other phenotype in respect to the pollen-producing individual. plant species monomorphic for the relative length of stamens and stylus are said to be homostylous. in amsinckia lehm. (boraginaceae), a genus which includes both homostylous and heterostylous species, heterostyly is produced by heterochrony (li & johnston 2010). the growth curve for the pistil is identical in the homostylous a. vernicosa hook. & arn. and in the thrum flowers of a related heterostylous species (a. furcata suksdorf); the growth curve of the free minelli a., biological systematics in the evo-devo era 11 stamen filament is identical in the homostylous and the pin flowers of the heterostylous species. within the genus, heterostyly has been reconstructed as the plesiomorphic condition, which is however derived in a wider, family-wide context. but a few amsinckia are homostylous, as a consequence of character reversal. the homostylous a. vernicosa has evolved from a heterostylous ancestor comparable to a. furcata, whereas the homostylous a. gloriosa eastw. ex suksdorf has evolved from a heterostylous ancestor comparable to a. douglasiana a. dc; thus, the relationship between a. vernicosa and a. gloriosa is an interesting case of heterochrony-based homoplasy. 5. phylogenetic signal of heterochronic patterns in the leguminosae evo-devo is probably the ultimate resource we can use to dispose of the last surviving remnants of the century-old idea that different kinds of characters are the most adequate to diagnose taxa of different levels. not so long ago as 1984, tucker adopted a naïve version of recapitulation to suggest an operational dichotomy between morphological and morphogenetic characters in the leguminosae. in his scheme, traits fixed early in ontogeny, such as the order of initiation and the position of organs, should be given value at higher (subfamily, tribe) levels, and traits fixed later in ontogeny, such as size, shape, and possible fusion of petals and other organs, should instead be given value at lower (genus and species) levels. this ignores heterochrony, of which there are many examples in this clade of plants. in the end, however, heterochrony turns out to be phylogenetically informative. still within the legumes, for example, grimes (1999) determined that the heterochronic separation of vegetative and reproductive growth, causing cauliflory, is a synapomorphy of the clade zygia, whereas the so-called inga alliance (cojoba britton & rose, cathormion (benth.) hassk., inga miller, macrosamanea britton & rose, zygia p. browne) is characterized by another heterochrony, i.e., the persistent activity of its meristems, although with character reversal in the subclade inga+cathormion. in another study, where the unusual flower of duparquetia orchidacea baill. was compared to those of other legumes (cercis canadensis l., petalostylis labicheoides r. br., labichea lanceolata benth., dialium guineense willd. and tamarindus indica l.), prenner & klitgaard (2008) demonstrated the effects on flower morphology of heterochronies such as anticipation or retardation of a whole whorl with respect to another, or of a single organ (e.g., a sepal or a petal) with respect to the other elements of its whorl. finally, citerne et al. (2006) have shown that late development can reverse a trend apparently determined at an early developmental stage. for example, in another legume (cadia purpurea (g. piccioli) aiton), a flower that initially seems to develop into a bilaterally symmetrical (zygomorphic) flower, such as one of lupinus l., ‘corrects’ its reduction in symmetry and eventually matures as a radially symmetrical (actinomorphic) flower. evo-devo revisitation of character reversal another critical point in phylogenetic analysis is the plausibility of character reversal. dollo’s (1893) macroevolutionary law states that structures that are lost in evolution will not be regained. if universally true, dollo’s law would provide an important constraint on our reconstructions of phylogeny. however, a formal test of this hypothesis is possibly unfeasible; at least, ancestral character state reconstruction does not seem to offer a reliable criterion to evaluate it (cunningham 1999). as a consequence, we should turn to empirical evidence. indeed, there are a number of recent phylogenetic studies suggesting exceptions to dollo’s law. wiens (2011) has offered strong evidence of the re-evolution of mandibular teeth in the frog genus gastrotheca fitzinger, 1843. based on a time-calibrated phylogeny for 170 amphibian species, mandibular teeth were apparently lost in the last common ancestor of modern frogs some 230 million years ago (mya) but have been regained in the gastrotheca lineage in the last ca. 5–17 million years. european journal of taxonomy 125: 1–23 (2015) 12 evo-devo and speciation speciation processes are seldom studied from the point of view of changes in developmental control and timing (e.g., naisbit et al. 2003), but this perspective deserves closer attention. developmental evolution and sympatric speciation in several plant and animal species, two or more well-defined ‘morphs’ occur in the same area (the same basin, in the case of aquatic organisms), with alternative phenotypes generally confined to different habitats (e.g., landry et al. 2007). the question is, how two or more morphs of the same species (up to four, e.g., the deep-, shallow-, riverand bay spawning morphs of the whitefish coregonus lavaretus (linnaeus, 1758) in lake femund, norway; østbye et al. 2005) can co-exist indefinitely in conditions of strict or approximate sympatry. far from being an exceptional case, this multiplicity of syntopic morphs is usual in the salmonids of the genus coregonus linnaeus, 1758 in lakes of eastern north america (pigeon et al. 1997) and the central alps (douglas et al. 2005). the morphs found in one basin have demonstrably differentiated independently of a parallel divergence in other basins. changes in the regulation of ontogeny are largely responsible for the differences between sympatric morphs. differences in the time required by the embryo to develop till hatching (rogers & bernatchez 2006) are often involved, together with differential gene expression throughout the entire embryonic and early post-embryonic development. after hatching, differences in gene expression between sympatric morphs are even more conspicuous (nolte et al. 2009). in another case, during embryonic development the transcriptome of two sympatric morphs was nearly identical, but defects in the hybrids between them suggested extensive disruption of regulatory control of gene expression, apparently affecting over 2000 genes, including crucial developmental genes (renaut & bernatchez 2011). developmental timing and speciation the importance of evolutionary developmental biology in addressing taxonomic questions is greatly increased if we open evo-devo’s scope from the exclusive consideration of development and evolution of morphology to the additional consideration of the developmental control and evolution of temporal phenotypes (minelli & fusco 2012). temporal differences in the life cycle are indeed often important in ensuring prezygotic isolation between newly divergent populations. alexander & bigelow (1960) used the term ‘allochronic speciation’ for those cases of sympatric speciation where temporal differences in the life cycles foster the isolation between two divergent populations. the plausibility of this scenario is strongly supported by well-documented cases of either individual or geographic variation in life history traits that would easily lead to temporal isolation of initially conspecific populations. for example, populations with egg diapause and populations without egg diapause co-exist in the north-american crickets eunemobius carolinus (de geer, 1773), miogryllus verticalis (serville, 1838) and teleogryllus commodus (f. walker, 1869). in other crickets (allonemobius fasciatus (de geer, 1773), oecanthus niveus (de geer, 1773) and oe. quadripunctatus beutenmüller, 1894) the number of generations per year varies between 1 and 3 according to the population (alexander 1968). individual variation in life history schedule is known for the north american cicadas of the genus magicicada davis, 1925, whose short adult season is usually reached at either 13 (e.g., m. tredecim (walsh & riley, 1868), m. tredecassini alexander & moore, 1962, m. tredecula alexander & moore, 1962) or 17 years of age (e.g., m. septendecim (linnaeus, 1758), m. cassinii (fisher, 1852), m. septendecula alexander & moore, 1962). interestingly, there are occasional records of 13-year cicadas emerging later than expected, but exactly 4 years later, that is, as 17-year cicadas (marshall et al. 2011). additionally, phylogeny suggests a 13-year cycle as the primitive condition in this clade, with three independent transitions to a 17-year cycle. an example of closely related, parapatrically distributed minelli a., biological systematics in the evo-devo era 13 species with different developmental length are the 17-year m. septendecim and the 13-year m. tredecim (marshall & cooley 2000). a detailed discussion of developmental plasticity and speciation in these cicadas was provided by west-eberhard (2003). phenologically distinct populations with parapatric distribution are also known for the pine bark bug aradus cinnamomeus panzer, 1806 (heliövaara et al. 1988). maturity is usually reached by this hemipteran at the age of 2 years and individual populations are known to mature synchronously, some in the odd, some in the even years. odd-year bugs are occasionally found in the even-year area, but are very rare, as are occasional even-year bugs in odd-year populations. parapatric populations reaching maturity in alternate years are thus virtually isolated, reproductively, but the whole picture is complicated by the existence of peripheral populations with a 3-year life cycle. in some flowering plants, seasonal dimorphism or polymorphism has been described, for example in several genera of the orobanchaceae such as odontites ludwig, euphrasia l., melampyrum l. and rhinanthus l. (e.g., de soó & webb 1972), but also in gentianella moench (gentianaceae) (pritchard & tutin 1972; lennartsson 1997) and in solidago l. (asteraceae) (e.g., pors & werner 1989). in rhinanthus, a discontinuity between earlyand late-flowering plants has long been interpreted as largely due to haymaking, at least in high altitude localities where this is done only once or twice per year; this may have helped in reducing the potential gene flow between earlyand late-flowering phenotypes, between which morphological differences would have rapidly accumulated (see, however, briggs & walters (1997) for a critical assessment of this case). flora europaea authors responsible for these genera (e.g., de soó & webb 1972; pritchard & tutin 1972) have provided keys and descriptions for all the best characterized seasonal phenotypes in these genera without attempting to assess their taxonomic status. in a more recent study on gentianella species, however, lennartsson (1997) suggests that one and the same (macro)species can be represented in some areas by a single, long flowering and morphologically variable form, in other areas by two sympatric forms, phenologically distinct and morphologically distinguishable. the latter condition may eventually represent an incipient stage of speciation. what matters in the context of our analysis is the fact that these alternative phenotypes are produced by a different developmental regulation of flowering time. acknowledgements i am very grateful to henrik enghoff and koen martens for their invitation to write this review paper and to an anonymous referee for the useful suggestions on a previous version. references abzhanov a., kuo w.p., hartmann c., grant b.r., grant p.r. & tabin c.j. 2006. the calmodulin pathway and evolution of elongated beak morphology in darwin’s finches. nature 442: 563–567. http://dx.doi.org/10.1038/nature04843 abzhanov a., protas m., grant b.r., grant p.r. & tabin c.j. 2004. bmp4 and morphological variation of beaks in darwin’s finches. science 305: 1462–1465. http://dx.doi.org/10.1126/science.1098095 alberch p. 1991. from genes to phenotype: dynamical systems and evolvability. genetica 84: 5–11. http://dx.doi.org/10.1007/bf00123979 alberch p., gould s.j., oster g. & wake d. 1979. size and shape in ontogeny and phylogeny. paleobiology 5: 296–317. alexander r.d. 1968. life cycle origins, speciation, and related phenomena in crickets. quarterly review of biology 43: 1–41. http://dx.doi.org/10.1038/nature04843 http://dx.doi.org/10.1126/science.1098095 http://dx.doi.org/10.1007/bf00123979 european journal of taxonomy 125: 1–23 (2015) 14 alexander r.d. & bigelow r.s. 1960. allochronic speciation in field crickets, and a new species, acheta veletis. evolution 14: 334–346. http://dx.doi.org/10.2307/2405976 allsopp p.g. & lambkin c.l. 2006. canegrubs and cladistics: what story do adult, larval and ecological characters tell? australian journal of entomology 45: 55–66. http://dx.doi.org/10.1111/j.14406055.2006.00511.x altenberg l. 1995. genome growth and the evolution of the genotype-phenotype map. in: banzhaf w. & eeckman f.h. 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botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark. http://dx.doi.org/10.1038/nrg2949 http://dx.doi.org/10.1186/1471-2148-9-82 http://dx.doi.org/10.1111/j.1558-5646.2011.01221.x http://dx.doi.org/10.1080/10635150590906037 http://dx.doi.org/10.1080/10635150590906037 http://dx.doi.org/10.1007/s13127-010-0020-4 http://dx.doi.org/10.1093/molbev/msg140 http://dx.doi.org/10.2307/2409308 http://dx.doi.org/10.2307/2399009 http://dx.doi.org/10.2307/2399009 http://dx.doi.org/10.1111/j.1096-3642.2005.00139.x european journal of taxonomy 291: 1–13 issn 2118-9773 https://doi.org/10.5852/ejt.2017.291 www.europeanjournaloftaxonomy.eu 2017 · legendre f. et al. this work is licensed under a creative commons attribution 3.0 license. d n a l i b r a r y o f l i f e , r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:59adaf1d-4704-49a2-9467-ade8813f75fa 1 molecular phylogeny of blaberidae (dictyoptera, blattodea), with implications for taxonomy and evolutionary scenarios frédéric legendre 1,*, philippe grandcolas 2 & france thouzé 3 institut de systématique, évolution, biodiversité isyeb – umr 7205 – mnhn cnrs upmc ephe, muséum national d’histoire naturelle, sorbonne universités, 57 rue cuvier, cp 50, 75005 paris, france. * corresponding author: legendre@mnhn.fr 2 email: pg@mnhn.fr 3 email: francethouze@gmail.com 1 urn:lsid:zoobank.org:author:8b1163c9-338e-425a-9107-c185d4ab415f 2 urn:lsid:zoobank.org:author:fccf6c9c-f929-453e-989b-c5bb4b04b54a 3 urn:lsid:zoobank.org:author:6692eb45-5e95-41ab-bb1f-59b3154b3402 abstract. in the present “tree-thinking” period, relying on accurate phylogenetic hypotheses is of paramount importance for biologists interested in an evolutionary perspective. in the blaberidae cockroaches, a well-defined monophyletic family comprising several model species, no such phylogenetic tree is available despite several earlier contributions. here, using six molecular markers (12s, 16s, 18s, 28s, coi and coii), we investigate the relationships of blaberidae and compare our results with the traditional morphology-based classification. this resulted in a broad spectrum of situations, from congruent and well-supported hypotheses (e.g., the monophyly of blaberidae, oxyhaloinae and (geoscapheiinae + panesthiinae)) to incongruent and weakly supported results (e.g., polyphyly of perisphaerinae). we emphasize that interesting and contrasted situations lie between the two extremities of this spectrum, especially concerning the genera thanatophyllum grandcolas, 1991, phoetalia stål, 1874, laxta walker, 1868 and pronauphoeta shelford, 1909. we also discuss the phylogenetic position of two incertae sedis genera (eustegasta gerstaecker, 1883 and gynopeltis gerstaecker, 1869). we conclude that in-depth signal analyses should be performed to better understand molecular evolution and its consequence on tree reconstruction for this group. as for phylogenetic relationships per se, new markers should be searched for, especially to decipher deeper relationships in blaberidae. keywords. biogeography, incongruence, genitalia, morphology, phylogenetic signal. legendre f., grandcolas p. & thouzé f. 2017. molecular phylogeny of blaberidae (dictyoptera, blattodea), with implications for taxonomy and evolutionary scenarios. european journal of taxonomy 291: 1–13. https://doi. org/10.5852/ejt.2017.291 introduction we are presently experiencing what has been called a “tree-thinking biology” era (grandcolas et al. 1994; o’hara 1997; baum et al. 2005; gregory 2008; omland et al. 2008; baum & smith 2013; losos https://doi.org/10.5852/ejt.2017.291 www.europeanjournaloftaxonomy.eu https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:59adaf1d-4704-49a2-9467-ade8813f75fa mailto:legendre%40mnhn.fr?subject= mailto:pg%40mnhn.fr?subject= mailto:francethouze%40gmail.com?subject= http://zoobank.org/urn:lsid:zoobank.org:author:8b1163c9-338e-425a-9107-c185d4ab415f http://zoobank.org/urn:lsid:zoobank.org:author:fccf6c9c-f929-453e-989b-c5bb4b04b54a http://zoobank.org/urn:lsid:zoobank.org:author:6692eb45-5e95-41ab-bb1f-59b3154b3402 https://doi.org/10.5852/ejt.2017.291 https://doi.org/10.5852/ejt.2017.291 european journal of taxonomy 291: 1–13 (2017) 2 et al. 2013). myriad phylogenies are produced every year (parr et al. 2012) and these hypotheses serve to investigate diverse evolutionary questions or to revise classifications. even though these numerous hypotheses might be contradictory and thus bewildering (chiapella et al. 2014), they are essential for present and future studies, especially when they involve model organisms from various disciplines. blaberid cockroaches comprise model species in research about biomimetics (e.g., nelson et al. 1997), physiology (li 2007), sociality (costa 2006; pellens et al. 2007) or reproductive behavior (roth &willis 1954; banerjee et al. 2016). these fields of research would benefit from a clear phylogenetic hypothesis or a phylogeny-derived robust classification. yet, such a phylogeny is still lacking despite several relevant contributions since saussure (1864) erected the blaberidae family. mckittrick (1964) was the first to define this family as we consider it today. she distinguished three complexes which she named blaberoid, epilamproid and panchloroid, but without performing any formal phylogenetic analysis. a cladistic analysis of her data, however, revealed that her blaberoid and panchloroid complexes were not monophyletic (legendre 2007). later, cockroach systematists identified several morphological synapomorphies supporting the monophyly of blaberidae (grandcolas 1996; klass & meier 2006; brood sac underneath the female genitalia, sclerite r3v small, first vein of vannus with many basal rami, tracheation very dense and tracheae vesicle-shaped, presence of a muscle connecting sclerite l2 and pouch lve). monophyly has also repeatedly been found with molecular data as well (maekawa et al. 2003; inward 2007; pellens et al. 2007; djernaes et al. 2012; legendre et al. 2014, 2015). within blaberidae, however, the phylogenetic relationships remain controversial, despite early morphological studies (e.g., grandcolas 1993, 1997). the only largely supported results are the close relationships between zetoborinae and blaberinae on the one hand, and panesthiinae and geoscapheinae on the other. the position of panchlorinae as the first diverging lineage within the family also seems to be consistently found lately even though often based on a limited taxon sample (legendre et al. 2014, 2015). the phylogenetic positions of the seven remaining subfamilies still need to be investigated further, as do the phylogenetic positions of 24 incertae sedis genera (according to the blattodea species file, bsf; beccaloni 2014). the taxonomy of cockroaches relies heavily on morphological characters, especially those of the genitalia. insect taxonomists often use genitalia for species identification as well as for classification, proposing synapomorphies for diverse subfamilies and genera (e.g., grandcolas 1993, 1994, 1996, 1997; roth 2003; klass & meier 2006). the current classification systems of cockroaches are thus based mainly on morphological characters (roth 2003). however, like any set of characters, morphological characters benefit from completion with other kinds of data, as already shown when combining molecular and behavioral characters for instance (legendre et al. 2014). in terms of geographical range, blaberidae have a pantropical distribution. some subfamilies are restricted to one continent, while others encompass several. blaberinae and zetoborinae are endemic to america (mainly south america), gyninae and oxyhaloinae are endemic to africa, geoscapheinae is endemic to australia, and paranauphoetinae is endemic to asia. panchlorinae is found in america and africa, pycnoscelinae is found in asia and south america, while diplopterinae and panesthiinae are found in asia and australasia. perisphaerinae is found in africa, asia and australasia, and epilamprinae is found in america, africa, asia and australasia (princis 1964; beccaloni 2014). in this paper, we investigate the phylogenetic relationships in blaberid cockroaches using six molecular markers. we then compare our results to the largely preor semi-phylogenetic current morphology-based classification of this family as presented in the csf (beccaloni 2014). we underline the differences found and interpret them in terms of potential morphological evolution and biogeography. we also propose future directions to improve our knowledge of the phylogenetics of blaberidae. legendre f. et al., molecular phylogeny of blaberidae 3 material and methods taxonomic and character sampling we sampled 136 taxa, 128 of them belonging to 11 out of the 12 currently distinguished subfamilies of blaberidae (according to the bsf; beccaloni 2014): 18 blaberinae (12 genera out of 23 genera recognized in the bsf, i.e. ~52%), one diplopterinae (one genus, 100%), 26 epilamprinae (at least 11 genera, ~24%), 10 geoscapheinae (four genera, 100%), five gyninae (one genus, 20%), 13 oxyhaloinae (11 genera, ~65%), three panchlorinae (one genus, 20%), 23 panesthiinae (five genera, ~71%), 18 perisphaerinae (at least seven genera, ~37%), two pycnoscelinae (one genus, ~33%), and nine zetoborinae (eight genera, 57%). we also included five specimens belonging to the incertae sedis genera eustegasta gerstaecker, 1883 and gynopeltis gerstaecker, 1869. outgroup taxa comprise one ectobiidae (paratropes sp. 1), one corydiidae (therea petiveriana linnaeus, 1758) and one blattidae (blatta orientalis linnaeus, 1758) as the rooting outgroup taxon. among ectobiidae, all subfamilies have been postulated as being the sister group of blaberidae, including pseudophyllodromiinae (grandcolas 1996; inward et al. 2007; pellens et al. 2007; djernaes et al. 2012; legendre et al. 2015). at first, we included several species of pseudophyllodromiinae (anisopygia sp., asemoblattana sp., euphyllodromia sp., latiblattella sp. 3) but we then discarded them because of their artifactual position within blaberidae (among epilamprinae, data not shown). all the species of pseudophyllodromiinae sequenced so far show highly modified sequences, which results in high percentages of divergence with ingroup species and may cause artifacts in tree reconstruction such as long branch attraction (wheeler 1990; bergsten 2005; legendre et al. 2015). also a couple of additional species of blattellinae were used as outgroup taxa in preliminary analyses (ischnoptera sp. and xestoblatta sp.), but were not kept because of their missing data, which might alter tree reconstruction (e.g., lemmon et al. 2013; roure et al. 2013). whatever the outgroup combinations used in our preliminary analyses, the supported results presented here were always retrieved. we used six molecular markers to reconstruct blaberid phylogenetic relationships: 12s rrna (~440 bp), 16s rrna (~440 bp), 18s rrna (~1900 bp), 28s rrna (~2350 bp), cytochrome oxidase subunit i (coi, 1179 bp) and cytochrome oxidase subunit ii (coii, 671 bp). primers and molecular techniques were described in legendre et al. (2008). we provided 175 new sequences, which were deposited in genbank with the following accession numbers: ky497575–ky497749. in average, taxa were documented for ca 4000 bp (for a total alignment of 6991 bp). we sampled 92% of the taxa for 12s, 57% for 16s, 95% for 18s, 76% for 28s, 38% for coi and 78% for coii. the supplementary file shows the details of the completion of our dataset. phylogenetic analyses all sequences were blasted prior to any analysis to check for potential contamination. then, we computed alignments using muscle (edgar 2004) as implemented in seaview v. 4 (gouy et al. 2010). for 28s, alignments were performed separately for each fragment and then concatenated, whereas fragments were concatenated before alignment for 18s. the 28s fragments do not or hardly overlap so that they could not be aligned altogether. for each marker, the proposed alignments were sometimes dubious, especially in the terminal regions because of fragments that were not always congruent in coverage. we corrected these problems by refining the alignment manually. morever, we ensured that alignments respect the codon protein reading frame for coi and coii. once we obtained separate alignments for the six markers, we used sequencematrix v. 1.7.8 (vaidya et al. 2011) to produce the full dataset composed of 6991 bp. we performed separate and combined analyses in a probabilistic framework. for separate analyses, we selected the best model that does not mix i (proportion of invariant sites) and g (heterogeneity of substitution rates) using jmrmodeltest v. 2.1.3 (darriba et al. 2012) with the aicc criterion. http://www.europeanjournaloftaxonomy.eu/index.php/ejt/rt/suppfiles/415/0 european journal of taxonomy 291: 1–13 (2017) 4 for combined analyses, we used partitionfinder v. 1.1.1 (lanfear et al. 2012; options: branchlengths= linked, model_selection=aicc and search=greedy) to select the best partition strategy among models supported by mrbayes. the best partitioning strategy involved nine partitions: one partition with a gtr + g model for each non-protein-coding marker; four distinct partitions for the first two positions of coi and coii with a gtr + g model; and a common partition for the third positions of coi and coii, also with a gtr + g model. in maximum likelihood (ml), we used raxml v. 8.2.4 (stamatakis 2014) to reconstruct phylogenetic hypotheses, whereas we used mrbayes v. 3.1.2 (ronquist & huelsenbeck 2003) in bayesian inferences (bi). ml analyses were performed using the rapid hill-climbing algorithm for 500 replicates. we then computed 500 bootstrap replicates. in bi, we launched two runs of four chains each until the average deviation of split frequencies reached a value inferior to 0.01 (options: stoprule=yes, stopval=0.01). we then assessed convergence using tracer v. 1.6 (rambaut et al. 2014), checking that ess values all exceed 200. results the best tree from the ml combined analysis is shown in fig. 1 and is very similar to the tree reconstructed in bayesian inference (note that convergence diagnostics and ess values were satisfying, except for the ess values of the rate multiplier parameters). blaberidae were monophyletic with a moderate bootstrap support and a maximal posterior probability (bs = 94, pp = 1). panchlorinae, gyninae, pycnoscelinae and oxyhaloinae were monophyletic (bs / pp = 100 / 1, 99 / 1, 90 / 0.9 and 50 / 0.93, respectively), while blaberinae, epilamprinae, perisphaerinae and zetoborinae were polyphyletic. the group (panesthiinae + geoscapheinae) was monophyletic (bs = 99, pp = 1), with australian panesthia nested within geoscapheinae. overall, this tree showed very weak support values for deeper relationships within blaberidae. nevertheless, some clades with geographic congruence were retrieved. african perisphaerinae, asian perisphaerinae, african epilamprinae and asian-oceanian epilamprinae (except rhabdoblatta cf. circumdata from singapore) were monophyletic with high to moderate support values (bs / pp = 100 / 1, 100 / 0.96, 100 / 1 and 81 / 0.9, respectively), and a neotropical clade composed of the blaberinae and most of the zetoborinae was found (bs / pp = 43 / 0.86; see exceptions below). the phylogenetic position of phoetalia pallida (brunner von wattenwyl, 1865) (blaberinae) as sister species to schultesia lampyridiformis roth, 1973 (zetoborinae) was highly supported (bs / pp = 100 / 1) and found in almost all the analyses performed (fig. 2). three other blaberinae species (lucihormetica subcincta (walker, 1868), hormetica sp. and blaptica sp.) were found closer to most species of zetoborinae than to other blaberinae. parasphaeria boleiriana grandcolas & pellens, 2002 (zetoborinae) branched outside of the neotropical clade composed of the blaberinae and most of the zetoborinae, but its phylogenetic position was unsupported (bs / pp = 11 / 0.81), whereas the position of thanatophyllum akinetum grandcolas, 1991 (zetoborinae) as sister group to some neotropical epilamprinae was highly supported (bs / pp = 99 / 1) and found in half of the separate analyses (fig. 2). within oxyhaloinae, the nauphoetini were monophyletic, whereas the gromphadorhini were monophyletic only if the incertae sedis genera brachynauphoeta van herrewege, 1975 and heminauphoeta saussure, 1891 are taken into account. the incertae sedis oxyhaloinae genus pronauphoeta was sister group to all other species of oxyhaloinae sampled here (bs / pp = 50 / 0.93), but with other positions in three one marker analyses (12s, 16s and 18s; fig. 2). concerning perisphaerinae, laxta sp., the only australian individual sampled, was sister group to african perisphaerinae (bs / pp = 32 / 0.93), a result almost only found with the combined dataset (fig. 2). legendre f. et al., molecular phylogeny of blaberidae 5 f ig . 1 . [ pa rt 2 o n ne xt p ag e] o pt im al p hy lo ge ne tic tr ee re co ns tr uc te d in m ax im um l ik el ih oo d w ith th e co m bi ne d da ta se t. b oo ts tr ap v al ue s a nd p os te ri or pr ob ab ili tie s ar e re po rt ed f or e ac h no de ( bo ot st ra p va lu es b el ow 2 5% a re n ot d is pl ay ed ). t he c ol or o f in te rn al b ra nc he s is p ro po rt io na l t o bo ot st ra p va lu es . g eo gr ap hi c or ig in o f th e sp ec im en s se qu en ce d is p ro vi de d in b ra ck et s af te r th e sp ec ie s na m es . i n pu rp le , m on op hy le tic g ro up c on gr ue nt w ith m or ph ol og ic al h yp ot he se s; in b lu e, m on op hy le tic g ro up w ith g eo gr ap hi c co ns is te nc y at th e co nt in en ta l l ev el ; i n br ow n, in ce rt ae se di s s pe ci es ; i n gr ee n, fo ur s pe ci es w ith c on fli ct in g an d su pp or te d po si tio ns ( th an at op hy llu m a ki ne tu m g ra nd co la s, 1 99 1 an d p ho et al ia p al lid a (b ru nn er v on w at te nw yl , 18 65 ) o r c on gr ue nt , b ut w ea kl y su pp or te d po si tio ns (l ax ta s p. a nd p ro na up ho et a cf . v ir id ul a (p al is ot d e b ea uv oi s, 1 80 5) ). t he fo ur la tte r s pe ci es a re di sc us se d in th e te xt (s ee a ls o fi g. 2 ). t he s ub fa m ili es in di ca te d on th e ri gh t o f t he tr ee a re d er iv ed fr om tr ad iti on al m or ph ol og yba se d cl as si fic at io ns . u ni ts fo r t he b ra nc h le ng th s ca le a t t he b ot to m ri gh t: nu m be r o f e xp ec te d su bs tit ut io ns p er s ite . p an ch lo ri na e (s o u th a m e r ic a ) o xy ha lo in ae (a fr ic a ) p er is ph ae ri na e p. p. 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(s o u th a m e r ic a ) ze to bo ri na e (s o u th a m e r ic a ) d ip lo pt er in ae 10 0/ 1 10 0/ 1 10 0/ 1 94 /1 50 /0 .9 3 10 0/ 156 /0 .8 6 9 4/ 1 30 /0 .5 9 /49 /0 .7 7 86 /1 98 /1 80 /0 .5 1 90 /0 .9 99 /1 10 0/ 1 -/ -/ 0. 8 -/ -/ 0. 67 -/ 0. 71 99 /1 88 /1 10 0/ 1 89 /0 .7 9 10 0/ 0. 96 35 /45 /55 /0 .9 6 47 /0 .9 4 10 0/ 0. 96 29 /98 /1 -/ 0. 48 -/ 0. 48 81 /0 .9 62 /0 .7 5 53 /0 .8 6 29 /0 .7 634 /0 .8 4 77 /0 .9 98 /1 52 /0 .7 71 00 /1 76 /0 .9 96 /1 50 /0 .9 7 10 0/ 1 1 00 /1 74 /1 72 /0 .7 6 -/ b oo ts tr ap v al ue s <2 5 10 0 0. 3 le oz eh nt ne ra m ax im a h em in au ph oe ta s p. 1 b ra ch yn au ph oe ta fo ul po in te en si s e lli pt or hi na c ho pa rd i a le ur op od a in si gn is g ro m ph ad or hi na p or te nt os a p rin ci si a va nw ae rb ek i g ro m ph ad or hi na o bl on go no ta a pt er a m un da a pt er a fu sc a p er is ph ae ru s sp .1 c or yd id ar um s p. ( in de ) p se ud og lo m er is s p. o pi st ho pl at ia o rie nt al is c al ol am pr a irr or at a r ha bd ob la tta fo rm os an a c al ol am pr a sp . p er is ph ae ru s sp .2 ( in do ne si a, f lo re s) p er is ph ae ru s sp .3 ( p ap ua n ew g ui ne a) p er is ph ae ru s sp .4 ( p hi lip pi ne s) p er is ph ae ru s sp .4 ( p hi lip pi ne s) cf . r ha bd ob la tta s p. 1 (s in ga po re ) cf . r ha bd ob la tta s p. 2 (i nd on es ia , j av a) e pi la m pr in ae ( la os ) cf . s tic to la m pr a sp . ( p ap ua n ew g ui ne a) cf . r hi cn od a sp . ( p ap ua n ew g ui ne a) cf . r hi cn od a sp . ( p ap ua n ew g ui ne a) cf . c yr to no tu la s p. ( in do ne si a, f lo re s) cf . c yr to no tu la s p. ( in do ne si a, l om bo k) m or ph na m ac ul at a (s in ga po re ) p se ud op ho ra sp is s p. ( in do ne si a, j av a) p se ud op ho ra sp is s p. n r. c or yd id ar um p yg m ae a p er is ph ae ru s fle xi co lli s (s in ga po re ) r hy pa ro bi a m ad er ae th an at op hy llu m a ki ne tu m t he re a pe tiv er ia na b la tta o rie nt al is p ar at ro pe s sp .1 c ol ap te ro bl at ta s ur in am a g al ib la tta c rib ro sa ( f re nc h g ui an a) p ro na up ho et a cf . v ir id ul a (g ha na ) n au ph oe ta c in er ea h en sc ho ut ed en ia fl ex iv itt a h en sc ho ut ed en ia s p. p yc no sc el us s ur in am en si s p yc no sc el us s p. ( in do ne si a, s um ba ) e us te ga st a af f. ca ra bi di na (t og o) p an ch lo ra n iv ea p an ch lo ra s p. 1 p an ch lo ra a zt ec a d ip lo pt er a pu nc ta ta cf . r ha bd ob la tta s p. 3 (p ap ua n ew g ui ne a) cf . r ha bd ob la tta s p. 4 (p ap ua n ew g ui ne a) cf . r ha bd ob la tta s p. 5 (p ap ua n ew g ui ne a) european journal of taxonomy 291: 1–13 (2017) 6 perisphaerinae p.p. (africa + australia) blaberinae + zetoborinae p.p. (south america) gyninae + gynopeltis (africa) panesthiinae + geoscapheiinae (asia and oceania) epilamprinae p.p. (south america) epilamprinae p.p. (asia) zetoborinae (south america) 32/0.93 100/1 100/1 100/1 55/0.92 82/1 -/100/1 -/-/0.86 -/0.81 97/1 97/1 43/0.86 89/0.9 99/1 97/0.92 44/0.39 55/0.92 71/0.98 49/0.75 60/94/1 23/53/0.81 100/1 100/1 100/1 100/1 -/100/1 53/0.91 45/0.87 63/0.83 100/1 100/1 100/1 100/1 -/0.4 25/0.48 89/0.69 99/1 99/1 74/0.61 62/0.99 68/0.98 58/0.78 100/1 100/1 98/1 83/1 64/0.99 61/0.98 38/0.8 68/99/1 67/1 47/0.88 99/1 99/0.9 99/0.9 95/0.79 92/1 100/1 90/1 84/0.94 92/1 56/0.68 88/1 59/96/1 27/100/1 100/1 83/1 -/30/77/1 99/1 67/1 bootstrap values <25 100 0.3 laxta sp. cyrtotria sp. epilampra sp. blaptica sp. blaptica dubia monastria biguttata eublaberus posticus eublaberus distanti byrsotria fumigata archimandrita tessellata blaberus discoidalis blaberus craniifer blaberus giganteus gyna capucina gyna lurida salganea raggei salganea gressitti salganea esakii salganea taiwanensis ryukyuanus miopanesthia deplanata panesthia transversa caeparia crenulata hormetica sp.1 zetobora sp. tribonium sp. petasodes sp. blaberus sp.1 rhabdoblatta cf. circumdata (singapore) minablatta sp.2 minablatta sp.1 perisphaerinae sp.1 perisphaerinae (south africa) perisphaeria sp. (south africa) perisphaerinae (south africa) perisphaerinae (swaziland) cf. bantua sp. (malawi) schizopilia cf. fissicolis (french guiana) nr bantua sp. parasphaeria boleiriana lanxoblatta emarginata phortioeca phoraspoides phortioeca nimbata paradicta rotunda epilampra egregia (french guiana) epilampra taira (french guiana) lucihormetica subcincta schultesia lampyridiformis phoetalia pallida gynopeltis sp.1 (mozambique) gynopeltis sp.2 (malawi) gynopeltis sp.4 (zambia) gynopeltis sp.3 (mozambique) gyna cf. costalis (benin) gyna cf. maculipennis (mozambique) gyna sp. (mozambique) panesthia lata panesthia ancaudellioides panesthia cribrata panesthia angustipennis spadica panesthia saussurii panesthia sp.2 (indonesia, flores) panesthia sp.3 (papua new guinea) panesthia sp.1 panesthia heurni geoscapheus robustus geoscapheus rugulosus geoscapheus woodwardi neogeoscapheus dahmsi neogeoscapheus hirsutus panesthia sloanei ancaudellia marshallae ancaudellia kheili ancaudellia shawi panesthiinae macropanesthia rothi parapanesthia pearsoni panesthia tryoni tryoni panesthia tryoni tegminifera parapanesthia gigantea macropanesthia saxicola macropanesthia rhinoceros fig. 1. part 2. see legend on preceding page. legendre f. et al., molecular phylogeny of blaberidae 7 two incertae sedis genera were included in the analyses: eustegasta aff. carabidina had asian perisphaerinae as its sister group (bs / pp = 99 / 1); gynopeltis spp. were monophyletic (bs / pp = 99 / 0.9) and sister group to gyninae (bs / pp = 89 / 0.69). discussion like for many different taxonomic groups, the traditional classification of blaberidae relies mainly on morphological characters. twelve subfamilies have been delimited so far, while 24 genera remain of unknown affinity. we sampled 11 of these subfamilies with molecular data to investigate the phylogenetic relationships of the family, which should be useful for classification and evolutionary studies in this cockroach family. we ended up with a broad-spectrum situation regarding congruence of our molecular phylogenetic hypothesis with the traditional morphology-based classification. on the one side lie molecular hypotheses highly supported and congruent with morphology-based hypotheses. these unsurprising results have been suggested several times and concern the monophyly of blaberidae, (geoscapheinae + panesthiinae) and oxyhaloinae (maekawa et al. 2003; inward 2007; pellens et al. 2007; djernaes et al. 2012; legendre et al. 2014, 2015). likewise, a strong support was found, here and in previous studies, for gyninae, panchlorinae and pycnoscelinae. it is worth noting though that for these subfamilies, only one genus was sampled, which is insufficient to test the monophyly of these subfamilies. on the other side of the spectrum lie conflicting but poorly supported hypotheses. we do not discuss these results in depth because any conclusion about these groups would be premature. indeed, given that deeper nodes were not supported, it is impossible to rule on the monophyletic status of the subfamily perisphaerinae, for instance. admittedly, african and asian perisphaerinae form two distinct and highly supported monophyletic groups. they were never found as sister groups, but none of their potential sister-group relationships were supported (bs < 35, pp < 0.73), so assessing that they cannot be sister groups would be ill-advised. similarly, epilamprinae is split into four clades (and one isolated species) with some geographic consistency and with moderate to high support, but its sister-group relationships are weakly supported (bs < 25, pp < 0.73). it is thus impossible to reach conclusions about these subfamilies with the molecular data at hand. fig. 2. conflicting results among different molecular markers. the phylogenetic relationships of four species are detailed for separate and combined analyses performed in maximum likelihood (ml) and bayesian inference (bi). boxes colored as ‘untested or unresolved’ refer to missing data and multifurcation, respectively. ml bi 12s 12s coi coi coii coii combined combined 16s 16s 18s 18s 28s 28s bi ml true untested or unresolved false phoetalia sister-species of zetoborinae thanatophyllum sister-species of neotropical epilamprinae pronauphoeta sister-species of oxyhaloinae laxta sister-species of african perisphaerinae european journal of taxonomy 291: 1–13 (2017) 8 interestingly, none of the molecular markers used here were informative enough to decipher the relationships among the major well supported blaberid clades, which remain poorly supported. this pattern does not seem to result from conflicting signals among molecular markers, because none of the separate analyses show robust deep relationships. instead, it suggests that the molecular markers used are inefficient for deciphering these phylogenetic relationships. they appear uninformative for this period, which could suggest that a radiation occurred early on in the evolution of blaberidae (probably during the jurassic; legendre et al. 2015), even though this remains to be demonstrated. searching for new markers, as well as performing signal analyses, should help in making progress on this complicated issue (whitfield & lockhart 2007). conversely, it is worth noting that most of the clades found have a strong geographical coherence. we already mentioned epilamprinae and perisphaerinae but, interestingly, geoscapheinae, which are australian, are not monophyletic because australian panesthia are nested among them. with their pantropical distribution and their ancestry tracking back to the mesozoic, blaberidae can be a relevant model to investigate complex biogeographic scenarios. nevertheless, the present analysis shows that, before elaborating on them, more robust knowledge of relationships of subfamilies is needed, an issue already raised before (legendre et al. 2014). between the two extremities of this spectrum are arguably the most interesting situations: conflicting but highly supported results on one hand and congruent but poorly supported results on the other. phoetalia stål, 1874 and thanatophyllum grandcolas, 1991 are two genera showing highly supported, but conflicting positions with morphology-based classifications, where they belong to the subfamilies blaberinae and zetoborinae, respectively. here, on the basis of 5654 bp belonging to five markers (coii is missing), thanatophyllum akinetum was found to be the sister species of some neotropical species of epilamprinae. this relationship was found in half of the separate analyses performed (for the other analyses, t. akinetum was found as the sister species of asian epilamprinae, perisphaerinae or zetoborinae). at the moment of description, t. akinetum was placed within zetoborinae (grandcolas 1991), a position later supported in a cladistic study using 74 morphological characters and sampling 10 genera of zetoborinae, 18 of blaberinae, five of gyninae and two of diplopterinae (grandcolas 1993). as for phoetalia pallida, it was found nested within blaberinae in grandcolas (1993). here it was found as the sister species of a species of zetoborinae (schultesia lampyridiformis), a result also found in five out of the six separate analyses. this disagreement between two studies with different taxon and character samplings does not allow us to decide which hypothesis, and thus which evolutionary scenario, should be favored. blaberinae are notably characterized by male genitalia with spines on the prepuce (roth 1970c). this character could be a synapomorphy of blaberinae (including p. pallida), a striking convergence between p. pallida and blaberinae (without p. pallida), or, depending on the phylogenetic structure of deeper nodes that remain to be deciphered, these spines could actually be a symplesiomorphy of a neotropical clade comprising blaberinae and most of the species of zetoborinae. solving these issues for these two species will require morphoanatomic work at the blaberidae scale, building on previous studies (e.g., grandcolas 1991, 1993; roth 1970a, 1970b, 1970c), and/or in-depth signal analyses of molecular data (thouzé et al. in prep.). two results are congruent with morphology-based classification but with low support in molecular analyses. pronauphoeta sp. (oxyhaloinae) and laxta sp. (australian perisphaerinae) were found to be the sister species of all other oxyhaloinae and african perisphaerinae, respectively. pronauphoeta shelford, 1909 was first placed by shelford (1909) as intermediate between panchlora burmeister, 1838 (panchlorinae) and nauphoeta burmeister, 1838 (oxyhaloinae) but was later suggested to be closer to oxyhaloinae (rehn 1932; kumar 1975) than to panchlorinae, a hypothesis supported here in the combined analyses and with the markers 28s and coii (with the other markers, pronauphaeta sp. was found as the sister species of some epilamprinae, perisphaerinae or in a much deeper position). at the tribe level, pronauphoeta is of unknown affinity and, if its phylogenetic position were to be confirmed, legendre f. et al., molecular phylogeny of blaberidae 9 would belong to a new tribe. as for laxta walker, 1868, grandcolas (1997) revised the perisphaerinae and inferred that it belonged to this subfamily given one external morphological character and four genitalic characters. molecular data suggest that laxta is closely related to african perisphaerinae. nevertheless, its phylogenetic position fluctuates in the different separate analyses, with, for instance, diverse deep positions within the tree based on 18s (ml and bi), 28s (bi), coi (bi) or coii (bi). this result is a remarkable example of disagreement between separate and combined analyses, where signal from the separate analyses disappears in the concatenated analysis (sousa et al. 2013; thiergart et al. 2014), a phenomenon that remains to be understood. sampling other australian species, notably from the genus neolaxta mackerras, 1968, will be necessary to confirm the phylogenetic position of laxta and later suggest biogeographic scenarios for this subfamily, presently known from three continents (asia, africa and oceania; grandcolas 1997). we provide here the first taxonomic affiliation based on molecular data for the incertae sedis genera eustegasta and gynopeltis. gynopeltis was found, with strong support, as closely related to gyninae. other genera of gyninae than gyna must be sampled, however, to assess whether gynopeltis belongs to the gyninae or if another subfamily (gynopeltinae) should be erected. eustegasta has been suggested as belonging to the perisphaerinae before it was removed from this subfamily (roth 1973; grandcolas 1997). here this african genus, only documented for half of the marker, is found with strong support as a sister group to asian perisphaerinae. by deciphering phylogenetic relationships in blaberidae with six molecular markers, we ended up with phylogenetic hypotheses disagreeing, in part, with morphology-based classification. while some situations should be easily resolved with an increase in taxonomic sampling or a decrease in missing data (e.g., congruent results, but with weak support), others would require analyzing the phylogenetic signal of each molecular marker more in depth (e.g., the phylogenetic position of laxta). saturation plots and analyses of bias in nucleotide composition should be performed with phylogenetic informativeness and network analyses (huson & bryant 2006; townsend 2007; philippe & roure 2011). also, given the long branches in the putative sister group pseudophyllodromiinae (legendre et al. 2015) and the results of our preliminary analyses and of previous studies on the effect of outgroup choice on tree reconstruction in dictyoptera (ware et al. 2008; djernaes et al. 2012), the impact of outgroup choice in the phylogenetics of blaberidae should be investigated further. analyzing these conflicting signals and their causes remains, however, a complicated task, so cockroach phylogenetics would above all else benefit from new appropriate markers that are now reachable with the development of next generation sequencing methods (metzker 2010). markers informative for the jurassic period should probably be targeted first to investigate blaberid phylogenetics. acknowledgments this work was supported by the network “bibliothèque du vivant” funded by the cnrs, and by the muséum national d’histoire naturelle, paris (mnhn) under the grants “atm – taxonomie moléculaire: dna barcode et gestion durable des collections” and “atm – biodiversité actuelle et fossile. crises, stress, restaurations et panchronisme: le message systématique” to f.l. we thank tony robillard for his help in several field expeditions, for collecting specimens in mozambique and for commenting on a previous version of this manuscript. we also thank the following people for providing specimens or for their assistance in the field: leonid anisyutkin (zoological institute of the russian academy of sciences, saint-petersburg, russia), rudolf meier (national university of singapore, singapore), yayuk suhardjono and oscar effendy (museum of bogor, indonesia), odette morvan and jean cerda (french guiana), eric guilbert (mnhn), and henri-pierre aberlenc (centre de biologie pour la gestion des populations, montpellier, france). we thank roseli pellens for commenting on a previous version of this manuscript. some specimens were collected in papua new guinea during the expedition “our planet reviewed papua-new-guinea 2012–2013” set up by pro-natura international, mnhn, the european journal of taxonomy 291: 1–13 (2017) 10 institut de recherche pour le développement (ird, france) in partnership with the royal belgian institute of natural sciences, the new guinea binatang research center, the university of papua new guinea and the divine word university of madang and with core funding from the prince albert ii of monaco foundation, the stavros niarchos foundation, the total foundation, the fondation d’entreprise edf, the fonds pacifique, spiecapag, entrepose contracting, the new caledonia government, the reef foundation and the belgian national lottery. the ibisca expert network, the patron for this project, prof. r.l. kitching, and all other participants in this collective 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botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands. legend for supplementary file. list of specimens used in this study, with information about the completion of our dataset. a color code is used to show the traditional taxonomic affiliation of the species sampled. http://biodiversitylibrary.org/page/33103293 https://doi.org/10.1098/rstb.2013.0088 https://doi.org/10.1093/bioinformatics/btu033 https://doi.org/10.1186/s12862-014-0266-0 https://doi.org/10.1080/10635150701311362 https://doi.org/10.1080/10635150701311362 https://doi.org/10.1111/j.1096-0031.2010.00329.x https://doi.org/10.1111/j.1096-0031.2010.00329.x https://doi.org/10.1111/j.1365-3113.2008.00424.x https://doi.org/10.1111/j.1096-0031.1990.tb00550.x https://doi.org/10.1016/j.tree.2007.01.012 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/rt/suppfiles/415/0 european journal of taxonomy 387: 1–17 issn 2118-9773 https://doi.org/10.5852/ejt.2017.387 www.europeanjournaloftaxonomy.eu 2017 · chattová b. et al. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e 1 the genus luticola (bacillariophyta) on ile amsterdam and ile saint-paul (southern indian ocean) with the description of two new species barbora chattová 1,*, marc lebouvier 2, myriam de haan 3 & bart van de vijver 3,4 1 department of botany & zoology, faculty of science, masaryk university, kotlarska 2, 61137 brno, czech republic. 2 umr 6553 ecobio cnrs, université de rennes 1, station biologique, f-35380 paimpont, france. 3 botanic garden meise, research department, domein van bouchout, b-1860 meise, belgium. 4 university of antwerp, department of biology, ecobe, universiteitsplein 1, b-2610 wilrijk, belgium. * corresponding author: barbora.chattova@gmail.com 2 email: marc.lebouvier@univ-rennes1.fr 3 email: myriam.dehaan@plantentuinmeise.be 3,4 email: bart.vandevijver@plantentuinmeise.be abstract. five species of the terrestrial diatom genus luticola d.g.mann were found during a taxonomic survey of two small volcanic islands, ile amsterdam and ile saint-paul (southern indian ocean). apart from the two already known luticola species l. beyensii van de vijver et al. and l. subcrozetensis van de vijver et al., two new species are described: l. ivetana chattová & van de vijver sp. nov. and l. vancampiana chattová & van de vijver sp. nov. finally, one, up to now unknown, luticola species is briefly discussed and illustrated. detailed morphological descriptions of these taxa are provided based on both light and scanning electron microscopy observations. morphological features of the new species are compared to morphologically similar taxa, and notes on their ecology and biogeography are added. keywords. bacillariophyta, diatoms, luticola, new species, morphology. chattová b., lebouvier m., de haan m. & van de vijver b. 2017. the genus luticola (bacillariophyta) on ile amsterdam and ile saint-paul (southern indian ocean) with the description of two new species. european journal of taxonomy 387: 1‒17. https://doi.org/10.5852/ejt.2017.387 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/3.0/ mailto:barbora.chattova%40gmail.com?subject= mailto:marc.lebouvier%40univ-rennes1.fr?subject= mailto:myriam.dehaan%40plantentuinmeise.be?subject= mailto:bart.vandevijver%40plantentuinmeise.be?subject= https://doi.org/10.5852/ejt.2017.387 european journal of taxonomy 387: 1–17 (2017) 2 introduction the genus luticola d.g.mann is one of the dominant genera in the antarctic terrestrial diatom flora. luticola taxa can be distinguished in having uniseriate striae, composed of rounded to transapically elongated areolae covered internally by perforated hymenes, a usually distinct isolated pore in the central area, a longitudinal canal positioned within the valve wall and by a simple, filiform raphe with variable raphe endings (round et al. 1990; levkov et al. 2013). however, the correct taxonomy and biogeography of this genus in the sub-antarctic region was hampered due to species drift and force fitting the antarctic species into european and north american names (tyler 1996). due to the lack of appropriate literature, the first studies mainly reported typical cosmopolitan taxa such as l. cohnii (hilse) d.g.mann (round et al. 1990) or l. nivalis (ehrenb.) d.g.mann (round et al. 1990) and taxa described in the early 20th century such as l. gaussii (heiden) d.g.mann (round et al. 1990) and l. suecorum (g.w.f.carlson) van de vijver (van de vijver & mataloni 2008). moreover, l. muticopsis (van heurck) d.g.mann (round et al. 1990) was used as a catch-all taxon for all capitate luticola specimens, while the non-capitate forms were usually force-fitted into l. mutica (kütz.) d.g.mann (round et al. 1990). this resulted in stretched biogeographical distributions of those species, making them the most widespread in the entire antarctic region. recently, all sub-antarctic luticola species have been the subject of a thorough taxonomic and morphologic revision, in order to unravel the correct diversity of this genus. this resulted in the description of more than 25 new antarctic luticola taxa, mainly from the maritime antarctic region and the antarctic continent (van de vijver et al. 2006, 2011; kopalová et al. 2011; levkov et al. 2013; zidarova et al. 2014; kohler et al. 2015). several new taxa were also found in the sub-antarctic region. in 2002, l. beyensii van de vijver et al. (van de vijver et al. 2002b) was described from ile saint-paul, whereas l. ledeganckii van de vijver (van de vijver et al. 2002a), l. robusta van de vijver et al. (van de vijver et al. 2002a), l. crozetensis van de vijver et al. (levkov et al. 2013), l. ipevii van de vijver & levkov (levkov et al. 2013), and l. subcrozetensis van de vijver et al. (levkov et al. 2013), were described from ile de la possession, the main island of the crozet archipelago. the present paper continues this revision of the genus luticola with an analysis of five taxa observed on ile amsterdam and ile saint-paul, two islands in the southern indian ocean. two luticola taxa are described as new based on detailed light and scanning electron microscopy whereas a fifth unidentified taxon could only be found in light micrograph and is illustrated and discussed but due to the lack of sufficient observations, it is still unclear whether it also represents a new species. material and methods during two short visits (1998 and 1999) and two fieldwork seasons on the volcanic islands ile amsterdam (77°30′ e, 37°50′ s) and ile saint-paul (77°31′ e, 38°43′ s) in 2007 and 2016, more than 300 samples were collected from various habitats including waterbodies, soils and moss vegetations. both islands are among the most remote islands in the world, situated in the southern indian ocean, north of the subantarctic islands, halfway between south africa and australia, and form a separate district within the terres australes et antarctiques françaises (taaf). in geological terms, these islands are very young, with the main part of ile amsterdam formed during the period 400–200 kyr bp and ile saint-paul being even younger (< 50 kyr). more information regarding climate, vegetation and geology can be found in lebouvier & frenot (2007). sampling sites were chosen in order to represent a maximum diversity of habitat types. the samples were collected in pvc bottles and fixed with 3% formalin. in order to determine the moisture content of the sample, the f-value, referring to the f-classification of jung (1936) was selected as representative for moisture and used for each sample. it is a humidity scale based on water content as follow: fi = chattová b. et al., luticola on ile amsterdam and ile saint paul 3 submerged mosses, fii = free floating mosses, fiii = very wet (water drips from the samples without pressure), fiv = wet (water drips with a slight pressure), fv = quasi-wet (water drips after moderate pressure), fvi = moist (little water produced after high pressure), fvii = quasi-dry (only a few drops of water can be squeezed out), fviii = dry (contains no water). samples were prepared for microscopy analysis following the method described in van der werff (1955): small parts of the samples were cleaned by adding 37% h2o2 and heating to 80°c for about 1 h. the reaction was completed by addition of an excessive amount of kmno4. following digestion and centrifugation (three times 10 min at 3700×g), cleaned material was diluted with distilled water to avoid excessive concentrations of diatom valves on the slides. cleaned diatom valves were mounted in naphrax®. light microscopy (lm) observations were done at olympus bx53 microscopes, equipped with differential interference contrast (nomarski) optics. lm micrographs were taken using olympus uc30 camera connected to the cell sense standard program. samples and slides are stored at the brcollection (botanic garden meise, belgium). for scanning electron microscopy (sem), parts of the oxidized suspensions were filtered through a 1 µm isopore™ polycarbonate membrane filter (merck millipore). the stubs were sputter-coated with a platinum layer of 2 nm and studied in a jeol jsm7100f sem microscope at 1 kv (botanic garden meise, belgium). morphological terminology follows round et al. (1990), van de vijver & mataloni (2008) and levkov et al. (2013). the morphology of the new species is discussed based on detailed light and scanning electron microscopy observations and has been compared with known luticola species using the most current literature: rumrich et al. (2000), van de vijver & mataloni (2008), kopalová et al. (2011), van de vijver et al. (2011), levkov et al. (2013), zidarova et al. (2014) and kohler et al. (2015). results five luticola taxa were found in the samples during the survey. two taxa, l. beyensii and l. subcrozetensis, could be identified using the currently available literature. based on a morphological analysis, two taxa are described as new species: l. ivetana chattová & van de vijver sp. nov. and l. vancampiana chattová & van de vijver sp. nov. observation of the material in lm revealed the presence of a fifth unknown luticola taxon with a highly asymmetrical central area. as the taxon could not be found during sem analysis, we have insufficient data regarding its morphological ultrastructure and therefore the decision was taken not to describe it as new to science. class bacillariophyceae haeckel emend. medlin & kaczmarska (medlin & kaczmarska 2004) subclass bacillariophycidae d.g.mann (round et al. 1990) order naviculales bessey (bessey 1907) family diadesmidaceae d.g.mann (round et al. 1990) genus luticola d.g.mann (round et al. 1990) luticola beyensii van de vijver, ledeganck & lebouvier figs 1–12 diatom research 17: 235–241 (van de vijver et al. 2002b). type ile saint paul: taaf, sub-antarctica, 16 dec. 1999, b. van de vijver sample a9 (holo-: cas 220051, california academy of science; iso-: slide no. br–4045, university of antwerp, belgium). european journal of taxonomy 387: 1–17 (2017) 4 description light microscopy (figs 1–10) valves rhombic-lanceolate with clearly convex margins. larger individuals with more or less rostrate apices, in smaller specimens apices more broadly rounded. valve dimensions (n = 25): length 14.5– 22.0 µm, width 6.0–8.5 µm. axial area relatively narrow, linear. central area with a large fascia, rarely reaching the valve margins, due to a series of small areolae bordering the central area near the margins. isolated pore solitary, round, located close to the valve margin, never connected to a stria. raphe filiform, straight, with simple, bent proximal raphe endings, away from the isolated pore. terminal raphe fissures clearly hooked. striae weakly radiate near the central area, becoming more radiate towards the apices, 22–24 in 10 µm. areolae well visible in lm. scanning electron microscopy (figs 11–12) striae composed of 2–4 rounded areolae (figs 11–12). occasionally areolae fused within one stria forming transapically enlarged areolae (fig. 11). terminal raphe fissures clearly hooked, first deflected towards the side opposite the isolated pore, then hooked into the other side, weakly continuing onto the valve mantle (fig. 12). ecology and associated diatom flora luticola beyensii was found in relatively dry, bare soils and on dry mosses (f-value vii-viii) on both islands. the samples with l. beyensii were dominated by several taxa of the genus humidophila r.l.lowe et al. [humidophila contenta (grunow) r.l.lowe et al. (lowe et al. 2014), humidophila brekkaensis figs 1–12. luticola beyensii van de vijver et al. light (lm) and scanning electron micrographs (sem) of a population from ile amsterdam. 1–10. lm of valve face views. 11–12. sem of external view of entire valve, showing raphe structure, position of the isolated pore and striae structure. scale bars: 1–10 = 10 µm; 11–12 = 5 µm. chattová b. et al., luticola on ile amsterdam and ile saint paul 5 (j.b.petersen) r.l.lowe et al. (lowe et al. 2014)], and pinnularia borealis ehrenb. (ehrenberg 1843) and hantzschia amphioxys (ehrenb.) grunow (cleve & grunow 1880). luticola subcrozetensis van de vijver, kopalová, zidarova & levkov figs 13–26 diatoms of europe: diatoms of the european inland waters and comparable habitats 7: 228, pl. 46 figs 15–31, pl. 47 figs 4–6 (levkov et al. 2013). figs 13–26. luticola subcrozetensis van de vijver et al. light (lm) and scanning electron micrographs (sem) of a population from ile saint-paul. 13–22. lm of valve face views. 23–24. sem of external view showing composition of striae and typical raphe structure. 25–26. sem girdle view. scale bars: 13–22 = 10 µm; 23–26 = 5 µm. european journal of taxonomy 387: 1–17 (2017) 6 type ile de la possession: crozet archipelago, sub-antarctica, 5 dec. 1999, b. van de vijver (holo-: slide no. br–4290; iso-: slides brmzu9/25, mkndc 006278/b). description light microscopy (figs 13–22) valves rhombic-lanceolate to elliptic-lanceolate and even elliptic in smaller specimens with clearly convex margins and broadly rounded, non-protracted apices. valve dimensions (n = 25): length 12– 29 µm, width 7.0–8.5 µm. axial area linear, slightly widening towards apices and central area. central area bow-tie shaped, wider opposite the isolated pore, bordered by shortened striae on both sides. an isolated pore present in the central area, located halfway between the valve centre and the valve margin. raphe branches straight, bent away from the isolated pore at both proximal and distal ends. striae radiate throughout the entire valve, 15–17 in 10 µm. scanning electron microscopy (figs 23–26) striae composed of 3–5 small, rounded to slightly elongated areolae at the apices, only 1–2 rounded areolae per stria, extending past the terminal raphe fissures on the valve face (fig. 23). a single row of rounded areolae present along entire valve mantle, interrupted at apices (fig. 26). isolated pore rounded, slightly elongated, not associated with a stria. central area usually bordered by one, rarely two, series of small, rounded areolae. raphe branches straight to slightly curved. proximal raphe endings bent to the side opposite the isolated pore, terminating into weakly developed pores. terminal raphe fissures bent to the same side as the proximal endings terminating on the valve face, not extending onto the mantle (fig. 24). ecology and distribution luticola subcrozetensis is a typical sub-antarctic and maritime antarctic species. originally described from ile de la possession (crozet archipelago), the species was also reported from livingston island (zidarova et al. 2016), one of the major south shetland islands, located in the southern atlantic ocean. the species was recently found in moss samples from gough island (vinšová, unpubl. res.). it was present in samples from both ile saint-paul and ile amsterdam. the largest population was observed in a soil sample collected near the rockhopper penguin rookery of ile saint-paul, at an altitude of 110 m, hidden in cracks in the rocks that border the penguin rookery. luticola ivetana chattová & van de vijver sp. nov. figs 27–43 etymology the new species was named after mrs. iveta chattová, mother of the first author, on the occasion of her 50th birthday. type ile amsterdam: entrecasteaux, taaf, sub-antarctica, 37°51′18.6″ s, 77°31′23.5″ w, 21 dec. 2016, b. van de vijver sample w030 (holo-: slide no. br–4495, fig. 27; iso-: slide plp–330; university of antwerp, belgium). description light microscopy (figs 27–36) chattová b. et al., luticola on ile amsterdam and ile saint paul 7 figs 27–43. luticola ivetana chattová & van de vijver sp. nov. light (lm) and scanning electron micrographs (sem) from the type population from entrecasteaux on ile amsterdam, b. van de vijver sample w030. 27–36. lm of valve face views. 37–38. sem of valve exterior. 39. sem girdle view. 40. sem of valve interior. 41. sem of external detail of hooked terminal raphe fissures. 42. sem of external detail view of central area showing the typical deflection of proximal raphe endings. 43. sem of external detail view of areolae structure. scale bars: 27–36 = 10 µm; 37–38, 40–43 = 1 µm; 39 = 5 µm. european journal of taxonomy 387: 1–17 (2017) 8 valves elliptic-lanceolate with convex margins and broadly rounded, non-protracted apices. valve dimensions (n = 50): length 11.0–25.5 µm, valve width 6.0–7.5 µm. axial area narrow, linear, almost not widening towards the apices and central area. central area forming a bow-tie shaped stauros. one isolated pore present in the central area, positioned close to the valve centre. raphe filiform, straight with weakly deflected simple proximal raphe endings and elongated terminal raphe fissures. striae radiate throughout the entire valve, 16–18 in 10 µm. scanning electron microscopy (figs 37–43) striae composed of 2–4 round to elongated areolae, clearly enlarged near the central area and the valve margins (figs 37, 43). mantle areolae very large, rounded, never slitlike (fig. 39). central area bordered by 1–2 rounded to weakly transapically elongated areolae. isolated pore elliptic, clearly isolated from the central striae (fig. 38). raphe branches straight with short proximal raphe endings bent towards the side with the isolated pore (fig. 42). terminal raphe fissures hooked, continuing onto the valve mantle (fig. 41). internally, poroids of valve face occluded by hymens forming a continuous strip on each stria. distinct stauros visible. internal proximal raphe endings straight, terminating on the edge of the stauros. distal raphe endings terminating onto small helictoglossae (fig. 40). ecology and distribution so far, l. ivetana sp. nov. was observed on ile amsterdam only. the type locality where a large population of this new species was observed, was a small crack in a rock face at entrecasteaux, clearly under the permanent influence of seaspray. a very thin film of water was present in the crack together with wet mud. the sample was taken by scraping off the mud and the water from the crack. another large population where l. ivetana sp. nov. was found is a lava cavern in the partly collapsed grand tunnel, running from the cratères vénus to the northern coast. the sample was taken from wet mosses (f-value iv-v), growing on the wall of the cavern, close to the entrance, in a population of blechnum australe l. the sample was dominated by ferocia setosa (greville) van de vijver & houk (van de vijver et al. 2017), orthoseira verleyenii van de vijver (lowe et al. 2013), sellaphora barae van de vijver & e.j.cox (van de vijver & cox 2013) and various humidophila species. luticola vancampiana chattová & van de vijver sp. nov. figs 44–71 etymology the species is named after prof. dr. karel van camp (university of antwerp, belgium), to thank him for his efforts in conserving the van heurck collection and his broad interest in diatom research. type ile saint-paul: conserverie, taaf, sub-antarctica, 38°42′52.0″ s, 77°31′55.5″ w, 24 nov. 2016, b. van de vijver sample s029 (holo-: slide no. br–4496, fig. 47; iso-: slide plp–331; university of antwerp, belgium). description light microscopy (figs 44–67) valves linear-lanceolate to elliptic-lanceolate, elliptic in the smallest specimens, with protracted, subcapitate to rostrate apices. in smaller specimens, valve ends not as protracted, only subrostrate or obtusely rounded. initial cells rhombic-lanceolate. valve dimensions (n = 35): length 10.0–26.0 μm, width 5.0–8.5 μm. axial area linear, narrow. single, rounded isolated pore present in the central area, located halfway between the valve centre and margin, sometimes attached to a stria. central area wide, bow–tie shaped, bordered on both sides with a series of 3–5 round areolae. raphe branches straight, chattová b. et al., luticola on ile amsterdam and ile saint paul 9 figs 44–71. luticola vancampiana chattová & van de vijver sp. nov. light (lm) and scanning electron micrographs (sem) from the type population from conserverie on ile saint-paul, b. van de vijver sample s029. 44–67. lm showing the variation in size and shape of the valve apices. 68–69. sem of valve exterior. 70. sem of valve interior. 71. sem girdle view. scale bars: 44–67 10 µm; 68–71 = 5 µm. european journal of taxonomy 387: 1–17 (2017) 10 deflected away from the isolated pore at both proximal and distal ends. transapical striae radiate throughout, 16–19 in 10 µm. scanning electron microscopy (figs 68–71) valve mantle with a single row of round areolae (fig. 71), interrupted at the apices. striae composed of 3–4 areolae, 1–2 areolae at the apices. areolae round to elongated, clearly enlarged and prolonged close to the valve margins (fig. 68). internally, areolae occluded by hymens, forming a continuous strip across the valve (fig. 70). isolated pore showing an elliptic external opening. internally, central nodule thickened, expanding into stauros. isolated pore with semi-lunar opening formed by tonguelike structure. raphe branches straight. proximal raphe endings short, deflected to side opposite to the isolated pore. terminal raphe fissures deflected to the same side as the proximal endings, not extending onto the mantle (fig. 69). internally, proximal raphe endings straight, while distally, raphe branches terminating on small helictoglossae. ecology and distribution so far l. vancampiana sp. nov. was found on both ile amsterdam and ile saint-paul. the largest population was found in a wet soil covered by green algae close to the ocean on ile saint-paul. the sampling site is frequently visited by fur seals and penguins, as was visible in the macroremains (feathers, excrements). the sample was almost entirely dominated by the new luticola species. other (smaller) populations were found in soil samples collected near penguin rookeries where similar conditions (considerable biotic influence, considerable salinity input) prevailed. luticola sp. figs 72–82 description light microscopy (figs 72–82) valves linear-lanceolate to elliptic-lanceolate in smaller specimens, with rounded, non-protracted apices. valve dimensions (n = 12): length 10.7–20.5 µm, width 4.5–6.5 µm. axial area narrow, linear. central area wide, bow-tie shaped, clearly asymmetrical with the side bearing the isolated pore markedly wider. isolated pore present in the central area, located halfway between the valve centre and margin. raphe branches straight, deflected away from the isolated pore at both proximal and distal ends. transapical striae punctate, radiate throughout the entire valve, 20–24 in 10 µm. remarks so far, this species has only been observed with light microscopy. further studies and sem observations will be necessary to determine its correct taxonomic position. the species was so far found in only two soil samples (a6 and a7) taken from ile saint-paul. both samples are dominated by l. beyensii, figs 72–82. luticola sp. light micrographs of a population from ile saint-paul. scale bar: 10 μm. chattová b. et al., luticola on ile amsterdam and ile saint paul 11 humidophila brekkaensis, pinnularia acidicola van de vijver & le cohu (van de vijver et al. 2002a) and hantzschia amphioxys. discussion the genus luticola is one of the dominant components of the diatom flora in terrestrial habitats on subantarctic islands. these habitats show a diverse and very characteristic terrestrial flora with high degree of regional endemism in the entire antarctic region, as is demonstrated by the large number of endemic taxa in typical terrestrial genera such as microfissurata lange-bertalot et al. (cantonati et al. 2009), orthoseira thwaites (van de vijver & kopalová 2008, lowe et al. 2013), humidophila (kopalová et al. 2015; chattová et al., 2017), hantzschia grunow (zidarova et al. 2010) and muelleria frenguelli (frenguelli) (van de vijver et al. 2010, 2014). the morphological and taxonomical analysis of the genus luticola on ile amsterdam and ile saint-paul revealed the presence of two new species. a wide combination of morphological features such as valve outline, shape of the valve apices, shape of the central and axial area, position and shape of the isolated pore, stria pattern and density and raphe and areola structure were used for comparison with all similar taxa known worldwide so far. sufficient morphological differences were found to justify the separation of both taxa as new to science. the main discriminative features between the new species and their morphologically similar taxa are summarized in table 1 (luticola ivetana sp. nov.) and table 2 (luticola vancampiana sp. nov.). luticola ivetana sp. nov. resembles several luticola species with elliptic lanceolate valves, however the new species can be easily distinguished by the proximal raphe endings typically bent towards the isolated pore. there are only a few species showing a similar deflection such as l. ledeganckii, l. frequentissima levkov et al. (levkov et al. 2013) and l. raynae zidarova & van de vijver (van de vijver et al. 2011). luticola raynae and l. ledeganckii have typical capitate apices, always lacking in l. ivetana sp. nov. the most similar is l. frequentissima, but the latter species can be differentiated by the shape of the central area, much smaller in l. ivetana sp. nov. and by its stria structure. luticola frequentissima has transapical striae composed of 4–6 areolae per stria contrary to l. ivetana sp. nov. that has only 2–4 areolae per stria. the two species also differ ecologically, l. ivetana sp. nov. seems to be an aerophilic and most likely even slightly brackish taxon given the habitat in entrecasteaux where it was found, while l. frequentissima always occurs in freshwater habitats. luticola mutica and l. imbricata (bock) levkov et al. (levkov et al. 2013) have longer proximal raphe endings curved doubly, a feature lacking in l. ivetana sp. nov. luticola imbricata additionally differs from the new taxon by the shape of its isolated pore (rounded vs elliptical) and by the apices that are narrowly rounded vs broadly rounded. luticola mutica does not only differ by the shape and deflection of the raphe endings but also by the different shape of central area and the structure of the striae. luticola australomutica van de vijver (van de vijver & mataloni 2008) is an antarctic species described from deception island, with a similar valve outline as l. ivetana sp. nov. nonetheless, both species can be distinguished without any difficulties based on the morphology of the distal and proximal raphe endings. luticola australomutica has short, almost straight distal raphe fissures, terminating before the last row of areolae, while in l. ivetana sp. nov. the distal raphe fissures are hooked, continuing onto the valve mantle. additionally, l. australomutica has striae composed of 4–5 areolae contrary to 2–4 areolae per stria in l. ivetana sp. nov. smaller specimens of l. ivetana sp. nov. also resemble l. pseudoimbricata levkov et al. (levkov et al. 2013) based on their valve shape, but can however be separated by the different shape and deflection of both raphe proximal and distal fissures. luticola similis levkov et al. can be differentiated from l. ivetana sp. nov. by the wide and asymmetrical central area, by its lanceolate valve outline and its narrowly rounded valve apices. european journal of taxonomy 387: 1–17 (2017) 12 ta xo n lu tic ol a iv et an a sp . n ov . lu tic ol a le de ga nc ki i lu tic ol a fr eq ue nt is si m a lu tic ol a ra yn ae lu tic ol a m ut ic a lu tic ol a im br ic at a lu tic ol a au st ra lo m ut ic a lu tic ol a ps eu do im br ic at a lu tic ol a si m ili s l en gt h (µ m ) 11 –2 5 14 –2 8 12 –2 7 15 –2 7 11 –2 8 10 –2 3 17 –3 5 9– 22 8– 33 w id th (µ m ) 6. 0– 7. 5 7. 0– 9. 0 6. 4– 9. 0 7. 0– 10 .0 6. 0– 9. 5 5. 0– 7. 0 5. 5– 8. 5 5. 0– 7. 0 5. 5– 7. 5 st ri ae (i n 10 µ m ) 16 –1 8 15 –1 8 12 –2 4 14 –1 5 16 –1 8 18 –2 2 16 –2 1 20 –2 4 21 –2 4 a re ol ae (p er s tr ia ) 2– 4 3– 4 4– 6 3– 5 3– 4 3– 4 4– 5 3– 4 3– 5 v al ve s ha pe el lip tic la nc eo la te el lip tic la nc eo la te el lip tic la nc eo la te to el lip tic lin ea rel lip tic to re ct an gu la r el lip tic la nc eo la te to lin ea rel lip tic lin ea rla nc eo la te to e lli pt ic la nc eo la te la nc eo la te to e lli pt ic la nc eo la te lin ea rel lip tic el lip tic la nc eo la te to el lip tic v al ve a pi ce s br oa dl y ro un de d ca pi ta te to su bc ap ita te ro un de d br oa dl y ro st ra te ro un de d na rr ow ly ro un de d sl ig ht ly ro st ra te ro un de d na rr ow ly ro un de d c en tr al a re a bo w -t ie sh ap ed re ct an gu la r t o el lip tic w id e, b ow -t ie sh ap ed tr an sv er sa lly el on ga te d na rr ow , el lip tic al bo w -t ie sh ap ed w ed ge to bo w -t ie sh ap ed re ct an gu la r t o bo w -t ie -s ha pe d w ed ge sh ap ed , as ym m et ri ca l a xi al a re a na rr ow , l in ea r na rr ow , l in ea r na rr ow , l in ea r na rr ow , lin ea r na rr ow , l in ea r na rr ow , l in ea r na rr ow , l in ea r na rr ow , l in ea r na rr ow , l in ea r pr ox im al ra ph e en di ng s ho ok ed , de fle ct ed to w ar ds th e is ol at ed p or e de fle ct ed to w ar ds th e is ol at ed p or e ho ok ed , de fle ct ed to w ar ds th e is ol at ed p or e ho ok ed do ub ly cu rv ed sh or t, de fle ct ed w ea kl y de fle ct ed w ea kl y cu rv ed be nt d is ta l r ap he en di ng s ho ok ed w ea kl y cu rv ed cu rv ed ho ok ed ho ok ed "? "sh ap ed sh or t, de fle ct ed sh or t, de fle ct ed "? "sh ap ed ta bl e 1. m or ph ol og ic al c om pa ri so n be tw ee n l. iv et an a sp . n ov . a nd m or ph ol og ic al ly s im ila r t ax a. chattová b. et al., luticola on ile amsterdam and ile saint paul 13 ta bl e 2. m or ph ol og ic al c om pa ri so n be tw ee n l. v an ca m pi an a sp . n ov . a nd m or ph ol og ic al ly s im ila r t ax a. ta xo n lu tic ol a va nc am pi an a sp . n ov . lu tic ol a ro bu st a lu tic ol a le de ga nc ki i lu tic ol a m on ta na lu tic ol a su ec or um lu tic ol a ip ev ii lu tic ol a tr un ca ta lu tic ol a m ut ic op si s lu tic ol a pe rm ut ic op si s l en gt h (µ m ) 10 –2 6 15 –4 2 14 –2 8 17 –3 8 20 –4 2 16 –3 4 17 –3 6 12 –2 2 17 –2 6 w id th (µ m ) 5. 0– 8. 5 7. 5– 11 .5 7. 0– 9. 0 7. 0– 10 .0 7. 5– 13 .0 6. 0– 9. 0 8. 3– 10 .7 7. 0– 9. 0 7. 0– 9. 2 st ri ae (i n 10 µ m ) 16 –1 9 13 –1 4 15 –1 8 19 –2 2 13 –1 4 14 –1 8 16 –1 9 15 –1 8 18 –2 2 a re ol ae (p er s tr ia ) 3– 5 3– 4 3– 4 5– 8 3– 5 4– 5 3– 5 3– 4 4– 6 v al ve s ha pe lin ea rla nc eo la te to e lli pt ic la nc eo la te lin ea rla nc eo la te to lin ea rel lip tic el lip tic la nc eo la te la nc eo la te to e lli pt ic la nc eo la te lin ea rla nc eo la te to e lli pt ic la nc eo la te el lip tic la nc eo la te la nc eo la te to li ne ar la nc eo la te lin ea rel lip tic lin ea rla nc eo la te to lin ea rel lip tic v al ve a pi ce s su bc ap ita te to ro st ra te ro st ra te to ca pi ta te ca pi ta te to su bc ap ita te br oa dl y ro st ra te br oa dl y ro un de d, ro st ra te br oa dl y ro un de d tr un ca te br oa dl y ca pi ta te br oa dl y ca pi ta te c en tr al a re a bo w -t ie -s ha pe d tr an sv er sa lly el lip tic re ct an gu la r t o el lip tic bo w -t ie sh ap ed tr an sv er sa lly el lip tic w ed ge sh ap ed re ct an gu la r t o el lip tic bo w -t ie sh ap ed tr an sv er sa lly el on ga te d a xi al a re a na rr ow , l in ea r w id e, li ne ar na rr ow , l in ea r na rr ow , l in ea r na rr ow , l in ea r na rr ow , l in ea r na rr ow , l in ea r w id e, li ne ar na rr ow , lin ea r pr ox im al ra ph e en di ng s sh or t, de fle ct ed di st in ct ly de fle ct ed de fle ct ed to w ar ds th e is ol at ed p or e sl ig ht ly cu rv ed do ub ly cu rv ed di st in ct ly de fle ct ed di st in ct ly de fle ct ed di st in ct ly de fle ct ed de fle ct ed d is ta l r ap he en di ng s de fle ct ed sh or t, de fle ct ed w ea kl y cu rv ed ho ok ed w ea kl y cu rv ed w ea kl y cu rv ed w ea kl y cu rv ed w ea kl y cu rv ed w ea kl y cu rv ed european journal of taxonomy 387: 1–17 (2017) 14 the largest specimens of l. vancampiana sp. nov. bear some similarities to l. robusta. however, both taxa significantly differ in stria density (13–14 in 10 µm in l. robusta vs 16–19 in 10 µm in l. vancampiana sp. nov.). additionally, l. robusta has a markedly wider axial area. luticola vancampiana sp. nov. may also resemble l. ledeganckii based on valve shape and stria density. the main difference between these two species can be observed in the direction of proximal raphe endings, which are deflected towards the isolated pore in l. ledeganckii. luticola montana levkov et al. (levkov et al. 2013) has a similar valve shape compared to l. vancampiana sp. nov., but can be easily differentiated from the latter species by its stria and areola density. luticola montana has finer (19–22 in 10 µm) striae composed of 5–8 areolae, contrary to l. vancampiana sp. nov. that has only 3–5 areolae per stria. additionally, the distal raphe fissures in l. montana continue onto the valve mantle, interrupting the row of areolae. luticola suecorum has similar valve shape and protracted apices. nevertheless, both species can be hardly confused and are easily distinguished by their valve dimensions and stria density. luticola suecorum is much larger (20–42 µm, 7.5–13 µm) with coarser striae (13–14 in 10 µm). additional differences can be observed in the sem. luticola suecorum has doubly hooked distal raphe endings that extend onto the valve mantle and an ornamented central area with surface depressions. luticola ipevii can be differentiated from l. vancampiana sp. nov. by its slender, elongated valve outline and non-protracted, broadly rounded apices. smaller valves of l. vancampiana sp. nov. are similar to l. truncata kopalová & van de vijver (kopalová et al. 2009). a clear distinction between the two taxa can be made due to the valve width. luticola vancampiana sp. nov. (w = 5.0–8.5 µm) has narrower valves for any given valve width when compared to l. truncata (w = 8.5–10.5 µm). finally, l. muticopsis and l. permuticopsis kopalová & van de vijver (kopalová et al. 2011) have both more capitate apices. moreover, l. permuticopsis has a higher striae density (18–22 in 10 µm). the last and unknown species of luticola, presented here as luticola sp., is characteristic by its asymmetrical central area, narrow valves and consistently linear-lanceolate outline, a combination of features that is almost never encountered within the genus luticola. based on the valve shape and striae density, the most similar species is l. frequentissima. both species can be differentiated by their proximal raphe endings which are deflected towards the isolated pore in l. frequentissima. moreover, luticola sp. has slightly narrower valves for any given valve length when compared to l. frequentissima. the central area of l. similis levkov et al. (levkov et al. 2013) is slightly asymmetrical, but always wider on the side opposite to the isolated pore. luticola mutica has a more variable valve outline, a narrower central area and coarser striae (16–18 in 10 µm). acknowledgements the authors would like to thank drs. pieter ledeganck, prof. dr. eric vidal, dr. jennie whinam, prof. dr. willem de smet, dr. germinal rouhan and dr. damien ertz for their help during the sampling campaigns, supported by the french polar institute (programmes ipev 136 and 1167). mrs. b. chattová benefited from an institutional support of masaryk university and an erasmus scholarship while studying at the botanic garden meise (belgium) and the university of antwerp (belgium). references bessey c.e. 1907. a synopsis of plant phyla. nebraska university studies 7: 275–373. cantonati m., van de vijver b. & lange-bertalot h. 2009. microfissurata gen. nov. 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region: maritime antarctica. iconographia diatomologica 24: 1–504. https://doi.org/10.2216/07-67.1 https://doi.org/10.1080/0269249x.2002.9705541 https://doi.org/10.1127/0029-5035/2006/0082-0069 https://doi.org/10.2216/09-27.1 https://doi.org/10.1127/0029-5035/2011/0092-0137 https://doi.org/10.5507/fot.2014.006 https://doi.org/10.11646/phytotaxa.332.1.2 https://doi.org/10.5091/plecevo.2010.402 https://doi.org/10.11646/phytotaxa.170.3.2 chattová b. et al., luticola on ile amsterdam and ile saint paul 17 manuscript received: 12 july 2017 manuscript accepted: 17 october 2017 published on: 22 december 2017 topic editor: frederik leliaert ph.d., koen martens desk editor: alejandro quintanar printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, madrid, spain. european journal of taxonomy 524: 1–15 issn 2118-9773 https://doi.org/10.5852/ejt.2019.524 www.europeanjournaloftaxonomy.eu 2019 · collado g.a. et al. this work is licensed under a creative commons attribution license (cc by 4.0). r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:6a434946-471e-4927-8235-a793f249d560 1 systematic evaluation of cryptic freshwater snails from central chile, including the enigmatic littoridina santiagensis (gastropoda, truncatelloidea) gonzalo a. collado 1*, karina p. aguayo 2, néstor j. cazzaniga 3, diego e. gutiérrez gregoric 4 , micaela de lucía 5 & martin haase 6 1,2 departamento de ciencias básicas, facultad de ciencias, universidad del bío-bío, avenida andrés bello 720, chillán, chile. 3 departamento de biología bioquímica y farmacia, universidad nacional del sur, san juan 670. 8000 bahía blanca, argentina. 4,5 división zoología invertebrados, museo de la plata, facultad de ciencias naturales y museo, universidad nacional de la plata, paseo del bosque s/n°, la plata, buenos aires, b1900wfa, argentina. 6 vogelwarte, zoological institute and museum, university of greifswald, soldmannstr. 23,17489 greifswald, germany. *corresponding author: gcollado@ubiobio.cl 2 email: karina.aguayo.rosales@gmail.com 3 email: ficazzan@criba.edu.ar 4 email: dieguty@fcnym.unlp.edu.ar 5 email: mec_dl@hotmail.com 6 email: mhaase@uni-greifswald.de 1 urn:lsid:zoobank.org:author:04354188-59c0-4d39-b418-f13320d21a17 2 urn:lsid:zoobank.org:author:3a2edef7-a8d6-4755-8535-75c8d0d53e8f 3 urn:lsid:zoobank.org:author:2ec19b39-ed0d-47af-95aa-4ac32c616751 4 urn:lsid:zoobank.org:author:97674ddc-a422-4e70-8106-2113d778eb0b 5 urn:lsid:zoobank.org:author:c8d15276-9582-4578-981f-18e40ed5a2cd 6 urn:lsid:zoobank.org:author:8ce4b347-14c5-464f-99c4-de2263fc55d5 abstract. walter biese described littoridina santiagensis biese, 1944 (cochliopidae) from estero dehesa in 1944 based exclusively on external shell features and a second allopatric population in yeso spring three years later. since 2011 different samplings have been carried out at the type locality and have only provided specimens of the morphologically similar invasive mudsnail potamopyrgus antipodarum gray, 1843 (tateidae), raising doubts about the identity of the species. the recent finding of two snail morphotypes in yeso spring, a thick shelled form congruent with type specimens of l. santiagensis and a slender one morphologically associable to p. antipodarum, allowed comparative studies, including the taxonomic analysis of additional populations with similar shell morphology occurring in central chile. a dna barcoding (coi) approach identified the slender form from yeso spring in maipo basin and a second population from the contiguous rapel basin indeed as the invasive p. antipodarum; however, https://doi.org/10.5852/ejt.2019.524 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/4.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:6a434946-471e-4927-8235-a793f249d560 mailto:gcollado%40ubiobio.cl?subject= mailto:karina.aguayo.rosales%40gmail.com?subject= mailto:ficazzan%40criba.edu.ar?subject= mailto:dieguty%40fcnym.unlp.edu.ar?subject= mailto:mec_dl%40hotmail.com?subject= mailto:mhaase%40uni-greifswald.de?subject= http://zoobank.org/urn:lsid:zoobank.org:author:04354188-59c0-4d39-b418-f13320d21a17 http://zoobank.org/urn:lsid:zoobank.org:author:3a2edef7-a8d6-4755-8535-75c8d0d53e8f http://zoobank.org/urn:lsid:zoobank.org:author:2ec19b39-ed0d-47af-95aa-4ac32c616751 http://zoobank.org/urn:lsid:zoobank.org:author:97674ddc-a422-4e70-8106-2113d778eb0b http://zoobank.org/urn:lsid:zoobank.org:author:c8d15276-9582-4578-981f-18e40ed5a2cd http://zoobank.org/urn:lsid:zoobank.org:author:8ce4b347-14c5-464f-99c4-de2263fc55d5 european journal of taxonomy 524: 1–15 (2019) 2 l. santiagensis was recovered among species of potamolithus pilsbry, 1896 (tateidae), justifying the potamolithus santiagensis (biese, 1944) comb. nov. besides recognition of three other populations as belonging to potamolithus, the molecular analysis also suggests trans-andean dispersal of this group of snails in the southern cone of south america. keywords. cryptic species, freshwater snails, cochliopidae, new combination, tateidae. collado g.a., aguayo k.p., cazzaniga n.j., gutiérrez gregoric d.e., de lucía m. & haase m. 2019. systematic evaluation of cryptic freshwater snails from central chile, including the enigmatic littoridina santiagensis (gastropoda, truncatelloidea). european journal of taxonomy 524: 1–15. https://doi.org/10.5852/ejt.2019.524 introduction the taxonomic placement of minute chilean freshwater snails solely described based on morphological characters, such as littoridina santiagensis biese, 1944, is rather controversial. this species was described in the genus littoridina souleyet, 1852 by walter biese from estero dehesa, a small tributary of the mapocho river located east of santiago in the maipo basin, based on characteristics of the shell and for which only a single drawing is available from the original description (biese 1944) (fig. 1). along with similar forms, biese related l. santiagensis to the littoridina hatcheri pilsbry, 1911 group, a species subsequently transferred to the genus heleobia stimpson, 1865 within the family cochliopidae (hershler & thompson 1992), but now recognized as member of the family tateidae, closely related to the genus potamolithus pilsbry, 1896 (koch et al. 2015). shortly after the original publication, biese (1947) discovered a second allopatric population in a spring tributary of the yeso river, 55 km south of the type locality, which he assigned to l. santiagensis using the same typological criteria. subsequent finds of the species have not been made, but it was cited by stuardo (1961) in one of the most important checklists of chilean freshwater mollusks of the 20th century. later, hershler & thompson (1992), in their influential work on cochliopinae, placed all the chilean species of littoridina, including l. santiagensis, in the genus heleobia, but without providing arguments for this translocation. this was perhaps one of the reasons why several authors continued to consider these species as members of littoridina (e.g., sielfeld 2001; valdovinos 1999, 2006, 2008). fig. 1. ‟littoridina” santiagensis biese, 1944. a. the only drawings of a specimen of the species taken from the original description by biese (1944). b. lectotype housed at the museo nacional de historia natural, santiago, chile (mnhncl) (after collado et al. 2011). https://doi.org/10.5852/ejt.2019.524 collado g.a. et al., systematic evaluation of cryptic freshwater snails 3 during a study of chilean freshwater snails performed by the first author in 2011, specimens of l. santiagensis were not found at its type locality but specimens attributable to the species were found in yeso spring (collado et al. 2011). based on anatomical characters of the male copulatory organ, the authors moved the chilean species of littoridina to the genus heleobia, although in the case of l. santiagensis this was not justified because no males were found among the specimens from that locality. more recently, a molecular phylogenetic analysis based on dna sequences of the mitochondrial gene cytochrome oxidase subunit i (coi) obtained from snails collected from estero dehesa identified these specimens as the new zealand mudsnail potamopyrgus antipodarum (gray, 1843) in the family tateidae (collado 2014), which is morphologically similar to l. santiagensis, suggesting either species displacement or an early misidentification of these snails as members of the genus littoridina. the absence of males in snails examined from yeso spring (collado et al. 2011) also contributed to raise doubts about the identity of this species, since the genera littoridina and heleobia contain dioecious species (e.g. hubendick 1955; gaillard & de castellanos 1976; cazzaniga 1980, 1982a, 1982b; hershler & thompson 1992; collado et al. 2011; ovando & de francesco 2011; collado 2012, 2015). new collections at the type locality (estero dehesa) of l. santiagensis in 2014, 2015 and 2017 have only been successful for p. antipodarum despite intensive sampling effort performed in different microhabitats (pers. obs.). on the other hand, two morphotypes were found in yeso spring, a slender form conferred to p. antipodarum and another thicker one provisionally assigned to l. santiagensis, taking into account the previous record by biese and its morphological similarity regarding the type specimens of this species studied previously (fig. 1) (collado et al. 2011). given the new material, in the present study we performed comparative studies between these snails and evaluated their systematic position based on morphological and molecular data, including other morphologically similar snail populations of uncertain taxonomic status found in central chile. material and methods the snails were collected from estero dehesa (33°21′58.49″ s, 70°31′15.70″ w) and yeso spring (33°45′29.36″ s, 70°10′00.41″ w), maipo basin, región metropolitana, chile. collections were also performed at three other localities harboring populations with a similar shell morphology, the irrigation canal lo carreño (34°36′45″ s, 70°55′47.8″ w) in the región de o’higgins, located about 143 km south of estero dehesa, two populations from the región del maule, viña casas del maule (35°33′04.37″ s, 71°40′12.25″ w), located about 265 km south of estero dehesa, and el colorado, located about 50 km southeast of viña casas del maule. to study the gross morphology, specimens were photographed using a motic smz–168 stereo microscope provided with a moticam 2000 digital camera and then compared with standard specimens of known taxonomic allocation (collado 2014, 2016), type specimens or drawings of them (pilsbry 1911; biese 1944, 1947). the following measurements were taken on 95 snails: shell length (sl), shell width (sw), aperture length (al), aperture width (aw), body whorl length (bwl) and spire length (spl). shell variables were logarithmically transformed for statistical analyses. since variables were partly non-normally distributed and had inhomogeneous variances, we performed non-parametric kruskal-wallis tests and post-hoc pairwise mann-whitney u-tests for comparisons in statistica ver. 7.0 (statsoft inc. 2004). a principal component analysis (pca) was performed to visualize the morphological variation of different populations. protoconchs, radulae and opercula were isolated and cleaned in a diluted sodium hypochlorite solution and then observed using a hitachi su3500 scanning electron microscope (sem). voucher specimens were deposited in the museo de ciencias naturales profesor pedro ramírez fuentes (mcnpprf 139–11 to 139–21), chillán, chile. dna extraction and amplification of partial sequences of the coi gene were performed following collado (2014) using the new pair of primers atctggtttagtrggraca (forward) and cctgcyaaaacaggyaaag (reverse). the new sequences were deposited in genbank (mh536524–mh536536). genetic uncorrected p-distances were estimated in mega v.7.0 (kumar european journal of taxonomy 524: 1–15 (2019) 4 et al. 2016). an initial blastn-analysis against the data base of genbank confirmed that the species in question belonged to tateidae. we also performed a bayesian tree-based barcoding approach (erickson & driskell 2012) to assess the familial/generic allocation of the species analyzed together with representative species of these families retrieved from genbank (hershler et al. 1999; haase 2005; kroll et al. 2012; hamada et al. 2013; wilke et al. 2013; koch et al. 2015; collado et al. 2016a; de lucía & gutiérrez gregoric 2017; colgan & da costa, unpublished data). as coi was highly saturated across both families as shown by the test of xia et al. (2003), our analysis was not supposed to reflect accurate relationships, but it allowed us to assign the chilean species to their respective families and genera. we constructed a bayesian tree using mrbayes ver.3.2.2 (ronquist et al. 2011) based on the hky + i + γ substitution model identified as best fitting by jmodeltest ver.2.1.4 (darriba et al. 2012) according to all four criteria implemented in this program. mrbayes was run for 2 mio generations with a burn-in of 25%. in order not to aggravate the saturation problem by introducing an outgroup, the tree was midpoint rooted in figtree ver.1.4.3 (http://tree.bio.ed.ac.uk/software/figtree/). for further comparison, we also obtained sequences of two specimens of potamolithus from puerto chico, llanquihue lake, región de los lagos (41°19′37.06″ s; 72°57′26.64″ w), the type locality of p. australis biese, 1944, the only species of the genus reported from chile. however, as it has been considered a nomen dubium (lópez armengol 1985), we treat these snails as potamolithus sp. until a formal redescription may be established. results species delimitation the classification of specimens based on external shell features was difficult to perform in the field. under magnification, two snail morphotypes could be recognized in yeso spring and lo carreño, a slender form morphologically associable to p. antipodarum and a thick-shelled one congruent with the type specimens of l. santiagensis, here transferred to potamolithus based on anatomical characters and dna barcode analysis (see below). the thicker-shelled specimens from el colorado and viña casas del maule were assigned to potamolithus as well (fig. 2). superfamily truncatelloidea gray, 1840 family tateidae thiele, 1925 genus potamolithus pilsbry, 1896 type species: paludina lapidum d’orbigny, 1835, by subsequent designation. potamolithus santiagensis (biese, 1944) comb. nov. figs 1–5 littoridina santiagensis biese, 1944: 187–188, fig. 21 (estero dehesa, cerro manquehue, barnechea, east to santiago, chile, type locality). littoridina santiagensis–stuardo, 1961: 17. —valdovinos, 1999: 128, 2006: 90. —sielfeld, 2001: 3. heleobia santiagensis (biese, 1944). hershler &thompson, 1992: 55. — collado et al., 2011: 51, 54–56, fig. 2r–s. type material lectotype chile • santiago; 200611, mnhncl (collado et al. 2011). http://tree.bio.ed.ac.uk/software/figtree/ collado g.a. et al., systematic evaluation of cryptic freshwater snails 5 description shell. ovate-conic, dark brown, relatively thick, with about 5.5 whorls (fig. 3a–d). shell length about 3.0 mm (table 1). periostracum brown. protoconch smooth (fig. 3e), with about 1.3 whorls and 400 μm length (± 10 μm of standard deviation, n = 9). aperture ovate, slightly angled adapically, lip thickened. umbilicus imperforate or absent. operculum ovate, thin, multispiral, light brown-transparent, with eccentric nucleus (fig. 3f–g); attachment scar occupying almost half of the internal surface (fig. 3g). mantle black with a conspicuous gray band on anterior margin, head black, somewhat depigmented in center (fig. 3h–i). foot black. females. with nuchal node, white lips, tentacles grayish or black (fig. 3h–i). some specimens with a white band at base of tentacles where eyes are located. males were not observed. radula. taenioglossate (formula 3-1-3), with two marginal teeth and a lateral tooth placed on each side of the central tooth (fig. 3j–k). central tooth trapezoidal, dorsally concave, ca. 20 μm wide; basal tongue u-shaped; with 5–6 lateral cusps on each side of median cusp and 3–4 pairs of basal cusps, first basal cusps arise from tooth face and are larger than those on cutting edge. median cusp of central tooth fig. 2. shells of truncatelloidean freshwater snails observed in the present study. a–b. slender morphotype from el yeso spring (a) and lo carreño (b) assigned to potamopyrgus antipodarum (gray, 1843). c–e. thicker morphotype from el yeso spring (c), lo carreño (d) and el colorado (e) assigned to potamolithus santiagensis (biese, 1944) comb. nov. f. thicker morphotype from viña casas del maule assigned to potamolithus sp. scale bar = 1 mm. european journal of taxonomy 524: 1–15 (2019) 6 well-developed and pointed. lateral tooth with 11 cusps and median cusp well-developed and pointed. inner marginal teeth with about 30 cusps (fig. 3k). fig. 3. potamolithus santiagensis (biese, 1944) comb. nov., yeso spring, chile. a. shell imaged using sem. b–d. shell of the same specimen photographed under a stereo microscope (frontal, dorsal, lateral views). e. protoconch. f–g. opercula of two specimens (outer, inner sides, respectively). h. head of a female. i. head of another female having a nuchal node. j. anterior-central section of radular ribbon. k. central teeth. abbreviations: f = foot; h = head; l = lip; lt = left tentacle; nn = nuchal node; rt = right tentacle. scale bars: a–d = 1.0 mm; e = 250 μm; f–g = 500 μm; h–i = 0.5 mm; j = 50 μm; k = 10 μm. collado g.a. et al., systematic evaluation of cryptic freshwater snails 7 ecology potamolithus santiagensis comb. nov. is a herbivorous-detritivorous species that inhabits small water bodies like springs and small streams. distribution yeso spring in cajón del maipo, región metropolitana, central chile (biese 1947; collado et al. 2011; present study). this spring is a small tributary of the yeso river, which empties in the maipo river. the species also inhabits small irrigation canals in the región de o’higgins (lo carreño) and región del maule (el colorado). snail collections made in 2011, 2014, 2015 and 2017 at the type locality estero dehesa have not provided specimens of the species. this ecosystem has been invaded by p. antipodarum (collado 2014) and physa sp. (unpublished data). in yeso spring, p. santiagensis comb. nov. coexists with p. antipodarum and snails of the genus chilina gray, 1828. morphometric analysis the mean values of the shell variables obtained for the freshwater snails inhabiting central chile are shown in table 1. although the kruskal-wallis tests considering the six variables analyzed were statistically significant, only 23 of 60 pairwise post-hoc comparisons among native potamolithus populations and p. antipodarum were significant, providing evidence of the difficulties in distinguishing these snails (table 2). however, in the pca the potamolithus populations were well separated from p. antipodarum fig. 4. pca of potamolithus populations and potamopyrgus antipodarum (gray, 1843) collected from central chile. european journal of taxonomy 524: 1–15 (2019) 8 shell variable n sw al aw bwl spl p. santiagensis comb. nov. el yeso 26 1.7 (± 0.1) 1.3 (± 0.1) 0.9 (± 0.1) 2.1 (± 0.1) 0.6 (± 0.1) p. santiagensis comb. nov. el colorado 17 1.9 (± 0.1) 1.6 (± 0.1) 1.1 (± 0.1) 2.4 (± 0.2) 0.9 (± 0.1) p. santiagensis comb. nov. lo carreño 9 1.7 (± 0.1) 1.3 (± 0.1) 1.2 (± 0.1) 2.1 (± 0.1) 0.8 (± 0.1) potamolithus sp. viña casas del maule 6 2.0 (± 0.1) 1.6 (± 0.2) 1.1 (± 0.1) 2.4 (± 0.2) 0.7 (± 0.1) p. antipodarum el yeso 30 1.8 (± 0.2) 1.4 (± 0.1) 1.0 (± 0.1) 2.2 (± 0.2) 1.2 (± 0.1) p. antipodarum lo carreño 7 2.6 (± 0.2) 2.1 (± 0.2) 1.4 (± 0.1) 3.2 (± 0.2) 1.7 (± 0.3) table 1. average shell dimensions (± standard deviation) of six linear variables used in the comparative study of potamolithus populations and potamopyrgus antipodarum (gray, 1843) from central chile. fig. 5. bayesian tree based on coi sequences. numbers at nodes indicate posterior probability values (only given if ≥ 0.95). names in bold refer to new sequences reported in this paper. numbers following taxa refer to genbank sequences. collado g.a. et al., systematic evaluation of cryptic freshwater snails 9 in the morphometric space (fig. 4). in this analysis, the first two components accounted cumulatively for 95.27% of the variance (pc1: 79.78%; pc2: 15.49%). molecular analysis the coi phylogenetic analysis grouped the original sequences and those downloaded from genbank into two main clades, tateidae and cochliopidae, respectively, both supported by posterior probabilities (p.p.) of 1.00 (fig. 5). our original sequences clustered in the first clade, either in the genus potamolithus or potamopyrgus, both groups inferred with high node support (0.97 and 1.00 p.p., respectively). the sequences of the slender morphotype from el yeso spring and lo carreño were identified as p. antipodarum, and those of the thicker one as representatives of potamolithus (fig. 5). the snails from el colorado, viña casas del maule, puerto chico, “heleobia” sp. from uspallata and “heleobia” hatcheri from aguas negras in argentina retrieved from genbank were also recovered among potamolithus species. the coi genetic distances among tateidae species/populations from south america ranged between 0.2% and 17.0% (table 3). the genetic distances between p. santiagensis comb. nov. and snails from puerto chico were estimated at 6.7–7.2%. in the bayesian tree, they were placed in different subclades. the population from viña casas del maule grouped with “heleobia” sp. from uspallata (1.00 p.p.); they differed genetically from p. santiagensis comb. nov. by about 5.0%. on the other hand, the sequence divergence between the population from el colorado and p. santiagensis comb. nov. from yeso spring and lo carreño was low (0.0–0.6%). discussion the minute native gilled freshwater snails in chile, at present placed in the genera littoridina, heleobia or potamolithus, have had a complicated taxonomic history, as they have been ascribed to the families amnicolidae, hydrobiidae, littoridinidae and cochliopidae (e.g., pilsbry 1911; biese 1944, 1947; hershler & thompson 1992; valdovinos 1999, 2006; figueroa et al. 1999; sielfeld 2001). in argentina, the taxonomic position of some species traditionally recognized as representatives of the genus heleobia (cochliopidae) as members of the family tateidae has recently been discussed by different authors (koch et al. 2015; de lucía & gutiérrez gregoric 2017). in the present study, the tree-based barcoding analysis provided evidence to transfer littoridina/heleobia santiagensis to the genus potamolithus (tateidae). the presence of a nuchal node, also observed in females of several species of this genus (davis & pons da silva 1984; lópez armengol 1996; pons da silva & veitenheimer-mendes 2004a), absent in heleobia, is a morphological character that also supports this change, although it is not a diagnostic character of the genus (núñez 2017). the three or four pairs of basal cusps on the rachidian tooth characteristic of the tateid radula also distinguish this group from heleobia species (pons da silva & veitenheimer-mendes 2004a; cazzaniga 2011; koch et al. 2015; de lucía & gutiérrez gregoric 2017), which usually have only one pair of basal cusps, rarely two (gaillard & de castellanos 1976; pons da silva & veitenheimer-mendes 2004b; collado et al. 2016b). the bayesian analysis revealed that part of the material from lo carreño and the population from el colorado grouped with sequences of p. santiagensis comb. nov. from el yeso spring, so we assigned these snails to this species. in lo carreño we also identified the new zealand mudsnail p. antipodarum, extending its invasive range to a new hydrological basin further south of the previously known distribution in the country (collado 2014, 2016). morphometric data subjected to pca also separated potamolithus populations and the new zealand mudsnail. taking into account that the presence of the new zealand mudsnail has also been reported at several type localities of endemic species of heleobia in the región de coquimbo (biese 1944, 1947; collado 2014), where they were previously confused (collado et al. 2011), these cryptic snails could potentially also be discriminated using this simple methodology. single shell dimensions may or may not be operational, though, since a number of comparisons were statistically undifferentiable. the bayesian analysis also inferred the sequences of “heleobia” sp. uspallata and european journal of taxonomy 524: 1–15 (2019) 10 “heleobia” hatcheri from argentina as members of the genus potamolithus, representing other cases of familial and generic misidentification. in the southern cone of south america, the close phylogenetic relationships found between samples of “heleobia” hatcheri from the central-west region of argentina and potamolithus sp. from llanquihue lake in the chilean patagonia suggests that dispersal across the andes may have played a role in the distribution of these species. the same may be inferred regarding the chilean potamolithus populations from viña casas del maule and “heleobia” sp. from uspallata, argentina, and the lowland populations of p. santiagensis comb. nov., which exhibit disjunct distributions in chile. the new zealand mudsnail has been considered as a highly invasive species worldwide (son 2008; alonso & castro-díez 2012; butkus et al. 2012), potentially able to seriously affect the native snail fauna in invaded aquatic ecosystems (richards 2002). after this species arrived in the snake river drainage in north america in 1987 (bowler 1991), five native species were recorded as either “threatened” or “endangered”, in part due to the introduction of the species (richards 2002). it is possible that something similar has happened in estero dehesa considering the failure to find specimens of p. santiagensis species/population shell variable p. santiagensis (el yeso) p. antipodarum (el yeso) p. santiagensis (el colorado) potamolithus sp. (vcm) p. antipodarum (el yeso) sl<0.001 < 0.001 p. santiagensis comb. nov. (el colorado) < 0.001 1.000 potamolithus sp. (viña casas maule) 0.068 1.000 1.000 p. santiagensis comb. nov. (lo carreño) 0.379 0.236 1.000 1.000 p. antipodarum (el yeso) sw<0.001 1.000 p. santiagensis comb. nov. (el colorado) 0.001 0.003 potamolithus sp. (viña casas maule) 0.016 0.044 1.000 p. santiagensis comb. nov. (lo carreño) 1.000 1.000 0.019 0.054 p. antipodarum (el yeso) al<0.001 0.374 p. santiagensis comb. nov. (el colorado) < 0.001 < 0.001 potamolithus sp. (viña casas maule) < 0.001 0.029 1.000 p. santiagensis comb. nov. (lo carreño) 1.000 1.000 0.022 0.151 p. antipodarum (el yeso) aw<0.001 1.000 p. santiagensis comb. nov. (el colorado) < 0.001 0.008 potamolithus sp. (viña casas maule) 0.058 0.271 1.000 p. santiagensis comb. nov. (lo carreño) < 0.001 < 0.001 0.848 1.000 p. antipodarum (el yeso) bwl <0.016 0.511 p. santiagensis comb. nov. (el colorado) 0.002 0.361 potamolithus sp. (viña casas maule) 0.013 0.371 1.000 p. santiagensis comb. nov. (lo carreño) 1.000 1.000 1.000 0.872 p. antipodarum (el yeso) spl<0.001 < 0.001 p. santiagensis comb. nov. (el colorado) < 0.001 0.038 potamolithus sp. (viña casas maule) 1.000 0.004 1.000 p. santiagensis comb. nov. (lo carreño) 0.382 0.003 1.000 1.000 table 2. p-values of kruskal-wallis (in column shell variable) and pairwise post-hoc mann-whitney u-tests of six linear variables. collado g.a. et al., systematic evaluation of cryptic freshwater snails 11 comb. nov. and the current occurrence of new zealand mudsnails at this locality. although without being alarmist, perhaps a “chronicle of a death foretold” could be written for the el yeso spring population based on the noticeable abundance of new zealand mudsnails in this habitat compared to p. santiagensis comb. nov. in this context, the finding of the new populations of this species in el colorado, where the new zealand mudsnail has not been introduced, acquires real significance giving a light of hope for the preservation of the species. acknowledgments we thank the staff of the museo de ciencias naturales y arqueología profesor pedro ramírez fuentes, chillán, chile. we also thank francis miño and nicolás villalobos for logistic support in the field and cristian suárez for assistance with the sem observations. mdl is a fellow in comisión de investigaciones científicas de la provincia de buenos aires and degg is a staff reasearcher in consejo nacional de investigaciones científicas y técnicas (conicet). njc is a staff reasearcher in comisión de investigaciones científicas de la provincia de buenos aires. gac is part of the grupo de biodiversidad y cambio global, universidad del bío-bío. the authors also thank the conicytspecies/population 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 1. potamopyrgus antipodarum el yeso spring 1 0.0 2. potamopyrgus antipodarum lo carreño 1 0.0 0.0 3. “heleobia” hatcheri 17.0 17.0 0.0 4. “heleobia” sp. uspallata 14.5 14.5 8.2 0.0 5. potamolithus agapetus 15.1 15.1 11.4 9.2 0.0 6. potamolithus ribeirensis 17.0 17.0 12.1 9.6 11.0 0.0 7. potamolithus buschii 15.1 15.1 10.6 9.2 7.6 10.2 0.0 8. potamolithus elenae 14191 15.3 15.3 9.0 7.0 8.8 9.4 6.8 0.0 9. potamolithus santiagensis el yeso spring 4 15.3 15.3 9.0 5.3 10.2 10.8 10.4 9.0 0.0 10. potamolithus santiagensis el yeso spring 35 15.3 15.3 9.0 5.3 10.2 10.8 10.4 9.0 0.0 0.0 11. potamolithus santiagensis el colorado 1 15.3 15.3 8.6 5.1 10.0 10.4 10.4 8.6 0.6 0.6 0.0 12. potamolithus santiagensis el colorado 2 15.3 15.3 9.0 5.3 10.2 10.8 10.4 9.0 0.0 0.0 0.6 0.0 13. potamolithus santiagensis lo carreño 9 15.1 15.1 8.6 5.1 9.8 10.4 10.4 8.6 0.6 0.6 0.4 0.6 0.0 14. potamolithus santiagensis lo carreño 1 15.3 15.3 9.0 5.3 10.2 10.8 10.4 9.0 0.0 0.0 0.6 0.0 0.6 0.0 15. potamolithus sp. viña casas del maule 1 14.3 14.3 8.0 0.2 9.0 9.4 9.0 6.8 5.1 5.1 4.9 5.1 4.9 5.1 0.0 16. potamolithus sp. puerto chico 6 15.5 15.5 5.7 6.1 9.2 10.0 8.2 6.5 7.2 7.2 7.2 7.2 6.8 7.2 5.9 0.0 17. potamolithus sp. puerto chico 14 15.3 15.3 5.5 5.9 9.0 9.8 8.0 6.3 7.0 7.0 7.0 7.0 6.7 7.0 5.7 0.2 table 3. percent of mean sequence variation (based on p-distance) of coi data among south american species/populations belonging to the tateidae. european journal of taxonomy 524: 1–15 (2019) 12 fondequip program (no. eqm-140088) and recognize the contribution of the anonymous reviewers. this paper was supported by conicyt-fondecyt 11130697. references alonso á. & castro-díez p. 2012. the exotic aquatic mud snail potamopyrgus antipodarum (hydrobiidae, mollusca): state of the art of a worldwide invasion. aquatic sciences 74: 375–383. https://doi.org/10.1007/s00027-012-0254-7 biese w.a. 1944. revisión de los moluscos terrestres y de agua dulce provistos de concha de chile. parte i, familia amnicolidae. boletín del museo nacional de historia natural 22: 169–190. biese w.a. 1947. revisión de los moluscos terrestres y de agua dulce provistos de concha de chile. parte ii, familia amnicolidae (continuación). boletín del museo nacional de historia natural 23: 63–77. bowler p.a. 1991. the rapid spread of the freshwater hydrobiid snail potamopyrgus antipodarum (gray) in the middle snake river, southern idaho. proccedings of the desert fishes council 21: 173–182. butkus r., šidagytė e. & arbačiauskas k. 2012. two morphotypes of the new zealand mud snail potamopyrgus antipodarum (j.e. gray, 1843) (mollusca: hydrobiidae) invade lithuanian lakes. aquatic invasions 7: 211–218. https://doi.org/10.3391/ai.2012.7.2.007 cazzaniga n.j. 1980. notas sobre los hidróbidos argentinos. i (gastropoda: rissoidea). acerca de littoridina occidentalis (doering, 1884). neotropica 26 (76): 187–191. cazzaniga n.j. 1982a. notas sobre los hidróbidos argentinos. 5. conquiliometría de littoridina parchappii (d’orbigny, 1835) (gastropoda rissoidea) referida a su ciclo de vida en poblaciones australes. iheringia série zoologia 61: 97–118. cazzaniga n.j. 1982b. nota sobre los hidróbidos argentinos. ii (gastropoda: rissoidea). una littoridina del “grupo parchappii” en península valdés (chubut). revista del museo de la plata (nueva serie 13), sección zoología 129: 11–16. cazzaniga n.j. 2011. heleobia stimpson, 1865: taxonomía. in: cazzaniga n.j. 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botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany. european journal of taxonomy 383: 1–21 issn 2118-9773 https://doi.org/10.5852/ejt.2017.383 www.europeanjournaloftaxonomy.eu 2017 · holovachov o. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:8d18e64d-6e01-43cc-9a97-bd5c4d2160e1 1 belgopeltula belgica (vincx & gourbault, 1992) gen. et comb. nov. and mudwigglus micramphidium sp. nov. from the west coast of sweden, and reappraisal of the genus pseudaraeolaimus chitwood, 1951 (nematoda: araeolaimida: diplopeltidae) oleksandr holovachov department of zoology, swedish museum of natural history, box 50007, se–104 05 stockholm, sweden. email: oleksandr.holovachov@nrm.se urn:lsid:zoobank.org:author:89d30ed8-cfd2-42ef-b962-30a13f97d203 abstract. two species of the nematode family diplopeltidae are described from skagerrak. the new genus belgopeltula gen. nov. is proposed for diplopeltula belgica vincx & gourbault, 1992 and is characterised by: amphidial fovea circular in female and double-loop-shaped in male; excretory pore located at the level of cephalic setae bases; oral opening on the dorsal side of the body; pharynx subdivided into strongly muscularised fusiform corpus and weakly muscularised narrow and long postcorpus; female didelphic with antidromously reflexed ovaries; supplements absent. mudwigglus micramphidium sp. nov. is characterised by: a body of 0.6 mm long; cephalic sensilla 1.5 µm long; amphidial fovea loop-shaped, 8 µm long and 3.5 µm wide; gymnostom without cuticularised ring; tail elongate conoid, with subcylindrical distal part; terminal setae absent; spicules 15 µm long; gubernaculum present; two midventral precloacal setae. it is distinguished from m. macramphidium leduc, 2013 in having shorter amphidial fovea, shorter spicules and presence of two precloacal setae. redescription of diplopeltis cylindricauda allgén, 1932 is provided based on type material. diplopeltula minuta vitiello, 1972 is transferred to the genus mudwigglus leduc, 2013. diplopeltis cylindricauda allgén, 1932, diplopeltula laminata vitiello, 1972 and diplopeltula cassidaignensis vitiello, 1972 are transferred to the genus pseudaraeolaimus chitwood, 1951. keywords. morphology, new genus, new species, diplopeltula, taxonomy. holovachov o. 2017. belgopeltula belgica (vincx & gourbault, 1992) gen. et comb. nov. and mudwigglus micramphidium sp. nov. from the west coast of sweden, and reappraisal of the genus pseudaraeolaimus chitwood, 1951 (nematoda: araeolaimida: diplopeltidae). european journal of taxonomy 383: 1–21. https://doi.org/10.5852/ejt.2017.383 introduction the genus diplopeltula gerlach, 1950 was proposed to accommodate two new species, d. breviceps gerlach, 1950 (type species) and d. longiceps gerlach, 1950, as well as previously known species diplopeltis cylindricauda allgén, 1932 (allgén 1932; gerlach 1950). this new genus was placed in the family diplopeltidae based on the morphology of the amphidial fovea (non-spiral), gubernaculum https://doi.org/10.5852/ejt.2017.383 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:8d18e64d-6e01-43cc-9a97-bd5c4d2160e1 mailto:oleksandr.holovachov%40nrm.se?subject= http://zoobank.org/urn:lsid:zoobank.org:author:89d30ed8-cfd2-42ef-b962-30a13f97d203 https://doi.org/10.5852/ejt.2017.383 european journal of taxonomy 383: 1–21 (2017) 2 (weakly developed) and caudal glands (opening through three separate outlets). the explicit generic diagnosis was not provided by the author (gerlach 1950), however, following diagnostic characters of the genus diplopeltula can be deduced based on the identification key included in the same publication: amphid is loop shaped, with equally long ventral and dorsal branches; amphid not located on the shieldlike cuticular plate. over the years, many new species were described in diplopeltula or transferred into it from other genera (gerlach 1954, 1956; vitiello 1972; juario 1974; boucher & helléouët 1977; jensen 1978), making a total of 23 species by 1992 (vincx & gourbault 1992). some of these species did not match the original set of diagnostic characters used by gerlach (1950), e.g. in having amphidial fovea with loop branches of unequal length, or amphid located on the shield-like cuticular plate. as a result, the genus diplopeltula became very heterogeneous morphologically, making it impossible to propose concise and unambiguous generic diagnosis (vincx & gourbault 1992). this morphological diversity of the genus was discussed in detail by holovachov et al. (2009), who subdivided diplopeltula into several distinct morphological groups without taxonomic status, as well as transferred a number of species to the genera diplopeltoides gerlach, 1962 (gerlach 1962) and pararaeolaimus timm, 1961 (timm 1961). a new genus mudwigglus leduc, 2013 was proposed within the family diplopeltidae for three new species as well as one known species, which was originally described in diplopeltula (leduc 2013). and most recently, in view of new morphological data, another six species were transferred from the genus diplopeltula to the genus diplopeltoides (holovachov & boström 2017). material, collected recently during the project “systematics of swedish free-living nematodes of the orders desmodorida and araeolaimida”, provides additional information on the morphology and diversity of the species currently assigned to the genus diplopeltula in particular and of the family diplopeltidae in general. material and methods bottom sediment samples were collected in several locations in the southern part of the skagerrak and in gullmarn fjord off the west coast of sweden. all samples were collected with a bottom dredge or box corer and further sieved in the laboratory before fixation. nematodes were extracted from samples using a decanting and sieving method (smallest mesh sizes: 45 µm or 70 µm). freshwater was used during sieving to induce osmotic shock in nematodes and separate them from the substrate (giere 2009). samples were immediately fixed in 4% formaldehyde. for light microscopy, formaldehyde-preserved specimens were transferred to pure glycerine using the rapid method of seinhorst (1959) as modified by de grisse (1969). permanent nematode mounts on glass slides were prepared using the paraffin wax ring method. all curved structures were measured along the curved median line. type and other specimens are deposited in the invertebrate collections of the department of zoology, swedish museum of natural history, stockholm, sweden (smnh). type specimen of diplopeltis cylindricauda (smnh type-3435) from the invertebrate collection of the swedish museum of natural history (boström & holovachov 2017) was included in this study. abbreviations used in tables are according to hunt & palomares-ruis (2012). results phylum nematoda potts, 1932 class chromadorea inglis, 1983 order araeolaimida de coninck & schuurmans stekhoven, 1933 family diplopeltidae filipjev, 1918 genus belgopeltula gen. nov. urn:lsid:zoobank.org:act:c016d9a9-f945-4320-a549-9180f7e3cf80 type and only species belgopeltula belgica (vincx & gourbault, 1992) gen. et comb. nov. = diplopeltula belgica vincx & gourbault, 1992 http://zoobank.org/urn:lsid:zoobank.org:act:c016d9a9-f945-4320-a549-9180f7e3cf80 holovachov o., genera belgopeltula, mudwigglus and pseudaraeolaimus 3 diagnosis cuticle transversely striated; striae without ornamentation. lateral alae absent. body pores and epidermal glands absent. somatic sensilla indistinct. labial region bluntly rounded; lips fused. inner labial sensilla invisible if present. outer labial sensilla papilliform, located on the anterior surface of lips. cephalic sensilla setiform; their bases located at the base of the labial region, anterior to amphid. subcephalic and cervical sensilla, deirid and ocelli absent. amphidial fovea unispural (circular) in female and double-loop-shaped in male (starting from the porus amphidialis, fovea extends anteriorly for a short distance, makes a loop towards ventral side of the body and extends posteriorly for the full length of the amphidial fovea, makes another loop towards dorsal side of the body and extends anteriorly almost to the level of the porus amphidialis). secretory-excretory system present; renette cell located opposite to the anterior part of intestine. cuticularised secretory-excretory duct very short, opens to the exterior at the level of cephalic setae bases. oral opening shifted towards the dorsal side of the body. buccal cavity small and narrow, cheilostom short, remaining part of stoma undifferentiated; its lining is uniform with the lining of the pharynx. pharyngeal tubes absent. pharynx subdivided into anterior corpus and posterior postcorpus; corpus fusiform, muscular, with evenly distributed myofilaments; postcorpus narrow and long, weakly muscularised; pharyngeal lumen uniform in thickness along the entire pharynx length; valves absent. pharyngeal gland orifices penetrate pharyngeal lumen as the base of the stoma. postcorpus is surrounded by six large pseudocoelomocyles. female reproductive system didelphic-amphidelphic, with equally developed branches, ovaries reflexed antidromously. spermatheca present. vulva equatorial. vagina straight; pars proximalis vaginae encircled by single sphincter muscle; pars refringens vaginae and epiptygmata absent. male reproductive system diorchic, testes opposed (anterior testis outstretched, posterior testis reflexed). spicules symmetrical, arcuate; gubernaculum present. supplements, precloacal and postcloacal sensilla absent. three caudal glands present, their cells and nuclei are incaudal. spinneret present.‘‘ etymology the name “belgopeltula” combines parts of the original binomen: “belg–” comes from species name belgica, and “–opeltula” – from genus name diplopeltula. relationships the new genus is proposed to accommodate the single species diplopeltula belgica and is particularly characterised by following unique characters: amphidial fovea circular in female and double-loop-shaped in male; secretory-excretory pore located at the level of cephalic setae bases; oral opening subterminal, located on the dorsal side of the body; pharynx subdivided into strongly muscularised fusiform corpus and weakly muscularised narrow and long postcorpus; female didelphic with antidromously reflexed ovaries; male diorchic with opposed testes; preand postcloacal supplements absent. the morphology of the pharynx in belgopeltula gen. nov. needs to be clarified. in the original publication, the pharynx is described as: “muscular with an elongated terminal bulb”. the drawing shows rather cylindrical anterior part of the pharynx that rapidly widens short distance posterior to the amphideal fovea; the nerve ring is drawn immediately in front of this pharyngeal expansion, at 54% of the pharynx length. the only other known record of the same species published by fadeeva & mordukhovich (2013) describes the pharynx as follows: “pharynx terminates in a small bulb-like expansion”. nerve ring is not depicted in the figure but is said to encircle pharynx at 77% of its length. videos of the pharyngeal region of the holotype of diplopeltula belgica were examined. morphology of the pharynx and surrounding tissues in the holotype confirms well with observations of pharyngeal region in recent specimens: fusiform corpus and narrow postcorpus; large pseudocoelomocyles surrounding anterior part of postcorpus and glandular cells surrounding posterior part of it. photographs european journal of taxonomy 383: 1–21 (2017) 4 of the specimen from the sea of japan also show most of the features observed in specimens from the coast of sweden, despite it being not in the best quality: distinct fusiform corpus and narrower postcorpus; large pseudocoelomocyles surrounding postcorpus along most of its length; basal part of the pharynx surrounded by small glandular cells. the unique shape of the male amphid (starting from the porus amphidialis, fovea extends anteriorly for a short distance, makes a loop towards ventral side of the body and extends posteriorly for the full length of the amphid, makes another loop towards dorsal side of the body and extends anteriorly almost to the level of the porus amphidialis) separates belgopeltula gen. nov. from all other genera classified in the family diplopeltidae (leduc 2013; fonseca & bezerra 2014), which have unispiral or single loopshaped amphid. belgopeltula gen. nov. and mudwigglus leduc, 2013 are the only two genera in the family diplopeltidae with antidromously reflexed ovaries, while the rest of the genera are characterised by outstretched ovaries. in this respect, both genera show similarities to the family diplopeltoididae tchesunov, 1990 (tchesunov 1990) (currently placed in the order plectida gadea, 1973 (gadea 1973)). however, as discussed in leduc (2013), the morphology of female ovaries (outstretched vs reflexed) may not always be consistent in all species within large taxonomic categories, and in some cases may not have diagnostic value. to make things more complicated, the morphology of female ovaries has not been described in large number of species placed in the genus diplopeltula, including its type species d. breviceps. in having oral opening located on the dorsal side of the body, the new genus resembles the genus campylaimus cobb, 1920 (cobb 1920) and some species of the genus diplopeltula. it can be easily separated from both genera in the morphology of the amphidial fovea (circular in female and characteristic double-loop-shaped in male vs single loop in both females and males in diplopeltula and campylaimus), the female reproductive system (ovaries reflexed in belgopeltula gen. nov. vs ovaries outstretched in diplopeltula and campylaimus), the pharynx (subdivided into fusiform corpus and narrow postcorpus in belgopeltula gen. nov. vs cylindrical in diplopeltula and campylaimus) and the position of excretory pore (located at the level of cephalic setae bases in belgopeltula gen. nov. vs at the level of posterior part of pharynx or anterior part of intestine in diplopeltula and campylaimus). belgopeltula belgica (vincx & gourbault, 1992) gen. et comb. nov. figs 1–2; table 1 diagnosis (based on all known populations) belgopeltula belgica gen. et comb. nov. is characterised by a body of 0.33–0.55 mm long; cephalic sensilla setiform, 0.5–4 µm long; amphidial fovea sexually dimorphic, circular in female and doubleloop-shaped in male; oral opening located on the dorsal side of the body; secretory-excretory pore located at the level of cephalic setae bases; tail elongate-conoid; ovaries antidromously reflexed; testes opposed; spicules 16–21 µm long, curved, with weakly defined manubrium and subcylindrical shaft; gubernaculum plate-like, with caudal apophysis; supplements absent. material examined sweden: 2 ♂♂, 2 ♀♀, skagerrak off the west coast of sweden, 58°20′06″ n, 11°09′24″ e, muddy sediment from 53 m deep, 9 aug. 2011, o. holovachov leg. (smnh 161699–161701); 1 ♂, skagerrak off the west coast of sweden, 58°18′06″ n, 11°05′22″ e, mud and gravel from 40–60 m deep, 9 aug. 2011, o. holovachov leg. (smnh 161702); 1 ♂, skagerrak off the west coast of sweden, 58°22′14″ n, 11°05′00″ e, gravel, mud and algae from 30–70 m deep, 9 aug. 2011, o. holovachov leg. (smnh 161703); 3 ♂♂, 3 ♀♀, skagerrak off the west coast of sweden, 58°21′35″ n, 11°08′30″ e, fine gravel holovachov o., genera belgopeltula, mudwigglus and pseudaraeolaimus 5 fig. 1. belgopeltula belgica (vincx & gourbault, 1992) gen. et comb. nov., female entire view. scale bar = 20 µm. european journal of taxonomy 383: 1–21 (2017) 6 table 1. measurements (in µm) of belgopeltula belgica (vincx & gourbault, 1992) gen. et comb. nov., mudwigglus micramphidium sp. nov. and pseudaraeolaimus cylindricauda (allgén, 1932) comb. nov. (presented as mean ± sd and (range) where appropriate). belgopeltula belgica mudwigglus micramphidium pseudaraeolaimus cylindricauda locality north sea sea of japan skagerrak skagerrak campbell island number of specimens holotype male 1 male 4 males 7 females holotype male lectotype male body length 550 381 386 ± 26 (358–418) 358 ± 28 (326–414) 594 985 body diameter (bd) 19 21.5 18.4 ± 2.1 (16.0–21.0) 25.0 ± 3.3 (21.5–31.0) 17.0 19 pharynx length 87 98 98 ± 6 (91–104) 91 ± 3 (86–96) 78 133 tail length 120 74 80 ± 2 (79–82) 71 ± 3 (67–78) 66 93 anal body diameter (abd) 18 14 14.8 ± 1.5 (13.5–16.5) 13.1 ± 1.8 (11.0–16.5) 14.0 18 a 28.9 17.7 21.1 ± 2.1 (18.1–22.8) 14.4 ± 1.4 (13.1–16.9) 34.7 51.8 b 6.3 3.9 4.0 ± 0.1 (3.9–4.0) 4.0 ± 0.3 (3.7–4.6) 7.6 7.4 c 4.6 5.2 4.8 ± 0.3 (4.6–5.2) 5.0 ± 0.3 (4.7–5.4) 8.9 10.6 c’ 6.7 5.3 5.4 ± 0.4 (5.0–5.8) 5.5 ± 0.7 (4.7–6.7) 4.7 5.2 v or t (%) ? ? 35.7 ± 2.0 (34.2–37.1) 49.4 ± 1.8 (47.1–52.3) 48.7 59.3 labial region diameter ? ? 5.7 ± 1.0 (4.0–6.5) 6.7 ± 0.7 (6.0–8.0) 8.0 5.5 oral opening from ant. end ? ? 2.1 ± 0.6 (1.5–3.0) 2.7 ± 0.7 (2.0–3.5) apical apical cephalic setae length 4 1.5 1.0 ± 0.3 (0.5–1.5) 0.8 ± 0.3 (0.5–1.5) 1.5 7.0 amphidial fovea from ant. end 18 19.7 18.0 ± 0.9 (17.0–19.0) 19.1 ± 1.9 (16.5–22.0) 11.5 5.5 amphidial fovea length (max) 26 37 15.9 ± 2.1 (13.5–18.5) 2.6 ± 0.7 (1.5–3.5) 8.0 11.0 amphidial fovea width 4 4 4.3 ± 0.8 (3.5–5.0) 2.6 ± 0.7 (1.5–3.5) 3.5 3.5 nerve ring from ant. end 47 76 ? ? 50 95 nerve ring from anterior end as % of pharynx length 54 77 ? ? 64 71 excretory pore from ant. end subapical ? subapical subapical 66 12.5 secretory-excretory pore from ant. end as % of pharynx length subapical ? subapical subapical 85 9 vagina or spicules length 21 16.5 17.4 ± 0.8 (16.5–18.0) 4.9 ± 0.9 (4.0–6.5) 15.0 17 rectum or gubernaculum length 9 4.1 7.1 ± 0.7 (6.5–8.0) 11.0 ± 1.5 (9.0–13.5) 6.0 8.0 vagina length / bd – – – 0.2 (0.1–0.3) – – rectum or spicules / abd – – 1.1 ± 0.1 (1.1–1.2) 0.8 ± 0.1 (0.7–1.0) 1.1 0.9 holovachov o., genera belgopeltula, mudwigglus and pseudaraeolaimus 7 from 20–60 m deep, 9 aug. 2011, o. holovachov leg. (smnh 161704); 6 ♂♂, 9 ♀♀, skagerrak off the west coast of sweden, 58°7′58″ n, 11°10′05″ e, coarse gravel from 30–50 m deep, 9 aug. 2011, o. holovachov leg. (smnh 161705–161707). description adult body fusiform in female and cylindrical in male, tapering anteriorly in the pharyngeal region and posteriorly in the tail region, ventrally curved upon fixation. cuticle finely striated along entire body, striae without longitudinal incisures or any other ornamentation visible under the light microscope. somatic setae indistinct if present. labial region rounded, lips fused. inner and outer labial sensilla not observed. cephalic sensilla small setiform, equal to 0.1–0.2 labial region diameters in length, their bases are located 3–7 µm from anterior end. amphidial fovea dissimilar in shape and size between sexes (see descriptions of each sex). oral opening shifted towards the dorsal side of the body, 1.5–3.5 µm from the anterior end. stoma very small, cheilostom short, remaining part of stoma undifferentiated, its lining is uniform with the lining of the pharynx. pharynx distinctly subdivided into anterior corpus and posterior postcorpus; corpus slightly fusiform, muscular; postcorpus narrow and long. pharyngeal gland nuclei indistinct. pharyngeal gland orifices penetrate pharyngeal lumen at the base of the stoma. postcorpus is surrounded by six large pseudocoelomocytes, two cells arranged longitudinally in three body sectors: dorsal, right-subventral and left-subventral. nerve ring is located at the posterior end of the pharynx, its exact position if obscured by small glandular cells (neuron bodies?). secretory-excretory system present; renette cell located opposite to the ventral side of the anterior part of intestine; secretory-excretory pore opens to the exterior just posterior to cephalic setae bases. tail elongate-conoid. caudal glands and spinneret present. female amphidial fovea unispiral (circular). reproductive system didelphic, amphidelphic; ovary branches symmetrical, antidromously reflexed. anterior ovary situated to either right or left of intestine; posterior ovary situated to either right or left of intestine. spermathecae present, filled with round spermathozoa. vulva located at mid-body. vagina straight, pars proximalis vaginae surrounded by a sphincter muscle; pars refringens vaginae absent. intrauterine egg not seen. rectum short. male amphidial fovea double-loop-shaped: starting from the porus amphidialis, fovea extends anteriorly for a short distance, makes a loop towards ventral side of the body and extends posteriorly for the full length of the amphid, makes another loop towards dorsal side of the body and extends anteriorly almost to the level of the porus amphidialis. amphid length (longest) to width ratio is 2.7–5.2. narrow space between amphidial branches not refractive and not ornamented. reproductive system diorchic, anterior testis outstretched; posterior testis reflexed. spicules paired and symmetrical, weakly curved, with weakly defined manubrium and subcylindrical shaft. gubernaculum plate-like, with strong dorsocaudal apophysis. supplements and caudal setae indistinct or absent. remarks both published descriptions of diplopeltula belgica are each based on single male specimens (vincx & gourbault 1992; fadeeva & mordukhovich 2013); females of this species were unknown until now. recent male specimens resemble type male from the north sea (vincx & gourbault 1992) in structure of amphid, digestive (see above) and excretory systems, with the exception of the following: shorter body (358–418 µm in recent specimens vs 550 µm in type male), shorter amphid (13.5–18.5 µm in recent specimens vs 26 µm in type male), shorter cephalic setae (0.5–1.5 µm in recent specimens vs 4 µm in type male), shorter (16.5–18 µm in recent specimens vs 21 µm in type male) and less curved (weakly arcuate in recent specimens vs strongly arcuate in type male) spicules. european journal of taxonomy 383: 1–21 (2017) 8 fig. 2. belgopeltula belgica (vincx & gourbault, 1992) gen. et comb. nov., male entire view. scale bar = 20 µm. holovachov o., genera belgopeltula, mudwigglus and pseudaraeolaimus 9 specimen from the sea of japan (fadeeva & mordukhovich 2013) is closer in size to specimens from skagerrak (table 1) – the only substantial differences are the length of the amphidial fovea (13.5– 18.5 µm in recent specimens vs 37 µm in the male from the sea of japan) and less curved (weakly arcuate in recent specimens vs strongly arcuate in the male from the sea of japan) spicules. greatest discrepancies between all three populations are in the assumed position of the nerve ring and the morphology of male gonad. in the original description, the nerve ring is drawn short distance behind the amphid, at 54% of the pharynx length (vincx & gourbault 1992). male specimen from the sea of japan is described to have the nerve ring at 77% of the pharynx length (fadeeva & mordukhovich 2013), but it is not drawn on the figure. in recent specimens, the position of the nerve ring can not be identified with confidence, and further observations are required to clarify this. regarding the morphology of the male gonad, type specimen is said to have both testes outstretched (vincx & gourbault 1992), while our observations clearly show that the posterior testis in this species is reflexed posteriad. genus mudwigglus leduc, 2013 type species mudwigglus patumuka leduc, 2013 diagnosis (emended after leduc 2013) cuticle transversely striated; striae without ornamentation. lateral alae absent. body pores and epidermal glands present. somatic sensilla present. labial region bluntly rounded; lips fused. inner labial sensilla invisible if present. outer labial sensilla papilliform, located on the anterior surface of lips. cephalic sensilla setiform; their bases located at the base of the labial region, anterior to amphid. subcephalic and cervical sensilla, deirid and ocelli absent. amphidial aperture loop-shaped (inverted u-shaped). secretory-excretory system present; renette cell located opposite to the posterior part of the pharynx and anterior part of intestine. cuticularised secretory-excretory duct very short, opens to the exterior at the level of the posterior part of pharynx. oral opening apical. buccal cavity small but distinctly subdivided into three sections: cheilostom narrow tubular; gymnostom barrel-shaped or truncate conoid, with weakly cuticularised walls; stegostom short conoid, its lining is uniform with the lining of the pharynx. pharyngeal tubes absent. pharynx subdivided into anterior corpus and posterior bulbus; corpus cylindrical, muscular, with evenly distributed myofilaments; bulbus muscular; pharyngeal lumen uniform in thickness along the entire pharynx length; valves absent. dorsal gland orifice penetrates pharyngeal lumen at the base of the stoma, ventrosublateral gland orifices indistinct. dorsal gland nucleus is visible in the basal bulbus. female reproductive system didelphicamphidelphic with equally developed branches, ovaries antidromously reflexed. spermatheca absent. vulva equatorial. vagina straight or oblique; pars proximalis vaginae encircled by single sphincter muscle; pars refringens vaginae and epiptygmata absent. male reproductive system diorchic; anterior testis outstretched, posterior testis outstretched, with posteriorly reflexed terminal section of the germinal zone. spicules symmetrical, arcuate; gubernaculum present. supplements and precloacal sensilla present. setae in caudal region of males arranged in subventral and subdorsal pairs along the tail. three caudal glands present, their nuclei are incaudal. caudal glands open separately via three pores (this structure is obscure in some species). valid species mudwigglus macramphidium leduc, 2013 mudwigglus micramphidium sp. nov. mudwigglus minutus (vitiello, 1972) comb. nov. = diplopeltula minuta vitiello, 1972 mudwigglus nellyae (vincx & gourbault, 1992) european journal of taxonomy 383: 1–21 (2017) 10 = diplopeltula nellyae vincx & gourbault, 1992 mudwigglus patumuka leduc, 2013 mudwigglus plebeius leduc, 2013 nomenclatorial change diplopeltula minuta vitiello, 1972 is here transferred to the genus mudwigglus based on its close morphological affinities to m. plebeius, from which it can only be separated by features of male accessory apparatus (one or two setae in m. minutus vs one setae and two tubular supplements in m. plebeius). mudwigglus micramphidium sp. nov. urn:lsid:zoobank.org:act:c5028b8c-5860-45b6-a1d0-dbf6e1b30ed2 fig. 3; table 1 diagnosis mudwigglus micramphidium sp. nov. is characterised by a body of 0.6 mm long; cephalic sensilla 1.5 µm long; amphidial fovea an inverted u-shape, 8 µm long and 3.5 µm wide; gymnostom without cuticularised ring; tail elongate conoid, with subcylindrical distal part; terminal setae absent; spicules 15 µm long; gubernaculum present; two midventral precloacal setae. etymology the specific epithet refers to the overall similarity between the new species and m. macramphidium, as well as one of the major differences between these two species (amphid size). material examined holotype sweden: ♂, skagerrak, 58°28′12.1″–19.5″ n, 10°37′01.1″–07.1″ e, soft bottom 180–216 m deep, 10 oct. 2012, “inventering bratten” leg. (smnh type-8889). description adult male body cylindrical, tapering posteriorly in the tail region, strongly ventrally curved upon fixation. cuticle finely striated along the entire body, annules without longitudinal striation or any other ornamentation visible under the light microscope. somatic setae present, sparsely distributed over entire body, with one ventrosublateral pair in the pharyngeal region just posterior to nerve ring, and three pairs along the tail: one dorsosublateral pair short distance posterior to cloacal opening, one ventrosublateral pair at about the middle of tail length, and one sublateral pair close to tail terminus. labial region truncate conoid; lips fused. inner and outer labial sensilla not observed. cephalic sensilla small setiform, equal to 0.2 labial region diameters in length, their bases are located 5 µm from anterior end. amphidial fovea an inverted u-shape with its dorsal branch slightly longer (8 µm) than ventral branch (6.5 µm). amphidial fovea length (longest) to width ratio is 2.2. narrow space between amphidial branches not refractive and not ornamented. distinct cell with large nucleus is located behind the amphid, and is connected to amphidial fovea. stoma very small, but distinctly subdivided into three sections: cheilostom narrow tubular, 3 µm long; gymnostom truncate conoid, narrowing towards anterior end, with weakly cuticularised walls, 3 µm long and 2 µm wide, without cuticularised ring; stegostom very short conoid, its lining is uniform with the lining of the pharynx. pharynx distinctly subdivided into anterior corpus and posterior basal bulbus; corpus subcylindrical, muscular; basal bulbus ovoid, muscular. dorsal pharyngeal gland nucleus located in the basal bulbus; dorsal pharyngeal glad orifice penetrates pharyngeal lumen at the base of the stoma. ventrosublateral pharyngeal gland nuclei and orifices indistinct. nerve ring encircling http://zoobank.org/urn:lsid:zoobank.org:act:c5028b8c-5860-45b6-a1d0-dbf6e1b30ed2 holovachov o., genera belgopeltula, mudwigglus and pseudaraeolaimus 11 pharynx at posterior third of its length. secretory-excretory system present; renette cell located opposite to the ventral side of the basal bulbus and anterior part of the intestine; secretory-excretory pore opens to the exterior at level of the basal bulbus. reproductive system diorchic; anterior testis outstretched, posterior testis outstretched, with posteriorly reflexed terminal section of the germinal zone. large spermatogonia are visible. spicules paired and symmetrical, arcuate, with weakly developed manubrium fig. 3. mudwigglus micramphidium sp. nov., holotype male (smnh type-8889) entire view. scale bar = 20 µm. european journal of taxonomy 383: 1–21 (2017) 12 and subcylindrical shaft. gubernaculum present, platelike. male accessory apparatus composed of two midventral precloacal setiform sensilla. tail elongate conoid, subcylindrical distally. terminal setae absent. caudal glands and their outlets obscure. female not found. remarks mudwigglus micramphidium sp. nov. closely resembles m. macramphidium described from new zealand (leduc 2013) in general body size (table 2), shape and length of the tail and absence of gubernaculum apophysis. the new species can be separated from m. macramphidium in having shorter amphid (amphid length/width ratio 2.2 in m. micramphidium sp. nov. vs 3.5 in m. macramphidium), shorter spicules (15 µm in m. micramphidium sp. nov. vs 23–25 µm in m. macramphidium), male accessory apparatus (two precloacal setae in m. micramphidium sp. nov. vs one precloacal setae and two tubular supplements in m. macramphidium). key to species of mudwigglus leduc, 2013 see table 2 for additional diagnostic characters. 1. gubernaculum apophysis absent ...................................................................................................... 2 – gubernaculum apophysis present ..................................................................................................... 3 2. amphid length/width ratio 3.5; spicules 23–25 µm long ..............m. macramphidium leduc, 2013 – amphid length/width ratio 2.2; spicules 15 µm long ............................ m. micramphidium sp. nov. 3. tail conoid with clavate tip; c’-ratio > 3 ........................................................................................... 4 – tail cylindrical with bluntly rounded tip; c’-ratio < 3 ...................................................................... 5 4. males with one precloacal setae and two tubular supplements .................. m. plebeius leduc, 2013 – males with one or two precloacal setae, tubular supplements absent ................................................. .............................................................................................. m. minutus (vitiello, 1972) comb. nov. 5. body longer than 0.7 mm; spicules 31–37 µm; vagina oblique .............. m. patumuka leduc, 2013 – body shorter than 0.4 mm; spicules 16–23 µm; vagina perpendicular .............................................. ................................................................................................m. nellyae (vincx & gourbault, 1992) genus pseudaraeolaimus chitwood, 1951 type species pseudaraeolaimus perplexus chitwood, 1951 diagnosis (emended after fonseca & bezerra 2014) cuticle visually smooth or finely striated; striae without ornamentation. lateral alae absent. somatic sensilla present. labial region rounded; lips fused. inner and outer labial sensilla indistinct. cephalic sensilla setiform; their bases located at the base of the labial region, anterior to amphidial fovea. cervical (paramphidial) sensilla present, located at the level of the amphid and posterior to it. deirid absent. ocelli present or absent. amphidial fovea loop-shaped (inverted u-shaped). secretory-excretory system present; renette cell located opposite to the anterior part of intestine. cuticularised secretory-excretory duct very short, opens to the exterior just posterior to cephalic setae bases. oral opening apical. buccal holovachov o., genera belgopeltula, mudwigglus and pseudaraeolaimus 13 cavity small and undifferentiated, its lining is uniform with the lining of the pharynx. pharyngeal tubes absent. pharynx subdivided into anterior corpus and posterior postcorpus; corpus cylindrical, muscular, with evenly distributed myofilaments; postcorpus glandular; pharyngeal lumen uniform in thickness along the entire pharynx length; valves absent. female reproductive system didelphic-amphidelphic with equally developed branches, ovaries outstretched. vulva equatorial. male reproductive system diorchic, anterior testis outstretched, posterior testis outstretched, with posteriorly reflexed terminal section of the germinal zone. spicules symmetrical, arcuate; gubernaculum present. precloacal sensillum present. setae in caudal region of males arranged in subventral and subdorsal pairs along the tail. three caudal glands present, their nuclei are incaudal. spinneret present. valid species pseudaraeolaimus cassidaigniensis (vitiello, 1972) comb. nov. = diplopeltula cassidaigniensis vitiello, 1972 pseudaraeolaimus cylindricauda (allgén, 1932) comb. nov. = diplopeltis cylindricauda allgén, 1932 = diplopeltula cylindricauda (allgén, 1932) pseudaraeolaimus laminata (vitiello, 1972) comb. nov. = diplopeltula laminata vitiello, 1972 pseudaraeolaimus perplexus chitwood, 1951 pseudaraeolaimus ocellatus kito, 1981 table 2. selected diagnostic characters of species of the genus mudwigglus leduc, 2013 based on literature data supplemented with recent observations. characters m. patumuka m. nellyae m. plebejus m. minutus m. macramphidium m. micramphidium sp. nov. body length (mm) 0.79–0.95 0.28–0.31 0.46–0.68 0.38–0.53 0.60–0.76 0.59 tail length (µm) 52–63 20–33 39–65 44–50 54–69 66 c’ ratio 1.8–2.7 1.3–2.4 2.8–4.3 3.8–4.4 3.5–4.6 4.7 amphidial fovea length/width ratio 2.5 2.0 2.0 2.3–2.7 3.5 2.2 vagina shape oblique* straight** oblique na*** straight na spicule length (µm) 31–37 16–23 18–26 16 23–25 15 gubernaculum apophysis present present present present absent absent precloacal supplements 3–4 2 2 0 2 0 precloacal setae 1 1 1 2 1 2 terminal setae 3 3 2 2 0 0 * – at an oblique angle to the body axis ** – at a straight angle to the body axis *** – not available, females are unknown european journal of taxonomy 383: 1–21 (2017) 14 remarks according to tchesunov & miljutina (2008), main diagnostic characters of the genus pseudaraeolaimus include: terminal (apical) oral opening, amphidial fovea loop-shaped, stoma small undifferentiated, pharynx glandular in its posterior part, excretory pore close to anterior end, ovaries paired, gubernaculum without apophyses. in addition, both pseudaraeolaimus perplexus and p. ocellatus have several pairs of strongly developed cervical setae. this last feature is also known in other genera of diplopeltidae, such as araeolaimus de man, 1888, diplopeltis cobb in stiles & hassall, 1905, metaraeolaimoides de coninck, 1936 and southerniella allgén, 1932, but not in type species of the genus diplopeltula, d. breviceps. nomenclatorial changes 1) diplopeltula cassidaigniensis vitiello, 1972 is transferred to the genus pseudaraeolaimus based on presence of terminal oral opening, undifferentiated stoma, loop-shaped amphidial fovea, strongly developed cervical setae. 2) diplopeltis cylindricauda allgén, 1932 is transferred to the genus pseudaraeolaimus based on presence of terminal oral opening, undifferentiated stoma, loop-shaped amphidial fovea, strongly developed cervical setae, excretory pore close to anterior end (fig. 4). 3) diplopeltula laminata vitiello, 1972 is transferred to the genus pseudaraeolaimus based on presence of terminal oral opening, undifferentiated stoma, loop-shaped amphidial fovea, strongly developed cervical setae. pseudaraeolaimus cylindricauda (allgén, 1932) comb. nov. fig. 4; table 1 diagnosis pseudaraeolaimus cylindricauda comb. nov. is characterised by a body of 1 mm long; cephalic sensilla 7 µm long; amphidial fovea an inverted u-shape, 11 µm long and 3.5 µm wide; two pairs of cervical setae at level with amphidial fovea and 4–5 pairs at level with midpharynx; ocelli absent; tail subcylindrical; spicules 17 µm long. material examined new zealand: campbell island: lectotype ♂ (smnh type-3435). description adult male body cylindrical, tapering posteriorly in the tail region, strongly ventrally curved upon fixation. cuticle finely striated along the entire body, striae without longitudinal incisures or any other ornamentation visible under the light microscope. somatic setae (except cervical setae, see below) are indistinct. labial region truncate conoid; lips fused. inner and outer labial sensilla not observed. cephalic sensilla setiform, equal to 1.3 labial region diameters in length, their bases are located 1.5 µm from anterior end. amphidial fovea an inverted u-shape with its dorsal branch slightly longer (11 µm) than ventral branch (10.5 µm). amphidial fovea length (longest) to width ratio is 3.1. narrow space between amphidial branches not refractive and not ornamented. stoma very small, undifferentiated, its lining is uniform with the lining of the pharynx. pharynx poorly preserved, cylindrical along most of its length. pharyngeal glands and nuclei indistinct. nerve ring encircling pharynx at posterior third of its length. secretory-excretory system present; renette cell located opposite to the ventral side of the and anterior part of the intestine; secretoryexcretory pore opens to the exterior at level with the amphid. reproductive system diorchic; anterior testis outstretched, posterior testis outstretched, with posteriorly reflexed terminal section of the germinal zone. holovachov o., genera belgopeltula, mudwigglus and pseudaraeolaimus 15 spicules paired and symmetrical, arcuate, with rounded manubrium and conoid shaft. gubernaculum present, plate-like and without apophyses. male accessory apparatus composed of single midventral precloacal papilliform sensillum. tail subcylindrical. caudal glands obscure, spinneret present. female not available. key to species of pseudaraeolaimus chitwood, 1951 see table 3 for additional diagnostic characters. 1. body 2 mm long; amphidial fovea with transverse striation .............................................................. ...............................................................................................p. laminata (vitiello, 1972) comb. nov. – body shorter than 1.5 mm long; amphidial fovea without transverse striation ................................ 2 2. ocelli present ..................................................................................................p. ocellatus kito, 1981 – ocelli absent ..................................................................................................................................... 3 3. amphidial fovea a broad loop, less than 1.5 times long as it is wide ...p. perplexus chitwood, 1951 – amphidial fovea a narrow loop, more than 3 times long as it is wide ............................................. 4 fig. 4. pseudaraeolaimus cylindricauda (allgén, 1932) comb. nov. lectotype male (smnh type-3435). a. anterior end. b. posterior end. scale bar = 20 µm. european journal of taxonomy 383: 1–21 (2017) 16 4. cervical setae short, only two pairs at level with amphidial fovea .................................................... ...................................................................................p. cassidaigniensis (vitiello, 1972) comb. nov. – cervical setae long, two pairs at level with amphidial fovea and 4–5 pairs at level with midpharynx ....................................................................p. cylindricauda (allgén, 1932) comb. nov. family diplopeltidae filipjev, 1918 diagnosis (emended after fonseca & bezerra 2014) cuticle visually smooth, finely striated or coarsely annulated. lateral alae absent. somatic sensilla present. inner and outer labial sensilla papilliform. cephalic sensilla setiform; their bases located at the base of the labial region, anterior to amphidial fovea. cervical (paramphidial) sensilla present in some genera, located at the level of the amphid and posterior to it. deirid absent. ocelli present or absent. amphidial fovea unispiral or loop-shaped. secretory-excretory system present. oral opening apical or shifted either to dorsal or ventral body side. buccal cavity tubular, cylindrical or undifferentiated. teeth-like structures absent. pharynx variable in shape; pharyngeal lumen uniform in thickness along the entire pharynx length; valves absent. female reproductive system didelphic-amphidelphic or monodelphic-prodelphic. ovaries outstretched or reflexed antidromously. male reproductive system diorchic. posterior testis is either anteriorly oriented and reflexed in the anterior section (reflexed terminal section of the germinal zone) or is posteriorly directed. spicules symmetrical; gubernaculum present. precloacal sensilla present in some genera. three caudal glands present, their nuclei are incaudal. caudal glands open separately via three pores, or together via a common spinneret. valid genera araeolaimus de man, 1888 = araeolaimoides de man, 1893 = coinonema cobb, 1920 = parachromagaster allgén, 1929 belgopeltula gen. nov. table 3. selected diagnostic characters of species of the genus pseudaraeolaimus chitwood, 1951 based on literature data supplemented with recent observations. characters p. perplexus p. ocellatus p. cylindricauda p. laminata p. cassidaigniensis body length (mm) 1.08–1.20 0.77–0.93 0.99 2.09 1.36 amphidial fovea shape broad loop narrow loop narrow loop narrow loop narrow loop amphid striation absent absent absent present absent ocelli absent present absent absent absent cervical setae two pairs five pairs 6-7 pairs six pairs two pairs spicule length (µm) 14 18–19 17 30 na* gubernaculum apophysis absent absent absent present na precloacal supplements 1 1 1 ? na * – not available, specimens of particular sex are unknown holovachov o., genera belgopeltula, mudwigglus and pseudaraeolaimus 17 campylaimus cobb, 1920 cylindrolaimus de man, 1880 diplopeltis cobb in stiles & hassal, 1905 = dipeltis cobb, 1891 = discophora villot, 1875 diplopeltula gerlach, 1950 edalonema andrássy, 2007 = adelonema holovachov & sturhan, 2003 intasia tchesunov & miljutina, 2008 metaraeolaimoides de coninck, 1936 morlaxia vincx & gourbault, 1988 mudwigglus leduc, 2013 pararaeolaimus timm, 1961 pseudaraeolaimus chitwood, 1951 southerniella allgén, 1932 key to genera of diplopeltidae see table 4 for additional diagnostic characters. 1. amphidial fovea a transverse slit; cuticle with ten longitudinal ridges ....edalonema andrássy, 2007 – amphidial fovea double-loop-shaped in male, unispiral in female .................. belgopeltula gen. nov. – amphidial fovea unispiral ................................................................................................................ 2 – amphidial fovea loop-shaped ........................................................................................................... 4 2. basal bulb absent; freshwater or terrestrial ........................................cylindrolaimus de man, 1880 – basal bulb present; marine ................................................................................................................ 3 3. single anterior ovary ................................................................intasia tchesunov & miljutina, 2008 – two ovaries (anterior and posterior) ....................................................... southerniella allgén, 1932 4. oral opening shifted to the ventral body side ............................ morlaxia vincx & gourbault, 1988 – oral opening apical or shifted to the dorsal body side ..................................................................... 5 5. basal pharyngeal bulb present; ovaries antidromously reflexed ............... mudwigglus leduc, 2013 – basal pharyngeal bulb absent; ovaries outstretched ......................................................................... 6 6. lateral alae present, in some species fused with the ventral limb of the amphid ............................... ................................................................................................................... campylaimus cobb, 1920 – lateral alae absent ............................................................................................................................ 7 7. amphid in a shape of rounded loop .................................................................................................. 8 – amphid in a shape of elongated loop ............................................................................................. 10 8. anterior body region abruptly narrowing; amphid at a distance equal to more than five labial region diameters from the anterior end .............................................metaraeolaimoides de coninck, 1936 – anterior body region cylindrical or subcylindrical; amphid at a distance equal to less than two labial region diameters from the anterior end ............................................................................................. 9 9. buccal cavity cup-shaped; pronounced sexual dimorphism in the amphid size (large in males and small in females) .................................................................................. pararaeolaimus timm, 1961 european journal of taxonomy 383: 1–21 (2017) 18 ta bl e 4. s el ec te d di ag no st ic c ha ra ct er s of g en er a of t he f am ily d ip lo pe lti da e fi lip je v, 1 91 8 ba se d on l ite ra tu re d at a su pp le m en te d w ith r ec en t ob se rv at io ns (m od ifi ed a ft er t ch es un ov & m ilj ut in a 20 08 ). a ra eo la im us b el go pe ltu la g en . n ov . c yl in dr ol ai m us c am py la im us d ip lo pe lti s d ip lo pe ltu la e da lo ne m a c ut ic ul ar or na m en ta tio n ab se nt ab se nt ab se nt lo ng itu di na l s tr ia e pr es en t o r a bs en t ab se nt lo ng itu di na l s tr ia e pr es en t o r a bs en t te n lo ng itu di na l ri dg es o ra l o pe ni ng ap ic al su bd or sa l ap ic al su bd or sa l ap ic al ap ic al o r s ub do rs al ap ic al a m ph id ea l f ov ea sh ap e ro un de d lo op do ub le lo op in m al e, un is pi ra l i n fe m al e un is pi ra l el on ga te d lo op el on ga te d lo op el on ga te d lo op tr an sv er se s lit su bc ut ic ul ar p la te su pp or tin g am ph id ab se nt ab se nt ab se nt ab se nt pr es en t pr es en t o r a bs en t ab se nt o ce lli pr es en t ab se nt ab se nt ab se nt pr es en t ab se nt ab se nt c er vi ca l s et ae pr es en t ab se nt ab se nt ab se nt pr es en t ab se nt ab se nt b uc ca l c av ity co ni ca l m in ut e tu bu la r m in ut e m in ut e cy lin dr ic al tu bu la r ph ar yn x w ith s w el lin g at an te ri or ⅓ w ith fu si fo rm c or pu s an d na rr ow p os tc or pu s cy lin dr ic al cy lin dr ic al cy lin dr ic al cy lin dr ic al cy lin dr ic al fe m al e go na d di de lp hi c di de lp hi c di de lp hi c or m on od el ph ic -p ro de lp hi c di de lp hi c di de lp hi c di de lp hi c m on od el ph ic pr od el ph ic o va ri es ou ts tr et ch ed re fle xe d ou ts tr et ch ed ou ts tr et ch ed ou ts tr et ch ed ou ts tr et ch ed ou ts tr et ch ed c au da l g la nd s op en vi a sp in ne re t sp in ne re t sp in ne re t sp in ne re t sp in ne re t th re e po re s ca ud al g la nd s ab se nt in ta si a m et ar ae ol ai m oi de s m or la xi a m ud w ig gl us p ar ar ae ol ai m us p se ud ar ae ol ai m us so ut he rn ie lla c ut ic ul ar or na m en ta tio n ab se nt ab se nt ab se nt ab se nt ab se nt ab se nt ab se nt o ra l o pe ni ng ap ic al ap ic al su bv en tr al ap ic al ap ic al o r su bd or sa l ap ic al ap ic al a m ph id ea l f ov ea sh ap e un is pi ra l ro un de d lo op ro un de d lo op el on ga te d lo op ro un de d lo op el on ga te d lo op un is pi ra l su bc ut ic ul ar p la te su pp or tin g am ph id ab se nt ab se nt ab se nt ab se nt ab se nt ab se nt ab se nt o ce lli ab se nt pr es en t ab se nt ab se nt ab se nt pr es en t o r a bs en t ab se nt c er vi ca l s et ae ab se nt pr es en t ab se nt ab se nt ab se nt pr es en t ab se nt b uc ca l c av ity tu bu la r m in ut e m in ut e ba rr el -s ha pe cu psh ap ed m in ut e tu bu la r ph ar yn x w ith b as al b ul b su bc yl in dr ic al w ith b as al b ul b w ith b as al b ul b cy lin dr ic al gl an du la r p os te ri or ly w ith b as al b ul b fe m al e go na d m on od el ph ic pr od el ph ic di de lp hi c m on od el ph ic -p ro de lp hi c di de lp hi c di de lp hi c di de lp hi c di de lp hi c o va ri es ou ts tr et ch ed ou ts tr et ch ed ou ts tr et ch ed re fle xe d ou ts tr et ch ed ou ts tr et ch ed ou ts tr et ch ed c au da l g la nd s op en vi a sp in ne re t ? sp in ne re t th re e po re s or sp in ne re t th re e po re s sp in ne re t sp in ne re t holovachov o., genera belgopeltula, mudwigglus and pseudaraeolaimus 19 – buccal cavity conical; no sexual dimorphism in the amphid size .......... araeolaimus de man, 1888 10. cervical (paramphideal) setae absent; caudal glands open via three separate pores .......................... ................................................................................................................. diplopeltula gerlach, 1950 – cervical (paramphideal) setae present; caudal glands open via common spinneret ........................11 11. amphid located on a well-developed refractive subcuticular plate .................................................... .......................................................................................... diplopeltis cobb in stiles & hassal, 1905 – refractive subcuticular plate absent .......................................... pseudaraeolaimus chitwood, 1951 discussion the family diplopeltidae currently includes 14 genera (including newly described belgopeltula gen. nov.) and about 120 species (leduc 2013; fonseca & bezerra 2014). most of the genera are unequivocally defined and easily identifiable, while the genus diplopeltula is a morphological and taxonomic “hodgepodge” used to hold species that did not fit the diagnoses of other taxa of this family. as discussed in holovachov et al. (2009), several factors are at play here. first of all, the description of the type species of the genus diplopeltula, d. breviceps is brief and does not include many features that are now considered to be diagnostic (structure of the pharynx, excretory system, female reproductive system). second, many species assigned to diplopeltula are either described based on specimens of one sex, of which those based on males are prevalent, or the descriptions, even if based on specimens from both sexes, do not include data on the morphology of female reproductive system and other taxonomically important characters. the third factor mentioned in vincx & gourbault (1992) is “rather wide generic diagnosis”, or more like an absence of proper generic diagnosis in the original description (gerlach 1950) or subsequent revision of the group (gerlach 1962; vitiello 1972; vincx & gourbault 1992). last, but not least, type material of d. breviceps was not kept and therefore cannot be re-examined because s. gerlach did not make permanent slides until 1964 (f. riemann, in littera 10 apr. 2001). consequently, until d. breviceps can be recollected from the type locality and properly redescribed, the taxonomy of the genus diplopeltula will remain unclear. acknowledgments this research was supported by two grants from the swedish taxonomy initiative: “taxonomy and distribution of free-living nematodes of the order plectida in sweden” and “systematics of swedish free-living nematodes of the orders desmodorida and araeolaimida”. sampling in the skagerrak was conducted using vessels (“skagerak” and “oscar von sydow”) and facilities of the sven lovén centre for marine sciences in kristineberg. the author wishes to thank wim bert and dominick verschelde (ghent university, belgium) and natalia fadeeva (far eastern federal university, russia) for providing videos and photographs of specimens described as diplopeltula belgica. references allgén c. 1932. weitere beiträge zur kenntnis der marinen nematodenfauna der campbell-insel. nyt magazin for naturvidenskaberne 70: 97–198. boström s. & holovachov o. 2017. the swedish marine nematologist carl allgén (1886-1960): a biobibliography and his collection. zootaxa 4232: 451–490. https://doi.org/10.11646/zootaxa.4232.4.1 boucher g. & helléouët m. 1977. nématodes des sables fins infralittoraux de la pierre noire (manche occidentale). iii. araeolaimida et monhysterida. bulletin du muséum national d’histoire naturelle 297: 85–122. https://doi.org/10.11646/zootaxa.4232.4.1 european journal of taxonomy 383: 1–21 (2017) 20 cobb n.a. 1920. one hundred new nemas. (type species of 100 new genera). contributions to a science of nematology 9: 217–343. de grisse a.t. 1969. redescription ou modifications de quelques techniques utilisées dans l’étude des nématodes phytoparasitaires. mededelingen rijksfakulteit landbouwwetenschappen gent 34: 351–369. fadeeva n. & mordukhovich v. 2013. some new and poorly known nematode species from the sea of japan. deep-sea research ii 86–87: 119–123. https://doi.org/10.1016/j.dsr2.2012.07.044 fonseca g. & bezerra t.n. 2014. 7.15 order araeolaimida de coninck & schuurmans stekhoven, 1933. in: schmidt-rhaesa a. (ed) handbook of zoology. gastrotricha, cycloneuralia, gnathifera. volume 2: nematoda: 467–486. de gruyter, göttingen. gadea e. 1973. sobre la filogenia interna de los nematodos. publicaciones del instituto de biología aplicada 54: 87–92. gerlach s.a. 1950. die diplopeltiden, eine gruppe freilebender nematoden. kieler meeresforschungen 7: 138–156. gerlach s.a. 1954. nématodes marins libres des eaux souterraines littorales de tunisie et d’algérie. vie et milieu 4: 221–237. gerlach s.a. 1956. diagnosen neuer nematoden aus der kieler bucht. kieler meeresforschungen 12: 85–109. gerlach s.a. 1962. freilebende meeresnematoden von den malediven. kieler meeresforschungen 18: 81–108. giere o. 2009. meiobenthology. the microscopic motile fauna of aquatic sediments. 2nd edition. springer-verlag, berlin & heidelberg. https://doi.org/10.1007/978-3-540-68661-3 holovachov o. & boström s. 2017. three new and five known species of diplopeltoides (nematoda, diplopeltoididae) from sweden, and a revision of the genus. european journal of taxonomy 369: 1–35. https://doi.org/10.5852/ejt.2017.369 holovachov o., tandingan de ley i., mundo-ocampo m., gingold r. & de ley p. 2009. nematodes from the gulf of california. part 3. three new species of the genus diplopeltoides gerlach, 1962 (nematoda: diplopeltoididae) with overviews of the genera diplopeltis gerlach, 1962 and diplopeltula gerlach, 1950. russian journal of nematology 17: 43–57. hunt d.j & palomares-ruis e. 2012. general morphology and morphometrics of plant-parasitic nematodes. in: manzanilla-lópez r.h. & marbán-mendoza n. (eds) practical plant nematology: 25– 64. mundi-prensa, madrid. jensen p. 1978. four nematoda araeolaimida from öresund, denmark, with remarks on the oesophageal structure in aegialoalaimus. cahiers de biologie marine 19: 221–231. juario j.v. 1974. new free-living nematodes from the sublittoral zone of the german bight. veröffentlichungen des instituts für meeresforschung in bremerhaven 14: 275–303. leduc d. 2013. mudwigglus gen. n. (nematoda: diplopeltidae) from the continental slope of new zealand, with description of three new species and notes on their distribution. zootaxa 3682: 351–370. https://doi.org/10.11646/zootaxa.3682.2.8 seinhorst j.w. 1959. a rapid method for the transfer of nematodes from fixative to anhydrous glycerin. nematologica 4: 67–69. https://doi.org/10.1163/187529259x00381 https://doi.org/10.1016/j.dsr2.2012.07.044 https://doi.org/10.1007/978-3-540-68661-3 https://doi.org/10.5852/ejt.2017.369 https://doi.org/10.11646/zootaxa.3682.2.8 https://doi.org/10.1163/187529259x00381 holovachov o., genera belgopeltula, mudwigglus and pseudaraeolaimus 21 tchesunov a.v. 1990. a critical analysis of the family aegialoalaimidae (nematoda, chromadoria), trends in evolutionary development of marine nematode pharynx and a proposal of two new families. zoologicheskii zhurnal 69: 5–18. tchesunov a.v. & miljutina m.a. 2008. a new free-living nematode intasia monohystera gen. n., sp. n. (nematoda, araeolaimida, diplopeltidae) from the barents sea and the white sea, with a key to genera of diplopeltidae. russian journal of nematology 16: 33–48. timm r.w. 1961. the marine nematodes of the bay of bengal. proceedings of the pakistan academy of sciences 1: 1–88. vincx m. & gourbault n. 1992. six new species of the genus diplopeltula (nematoda: diplopeltidae) with remarks on the heterogeneity of the taxon. hydrobiologia 230: 165–178. https://doi.org/10.1007/bf00036563 vitiello p. 1972. sur quelques espèces de diplopeltula (nematoda, araeolaimida). vie et milieu 21: 535–544. manuscript received: 8 february 2017 manuscript accepted: 16 june 2017 published on: 19 december 2017 topic editor: rudy jocqué desk editor: pepe fernández printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain. https://doi.org/10.1007/bf00036563 european journal of taxonomy 487: 1–17 issn 2118-9773 https://doi.org/10.5852/ejt.2018.487 www.europeanjournaloftaxonomy.eu 2018 · savatenalinton s. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:b66e895b-c8aa-4e12-b4a7-11907e72d478 1 new genus of subfamily cypridopsinae kaufmann, 1933 (crustacea: ostracoda) from thailand sukonthip savatenalinton department of biology, plant and invertebrate taxonomy and its applications research unit, faculty of science, mahasarakham university, maha sarakham 44150, thailand. email: sukonthip.s@msu.ac.th urn:lsid:zoobank.org:author:254994a1-c8e1-4242-adf8-184661366b2d abstract. the new genus thaicypridopsis gen. nov. described here belongs to the tribe zonocypridini higuti & martens, 2012 in the subfamily cypridopsinae kaufmann, 1900. it is the first record of the  recent representatives within this tribe in southeast asia and oriental regions. thaicypridopsis gen. nov. has a number of remarkable characters that distinguish it from other genera in the subfamily: a submarginal row of distinctive tubercles on the postero-ventral part of the right valve in the interior view, a strong serrated claw g2 on the second antenna with an apical concavity, large bristles with hirsute endings and spatula-shaped apexes on all three maxillular endites and a bifurcate tip on the terminal claw of the second thoracopod. one new species, thaicypridopsis longispinosa sp. nov., is described under this new genus in the present contribution. apart from the above diagnostic characters, the new species also has long spines on the valve surface as its outstanding feature. the new taxon is morphologically compared with other related species and genera within and outside the subfamily cypridopsinae. in addition, due to the new genus belonging to a group of genera with the left valve overlapping the right valve in the subfamily, a key to the genera within this group is given. keywords. oncocypridinae, microcrustacean, taxonomy, diversity, southeast asia. savatenalinton s. 2018. new genus of subfamily cypridopsinae kaufmann, 1933 (crustacea: ostracoda) from thailand. european journal of taxonomy 487: 1–17. https://doi.org/10.5852/ejt.2018.487 introduction cypridopsinae kaufmann, 1900 which, is the most diverse subfamily in the family cyprididae baird, 1845, is presently composed of 18 genera: austrocypridopsis mckenzie, 1982; bryocypris røen, 1956; cabelodopsis higuti & martens, 2012; cavernocypris hartmann, 1964; cypridopsis brady, 1867; kapcypridopsis mckenzie, 1977; klieopsis martens et al. 1991; martenscypridopsis karanovic, 2000; neocypridopsis klie, 1940; plesiocypridopsis rome, 1965; potamocypris brady, 1870; pseodocypridopsis karanovic, 1999; sarscypridopsis mckenzie, 1977; siamopsis savatenalinton, 2017; tanganyikacypridopsis martens, 1985; thermopsis külköylüoğlu et al., 2003; tungucypridopsis victor & fernando, 1983; and zonocypris müller, 1898 (martens & savatenalinton 2011; higuti & martens 2012; savatenalinton 2017). however, the taxonomic position of the genus neocypridopsis is contentious as some authors have placed it in the subfamily cypridopsinae (e.g., martens & https://doi.org/10.5852/ejt.2018.487 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:b66e895b-c8aa-4e12-b4a7-11907e72d478 mailto:sukonthip.s%40msu.ac.th?subject= http://zoobank.org/urn:lsid:zoobank.org:author:254994a1-c8e1-4242-adf8-184661366b2d https://doi.org/10.5852/ejt.2018.487 european journal of taxonomy 487: 1–17 (2018) 2 savatenalinton 2011; higuti & martens 2012), while others (e.g., karanovic 2012) have defined it as  a genus in the subfamily oncocypridinae (family notodromadidae). there are three tribes within the cypridopsinae: potamocypridini (ghetti & mckenzie, 1981), zonocypridini higuti & martens, 2012 and cypridopsini kaufmann, 1900 (higuti & martens 2012). only the genus potamocypris belongs to the tribe potamocypridini and the members of the tribe zonocypridini are cabelodopsis and zonocypris while the tribe cypridopsini occupies the largest fraction (the remaining genera). in the cypridopsinae, six  genera  are  monospecific,  four  of  which  are  endemic  to  their  specific  area.  on  the  contrary,  cypridopsis is the dominant group with c. 77 species and is widely distributed in many zoogeographical regions (martens & savatenalinton 2011). the original descriptions of many cypridopsine species were incomplete and even questionable in some features, resulting in incorrect morphological comparisons and taxonomic positions. recently, the reexamination of zonocypris hispida (sars, 1901) based on new material from brazil revealed that this species is quite different from other cypridopsine taxa, hence a new genus, cabelodopsis, was erected for it (higuti & martens 2012). this is merely an example to show that a revision is needed in the subfamily. in addition, newly surveyed materials from poorly studied areas also harbored taxa, such as the genus siamopsis with five species, which were recently described  from thailand (savatenalinton 2017). there seems to be no doubt that there is an underestimation of the exact number of species in the cypridopsinae. thus far, six cypridopsine species have been recorded in thailand (savatenalinton & suttajit 2016; savatenalinton, 2017). in the present work, one new genus based on one new species belonging to the tribe zonocypridini of the subfamily cypridopsinae is described from thailand. material and methods sample collections were obtained from several types of water body in the central, northern and northeastern parts of thailand during the period 2005–2011. in total, 293 localities were examined: 3, 54 and 236 localities from the central, northern and northeastern parts, respectively. samples were taken with a hand net (mesh size 200 µm), preserved in 70% ethanol and later sorted using an olympus sz-pt stereo microscope. soft parts were dissected in glycerine and sealed on glass slides. valves were stored dry on micropalaeontological slides. drawings of the soft parts were made with the aid of a camera lucida. carapaces and valves were observed and illustrated using a scanning electron microscope (jeol jsm6460lv – at the faculty of science, mahasarakham university (msu)). the chaetotaxy of the limbs follows the model proposed by broodbakker & danielopol (1982), revised for the a2 by martens (1987) and for the thoracopods by meisch (2000). all type materials of the species were deposited in the natural history museum. non-type material is catalogued by its collection locality and held in the author's personal collection. institutional abbreviations msu = mahasarakham university, maha sarakham, thailand msu-zoc = ostracod collection of the natural history museum, mahasarakham university, maha sarakham, thailand abbreviations used in material citations n = northern part of thailand ne = northeastern part of thailand ne (with numbers) = ostracod samples collected from southern part of northeast thailand nee = ostracod samples collected from eastern part of northeast thailand nen = ostracod samples collected from northern part of northeast thailand th = ostracod samples collected from northern thailand savatenalinton s., new cypridopsine genus from thailand 3 abbreviations used for valves and carapace cp/cpd = carapace/carapace in dorsal view h = height of valves l = length of valves lv/lvi/lve = left valve/left valve internal view/left valve external view rv/rvi/rve = right valve/right valve internal view/right valve external view w = width of carapace abbreviations used for limbs a1  = first antenna a2 = second antenna md = mandibula mx1 = maxillula t1  = first thoracopod (maxilliped) t2 = second thoracopod (walking leg) t3 = third thoracopod (cleaning leg) cr = caudal ramus. results class ostracoda latreille, 1802 subclass podocopa g.o. sars, 1866 order podocopida g.o. sars, 1866 suborder cypridocopina jones, 1901 superfamily cypridoidea baird, 1845 family cyprididae baird, 1845 subfamily cypridopsinae kaufmann, 1900 tribe zonocypridini higuti & martens, 2012 thaicypridopsis gen. nov. urn:lsid:zoobank.org:act:9b2d9515-8ce7-4c39-b458-2f9be057599a type species thaicypridopsis longispinosa gen. et sp. nov. (here designated). etymology the genus is named after the country, thailand, where the new taxon was discovered. the name is a combination between the abbreviated country name (“thai”) and the existing generic name “cypridopsis”. diagnosis lv overlapping rv. rvi with a row of distinctive submarginal tubercles on postero-ventral part. valve surface with shallow pitted setae and/or spines. a2 with one strong serrated claw g2. largest seta on terminal segment of md-palp apically equipped with hairs. terminal segment of mx1-palp cylindrical and elongated, largest setae (bristles) on all three endites with hirsute endings and spatula-shaped apexes. seta d2 on t2 present (d1 absent), terminal claw with bifurcate ending. terminal segment of t3 with apical pincer. caudal ramus reduced, flagellum-like with triangular base. http://zoobank.org/urn:lsid:zoobank.org:act:9b2d9515-8ce7-4c39-b458-2f9be057599a european journal of taxonomy 487: 1–17 (2018) 4 differential diagnosis thaicypridopsis gen. nov. is generally similar to zonocypris and cabelodopsis due to the presence of a strong serrated claw g2 on the a2. it can be distinguished from the other genera of the subfamily cypridopsinae by the morphology of the rv in the interior view, which appears as a submarginal row of distinctive tubercles on the postero-ventral part, the presence of the robust claw on the terminal segment of the md-palp, which is apically equipped with hairs, the presence of large bristles with hirsute endings and spatula-shaped apexes on the first to the third mx1endites and the slightly bifurcated ending of the  terminal claw of the t2. these features are diagnostic characters of the new genus and the combination of these aspects has not yet been noted in any other genus of the cypridopsinae. distribution thailand (present study). thaicypridopsis longispinosa gen. et sp. nov. urn:lsid:zoobank.org:act:8c42600e-3232-40ef-bc77-765e1cc15953 figs 1–4 etymology the specific epithet ‘longispinosa’ refers to the outstanding appearance of long spines on valve surface.  this is the most prominent character of the new species and can be easily recognized from external morphology. type material holotype thailand – buriram province (ne) • ♀ (soft parts dissected in glycerine on a sealed glass slide  and valves stored dry on a micropalaeontological slide); huai jorakae mak (reservoir), muang district; 14°54΄41˝ n, 103°03΄07˝ e; 4 oct. 2010; msu-zoc.257. paratypes thailand – buriram province (ne) • 3 ♀♀ (stored as the holotype); same locality as for holotype;  4 oct. 2010; msu-zoc.258, 261 to 262 • 2 ♀♀ (stored as the holotype); kwan phayao (lake); 6 oct.  2007; msu-zoc.263 to 264 • 2 ♀♀ (carapaces stored dry on micropalaeontological slides); same locality  as for holotype; 4 oct. 2007; msu.zoc.259 to 260 • 1 ♀ (carapace stored dry on micropalaeontological  slide); kwan phayao (lake); 6 oct. 2007; msu.zoc.265 • 9 ♀♀ (in 70% ethanol); same locality as for  holotype; 4 oct. 2010; msu.zoc.266. type locality a total of 15 female specimens (1 holotype, 14 paratypes) were collected at the type locality on 4 oct. 2010 and 3 female specimens (in 70% ethanol) were collected at the type locality on 8 feb. 2011. accompanying ostracod fauna (4 oct. 2010): alicenula sp., bradleystrandesia weberi (moniez, 1892), bradleytriebella lineata (victor & fernando, 1981), cypretta sp.1, cypretta sp.2, cypridopsine sp., dentocypria chantaranothaii savatenalinton, 2017, metacypris srisumonae (savatenalinton et al., 2008), pseudocypretta maculata klie, 1932, pseudostrandesia mamarilorum (victor & fernando, 1981), pseudostrandesia striatoreticulata (klie, 1932), stenocypris sp., strandesia hornei savatenalinton & martens, 2010, strandesia kraepelini (müller, 1906); accompanying ostracod fauna (8 feb. 2011): cypretta sp.1, cypretta sp.2, cypridopsine sp., bradleystrandesia weberi, pseudocypretta maculata, pseudostrandesia mamarilorum, pseudostrandesia striatoreticulata, stenocypris sp., strandesia kraepelini. http://zoobank.org/urn:lsid:zoobank.org:act:8c42600e-3232-40ef-bc77-765e1cc15953 savatenalinton s., new cypridopsine genus from thailand 5 fig. 1. thaicypridopsis longispinosa sp. nov., ♀♀. a. lve (msu-zoc.257). b. rve (idem). c. cpd (msu-zoc.259). d. valve surface (msu-zoc.257). e. muscle scars of rv (msu-zoc.258). f. lvi (idem). g. rvi (idem). h. posterior part of lvi (idem). i. anterior part of lvi (idem). j. anterior part of rvi (idem). k–l. posterior part of rvi (idem). scale bars: a–c, f–g = 100 µm; d, l = 10 µm; e = 13 µm; h–k = 58 µm. european journal of taxonomy 487: 1–17 (2018) 6 other material examined thailand – phayao province (n) •  7  ♀♀;  locality  1;  kwan  phayao  (lake),  muang  district;  19°09΄33˝  n,  99°54΄36˝  e;  6  oct.  2007;  accompanying  ostracod  fauna: astenocypris papyracea (sars, 1903), bradleystrandesia weberi, bradleytriebella lineata, cypretta sp., cypridopsis vidua, cypridopsine sp., cypris subglobosa sowerby, 1840, dentocypria smithi savatenalinton, 2017, pseudostrandesia calapanensis (tressler, 1937), pseudostrandesia mamarilorum, stenocypris malayaica victor & fernando, 1981, strandesia kraepelini, tanycypris siamensis savatenalinton & martens, 2009; th142. – roi-et province (ne) • 19 ♀♀; locality 2; lam phayom (reservoir); 16°17΄17˝ n,  103°59΄46˝ e; 25 oct. and 20 feb. 2010; accompanying ostracod fauna:  bradleystrandesia weberi, bradleytriebella lineata, cypridopsine sp., dentocypria aequiloba, dentocypria chantaranothaii, limnocythere sp., pseudocypretta maculata, pseudostrandesia thailandensis savatenalinton & martens, 2010, stenocypris derupta vávra, 1906, strandesia kraepelini, strandesia sexpunctata klie, 1932, strandesia sp.; ne067. – chaiyaphum province (ne) • 5 ♀♀; locality 3; nong pagpung (swamp),  phu kiew district; 16°22΄27˝ n, 102°08΄02˝ e; 30 oct. 2010; accompanying ostracod fauna: candona sp., cypridopsine sp., dentocypria aequiloba, pseudocypretta maculata, stenocypris sp., strandesia kraepelini, strandesia sexpunctata; ne073. – chaiyaphum province (ne) • 2 ♀♀; locality 4; nong  sammuen  (swamp),  phu  kiew  district;  16°24΄27˝  n,  102°02΄02˝  e;  30  oct.  2010;  accompanying  ostracod fauna: metacypris srisumonae, alicenula sp., bradleystrandesia weberi, cypretta sp., dentocypria aequiloba, dentocypria chantaranothaii, pseudostrandesia mamarilorum, strandesia kraepelini, strandesia sexpunctata; ne074. – chaiyaphum province (ne)  •  2  ♀♀;  locality  5;  lam nam kam (rivulet), muang; 15°46΄37˝ n, 102°13΄18˝ e; 3 oct. 2010; accompanying ostracod  fauna: bradleystrandesia weberi, chrissia sp., cypretta sp. 2, cypridopsis vidua, cypridopsine sp., dentocypria aequiloba, dentocypria chantaranothaii, pseudocypretta maculata, pseudostrandesia gaetani savatenalinton & martens, 2010, pseudostrandesia mamarilorum, strandesia kraepelini, strandesia sexpunctata; ne007. – chaiyaphum province (ne) • 2 ♀♀; locality 6; huai pao (river),  kaset  sombun  district,  16°23΄03˝  n,  101°58΄47˝  e;  9  oct.  2007;  accompanying  ostracod  fauna:  cypretta sp., cypris subglobosa, dentocypria chantaranothaii, dentocypria smithi, dolerocypris sisaketensis savatenalinton & suttajit, 2016, pseudostrandesia mamarilorum (victor & fernando, 1981), strandesia kraepelini (müller, 1906), strandesia sexpunctata klie, 1932; th152. – ubon ratchathani province (ne) • 2 ♀♀; locality 7; kang sapue (river), phiboon mangsahan district;  15°14΄37˝ n, 105°14΄37˝ e; 24 oct. 2010; accompanying ostracod fauna: chrissia sp., cypridopsine sp., pseudocypretta maculata, pseudostrandesia calapaensis (tressler, 1937), pseudostrandesia gaetani, strandesia kraepelini, strandesia sexpunctata; ne057. – buriram province (ne) • 2 ♀♀; locality 8;  chon pratan reservoir, lampaimas district; 15°01΄05˝ n, 102°50΄29˝ e; 8 feb. 2011; accompanying  ostracod fauna: bradleystrandesia weberi, dentocypria aequiloba, pseudostrandesia thailandensis, stenocypris derupta, stenocypris sp., strandesia kraepelini; ne094. – udon thani province (ne) • 2 ♀♀; locality 9; nong kae (swamp), muang district; 17°27΄59˝ n, 102°48΄39˝ e; 31 jan. 2011;  accompanying ostracod fauna: alicenula sp., bradleystrandesia weberi, chrissia sp., cypretta sp., cypridopsis vidua, cypridopsine sp., dentocypria aequiloba, fabaeformiscandona sp., metacypris srisumonae, pseudocypretta maculata, pseudostrandesia mamarilorum, stenocypris derupta, stenocypris sp., strandesia hornei, strandesia kraepelini; nen029. – udon thani province (ne) •  2 ♀♀; locality 10; huai luang (reservoir), muang district; 17°28΄20˝ n, 102°49΄12˝ e; 31 jan. 2011;  accompanying ostracod fauna: alicenula sp., chrissia sp., cypretta sp. 1, cypretta sp. 2, dentocypria aequiloba, pseudocypretta maculata, strandesia kraepelini; nen030. – mukdahan province (ne) • 10 ♀♀; locality 11; nong lom (swamp), don tan district; 16°24΄18˝ n, 104°51΄30˝ e; 19 dec.  2011; accompanying ostracod fauna: cypridopsis vidua, cypridopsine sp., dentocypria aequiloba, dolerocypris sp., ilyocypris sp., limnocythere sp., pseudocypretta maculata, strandesia kraepelini, strandesia perakensis, strandesia sexpunctata; nee003. savatenalinton s., new cypridopsine genus from thailand 7 fig. 2. thaicypridopsis longispinosa  sp. nov., ♀♀.  a. a1 (msu-zoc.264). b. rome organ of a1 (msu-zoc.257). c. a2 (idem). d. distal segment of a2 (idem). e. claw g2 of a2 (idem). f. md-palp (msu-zoc.263). scale bars: 50 µm. european journal of taxonomy 487: 1–17 (2018) 8 diagnosis cp in lateral view subovate (length c. 1.5 times height) with maximum height situated at mid-length, anterior and posterior ends subequally rounded. cpd elliptical with maximum width situated slightly behind mid-length. lv overlapping rv all free margins. lv posterior part with double inner list, appearing well developed, but not oblique. rvi with a submarginal row of distinctive tubercles (c.18–20 tubercles) on postero-ventral part. valve surface equipped with shallow pitted, dispersedly long setae and markedly long spines. claw g2 on a2 strongly serrated with apical concavity, natatory setae long (far beyond end of terminal claw), four t-setae present in females with one seta claw-like appearance. robust claw on md-palp terminal segment with apical group of hairs. terminal segment of mx1-palp cylindrical and elongated, large bristles on three endites claw-like with hirsute endings and spatulashaped apexes, number of large bristles on first  to third endite being 1, 1, 2, respectively. t1 with  two a-setae (b-, cand d-setae absent). seta d2 on t2 present (d1 absent), terminal claw with slightly bifurcate ending. cr reduced, flagellum-like with triangular base. differential diagnosis as thaicypridopsis gen. nov. is a monospecific genus, the new species is compared with related species  in other genera within the cypridopsinae (also see discussion below). the new species is superficially  similar to cabelodopsis hispida (sars, 1901) higuti & martens, 2012 and zonocypris costata (vávra, 1897) due to the presence of a strongly serrated claw g2 on the a2 and of the tubercles on the posteroventral part of the rvi. the new species can be distinguished mainly by the robust apical claw on the terminal segment of the md-palp with hairy ending (smooth and pointed endings in c. hispida and z. costata), the morphology of the large bristles on three mx1-endites, which appear as hirsute endings and spatula-shaped apexes, and the presence of a bifurcated tip on the t2 terminal claw (smooth tip in c. hispida and z. costata). measurements (in μm) cp (n = 2), l = 528–584, w = 334–378; lv (n = 3), l = 534–588, h = 349–408; rv (n = 3), l = 507–578, h = 346–397. description female cp in lateral view. subtriangular, anterior margin rounded, posterior margin slightly wider rounded, dorsal margin strongly arched, greatest height situated at mid-length, lv overlapping rv all free margins, valve surface pitted and equipped with setae and long spines (fig. 1a–b), surface of long spines slightly serrated (fig. 1d). cpd (fig. 1c). subovate with evenly curved lateral margins, greatest width situated slightly behind midlength, posterior extremity round, anterior extremity more pointed. lvi (fig. 1f, h–i). both anterior and posterior margins subequally rounded, ventral margin sinuous at mid-length, submarginal selvage anteriorly, anterior calcified inner lamella wide with two inner lists,  posterior calcified inner lamella narrow with double inner list being well developed but not oblique,  postero-ventral part with submarginal shallow sockets or pits (conforming to tubercles on rv). rvi (fig. 1e, g, j–l). both anterior and posterior margins subequally rounded, ventral margin sinuous at mid-length, submarginal selvage anteriorly, anterior calcified inner lamella wide, posterior calcified  inner lamella narrow with two inner lists, postero-ventral part with c.18–20 prominent tubercles. savatenalinton s., new cypridopsine genus from thailand 9 fig. 3. thaicypridopsis longispinosa sp. nov., ♀♀.  a. md-coxa (msu-zoc.257). b. mx-1 (msuzoc.264). c. t1 (idem). d. endopodite of t1 (idem). scale bars: 50 µm. european journal of taxonomy 487: 1–17 (2018) 10 a1 (fig. 2a–b). seven-segmented, first segment with one short dorso-subapical seta (reaching tip of  next segment) and two long ventro-apical setae. second segment slightly wider than long, with one short dorso-apical seta (reaching half of next segment) and rome organ. third segment bearing two setae: one long dorso-apical (reaching beyond tip of penultimate segment) and one short ventro-apical setae (reaching slightly beyond half of next segment). fourth segment with two long dorsal setae and two short ventral setae (both reaching slightly beyond tip of next segment). fifth segment dorsally with two long setae, ventrally with two (one long, one short) setae, short one reaching slightly beyond tip of next segment. penultimate segment with four long apical setae. terminal segment with three (two long, one short) apical setae and long aesthetasc ya, length of aesthetasc ya c. length of last four segments, length of short seta slightly less than half of that of aesthetasc ya. fig. 4. thaicypridopsis longispinosa sp. nov., ♀♀. a. t2 (msu-zoc.257). b. distal part of t2 terminal claw (idem). c. t3 (idem). d. cr (msu-zoc.264). scale bars: a, c = 50 µm; b, d = 20 µm. savatenalinton s., new cypridopsine genus from thailand 11 a2 (fig. 2c–e). basal segment with two proximal setae and one long ventro-apical seta. exopodite with three (one long, two short) setae, long one reaching slightly beyond half of penultimate segment. first endopodal segment with five very long (reaching far beyond tips of terminal claws) and one short  natatory setae, length of the shortest seta reaching three fourth of penultimate segment, aesthetasc y long, ventro-apical seta long, extending beyond tip of terminal segment. penultimate segment undivided, distally with three serrated claws (g1–g3), g1 and g2 large, g2 strongly serrated apically with concave apex, g3 slender claw, aesthetasc y2 long (not reaching tip of terminal segment), z1–z3 setae long, z1 slightly larger than other z setae, z2 longest (beyond tip of claws g1–g3); this segment medially with two subequally long dorsal setae, four ventral setae of unequal length (t1–t4), one t seta clawlike appearance. terminal segment distally with two serrated claws (gm and gm), length of gm c. ⅔  of that of gm; medially with short g seta and ventral aesthetasc y3, length of aesthetasc y3 c. half of accompanying seta. md-palp (fig. 2f). first segment with two large setae (s1 and s2), one long and slender seta, and a short, smooth α-seta. second segment dorsally with three unequal long apical setae, shortest seta reaching tip  of next segment; ventrally with group of three long hirsute setae, one shorter hirsute seta and plumose, cone-shaped β-seta with pointed tip. penultimate segment consisting of three groups of setae: dorsally  with group of four unequal, long, subapical setae; laterally with apical γ-seta and three further apical  smooth setae, the former slightly plumose (length c. 2.4 times of terminal segment); ventrally with two subapical setae, one long (reaching beyond tip of terminal segment), one short (c. half of terminal segment). terminal segment bearing three claws and two shorter setae, largest claw appearing strongly robust with hairy ending. mx1 (fig. 3b). with two-segmented palp, basal segment of palp dorsally with group of six long, unequal apical setae; ventrally with one long subapical seta (reaching slightly beyond tip of terminal segment). terminal segment elongated (length c. 2 times of width), apically with three claws and two setae. large bristles on endites claw-like, with hirsute ending and spatula-shaped apex, number of large bristles on first to third endites being 1, 1, 2, respectively. sideways-directed bristles on first endite short, subequal  in length. t1 (fig. 3c–d). protopodite with two unequally short a-setae, b-, cand d-setae absent, distally with 12 hirsute apical setae of unequal length. endopodite weakly built palp with one very long, hirsute seta and two unequally shorter apical setae, length of shortest one less than ⅓ of long one.  t2 (fig. 4a–b). with d2 seta (d1 absent). second segment with long e-seta (length c. ⅔ of penultimate  segment). penultimate segment divided, proximal segment bearing long f-seta (reaching beyond tip of terminal segment), distal segment with short apical g-seta (reaching tip of terminal segment). terminal segment with two (one dorsally, one ventrally) apical h1 and h3 setae (length of the former less than ⅓  of claw, the latter short) and serrated claw (h2) with slightly bifurcated tip, length of h2 longer than that of penultimate segment. t3 (fig. 4c). a cleaning limb. first segment with subequal long d1, d2, dp setae. second segment with long apical e-seta (reaching half of next segment). third segment with short f-seta (not reaching tip of segment). terminal segment with an apical pincer and one reflexed subapical seta, length of the latter  slightly shorter than that of third segment. cr (fig. 4d). reduced, flagellum-like with triangular base, laterally with short subapical seta, length of  long seta c. 1.7 times of that of ramus and c. 3.8 times of that of short subapical seta. male unknown. european journal of taxonomy 487: 1–17 (2018) 12 key to genera with lv overlapping rv in the subfamily cypridopsinae 1. respiratory plate of t1 absent ................................................... austrocypridopsis mckenzie, 1982 – respiratory plate of t1 present ......................................................................................................... 2 2. claw g2 on a2 strongly serrated ...................................................................................................... 3 – claw g2 on a2 normal developed .................................................................................................... 5 3. terminal claw of t2 with bifurcate end, large setae on mx1endites with spatula-shaped apex, valve surface without circular ridges ............................................................ thaicypridopsis gen. nov. – terminal claw of t2 without bifurcate end, large setae on mx1 endites without spatula-shaped apex ................................................................................................................................................... 4 4. hinge simple and adont, valve surface without circular ridges .......................................................... ................................................................................................ cabelodopsis higuti & martens, 2012 – hinge well developed, valve surface with prominent circular ridges ......... zonocypris müller, 1898 5. pincer organ on t3 absent ........................................................................ neocypridopsis klie, 1940 – pincer organ on t3 present ............................................................................................................... 6 6. cr absent in female and male .................................................. pseudocypridopsis karanovic, 1999 – cr present in female ........................................................................................................................ 7 7. carapace elongated, marginal fused zone somewhat broad, a2 natatory setae short ........................ ...........................................................................................................cavernocypris hartmann, 1964 – carapace more ovated, marginal fused zone (if present) narrow, mostly a2 natatory setae long .... 9 8. right prehensile palp with wing-like expansion on terminal segment, a2 aesthetasc y 2-segmented ........................................................................................................... tungucypridopsis victor, 1983 – right prehensile palp without wing-like expansion on terminal segment, a2 aesthetasc y 3or 4-segmented .................................................................................................cypridopsis brady, 1867 discussion morphology and taxonomy the subfamily cypridopsinae, based on the valve overlap, is divided into two groups: the lv overlapping the rv and the rv overlapping the lv. the former group which comprises nine genera, including the new genus, is also united by the cylindrical and elongated shape of the terminal segment of the mx1palp. however, it should be emphasized that this group is not monophyletic because it includes two tribes: zonocypridini and cypridopsini (see higuti & martens 2012). the grouping standing on the valve overlap is merely suitable for the identification key as the valve overlap is primarily the most  easily recognized character. among the cypridopsine species, the new species seems to resemble the cypridopsis representatives, but its limbs’ morphology is different in many aspects, especially the strongly serrated claw g2 on  the a2, the large bristles with spatula-shaped apex on the mx1-endites and the t2 terminal claw with bifurcate tip. these features of the new species are not found in the cypridopsis. the presence of a submarginal row of the tubercles on the postero-ventral part of the rvi is a prominent character in the new genus. this aspect leads to the affinity between the new genus and a few other cypridopsine  genera, namely cabelodopsis, klieopsis, siamopsis and zonocypris. at an identical position, the first  three genera have septa-like or tiny tubercle(-like) structures (see martens et al. 1991; higuti & martens savatenalinton s., new cypridopsine genus from thailand 13 2012; savatenalinton 2017), while a submarginal row of prominent tubercles appears in the new genus and zonocypris costata (vávra, 1897) (müller 1898, as zonocypris madagascarensis). the presence of these tubercles and structures is thus a homologous character. among these five genera, thaicypridopsis gen. nov. and z. costata are likely more advanced due to the most well-developed form of tubercles. thaicypridopsis gen. nov. and z. costata have the lv overlapping the rv and the obvious tubercles occur on the smaller valve. this is similar to the case of the subfamily cyprinotinae, where the smaller valve also has tubercles (see, for example, broodbakker 1983; martens 1984; martens & wouters 1985; meisch 2000; savatenalinton & martens 2008). nevertheless, the position of tubercles is obviously different. they appear between the valve margin and the inner list in z. costata and the new genus whereas in cyprinotinae, they are on the valve margin. this indicates that these tubercles are not homologous structures in these two groups. due to the presence of tubercles on the smaller valve and complementary shallow sockets (pits) on the larger valve (see fig 1h, k–l), these structures could possibly relate to the valve closure. the septa-like or tubercle-like structures appearing on the larger valve, as in the case of klieopsis and siamopsis, would imply that these features seem to display different functions. to date, there are no such tubercles recognized on the lv in the cypridopsinae. based on the morphology of the soft parts, especially the strong serrated claw g2 of the a2, the new genus belongs to the tribe zonocypridini higuti & martens, 2012 accompanying zonocypris and cabelodopsis (see higuti & martens 2012). the strong serration on the claw g2 of the a2 is regular (without an apical cancavity), as seen in zonocypris species (see, for example, müller 1898; rome 1969; klie 1936, 1944) while in the new genus, the serrated margin is concave apically (see fig 2c, e). such irregular serration has not yet been mentioned in the subfamily cypridopsinae, except for the new genus. the claws on the terminal segment of the md-palp are smooth with pointed ends in the cypridopsine genera. the exception occurs in tanganyikacypridopsis (see martens 1985), thermopsis (see külköylüoğlu et al. 2003) and thaicypridopsis gen. nov., in which one large claw is apically equipped with a group of hairs. it should be noted that this feature should be used with caution as it would not be illustrated, even if it exists, in many older descriptions. the morphology of the mx1-endites in the new genus is remarkable as the large setae (bristles) are apically hirsute and spatula-shaped and they appear on all three endites. these striking setae are indicative and have not been recognized in any cypridopsine genera. most genera in the cypridopsinae have large bristles (zahnborsten) only on the third endite. an exceptional case was encountered in z. costata, in which large bristles were also found on the second one. however, the ending of the bristles is pointed in. z. costata, whereas the spatula-shaped ending represents the new genus. the mx1-endites are known as the feeding apparatus as they possess a number of strong apical setae. in the cyprididae species, the first two endites are equipped with monomorphic masticatory apical setae with different length, while  the heteromorphic setae only occur on the third one, typically exhibiting two large bristles together with the normal setae. the large bristles can be either smooth or serrated apically and their number can be more than two in some genera, such as oncocypris (see, for example, green 1973; rome 1962; klie 1938, 1939; savatenalinton 2015). such qualitative and/or quantitative modification strongly reinforces  the masticatory function of the mx1. in the new genus, based on the morphology of the modified setae  on the mx1-endites, which look like brushes, their function is unlikely restricted to mastication. the tip of the t2 terminal claw (h2) is slightly bifurcated in thaicypridopsis gen. nov. it can be easily recognized under medium maginification (c. 100 times). before the present study, this feature had never  been noticed in cypridopsinae, while it is one of the diagnostic characters of the genus oncocypris. the morphology of the t2 terminal claw is very slightly different between thaicypridopsis gen. nov. and oncocypris. whereas the outer margin of the claw is slightly sinuated in some oncocypris, such as o. debundshae (green 1973), o. euglypha (rome 1962) and o. rostrata (savatenalinton 2015), it is european journal of taxonomy 487: 1–17 (2018) 14 smooth in the new genus. in thaicypridopsis gen. nov., the terminal segment of the t3 possesses the pincer organ, which is identical to that of other cypridopsine genera. as the new genus shares some characters with oncocypris, this allows to further discuss the taxonomic position of the subfamily oncocypridinae which has oncocypris as the type genus. the main characters which keep the subfamily oncocypridinae in the family notodromadidae are the presence of the separate eye cups and the four or five large bristles (zahnborsten) on the third mx1 endite (for example,  see savatenalinton 2015) whereas the reduced cr seems to place oncocypridinae in the subfamily cypridopsinae. one conceivable solution of such a situation is that oncocypridinae could be the fourth tribe in the subfamily cypridopsinae. this viewpoint can also be supported by the morphology of the t2 which relates oncocypris to the cypridopsine genus, thaicypridopsis gen. nov. in this way, the family notodromadidae is no longer a polyphyletic group (see diaz & martens 2018). consequently, thaicypridopsis gen. nov. shares some valve and soft part characters mostly with cabelodopsis, zonocypris, klieopsis, siamopsis and oncocypris. the distinguishing features of the new genus mentioned above are taxonomically important differences supporting its position as a separate genus within the subfamily cypridopsinae. ecology and distribution thaicypridopsis longispinosa sp. nov. was encountered in reservoirs, lakes, swamps and rivers in the northern and northeastern parts of thailand. as large populations occur in reservoirs and lakes (huai jorakae mak, kwan phayao and lam phayom), in which the peripheral zones were densely occupied by aquatic plants, this could be its favorable habitats. the distribution of the new species was wide in the northeastern part while in the northern part it was encountered only in one locality (kwan phayao, point 1 in fig. 5), and there was no record in the central part. it should also be noted that the number of sampling localities from the northeastern part was high with 236 localities from 17 provinces whereas there were 54 localities from 13 provinces in the northern part and three localities from only one province in the central part. at present, t. longispinosa sp. nov. is a rare species in the country as it was found in fig. 5. distribution map of thaicypridopsis longispinosa sp. nov. (* = type locality; numbers = localities mentioned in “other material examined” section). savatenalinton s., new cypridopsine genus from thailand 15 12 of 293 surveyed localities (fig. 5). the discovery of t. longispinosa sp. nov. is the first record of the  recent representative of the tribe zonocypridini in southeast asia and the oriental region. according to the recent materials and the fossil evidence, there seems to be four lineages within zonocypridini with respect to the zoogeographical regions, namely afrotropical, palaearctic, neotropic and oriental lineages. this tribe could originate from the afrotropical region because it is the center of origin. in addition, zonocypris which is the most representatives of the tribe, is presently found on the african continent with high endemicity. the oriental lineage could have evolved from the african ancestor after the upper cretaceous due to the fact that the indian subcontinent was connected to east africa during the upper cretaceous. however, the current data on fossil records show the different phenomenon. the oldest fossils of zonocypris are from the cretaceous of europe, china, india and south america, while the fossils of african zonocypris, which were recovered from the miocene sediments, are more recent (mazzini 2011). nonetheless, their actual evolutionary pathways should be logically judged after extensive surveys of ostracod fossils are accomplished in these regions. acknowledgements this work was financially supported by the thailand research fund (trf, contract number mrg5380256)  and mahasarakham university, thailand (contract numbers 5405021, 5505004, 5705022 and 5808010). prof. dr. koen martens (rbins, brussels) and an anonymous reviewer provided useful comments. dr. akeapot srifa and nual-arnong nakkong (msu, thailand) offered technical assistance with the distribution map and the scanning electron micrographs, respectively. references broodbakker n.w. 1983. the genus hemicypris (crustacea, ostracoda) in the west indies. bijdragen tot de dierkunde 53 (1): 135–157. http://www.repository.naturalis.nl/document/547844 broodbakker n.w. & danielopol d.l. 1982. the chaetotaxy of cypridacea (crustacea, ostracoda) limbs: proposals for a descriptive model. bijdragen tot de dierkunde 52 (2): 103–120. http://www.repository.naturalis.nl/document/547666 diaz a. & martens k. 2018. on argentodromas bellanella gen. nov., sp. nov. 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(eds) süßwasserfauna von mitteleuropa. 8 (3): 1–522. spektrum akademischer verlag, heidelberg/ berlin. müller g.w. 1898. ergebnisse einer zoologischen forschungsreise in madagaskar und ost-afrika 1889– 1895 von dr. a. voeltzkow: die ostracoden. abhandlungen der senckenbergischen naturforschenden gesellschaft 21: 255–296. pinto r.l., rocha c.e.f. & martens k. 2005. on new terrestrial ostracods (crustacea, ostracoda) from brazil, primarily from são paulo state. zoological journal of the linnean society 145: 145–173. https://doi.org/10.1111/j.1096-3642.2005.00185.x rome d.r. 1962. exploration hydrobiologique du lac tanganyika (1946–1947) 3. ostracodes. royal belgian institute of natural sciences, brussels. rome d.r. 1969. contribution à l’étude des eaux douces de l’ennedi: 5. crustacés, ostracodes. bulletin de l’ institut fondamentale d’afrique noire, series a 31: 1074–1104. savatenalinton s. 2015. on three new species of non-marine ostracods (crustacea: ostracoda) from northeast thailand. zootaxa 3914 (3): 275–300. https://doi.org/10.11646/zootaxa.3914.3.3 savatenalinton s. 2017. siamopsis gen. nov. and five new species of  the subfamily cypridopsinae  kaufmann, 1900 (crustacea: ostracoda) from thailand. european journal of taxonomy 384: 1–39. https://doi.org/10.5852/ejt.2017.384 savatenalinton s. & martens k. 2008. redescription of hemicypris mizunoi okubo, 1990 (crustacea, ostracoda) from thailand, with a reassessment of the validity of the genera hemicypris and heterocypris. bulletin van het koninklijk belgisch instituut voor natuurwetenschappen, biologie 78: 17–27. https://doi.org/10.1023/a:1026308514466 https://doi.org/10.1007/bf00025786 https://doi.org/10.1111/j.1463-6409.1985.tb00192.x http://dx.doi.org/10.1111/j.1096-3642.2005.00185.x https://doi.org/10.11646/zootaxa.3914.3.3 https://doi.org/10.5852/ejt.2017.384 savatenalinton s., new cypridopsine genus from thailand 17 savatenalinton s. & martens k. 2013. on callistocypris thailandensis sp. nov. (ostracoda, crustacea) from thailand. zootaxa 3686 (5): 578–586. https://doi.org/10.11646/zootaxa.3686.5.6 savatenalinton s. & suttajit m. 2016. a checklist of recent non-marine ostracods (crustacea: ostracoda) from thailand, including descriptions of two new species. zootaxa 4067 (1): 001–034. https://doi.org/10.11646/zootaxa.4067.1.1 manuscript received: 10 august 2018 manuscript accepted: 12 november 2018 published on: 17 december 2018 topic editor: rudy jocqué desk editor: eva-maria levermann printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; meise botanic garden, belgium;  royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain; zoological research museum alexander koenig, bonn, germany. https://doi.org/10.11646/zootaxa.3686.5.6 https://doi.org/10.11646/zootaxa.4067.1.1 european journal of taxonomy 287: 1–20 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2017.287 www.europeanjournaloftaxonomy.eu 2017 · prigent s.r. et al. this work is licensed under a creative commons attribution 3.0 license. d n a l i b r a r y o f l i f e , r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:41d67a95-553f-4a8e-bc50-33baa1fa5b0a 1 connecting systematic and ecological studies using dna barcoding in a population survey of drosophilidae (diptera) from mt oku (cameroon) stéphane r. prigent 1, arnaud suwalski 2 & michel veuille 3,* institut systématique evolution biodiversité (isyeb) umr 7205 cnrs-mnhn-upmc-ephe, psl university, paris, france. * corresponding author: veuille@mnhn.fr 1 email: sprigent@mnhn.fr 2 email: suwalski@mnhn.fr 1 urn:lsid:zoobank.org:author:79a08f29-38da-4bd9-8f22-fe718d5fdd8d 2 urn:lsid:zoobank.org:author:e053ba9e-6371-4461-ae9d-fd5c823b9f2d 3 urn:lsid:zoobank.org:author:784ec85c-4d84-49dc-98f3-5ce406fc2ac4 abstract. the characters used in taxonomy to describe new species cannot always be used to identify species in population surveys involving large samples. we used dna barcoding to validate the taxonomic status of the morphospecies used in an ecological study involving 11 000 individual african drosophilids which had been determined without dissection. some taxonomic information had been lost by not discriminating between rare species or by mistakenly splitting a morphologically variable species into two groups. however, the original ecological dataset provided a reliable picture of species diversity and the conclusions based on the original dataset are still supported by the molecular data. keywords. barcode, drosophila, zaprionus, sophophora, cameroon. pringent s.r., suwalski a. & veuille m. 2017. connecting systematic and ecological studies using dna barcoding in a population survey of drosophilidae (diptera) from mt oku (cameroon). european journal of taxonomy 287: 1–20. http://dx.doi.org/10.5852/ejt.2017.287 introduction reliable procedures to evaluate species diversity are required given the global loss of biodiversity and its consequences for environmental policies. however, environmental studies generally need datasets involving samples containing large numbers of individuals in which species cannot always be determined using the wealth of information provided by taxonomists when describing new taxa, since their methods cannot routinely be applied to a large number of specimens. this occurs frequently in insects, for example in beetles (oliver & beattie 1996), mayflies (bauernfeind & moog 2000) and chironomids (carew et al. 2013), in which species often need to be distinguished using cryptic characters like male genitalia and internal parts observed through dissection. this is especially challenging in tropical environments http://dx.doi.org/10.5852/ejt.2017.287 www.europeanjournaloftaxonomy.eu https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:41d67a95-553f-4a8e-bc50-33baa1fa5b0a mailto:veuille%40mnhn.fr?subject= mailto:sprigent%40mnhn.fr?subject= mailto:suwalski%40mnhn.fr?subject= http://zoobank.org/urn:lsid:zoobank.org:author:79a08f29-38da-4bd9-8f22-fe718d5fdd8d http://zoobank.org/urn:lsid:zoobank.org:author:e053ba9e-6371-4461-ae9d-fd5c823b9f2d http://zoobank.org/urn:lsid:zoobank.org:author:784ec85c-4d84-49dc-98f3-5ce406fc2ac4 http://dx.doi.org/10.5852/ejt.2017.287 european journal of taxonomy 287: 1–20 (2017) 2 in which many species remain to be described. as a compromise, environmental studies often use operational taxonomic units (otu) at a lower resolution than the species level, and assume they are sufficiently informative for analysis. these otus retain biological meaning as long as each consists of a monophyletic unit containing species sharing morphological, physiological and behavioral characters. the accepted risk is a loss of resolution of the study, since two related species can diverge in a number of characters. this approximation cannot, however, be assessed as long as the ability of the researcher to recognize species has not been evaluated using molecular analysis, inter-specific crosses or examination of museum types. a useful tool to clarify this point is dna barcoding, which was designed as a reliable technique for identifying species (hebert et al. 2003; monaghan et al. 2005; janzen et al. 2009; lukhtanov et al. 2009) and detect cryptic species (hebert et al. 2004; yassin et al. 2008). it may also be used for biodiversity assessment in groups where taxonomic information is incomplete or difficult to use (blaxter et al. 2005; passmore et al. 2006; frezal & leblois 2008) or for large inventories (telfer et al. 2015). here we apply dna barcoding to a sample derived from a study carried out on a drosophilid community from the central african montane forest of mt oku in cameroon (prigent et al. 2013). drosophilids, an important model in evolutionary biology (powell 1997; ashburner et al. 2004; markow & o’grady 2005), are also considered a useful marker in ecosystem dynamics studies (parsons 1991; davis & jones 1994; mata et al. 2008). they cover a large diversity of habitats, and species assemblages vary according to environmental changes in time and space (krijger & sevenster 2001; avondet et al. 2003; tidon 2006; prigent et al. 2013). standardized traps can be used to collect them efficiently in large numbers. the study took place from october 2008 to october 2009, in the montane forest of mt oku, on the cameroon volcanic line (prigent et al. 2013). about 11 000 specimens were caught using banana traps over a year at different altitudes between 2200 m and 2800 m, and preserved in 70% ethanol until further determination. drosophilid identification involves a large number of characters including relative size and shape of body parts, color pattern, wing venation, chaetotaxy and form of the male genitalia. the male genitalia are, in fact, the main structure used to discriminate between taxa. these minute organs usually need to be dissected and are often difficult to examine in ethanol-fixed individuals. also, in a large survey it is not logistically feasible to dissect every specimen. moreover, several hundred species are known in africa (537 valid species according to the taxodros database; www.taxodros.uzh.ch) and species were described with varying precision. old descriptions often lack detail (coquillett 1902; adams 1905; séguy 1933). some revisions of african taxa include new descriptions for formerly described species within a group (tsacas 1972, 1980a), but no comprehensive comparative survey is available for the complete african fauna. the most important works providing a key were carried out on the drosophilid faunas of ivory coast (burla 1954b), south africa (tsacas 1990) and malawi (chassagnard et al. 1997). identification remains difficult, since drosophila includes several species complexes, each containing morphologically similar species (tsacas 1980b; tsacas & lachaise 1981; tsacas & chassagnard 1994; tsacas 2002). the material was therefore sorted on the basis of visible characters. morphospecies (i.e., otus) were then defined as groups that could be separated from other groups by their external morphology. determining females is a real challenge given the prevalent use of male genitalia as taxonomic criteria. females might be expected to share characters with conspecific males. however, coloration patterns are sexually dimorphic in many species and in several species from the very speciose montium subgroup (or group, see da lage et al. 2007), females are dimorphic for abdominal color patterns. sorting the specimens resulted in 62 recognizable morphospecies that were assigned, where possible, to known species according to published descriptions of external morphology (prigent et al. 2013). sample size http://www.taxodros.uzh.ch prigent s.r. et al., barcoding cameroon drosophilidae 3 was highly variable across species. one of them, zaprionus vittiger coquillett, 1901, made up 81% of the catches, whereas many other species were rare. the spatial and temporal distribution of dominant species was a clear illustration of the high dependence of the drosophilid community on climatic factors. it was therefore essential to assess the taxonomic reliability of this study. using dna barcoding confirmed the correct determination of dominant species. we detected some problems in the correct identification of cryptic species and in the allocation of females to the right species in groups of related species. the delimitation of species remained uncertain in some species due to low divergence between them. mitochondrial introgression has been reported in some species and thus cannot be excluded in our sample. the conclusions of the environmental study are not modified, given the co-occurrence and low abundance of the species involved in these uncertainties. dna barcoding actually helped in clarifying temporal or distributional patterns of cryptic species. it should allow comparison with further studies. material and methods we used specimens collected in the framework of the ecological study carried out on mount oku (prigent et al. 2013) along with an additional sample, involving the same morphospecies, collected at koh kesoten, a locality close to the sampling place. we extracted dna from 125 individuals using the qiagen® dna extraction kit. whole photographs of the individuals were taken under a leica m165c binocular stereo microscope with coupled numerical imaging, and the legs, wings and genitalia were kept apart as vouchers. the rest of the body of the flies was used for dna extraction. forty additional extractions were made from one leg, allowing us to preserve the rest of the specimen as a voucher. all of the material will be deposited at the muséum national d’histoire naturelle (mnhn), paris, when fully identified. amplification and sequencing were carried out using already published primer sequences for coi (bouiges et al. 2013). barcoding was applied to both males and females for each morphospecies, when they were available. each sequence was blasted against the ncbi sequence database. all available drosophilid coi sequences were downloaded (more than 3700 sequences, including more than 600 species mostly from drosophila and closely affiliated genera), aligned with our data and analyzed using mega5. the molecular phylogeny of drosophila has been thoroughly studied using a number of markers (e.g., pelandakis & solignac 1993; remsen & o’grady 2002; da lage et al. 2007; van der linde et al. 2010), including whole genomes (drosophila 12 genomes consortium 2007). these studies were searching deep nodes in order to identify the main groups in this very large genus. their results were of relatively little help to us. there were two reasons for this. first, there was little overlap between the species used in these studies and those from our sample, and thus they could not be used as barcode standards. second, the purpose of a taxonomic study like ours is not to identify deep nodes but shallow ones. this is a technical issue. hajibabaei et al. (2007) addressed the question of the best method being used depending on the kind of study being carried out. these authors stated that “the analysis of dna barcoding data is usually performed by a clustering method, such as distance-based neighbor-joining (nj)”, whereas the construction of phylogenetic trees is carried out “by using optimality criteria such as maximum likelihood, maximum parsimony, or bayesian analysis”. austerlitz et al. (2009) compared several phylogenetic and classification methods in analyses using known experimental datasets or data obtained from simulations using known parameter values. they confirmed the good performance of methods aimed at identifying closely related groups, such as nearest neighbor and neighbor-joining. for this reason, we analyzed our data using neighbor-joining, but we also ran a maximum-likelihood analysis on the same data, using the same software (mega5) in order to assess congruency between two methods that are widely divergent in their principles of analysis. we used a kimura-2-p substitution model for neighbor-joining, and a gtr + g + i model for maximum likelihood. european journal of taxonomy 287: 1–20 (2017) 4 results sequencing success we obtained 111 sequences out of 165 specimens (by body or leg dna extraction). our success in dna amplification and sequencing was very heterogeneous across samples, but was not linked to time, seasonality or sample size (fig. 1). we obtained a coi sequence for 52 of the 62 morphospecies (table 1). six of the ten unsuccessful assays were otus represented by only one individual. both male and female specimens were successfully sequenced in 40 morphospecies. only one sex was available in seven other morphospecies (prigent et al. 2013). in the five remaining morphospecies only one sex was successfully sequenced, although both sexes were available. fig. 1. percent success and failure in obtaining a coi sequence from specimens. a. total number of flies in the sample. b. date of field collection (month indicated in lowercase roman numerals). n = number of specimens used. prigent s.r. et al., barcoding cameroon drosophilidae 5 ta bl e 1. l is t o f t he m or ph os pe ci es re co gn iz ed in p ri ge nt e t a l. (2 01 3) a nd th ei r t ax on om ic in te rp re ta tio n ba se d on d n a b ar co de a na ly si s. c la ss ifi ca tio n: th e po si tio n of th e ta xo n in th e cl as si fic at io n of d ro so ph ili da e; th e lo ic ia na a nd th e nu tr ita s pe ci es c om pl ex es w er e es ta bl is he d by t sa ca s (2 00 2) a nd ts ac as & c ha ss ag na rd ( 19 94 ), re sp ec tiv el y. t he a da m si , b ra ch yt ar sa a nd d ya ra m an ka na s pe ci es g ro up s ar e no t fo rm al ly d efi ne d; t he y ar e si m pl y pu ta tiv e gr ou ps a cc or di ng t o pr ig en t et a l. (2 01 3) . m or ph os pe ci es : th e id en tifi ca tio n of t he t ax on g iv en i n ta bl e 1 of p ri ge nt e t al . ( 20 13 ). c lo se st re la tiv e: a s de te rm in ed b y bl as tin g ag ai ns t t he n c b i da ta ba se . p er ce nt s im ila ri ty ( % s im il. ): w ith th e cl os es t r el at iv e. r el at ed ne ss : w ith th e cl os es t si st er ta xo n or s is te r g ro up e ith er fr om th e sa m pl e or fr om th e n c b i d at ab as e un de r n ei gh bo rjo in in g an al ys is ; i n pa re nt he se s th e be st b oo ts tr ap s co re ob se rv ed o ve r s ev er al a na ly se s. c o i b ar co de : g en b an k ac ce ss io n nu m be rs . n .a . = n ot a va ila bl e. [c on tin ue d on n ex t 3 p ag es ] c la ss ifi ca ti on m or ph os pe ci es c lo se st r el at iv e % s im il. r el at ed ne ss (b oo ts tr ap v al ue ) c o i ba rc od e g en us d ro so ph ila su bg en us d or si lo ph a 1 d . b us ck ii d . b us ck ii 10 0 d . b us ck ii (1 00 ) k x 77 10 77 –7 8 su bg en us d ro so ph ila im m ig ra ns g ro up 2 d . i m m ig ra ns d . i m m ig ra ns 10 0 d . i m m ig ra ns (1 00 ) k x 77 10 79 lo ic ia na c om pl ex 3 d . a llo ch ro a d . f al le ni 90 sc ap to dr os op hi la (4 5) k x 77 10 80 –8 1 4 d . a ff . a llo ch ro a n. a. n. a. n. a. n. a. nu tr ita c om pl ex 5 d . a ff . n ut ri ta d . e re ct a 90 “d ya ra m an ka na g ro up ” (3 4) k x 77 10 82 un gr ou pe d 6 d . a ff . n iti da d . m un da 91 qu in ar ia g ro up (5 0) k x 77 10 83 –8 4 7 d ro so ph ila s p. 1 sc ap to m yz a fr us tu lif er a 90 d . f ra bu ru (3 5) k x 77 10 85 –8 6 “a da m si g ro up ” 8 d . a da m si d . p un ct at on er vo sa 92 d . p un ct at on er vo sa (8 8) k x 77 10 87 –8 8 9 d . a ff . a da m si d . p un ct at on er vo sa 92 sa m e as 8 k x 77 10 89 –9 0 10 d . a ca nt ho m er a d . m oj av en si s 89 h aw ai ia n d ro so ph ila (3 5) k x 77 10 91 –9 2 “b ra ch yt ar sa g ro up ” 11 d . a ff . b ra ch yt ar sa d . n ig el la 91 “b ra ch yt ar sa g ro up ” (8 9) k x 77 10 93 12 d ro so ph ila s p. 2 n. a. n. a. n. a. n. a. 13 d ro so ph ila s p. 3 n. a. n. a. n. a. n. a. european journal of taxonomy 287: 1–20 (2017) 6 c la ss ifi ca ti on m or ph os pe ci es c lo se st r el at iv e % s im il. r el at ed ne ss (b oo ts tr ap v al ue ) c o i ba rc od e 14 d ro so ph ila s p. 4 d . s p jf g -2 01 0 91 “b ra ch yt ar sa g ro up ” (8 9) k x 77 10 94 –9 5 15 d ro so ph ila s p. 5 d . h al ea ka la e 89 “b ra ch yt ar sa g ro up ” (8 9) k x 77 10 96 –9 7 “d ya ra m an ka na g ro up ” 16 d . d ya ra m an ka na le uc op he ng a sp . h w c -2 01 2l 90 “d ya ra m an ka na g ro up ” (2 1) k x 77 10 98 –9 9 17 d . a ff . d ya ra m an ka na s p. 1 n. a. n. a. n. a. n. a. 18 d . a ff . d ya ra m an ka na s p. 2 le uc op he ng a sp . h w c -2 01 2f 88 “d ya ra m an ka na g ro up ” (2 1) k x 77 11 00 su bg en us s op ho ph or a de nt is si m a gr ou p 19 a d . l am ot te i ( ♂ ) d . e re ct a 92 de nt is si m a gr ou p (7 9) k x 77 11 01 19 b d . l am ot te i ( ♀ ) d . s ub pu lc hr el la 93 de nt is si m a gr ou p (7 9) k x 77 11 02 20 d . a ff . m at ile i d . b oc ki 89 de nt is si m a gr ou p (7 9) k x 77 11 03 fim a gr ou p 21 d . a ff . k ul an go n. a. n. a. n. a. n. a. 22 d . m ic ro ra lis d . m ay ri 91 fim a gr ou p (1 00 ) k x 77 11 04 –0 5 23 d . a ff . m ic ro ra lis d . m ay ri 91 sa m e as 2 2 k x 77 11 06 –0 7 m el an og as te r gr ou p m el an og as te r su bg ro up 24 a d . e re ct a (♂ ) d . e re ct a 99 d . e re ct a (1 00 ) k x 77 11 08 24 b d . e re ct a (♀ ) d . y ak ub a / d . s an to m ea 10 0 sa m e as 2 7 k x 77 11 09 25 d . a ff . o re na n. a. n. a. n. a. n. a. 26 a d . t ei ss ie ri (♂ ) d . y ak ub a / d . s an to m ea 99 d . t ei ss ie ri (8 1) k x 77 11 11 26 b d . t ei ss ie ri (♀ ) d . y ak ub a / d . s an to m ea 10 0 sa m e as 2 7 k x 77 11 10 27 d . y ak ub a d . y ak ub a / d . s an to m ea 10 0 d . y ak ub a (9 5) k x 77 11 12 –1 3 m on tiu m s ub gr ou p 28 a d . b ak ou e (♂ ) d . n ik an an u 96 m on tiu m s ub gr ou p (9 9) k x 77 11 14 28 b d . b ak ou e (♀ ) d . s uz uk ii 92 sa m e as 1 9a k x 77 11 15 29 a d . b oc qu et i ( ♂ ) d . b ur la i 98 ne ar d . b ur la i ( 99 ) k x 77 11 16 29 b d . b oc qu et i ( ♀ ) d . s im ul an s 92 de nt is si m a gr ou p (7 9) k x 77 11 17 prigent s.r. et al., barcoding cameroon drosophilidae 7 c la ss ifi ca ti on m or ph os pe ci es c lo se st r el at iv e % s im il. r el at ed ne ss (b oo ts tr ap v al ue ) c o i ba rc od e 30 a d . a ff . b oc qu et i ( ♂ ) d . b ur la i 98 sa m e as 2 9a k x 77 11 18 30 b d . a ff . b oc qu et i ( ♀ ) d . t sa ca si 97 sa m e as 3 4a k x 77 11 19 31 d . b ur la i d . b ur la i 99 d . b ur la i ( 99 ) k x 77 11 20 32 a d . a ff . c ha uv ac ae s p. 1 (♂ ) d . b ur la i 99 sa m e as 3 1 k x 77 11 21 32 b d . a ff . c ha uv ac ae s p. 1 (♀ ) d . b ur la i 98 sa m e as 2 9a k x 77 11 22 33 d . a ff . c ha uv ac ae s p. 2 (♂ ) d . b ur la i 10 0 sa m e as 3 1 k x 77 11 23 34 a d . a ff . m eg ap yg a sp . 1 (♂ , ♀ 1 ) d . t sa ca si 97 m on tiu m s ub gr ou p (9 9) k x 77 11 24 –2 5 34 b d . a ff . m eg ap yg a sp . 1 (♀ 2 ) d . t sa ca si 96 d . d ip la ca nt ha (3 2) k x 77 11 26 35 a d . a ff . m eg ap yg a sp . 2 (♂ ) d . t sa ca si 97 sa m e as 3 4a k x 77 11 27 35 b d . a ff . m eg ap yg a sp . 2 (♀ ) d . n ik an an u 96 sa m e as 2 8a k x 77 11 28 36 d . n ik an an u d . n ik an an u 10 0 d . n ik an an u (1 00 ) k x 77 11 29 37 a d . s eg uy i ( ♂ ) d . t sa ca si 96 m on tiu m s ub gr ou p (9 9) k x 77 11 30 37 b d . s eg uy i ( ♀ ) d . t sa ca si 97 sa m e as 3 4a k x 77 11 31 g en us l is so ce ph al a 38 l. a ff . d io la d . p ar ab ip ec tin at a 91 so ph op ho ra (5 7) k x 77 11 32 g en us m ic ro dr os op hi la 39 m . a ff . m am ar u z. d av id i 91 w ith in z ap ri on us (4 6) k x 77 11 33 g en us s ca pt od ro so ph ila 40 s. la tif as ci ae fo rm is st eg an a sp . t l -2 01 2 90 d . a llo ch ro a (4 5) k x 77 11 34 41 s. a ff . n ic ol ae s p. 1 p ar as te ga na fe m or at a 90 p ho rt ic a so bo do (1 1) k x 77 11 35 –3 6 42 s. a ff . n ic ol ae s p. 2 p ho rt ic a sp . 1 h c -2 01 4 89 p ho rt ic a so bo do (7 ) k x 77 11 37 –3 8 43 sc ap to dr os op hi la s p. 1 d . t ra pe zi fr on s 88 p ho rt ic a so bo do (1 1) k x 77 11 39 –4 0 44 sc ap to dr os op hi la s p. 2 p ho rt ic a sp . 1 h c -2 01 4 89 sa m e as 4 2 k x 77 11 41 –4 2 45 sc ap to dr os op hi la s p. 3 p ho rt ic a fo lii se ta / d . j am bu lin a 91 d . i m m ig ra ns (7 6) k x 77 11 43 g en us s ca pt om yz a 46 sc ap to m yz a sp . 1 n. a. n. a. n. a. n. a. european journal of taxonomy 287: 1–20 (2017) 8 c la ss ifi ca ti on m or ph os pe ci es c lo se st r el at iv e % s im il. r el at ed ne ss (b oo ts tr ap v al ue ) c o i ba rc od e g en us z ap ri on us ar m at us g ro up 47 a z. s pi ne us (♂ ) z. c am pe st ri s 95 z. c am pe st ri s (9 9) k x 77 11 44 47 b z. s pi ne us (♀ ) z. li to s 93 w ith in z ap ri on us (4 6) k x 77 11 45 in er m is g ro up 48 a z. b ad yi (♂ ) z. d av id i 91 w ith in z ap ri on us (4 6) k x 77 11 47 48 b z. b ad yi (♀ ) z. c am pe st ri s 91 sa m e as 5 0a k x 77 11 46 49 z. in er m is n. a. n. a. n. a. n. a. 50 a z. m om or tic us (♂ ) z. c am pe st ri s 91 w ith in z ap ri on us (4 6) k x 77 11 49 50 b z. m om or tic us (♀ ) z. ta ro nu s 91 w ith in z ap ri on us (4 6) k x 77 11 48 51 a z. tu be rc ul at us z. tu be rc ul at us 99 z. tu be rc ul at us c om pl ex (9 9) k x 77 11 50 –5 1, -5 5 51 b z. tu be rc ul at us z. s ep so id es 10 0 z. s ep so id es c om pl ex (9 9) k x 77 11 53 –5 4, -5 6– 58 51 c z. tu be rc ul at us z. m as ca ri en si s 99 z. m as ca ri en si s (9 9) k x 77 11 52 , 59 ne gl ec tu s gr ou p 52 z. n eg le ct us n. a. n. a. n. a. n. a. vi tti ge r gr ou p 53 z. a ff . c am er ou ne ns is n. a. n. a. n. a. n. a. 54 a z. d av id i z. d av id i 99 z. d av id i ( 99 ) k x 77 11 60 , 62 –6 3 54 b z. d av id i z. p ro xi m us 98 z. p ro xi m us (9 9) k x 77 11 61 , 64 –6 5 55 z. in di an us z. a fr ic an us 98 z. a fr ic an us (8 6) k x 77 11 66 –6 8 56 z. k or ol eu z. c am er ou ne ns is 95 z. c am er ou ne ns is (8 7) k x 77 11 69 –7 0 57 z. a ff . l ac ha is ei z. la ch ai se i 99 z. la ch ai se i ( 99 ) k x 77 11 71 –7 2 58 z. o rn at us z. m eg al or ch is 99 z. o rn at us (9 9) k x 77 11 73 –7 4 59 z. a ff . o rn at us z. ta ro nu s 99 sa m e as 6 0a k x 77 11 75 –7 6 60 a z. ta ro nu s z. ta ro nu s 99 z. ta ro nu s (9 9) k x 77 11 78 60 b z. ta ro nu s z. s pi ni pi lu s 95 z. v itt ig er (6 6) k x 77 11 77 61 a z. v itt ig er z. s pi ni pi lu s 99 z. v itt ig er (1 00 ) k x 77 11 79 –8 3, -8 5 61 b z. v itt ig er z. s pi ni pi lu s 95 sa m e as 6 0b k x 77 11 84 62 z. a ff . v itt ig er z. s pi ni pi lu s 99 sa m e as 6 1a k x 77 11 86 –8 7 prigent s.r. et al., barcoding cameroon drosophilidae 9 species identification using dna databases all coi sequences were blasted against the ncbi sequence database (table 1). percent divergence from the closest sequence ranged from 0 to 12% (fig. 2). the distribution of matches was bimodal, with a first peak around 0–1% divergence and a second one around 8–9%, meaning that some of our taxa were present in the database, whereas others belonged to groups which were virtually absent from it. our identification of species (table 1) was correct with 100% sequence similarity for drosophila busckii coquillett, 1901, d. immigrans sturtevant, 1921, d. yakuba burla, 1954 and d. nikananu burla, 1954; and with 99% sequence similarity for d. erecta tsacas & lachaise, 1974, d. burlai tsacas & lachaise, 1974, zaprionus tuberculatus malloch, 1932, z. davidi chassagnard & tsacas, 1993, z. ornatus séguy, 1933, z. taronus chassagnard & tsacas, 1993 and z. vittiger. note that z. megalorchis chassagnard & tsacas, 1993 is a synonym for z. ornatus, and z. spinipilus chassagnard & mcevey, 1992 is a synonym for z. vittiger (yassin & david 2010; yassin et al. 2010). some of our morphospecies were actually a pool of several cryptic species, as shown by the fact that some individuals matched the reference sequence, while some others were substantially divergent. this occurred in the z. tuberculatus and z. davidi morphospecies. some specimens of z. tuberculatus matched z. sepsoides duda, 1939 (100% similarity) or z. mascariensis tsacas & david, 1975 (99% fig. 2. percent divergence of the morphospecies dna barcode from the closest neighbor found in the barcode database. european journal of taxonomy 287: 1–20 (2017) 10 similarity), two related species known to be hardly distinguishable from z. tuberculatus (tsacas et al. 1977). half of the sequences of the z. davidi morphospecies matched z. proximus collart, 1937 (98% similarity), suggesting the presence of a second species. our results also suggested that d. aff. chauvacae sp. 1 and sp. 2 are in fact d. burlai (99–100% similarity), that z. indianus gupta, 1970 could be z. africanus yassin & david, 2008 (98% similarity), that z. aff. lachaisei is z. lachaisei yassin & david, 2010 (99% similarity), that z. aff. ornatus is z. taronus (99% similarity), and that z. aff. vittiger is z. vittiger (99% similarity). misidentification was observed for females of the melanogaster subgroup. while males of this subgroup are easily determined using genitalia, females are morphologically very similar to each other and their identification requires dissection (yassin & orgogozo 2013). introgression of mitochondrial dna across species boundaries has been documented between several related species, including d. yakuba, d. teissieri tsacas, 1971 and d. santomea lachaise & harry, 2000 (llopart et al. 2005; bachtrog et al. 2006). in this group, females thought to belong to d. erecta and d. teissieri matched the d. yakuba sequence (100% similarity) and the sequence of the d. teissieri male matched that of its sister species d. yakuba (99% similarity). the differences in abdominal color pattern used to distinguish morphospecies overlapped the polymorphism of d. yakuba and were thus useless for species identification. in all remaining morphospecies, similarity with available data was so low that it was irrelevant for this study. for example, the best match of drosophila dyaramankana burla, 1954, a drosophiline, was with the steganine leucophenga sp. (90% similarity), and drosophila sp. 5 matched several hawaiian drosophila (89%). overall, two thirds of our morphospecies corresponded to poorly represented taxa in the coi database, either in terms of species or in terms of higher taxonomic units. phylogenetic relatedness a neighbor joining tree using the whole dataset (not shown) proved to be unreliable for deep nodes, since major groups from the known phylogeny of drosophilids were disrupted and showed incorrect connections. this was expected given the relatively poor information provided by the short barcode fragment, which was designed for identifying purposes by searching the closest neighbors of unidentified taxa. moreover, the genus drosophila is a very large taxonomic ensemble which is fragmented into smaller “groups” and “subgroups”, involving some endemism. in addition, some distinct genera, including zaprionus, are actually internal branches of the drosophila tree. most groups are not represented in africa, and a reliable topology of drosophilids can hardly be inferred from a geographically limited sample. a neighbor-joining tree was constructed for the species of zaprionus and microdrosophila aff. mamaru (burla, 1954) (fig. 3) and a second for the sophophora lineage of drosophila and lissocephala aff. diola tsacas & lachaise, 1979 (fig. 4). a maximum-likelihood tree was run on the same datasets. a remarkable result is that in each analysis, the two trees were fully congruent for all nodes supported by a 50% bootstrap value or more. the two series of bootstrap values are shown on figs 3 and 4 (on the neighbor-joining tree), except for one case where the two values were very close, although above and below this threshold: 48% and 52%, respectively. below we present and discuss the results obtained for each group of taxa separately. the most abundant genus in the mt oku forest is zaprionus (fig. 3), of which we found 16 morphospecies. coi sequences were obtained for 13 of them. nine of them belonged to the vittiger species group. the z. indianus morphospecies departed from z. indianus and was related to z. africanus, although with a node supported only by an 84–86% bootstrap value in phylogenetic analysis (fig. 3). the z. davidi morphospecies was split into two different clusters, as seen above, one of them being the true z. davidi, the other being closer to z. proximus. zaprionus aff. ornatus clustered with z. taronus, but was prigent s.r. et al., barcoding cameroon drosophilidae 11 fig. 3. phylogenetic analysis of the genus zaprionus and microdrosophila aff. mamaru (burla, 1954). this tree is the neighbor-joining tree. the maximum likelihood tree gives the same topology. nodes with a bootstrap value lower than 50% were merged. bootstrap values were calculated over 1000 repeats. above nodes: bootstrap values for maximum likelihood using a gtr + g + i model. below nodes: bootstrap values for neighbor-joining using the kimura-2p distance. european journal of taxonomy 287: 1–20 (2017) 12 fig. 4. phylogenetic analysis of the subgenus sophophora and lissocephala aff. diola tsacas & lachaise, 1979. conventions as for fig. 3. prigent s.r. et al., barcoding cameroon drosophilidae 13 not closely related to the reference sequence for z. ornatus. zaprionus aff. lachaisei matched the true z. lachaisei. two specimens misidentified as z. taronus and z. vittiger clustered together (“zaprionus sp. 1”) and matched the z. aff. vittiger sequence of bouiges et al. (2013). the z. aff. vittiger morphospecies of prigent et al. (2013) branched within sequences from z. vittiger, suggesting they belong to the same species. our two sequences for z. koroleu burla, 1954 made up a new lineage related to z. camerounensis chassagnard & tsacas, 1993 and z. nigranus yassin et al., 2008. this taxon, however, is different from z. beninensis chassagnard & tsacas, 1993, which is considered a synonym of z. koroleu (yassin & david 2010). z. tuberculatus belongs to a closely related group of species within the inermis group. our sequences for z. tuberculatus were split into three clusters corresponding to z. tuberculatus, z. sepsoides and z. mascariensis (fig. 3). this was expected, since our ecological study made no attempt to distinguish these three species, which can only be recognized by dissection. our results indicate that the three of them were present in the study. the three remaining morphospecies, z. spineus tsacas & chassagnard, 1990, z. momorticus graber, 1957 and z. badyi burla, 1954, gave no consistent results. the sequences for male and female for each of them were separate in the tree, suggesting that each individual belonged to a different species. only two sequences clustering together indicated that the dark male of the z. momorticus morphospecies belonged to the same species as the yellowish female of the z. badyi morphospecies. moreover, our two sequences for the z. badyi morphospecies lie outside the tuberculatus cluster, in which the database reference for z. badyi is found. for the sophophora subgenus of drosophila, morphospecies from mt oku clustered into four distinct lineages (fig. 4). they correspond to the melanogaster subgroup, the montium subgroup (both in the melanogaster group), and to two lineages for which no coi sequence has hitherto been available: the dentissima group and the fima group. the two morphospecies recognized for the fima group probably belong to the same species, d. microralis tsacas, 1981. in the dentissima group two morphospecies were recognized, d. lamottei tsacas, 1981 and d. aff. matilei tsacas, 1975. two females that we thought belonged to the montium subgroup were actually members of the dentissima group. the three d. melanogaster subgroup species from the mt oku forest, d. erecta, d. teissieri and d. yakuba, were correctly identified. we used the sequences of the montium subgroup found in databases and found that african species cluster in a sub-section of the tree (data not shown), also including the sequences of this group from mt oku. the d. burlai, d. aff. chauvacae sp. 1 and d. aff. chauvacae sp. 2 morphospecies clustered with the d. burlai database sequence (fig. 4). another cluster contained d. bocqueti tsacas & lachaise, 1974 and an undetermined morphospecies. the d. aff. megapyga sp. 1 and d. aff. megapyga sp. 2 morphospecies made up a third cluster. the d. seguyi smart, 1945 and d. bakoue tsacas & lachaise, 1974 morphospecies made up a fourth cluster. six morphospecies were represented by both males and females. in five morphospecies, all females formed clades apart from the males we identified as conspecific. thus, their determination was erroneous. of two females sequenced for the d. aff. megapyga sp. 1 morphospecies, one was wrong and the other correct. thus, females of similar morphology belonged to different species, confirming that the females of this group are difficult to determine. among the remaining species, identification was correct in d. immigrans and d. busckii, but unsettled in other species, since sequences did not generally cluster with available sequences. we did find some unexpected associations where otus clustered with supposed unrelated taxa. these were those between microdrosophila aff. mamaru and the genus zaprionus (fig. 3) and between lissocephala aff. diola and the fima group of the subgenus sophophora (fig. 4). however, the bootstrap values were always very low (40–60%). thus, these results may be artefacts resulting from the lack of appropriate barcode references and from the low resolving power of dna barcodes when divergence is high. european journal of taxonomy 287: 1–20 (2017) 14 the remaining morphospecies confirmed this conclusion. the closest relatives in the tree were connected with very low support. for example, d. acanthomera tsacas, 2001 clustered with a group of hawaiian species with a support of only 35%. none of the scaptodrosophila morphospecies clustered with a species of this genus which had already been sequenced except for scaptodrosophila sp. 3. this otu grouped either with s. riverata singh & gupta, 1977 (45%) or with d. immigrans (76%). at such a low level of discrimination and as a result of homoplasy, branching patterns and bootstrap values above the specific level are highly dependent on the size and on the composition of the sample of species used to construct the tree. they are unreliable and do not deserve further consideration. consequences for the mt oku ecological survey our ecological survey included 62 morphospecies (prigent et al. 2013). barcodes were obtained for 52 of them. the clustering of the specimens by their barcode suggests that nine morphospecies were duplicates of another species in the sample, as a probable result of color polymorphism within biological species. this does not substantially affect the conclusions of the survey, since the two incorrectly split subsamples occurred together in all cases. moreover, such cases did not generally involve numerically important species. the fusion of d. aff. adamsi with d. adamsi wheeler, 1959 and that of d. aff. microralis with d. microralis add up to a sample size of 20 individuals. similarly, the z. aff. vittiger sample size represents only 1.1% of the z. vittiger catches. the most important changes occurred within the montium subgroup. pooling the d. aff. chauvacae sp. 1, d. aff. chauvacae sp. 2 and d. burlai morphospecies made up a total of 101 individuals. in the same way, pooling d. aff. megapyga sp. 1 and d. aff. megapyga sp. 2 made up 77 individuals. in view of the total collection of 10 839 specimens, these changes had minor consequences for the statistics of abundance. the coi barcode also showed that some morphospecies actually included several species. this was expected in some well-documented cases, for which external morphology is known to be insufficient to distinguish closely related species. dna barcoding confirms the presence of two species in each of the z. davidi, z. vittiger and z. taronus morphospecies, and of three species in the z. tuberculatus morphospecies. this was an accepted lack of precision of the ecological survey, since these species were too numerous to allow for the dissection of all specimens. barcoding a stratified sample of morphospecies allowed us to check hypotheses as to underlying species assemblages in species complexes. the ecological distribution of the z. tuberculatus morphospecies (which is made up of three biological species) had shown two distinct peaks in altitudinal distribution with an absence of flies at intermediate elevations (prigent et al. 2013). we wondered whether this was due to habitat heterogeneity or to heterogeneity in the specific composition of the samples, given our inability to discriminate the three biological species by external morphology. the sequenced specimens from an elevation of 2300 m belonged to the three species; those from 2700 m belonged to two of them. this suggests that the gap in their distribution probably results from habitat heterogeneity, rather than from taxonomic confusion. the z. indianus morphospecies can similarly include three species in this geographical area. it shows a bimodal distribution around the year. a specimen collected in october belongs to the same species as two specimens collected in april. thus, barcoding does not suggest that distribution heterogeneity results from species heterogeneity, even though appropriate sampling would be necessary to reach a firm conclusion. finally, barcoding allowed us to validate the determination of 43 morphospecies. nine cryptic species had to be added to this count (table 1), making up a total number of 52 barcoded species. prigent s.r. et al., barcoding cameroon drosophilidae 15 discussion this study took place on the cameroon volcanic line, which belongs to an endangered biodiversity hotspot (myers et al. 2000). this concentration of species has been interpreted as resulting from successive cycles of extension and regression of the forest having taken place during the quaternary (maley 1996; plana 2004). during dry periods montane forests were restricted to montane refuges and to gallery forests. during rainy periods they could extend more widely and connect with each other to make up a continuous forest. drosophilids are an important component of this rich biodiversity. this study, together with an earlier one (prigent et al. 2013), was designed to record the response of drosophilid species to physical factors contributing to climate variation, including changes in habitat according to season and altitude. in this study the validity of the recognized morphospecies was tested using barcoding. consistent with the principle of dna barcoding, the study was very informative when closely related reference sequences were present in dna banks and almost intractable when an entire natural group of species was not represented. this occurred on several occasions, since african drosophila have not been as well studied as those from most other areas. overall the classification of our morphospecies was correct in the zaprionus genus and the sophophora subgenus of drosophila. it was also correct for the two cosmopolitan species, d. busckii and d. immigrans. morphospecies classification was more challenging in the drosophila subgenus and in other genera, probably due to the fact that none of these species were included in the barcode database, and due to a lack of information on related taxa. four taxa from our study were not represented in the database. the fima group belongs to the subgenus sophophora but its branching position was uncertain due to a low bootstrap value. similarly, the dentissima group was associated with the melanogaster subgroup (although with a low bootstrap value of 40–41%, fig. 4) even though the dentissima group is considered to be distinct from the melanogaster group (tsacas 1980). this result is, however, in agreement with the proposition of raising the ananassae and montium species subgroups to species groups (da lage et al. 2007). a barcode in the genus lissocephala was sequenced for the first time and branched within the subgenus sophophora, an unexpected observation needing confirmation, since this genus is generally given a basal position in drosophilinae based on morphology (throckmorton 1975) or molecular data (harry et al. 1996, 1998; yassin 2013). a species which we identified as belonging to the genus microdrosophila, based on burla’s (1954b) key to drosophilids from ivory coast, branches in the genus zaprionus, an unexpected result since the morphology of this morphospecies differs from that of zaprionus, which is very homogeneous across species. the lack of definition of barcoding in some drosophilid genera may result from the relatively poor record of african drosophilids in databases, a requirement for such studies (davison et al. 2009). thus, the lack of similar works constitutes an extrinsic limitation of our study. since drosophila taxonomy relies heavily upon male genitalia, and since most species are sexually dimorphic in color patterns, our results illustrate the poor correspondence between males and females across a number of species. several members of the melanogaster group show dimorphic females with black and yellow forms (burla 1954a), presumably evolved through sexual selection (yassin et al. 2016). furthermore, females from different species of the montium subgroup are practically indistinguishable. hence, some females thought to belong to the montium subgroup eventually appeared to actually belong to the dentissima group. the main conclusion of this study is that dna barcoding is a great help in making a link between taxonomic and ecological studies. this provides a means to assess our ability to identify species despite the methodological constraints brought about by the quantitative requirements of ecological studies. the results of our former ecological study (prigent et al. 2013) were practically unaffected by the fact that nine morphospecies represented excessive splitting, and that nine true species had not been recognized. the misidentified species happened to be rare ones. as a rule, abundant species were correctly determined. in three cases we had consciously pooled groups of two or three related species european journal of taxonomy 287: 1–20 (2017) 16 under a common heading (“zaprionus tuberculatus”, “z. indianus” and “z. davidi”), since these taxa could be distinguished only through dissection, and 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botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands. http://dx.doi.org/10.1371/journal.pone.0057158 http://dx.doi.org/10.1371/journal.pone.0057158 http://dx.doi.org/10.1111/j.1471-8286.2007.02020.x http://dx.doi.org/10.1038/ncomms10400 european journal of taxonomy 285: 1–20 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2017.285 www.europeanjournaloftaxonomy.eu 2017 · veron g. et al. this work is licensed under a creative commons attribution 3.0 license. d n a l i b r a r y o f l i f e , r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:eb6af6d6-36e7-4677-87e8-d72c42a273af 1 molecular systematics and biogeography of the hemigalinae civets (mammalia, carnivora) géraldine veron 1,*, céline bonillo 2, alexandre hassanin 3 & andrew p. jennings 4 1,3 institut de systématique, evolution, biodiversité, umr 7205 isyeb, cnrs mnhn upmc ephe, muséum national d’histoire naturelle, sorbonne universités, cp 51, 57 rue cuvier, 75231 paris cedex 05, france. 2 service de systématique moléculaire, ums cnrs 2700, muséum national d’histoire naturelle, cp 26, 57 rue cuvier, 75231 paris cedex 05, france. 4 small carnivores – research and conservation, 83 st. lawrence street, portland, 04101 maine, usa. * corresponding author: veron@mnhn.fr 2 email: bonillo@mnhn.fr 3 email: hassanin@mnhn.fr 4 email: smallcarnivores@yahoo.com 1 urn:lsid:zoobank.org:author:de2a6a51-29f0-4a5e-a60b-c61ad5e1842e 2 urn:lsid:zoobank.org:author:7333d242-0714-41d7-b2db-6804f8064b13 3 urn:lsid:zoobank.org:author:0dcc3e08-b2ba-4a2c-ada5-1a256f24daa1 4 urn:lsid:zoobank.org:author:3978ac9e-2c27-4b68-99da-038b502afda8 abstract. due to the difficulty in obtaining samples, the systematics of the hemigalinae civets has not been fully resolved. the aim of this study was to clarify the relationships of the species and the intraspecific diversity within this subfamily, and to explore the environmental factors that might have affected its evolution. using two mitochondrial and two nuclear markers, we confirmed that the hemigalinae comprises owston’s civet, the otter civet, hose’s civet and the banded civet, but also the sulawesi palm civet (formerly included in the paradoxurinae). our study showed that the banded and owston’s civets are sister species, and suggested that hose’s civet is sister to these two. within the banded civet, we observed a high divergence between individuals from the mentawai islands and those from sumatra and borneo (while the latter two were not strongly divergent), likely due to the deep sea channel between the mentawai islands and sumatra. unexpectedly, the sumatran and peninsular malaysian individuals were not closely related, despite the fact that these two regions have repeatedly been connected during the last glaciations. no high polymorphism was found within owston’s civet, although three groups were obtained: southern china, northern vietnam and central vietnam, which might be related to pleistocene climatic fluctuations. keywords. southeast asia, phylogeography, civets, viverridae, molecular systematics. veron g., bonillo c., hassanin a. & jennings a.p. 2017. molecular systematics and biogeography of the hemigalinae civets (mammalia, carnivora). european journal of taxonomy 285: 1–20. http://dx.doi.org/10.5852/ ejt.2017.285 http://dx.doi.org/10.5852/ejt.2017.285 www.europeanjournaloftaxonomy.eu https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:eb6af6d6-36e7-4677-87e8-d72c42a273af mailto:veron%40mnhn.fr?subject= mailto:bonillo%40mnhn.fr?subject= mailto:hassanin%40mnhn.fr?subject= mailto:smallcarnivores%40yahoo.com?subject= http://zoobank.org/urn:lsid:zoobank.org:author:de2a6a51-29f0-4a5e-a60b-c61ad5e1842e http://zoobank.org/urn:lsid:zoobank.org:author:7333d242-0714-41d7-b2db-6804f8064b13 http://zoobank.org/urn:lsid:zoobank.org:author:0dcc3e08-b2ba-4a2c-ada5-1a256f24daa1 http://zoobank.org/urn:lsid:zoobank.org:author:3978ac9e-2c27-4b68-99da-038b502afda8 http://dx.doi.org/10.5852/ejt.2017.285 http://dx.doi.org/10.5852/ejt.2017.285 european journal of taxonomy 285: 1–20 (2017) 2 introduction the subfamily hemigalinae (viverridae) comprises four species: owston’s civet chrotogale owstoni thomas, 1912, the otter civet cynogale bennettii gray, 1837, hose’s civet diplogale hosei (thomas, 1892), and the banded civet hemigalus derbyanus (gray, 1837), that are found in southeast asia and southern china (jennings & veron 2009; fig. 1). very little is known about their ecology, but they seem to be nocturnal, mainly terrestrial, and are primarily found in tropical forests (veron et al. 2006; jennings & veron 2009; jennings et al. 2013). the banded civet and otter civet occur mainly in the lowlands, while hose’s civet is primarily found at higher elevations (veron et al. 2006; jennings et al. 2013); owston’s civet seems to be found both in lowland and montane forests (jennings & veron 2009). these species are of conservation concern, with the otter civet listed as endangered and the other three as vulnerable by the iucn (2015), although there is a scarcity of data about their current population trends and sensitivity to environmental disturbance (schreiber et al. 1989; jennings & veron 2009; jennings et al. 2013). the destruction and degradation of tropical forests are clearly major threats to this group (meijaard & sheil 2008; corlett 2009; jennings et al. 2013). despite these threats, little molecular systematics work has been done on this group; knowledge of their phylogenetic diversity, intraspecific genetic diversity and structure, and population differentiation is crucial for defining conservation priorities and for the identification of evolutionarily significant units (esus, ryder 1986; moritz 1994; haig 1998; isaac et al. 2007). fig. 1. map showing the distribution of the samples of hemigalinae used in this study. the gray shadings show the range of chrotogale owstoni thomas, 1912, cynogale bennettii gray, 1837 and hemigalus derbyanus (gray, 1837); diplogale hosei (thomas, 1892) is only found on borneo and macrogalidia musschenbroekii (schlegel, 1879) only occurs on sulawesi. the size of the sample symbols corresponds to the number of samples from each area (the smallest equals 1, and the largest equals 12–14). veron g. et al., systematics and biogeography of the hemigalinae civets 3 the systematics of the hemigalinae had previously only been assessed using morphological data (pocock 1933; gregory & hellman 1939; wozencraft 1989; veron 1994, 1995) until patou et al. (2008) proposed a molecular phylogeny of the hemigalinae and paradoxurinae. patou et al. (2008) found that the banded civet and owston’s civet were sister species, with the otter civet as the sister taxon to these two (as also obtained later by agnarsson et al. 2010). unfortunately, hose’s civet and the sulawesi palm civet macrogalidia musschenbroekii (schlegel, 1879) were missing from these molecular studies. wilting & fickel (2012) provided data for these two species; they confirmed that hose’s civet is a member of the hemigalinae, and found that the sulawesi palm civet grouped with this subfamily (whereas it was previously classified within the paradoxurinae; wozencraft 2005). however, their phylogenetic trees showed very low branch support for some taxa, and the relationships between the species of hemigalinae were not resolved. very few studies have explored intraspecific variation within the species of hemigalinae. veron et al. (2004) conducted a molecular study on owston’s civet and obtained two main groups: northern and central vietnam; however, samples from china were missing. based on morphological features, two species for the otter civet have been proposed, cynogale lowei pocock, 1933 and c. bennettii, but examination of the only specimen available for c. lowei, and doubt about its geographic origin, led veron et al. (2006) to consider c. lowei as a synonym of c. bennettii. four subspecies of the banded civet were proposed by wozencraft (2005): hemigalus derbyanus derbyanus (gray, 1837) (myanmar, peninsular malaysia, sumatra); h. d. boei muller, 1838 (borneo); h. d. minor miller, 1903 (south pagai island, mentawai islands); and h. d. sipora chasen & kloss, 1927 (sipora island, mentawai islands). however, pocock (1933) had previously proposed that h. d. derbyanus ranged from myanmar to the malay peninsula, sumatra and borneo, and that h. d. minor comprises both the south pagai and sipora island populations. corbet & hill (1992) did not find any morphological support for these subspecies, and there has been no recent taxonomic revision of h. derbyanus. on the basis of a short mitochondrial fragment, wilting et al. (2012) suggested a split of ca 2.7 million years between h. d. derbyanus and h. d. minor; unfortunately, they did not provide details about their dataset and results, and their sequences are not available in genbank. tectonic movements, geographical and ecological barriers, and climatic variations across southeast asia have played important roles in the speciation and population structure of forest-dependent species (meijaard 2009; esselstyn et al. 2010; shekelle et al. 2010; woodruff 2010; veron et al. 2014, 2015b, 2015c), and therefore, these might have affected the distribution and speciation of the hemigalinae civets. the distinction between the indochinese and sundaic faunas might have arisen as a result of the rapid sea-level changes along the thai-malay peninsula during the last five million years, which caused local extirpations (woodruff & turner 2009), or it has been maintained by the ecological differences between these two subregions (meijaard 2009). within the sundaic region, climate-induced sea-level changes have altered the topography repeatedly, exposing and flooding land corridors between the mainland and different islands, and ecological factors played a role in restricting movements of species across this region. for instance, during the last glacial maximum (lgm), the central part of the sundaland might have been covered by open habitat (bird et al. 2005; meijaard 2003; de bruyn et al. 2014; but see also woodruff 2010), which could have limited the exchanges of forest-dependent species between borneo and the rest of the sundaic region (veron et al. 2014; 2015c), and this could have affected the hemigalinae civets. the aims of this study were to further clarify the relationships of the species within the hemigalinae, to study the intraspecific diversity and genetic structure within the banded and owston’s civets, and to explore the role of geographical barriers and environmental fluctuations in shaping the structure between and within species. we analysed three mitochondrial fragments, cytochrome b, control region and nadh dehydrogenase subunit 2, and two nuclear genes, beta-fibrinogen intron 7 and intereuropean journal of taxonomy 285: 1–20 (2017) 4 photoreceptor retinoid binding protein (iprb). due to the elusive nature of the hemigalinae civets and the difficulties of obtaining fresh samples, the use of museum samples was crucial for this study. material and methods sampling, extraction, pcr and sequencing we collected fresh (hair or tissue) and museum samples (from skins and tissues taken from skulls) from 51 individuals of nine species of hemigalinae and paradoxurinae (table 1, fig. 1). total genomic dna was isolated following a cetyltrimethyl ammonium bromide (ctab)-based protocol (winnepenninckx et al. 1993). for museum samples, we added dithiothreitol (dtt 1m, ca 15µl per extract) during tissue lysis to break up disulfide bonds and we increased the lysis time (up to 72 hours). we sequenced three mitochondrial fragments: cytochrome b gene (cytb), the control region (cr; hypervariable region 1) and nadh dehydrogenase subunit 2 (nd2), using the primers from veron et al. (2004, 2014, 2015a), patou et al. (2008, 2009) and wilting & fickel (2012). we sequenced the nuclear marker intron 7 of beta-fibrinogen (fgb) using the primers of yu & zhang (2005). irbp sequences were from previous studies (see table 1). polymerase chain reactions (pcrs) were performed as in patou et al. (2010), with annealing temperatures of 50°c for cytb and nd2, 61°c for cr, and 59°c for fgb. pcr products were visualized on a 1.5% agarose gel. pcr products were then purified and sequenced in both directions on an automated dna sequencer by genoscope and eurofins. sequences obtained from dna extracted from museum samples were amplified and sequenced twice to ensure their quality and authenticity. sequences were edited, assembled and aligned manually using bioedit, version 7 (hall 1999). phylogenetic and haplotype network analyses phylogenetic analyses were performed using neighbour-joining (nj), maximum likelihood (ml) and maximum parsimony (mp) as implemented in mega6 (tamura et al. 2013), and bayesian inference (bi) using mrbayes 3.2 (ronquist et al. 2012). we rooted the phylogenetic analyses with three viverrinae (genetta servalina pucheran, 1855, viverricula indica (saint-hilaire, 1803) and viverra tangalunga gray, 1832), one species of felidae (panthera leo (linnaeus, 1758)) and two species of herpestidae (urva auropunctata (hodgson, 1836) and u. javanica (saint-hilaire, 1818)). individual loci were first analysed independently and then combined. for ml, the best-fitting model was estimated prior to the analyses using mega6, following the akaike information criterion (aic). the selected model was then implemented in the ml analyses, using the nearest-neighbor-interchange heuristic method and node robustness was assessed through 1000 bootstrap replicates. for the bi analysis combining the four genes (cytb, nd2, fgb and irbp), jmodeltest v. 2.1.7 (darriba et al. 2012) was used to evaluate the models of nucleotide substitution for each of the ten partitions (fgb and the three codon-positions for cytb, nd2 and irbp), following the akaike information criterion. bayesian analyses were then conducted using the selected gtr + i + g model for cytb-1 and cytb-3, gtr + g model for cytb-2 and nd2-3, gtr model for fgb, hky + i + g model for nd2-1, hky + g model for nd2-2, hky model for irbp-1 and irbp-3, and k80 model for irbp-2. the posterior probabilities (pp) were calculated using four independent markov chains run for 10 000 000 metropolis-coupled mcmc generations, with trees sampled every 1000 generations and a burn-in of 25%. trees were visualized and edited using figtree v. 1.4.0 (rambaut 2012). we compared resulting topologies and their node support; nodes were considered as supported when posterior probabilities were ≥ 0.95 and bootstrap values were ≥ 70% (see alfaro et al. 2003). veron g. et al., systematics and biogeography of the hemigalinae civets 5 ta bl e 1. l is t of t he s am pl es i nc lu de d in t hi s st ud y. f or e ac h sa m pl e, w e re po rt t he i de nt ifi ca tio n nu m be r, th e sa m pl e/ sp ec im en n um be r (a h n u = a nh ui n or m al u ni ve rs ity , w uh u, c hi na ; b z m = m us eu m fü r n at ur ku nd e, b er lin , g er m an y; f m n h = f ie ld m us eu m o f n at ur al h is to ry , c hi ca go , i l , u sa ; i se m = i ns tit ut d es s ci en ce s de l’ e vo lu tio n, m on tp el lie r, fr an ce ; m c z = h ar va rd m us eu m o f c om pa ra tiv e z oo lo gy , c am br id ge , m a , u sa ; m n h n = m us éu m n at io na l d ’h is to ir e na tu re lle , p ar is , f ra nc e; r m n h = r ijk sm us eu m v an n at uu rl ijk e h is to ri e, n at ur al is , l ei de n, th e n et he rl an ds ), th e g en b an k nu m be rs fo r t he fi ve g en es , a nd lo ca lit y (o c p = sp ec im en s he ld a t t he o w st on ’s p al m c iv et c on se rv at io n pr og ra m , c uc p hu on g n at io na l pa rk , v ie tn am ; n d = n o da ta ). id en tifi ca tio n nu m be rs i n bo ld i nd ic at e sa m pl es u se d in t he c yt b + n d 2 + fg b + ir b p da ta se t fo r th e ph yl og en et ic an al ys es . g en b an k nu m be rs in b ol d ar e ne w s eq ue nc es fr om th is s tu dy ; o th er s eq ue nc es a re fr om th e fo llo w in g so ur ce s: c ra cr af t e t a l. 19 98 ; v er on & h ea rd 2 00 0; y od er e t a l. 20 03 ; g au be rt e t a l. 20 04 a, 2 00 4b ; g au be rt & c or de ir oe st re la 2 00 6; v er on e t a l. 20 04 ; y u et a l. 20 04 ; m as ud a et a l. 20 08 ; fa ng & w u 20 05 (u np ub l.) ; y u & z ha ng 2 00 5; p at ou e t a l. 20 08 , 2 00 9, 2 01 0; w ilt in g & f ic ke l 2 01 2; v er on e t a l. 20 15 b. s eq ue nc es < 2 00 b p ca nn ot be s ub m itt ed to g en b an k (a va ila bl e on re qu es t t o g v ). [c on tin ue d on n ex t t w o pa ge s] sp ec ie s id en ti fic at io n sa m pl e/ sp ec im en c yt b c r n d 2 f g b ir b p l oc al it y # # g en b an k # g en b an k # g en b an k # g en b an k # g en b an k # a rc tic tis b in tu ro ng a bt c 44 4 m n h n t c -4 44 k p 98 64 69 – – – – n d (s er vi on z oo ) a rc tic tis b in tu ro ng a bt c 94 m n h n t c -9 4 – – k m 81 95 58 e f6 80 50 3 d q 68 31 25 n d (c ar ni vo re p re se rv at io n tr us t) a rc to ga lid ia tr iv ir ga ta a tc 15 5 m n h n 2 00 149 5 k m 81 95 40 – k m 81 95 59 k m 81 95 23 – n d (p ar c z oo lo gi qu e de p ar is , m n h n ) a rc to ga lid ia tr iv ir ga ta a tt c 1 m n h n 1 99 819 70 – – – – d q 68 31 26 n d (m én ag er ie d u ja rd in d es p la nt es , m n h n , p ar is ) c hr ot og al e ow st on i c od q 28 67 75 “a h n u 03 10 ” d q 28 67 75 – – – – c hi na c hr ot og al e ow st on i c os h 1 m n h n s h -1 a f1 25 14 2 – – – – v ie tn am , n or th (f ro m tr af fic , o c p) c hr ot og al e ow st on i c os h 10 m n h n s h -1 0 a y 15 52 48 – – – – v ie tn am (h ôc hi -m in hv ill e z oo ) c hr ot og al e ow st on i c os h 11 m n h n s h -1 1 a y 15 52 63 – – – – v ie tn am (h ôc hi -m in hv ill e z oo ) c hr ot og al e ow st on i c os h 12 m n h n s h -1 2 a y 15 52 69 – – – – v ie tn am (h ôc hi -m in hv ill e z oo ) c hr ot og al e ow st on i c os h 13 m n h n s h -1 3 a y 15 52 66 – – – – v ie tn am (h ôc hi -m in hv ill e z oo ) c hr ot og al e ow st on i c os h 14 m n h n s h -1 4 a y 15 52 52 – – – – v ie tn am (h ôc hi -m in hv ill e z oo ) c hr ot og al e ow st on i c os h 15 m n h n s h -1 5 a y 15 52 53 – – – – v ie tn am (h ôc hi -m in hv ill e z oo ) c hr ot og al e ow st on i c os h 16 m n h n s h -1 6 a y 15 52 59 – – – – v ie tn am (h ôc hi -m in hv ill e z oo ) c hr ot og al e ow st on i c os h 3 m n h n s h -3 a y 15 52 51 – – – – v ie tn am (o c p) c hr ot og al e ow st on i c os h 4 m n h n s h -4 a y 15 52 58 – – – – v ie tn am , n or th (o c p) c hr ot og al e ow st on i c os h 6 m n h n s h -6 a y 15 52 64 – – – – v ie tn am , n or th (o c p) c hr ot og al e ow st on i c os h 8 m n h n s h -8 a f1 25 14 4 – – – – v ie tn am (h ôc hi -m in hv ill e z oo ) c hr ot og al e ow st on i c os h 9 m n h n s h -9 a y 15 52 49 – – – – v ie tn am (h ôc hi -m in hv ill e z oo ) european journal of taxonomy 285: 1–20 (2017) 6 sp ec ie s id en ti fic at io n sa m pl e/ sp ec im en c yt b c r n d 2 f g b ir b p l oc al it y # # g en b an k # g en b an k # g en b an k # g en b an k # g en b an k # c hr ot og al e ow st on i c ot 20 92 m n h n t -2 09 2 a y 15 52 62 – d q 68 39 84 e f6 80 50 5 d q 68 31 27 v ie tn am , n or th c hr ot og al e ow st on i c ot c 18 6 m n h n t c -1 86 a y 15 52 61 – – – – v ie tn am , n or th (o c p) c hr ot og al e ow st on i c ot c 18 7 m n h n t c -1 87 a y 15 52 67 – – – – v ie tn am , n or th (o c p) c hr ot og al e ow st on i c ot c 18 8 m n h n t c -1 88 a y 15 52 50 – – – – v ie tn am , n or th , k im b oi , h oa b in h (o c p) c hr ot og al e ow st on i c ot c 18 9 m n h n t c -1 89 a y 15 52 68 – – – – v ie tn am (o c p) c hr ot og al e ow st on i c ot c 19 0 m n h n t c -1 90 a y 15 52 70 – – – – v ie tn am (o c p) c hr ot og al e ow st on i c ot c 19 1 m n h n t c -1 91 a y 15 52 60 – – – – v ie tn am ( ca pt iv e bo rn , o c p) c hr ot og al e ow st on i c ot c 19 3 m n h n t c -1 93 a y 15 52 56 – – – – v ie tn am ( ca pt iv e bo rn , o c p) c hr ot og al e ow st on i c ot c 19 7 m n h n t c -1 97 a y 15 52 71 – – – – v ie tn am ( o c p) c hr ot og al e ow st on i c ot c 20 4 m n h n t c -2 04 a y 15 52 55 – – – – v ie tn am , c en te r, b in h d in h (o c p) c hr ot og al e ow st on i c ot c 20 5 m n h n t c -2 05 a y 15 52 54 – – – – v ie tn am , n or th , n gh e a n (o c p) c hr ot og al e ow st on i c ot c 20 6 m n h n t c -2 06 a y 15 52 65 – – – – v ie tn am , n or th , n gh e a n (o c p) c hr ot og al e ow st on i c ot c 20 7 m n h n t c -2 07 a y 15 52 57 – – – – v ie tn am , n or th , v an b an , l ao c ai p ro vi nc e c yn og al e be nn et tii c bm 19 62 m n h n 1 96 217 0 k p 98 64 70 – k p 98 64 85 – – m al ay si a, m al ac ca c yn og al e be nn et tii c bt c 14 7 m n h n t c -4 17 d q 68 39 92 – d q 68 39 83 – – b or ne o, s ar aw ak (n eg ar a z oo , k ua la l um pu r) d ip lo ga le h os ei d hf 88 29 8 fm n h 8 82 98 k p 98 64 71 – k p 98 64 86 – – m al ay si a, b or ne o, s ar aw ak , k el ab it pl at ea u, p a u m ur d ip lo ga le h os ei d hj q 21 91 12 r m n h 3 45 68 jq 21 91 12 – jq 21 91 10 – – m al ay si a, b or ne o, s ar aw ak , s al ek an m t d ip lo ga le h os ei d hj q 21 91 11 b z m 1 42 91 – – jq 21 91 11 – – b ru ne i, m ar ab ok m t h em ig al us d er by an us h dc 38 64 7 m c z 3 86 47 k p 98 64 72 – – – – in do ne si a, s um at ra , e as t, l itt le s ia k ri ve r h em ig al us d er by an us h dm 24 m n h n 1 88 624 k u 69 64 24 – – – – in do ne si a, s um at ra , p al em ba ng h em ig al us d er by an us h dm 48 9 m n h n 1 86 548 9 k u 69 64 25 – – – – m al ay si a, p en in su la r m al ay si a, p en an g is la nd h em ig al us d er by an us h dr 34 64 3 r m n h 3 46 43 k u 69 64 26 – – – – in do ne si a, b or ne o h em ig al us d er by an us h dr 51 09 r m n h 5 10 9 k u 69 64 27 – – – – in do ne si a, s um at ra , a tje h, r am pa ih h em ig al us d er by an us h dt c 09 m n h n t c -0 9 a f1 25 14 3 k p 98 64 68 a y 17 00 52 k p 98 64 82 a y 17 00 82 n d (s in ga po re z oo ) h em ig al us d er by an us h dt c 20 m n h n t c -2 0 k p 98 64 73 – – – – m al ay si a, b or ne o, s ar aw ak (c in ci nn at i z oo ) h em ig al us d er by an us h dt c 21 m n h n t c -2 1 <2 00 bp k p 98 64 63 – – – m al ay si a, b or ne o, s ab ah , d an um v al le y veron g. et al., systematics and biogeography of the hemigalinae civets 7 h em ig al us d er by an us h dt c 22 m n h n t c -2 2 a y 15 52 73 – d q 68 39 87 k p 98 64 79 – m al ay si a, b or ne o, s ab ah , d an um v al le y h em ig al us d er by an us h dt c 23 m n h n t c -2 3 a y 15 52 72 k p 98 64 64 – e f6 80 50 8 – m al ay si a, b or ne o, s ab ah , d an um v al le y h em ig al us d er by an us h dt c 28 6 m n h n t c -2 86 k p 98 64 74 k p 98 64 65 – k p 98 64 80 – in do ne si a, s ib er ut is ., m en ta w ai is ., n m ua ra si be ru t h em ig al us d er by an us h dt c 30 0 m n h n t c -3 00 k p 98 64 75 k p 98 64 66 – k p 98 64 81 – in do ne si a, b or ne o, w k al im an ta n, k er ap a b ej al i h em ig al us d er by an us h dt c 41 1 m n h n t c -4 11 k p 98 64 76 k p 98 64 67 – k u 69 64 29 – m al ay si a (n eg ar a z oo , k ua la l um pu r) h em ig al us d er by an us h dt c 75 6 m n h n t c -7 56 k u 69 64 28 – – – – su m at ra ? (b at u se cr et z oo & m ah ar an i z oo , j av a) m ac ro ga lid ia m us sc he nb ro ek ii m m jq 21 91 16 r m n h a jq 21 91 16 – jq 21 91 13 – – in do ne si a, s ul aw es i, m an ad ok in ilo m ac ro ga lid ia m us sc he nb ro ek ii m m jq 21 91 14 r m n h 3 22 44 – – jq 21 91 14 – – in do ne si a, s ul aw es i, a m ur an g m ac ro ga lid ia m us sc he nb ro ek ii m m jq 21 91 15 r m n h 3 22 43 – – jq 21 91 15 – – in do ne si a, s ul aw es i, a m ur an g m ac ro ga lid ia m us sc he nb ro ek ii m m m 18 68 m n h n 1 86 813 27 k p 98 64 77 – k p 98 64 87 – – in do ne si a, s ul aw es i p ag um a la rv at a p la b 30 39 59 “s e 4” a b 30 39 59 – – – – t ha ila nd p ag um a la rv at a p la y 52 50 40 – – – – – a y 52 50 40 c hi na , y un na n pr ov in ce p ag um a la rv at a p lc 72 m n h n c -7 2 – – d q 68 39 90 – – n d p ag um a la rv at a p ll 92 m n h n l -9 2 – – – k p 98 64 83 – c hi na , g ua ng xi p ar ad ox ur us p hi lip pi ne ns is p hf 31 67 fm n h l r h 3 16 7 a y 17 01 13 fj 88 15 43 a y 17 00 56 e f6 80 51 0 a y 17 00 86 ph ili pp in es , l ey te is la nd g en et ta s er va lin a g s is e m t -4 62 0 a f5 11 05 3 – a y 17 00 58 e f6 80 50 7 a y 17 00 88 g ab on vi ve rr ic ul a in di ca v i m n h n t c -2 5 – – – – d q 26 75 68 m ad ag as ca r vi ve rr ic ul a in di ca v i m n h n t c -1 77 a y 24 18 90 – – – – ta iw an vi ve rr ic ul a in di ca v i m n h n t c -2 56 – – d q 68 39 91 – – t ha ila nd vi ve rr a ta ng al un ga v t fm n h l r h 4 12 1 a y 17 01 12 – a y 17 00 55 – a y 17 00 85 ph ili pp in es vi ve rr a ta ng al un ga v t m n h n t c -5 34 – – – e f6 80 51 2 – m al ay si a p an th er a le o p le o – a f0 53 05 2 – a y 17 00 43 a y 63 43 74 a y 17 00 73 n d u rv a au ro pu nc ta ta u a “n el lis ” a y 17 01 08 – a y 17 00 51 – a y 17 00 81 c ar ib be an is la nd s u rv a au ro pu nc ta ta u a m n h n t c -3 40 – – – f j3 91 19 4 – c ro at ia , k or cu la is la nd u rv a ja va ni ca u j m n h n t c -2 58 k p 98 64 78 – fj 39 12 34 k p 98 64 84 – t ha ila nd sp ec ie s id en ti fic at io n sa m pl e/ sp ec im en c yt b c r n d 2 f g b ir b p l oc al it y # # g en b an k # g en b an k # g en b an k # g en b an k # g en b an k # european journal of taxonomy 285: 1–20 (2017) 8 we used dnasp v. 5.10 (librado & rosas 2009) to define haplotypes and to compute genetic diversity (haplotype and nucleotide diversity), and network v. 4.6 (www.fluxus-engineering.com) to construct haplotype median-joining networks (bandelt et al. 1999). we computed genetic distances (within and between groups) and genetic diversity (haplotype and nucleotide diversity) using mega6 and dnasp v. 5.10. results all new sequences were deposited in genbank (accession numbers: kp986463 to kp986487 and ku696424 to ku696429; table 1). due to the degraded nature of dna retrieved from samples taken from museum specimens and poorly preserved tissue, only partial sequences could be obtained from some samples and nuclear sequences were not retrieved from museum specimens. for the diplogale hosei cytb sequence from genbank (jq219112; wilting & fickel 2012), we replaced the nucleotides in position 327 to 344 by ‘n’, as we strongly suspected that these nucleotides were those of the primer sequence (their primer cytbviv0323f), based on a comparison within the alignment. the cytb and nd2 fragments that we obtained from a d. hosei specimen (fmnh 88298) differed by 0.5% and 1.5% to the two individuals sequenced by wilting & fickel (2012). the cytb sequence of a specimen of cynogale bennettii (mnhn 1962-170) did not differ from the sequence of the individual sequenced by patou et al. (2008), while the nd2 sequence differed by 0.7%. the cytb fragment of a macrogalidia musschenbroekii specimen (mnhn 1868-1327) did not differ from the three sequences obtained by wilting & fickel (2012), while its nd2 sequence differed by 0.0–0.6%. any differences we found may reflect sequencing errors, which can happen with ancient dna, or were due to individual variations (and these differences were not higher than the intraspecific variations observed within other species of viverridae). all gene fragments were analysed both individually and combined, with the different phylogenetic methods. the phylogenetic analyses of the dataset combining cytb, nd2, fgb, and irbp (length of the alignment, number of variable positions and number of parsimony-informative sites: 3342/1226/801, n = 15, including 6 outgroups) confirmed the monophyly of the hemigalinae and paradoxurinae (fig. 2). within the hemigalinae, hemigalus and chrotogale are sister species (with a bayesian posterior probability (bpp) of 0.91; ml boostrap = 96%), and diplogale is sister to these two species in the bi tree (bpp = 1; fig. 2), as well as in the mp tree (mp boostrap > 50%), while in the ml tree, the position of diplogale within this subfamily was unresolved (fig. 2). macrogalidia clusters within the hemigalinae in all analyses (bpp = 1; ml boostrap = 99%; fig. 2); it is sister to cynogale in the bi tree (bpp = 0.90), but its position (and that of cynogale) within this subfamily were unresolved in the ml and mp trees. for the complete cytb (1140 bp, n = 46), the distances between the two subfamilies (as previously defined) for hemigalinae (without macrogalidia) and paradoxurinae (including macrogalidia) ranged from 12.5% to 17.6%. the average cytb distance between the two clades obtained in our study, hemigalinae including macrogalidia and paradoxurinae without macrogalidia, was 15.9%, and the mean within group distances were 7.2% and 12.3%, respectively. the distances between macrogalidia and the species of hemigalinae ranged from 12.0% to 14.9%, and to the species of paradoxurinae from 14.4% to 15.9%. the mean distances between the remaining four species of the hemigalinae clade ranged from 11.9% to 16.7%; the most distant species to the other three was cynogale, and the smallest mean distance was between hemigalus and chrotogale. we analysed a fragment of cytb (895 bp, positions 123–1015 in complete cytb alignment) for h. derbyanus and c. owstoni in order to search for geographical structure within these two species (895 / 241 / 142; n = 39; model gtr + i + g; fig. 3). h. derbyanus was structured into two main clades: www.fluxus-engineering.com veron g. et al., systematics and biogeography of the hemigalinae civets 9 one containing an individual from siberut island (mentawai islands) and a second clade that comprised all other individuals (fig. 3). the overall mean distance within h. derbyanus was 1.6%; the mean distances of the siberut island individual to those from sumatra and borneo were 4.7% and 5.5%, respectively, whereas it ranged from 1.0% to 1.3% between sumatra, borneo and peninsular malaysia. c. owstoni formed one group from northern vietnam (although its monophyly was poorly supported or not retrieved in some analyses), that is sister group to another from central vietnam, whereas the individual from china was distant to these two (fig. 3). the overall mean distance within c. owstoni was 1.1%; the mean distance between the two vietnam groups was 1.4%, while the chinese individual was separated from the latter groups by a mean distance of 2.1% to 3.4%. a fragment of cr was obtained for a few individuals of h. derbyanus (592 / 18 / 0, n = 6) and provided a similar phylogenetic structure (not shown) as cytb, with the same two main clades; the mean distance between the siberut island individual and the others was 4.7%. we obtained four cr haplotypes (195 bp excluding missing data; haplotype diversity, hd: 0.8; nucleotide diversity, pi: 0.01880; average number of nucleotide differences, k: 3.667), with the siberut island individual separated by eight to ten mutations to any other haplotype. we also used a cytb fragment (834 bp, positions 129–962 in cytb alignment) to compute haplotype networks for h. derbyanus and c. owstoni. for h. derbyanus, we obtained eight haplotypes (hd: 0.972; pi: 0.01952; k: 16.278; n=9; fig. 4), with the siberut island haplotype (h6) separated by 39 mutations from the other individuals. one sumatra individual (h1) was separated from a haplogroup of borneo and zoo individuals. for c. owstoni, we obtained 19 haplotypes (hd: 0.9692; pi: 0.00886; k: 7.345; fig. 2. bayesian tree reconstructed from a combined dataset of cytb + nd2 + fgb + irbp (3342 bp). the values on the branches are bayesian posterior probabilities for the partitioned analysis (see text for models) and bootstrap proportions obtained from ml analysis (model: gtr + i + g). european journal of taxonomy 285: 1–20 (2017) 10 fig. 3. phylogenetic tree obtained with nj for a fragment of cytb (893 bp). the values over the branches are the bootstrap proportions for nj, and below the branches are those for ml. co = chrotogale owstoni thomas, 1912; hd = hemigalus derbyanus (gray, 1837); dh = diplogale hosei (thomas, 1892); cb = cynogale bennettii gray, 1837. veron g. et al., systematics and biogeography of the hemigalinae civets 11 n = 26; fig. 5), forming two groups for northern and southern vietnam, separated by eight mutations (the chinese individual could not be used in this analysis due to missing data). we also computed haplotypes using smaller cytb fragments in order to include additional individuals of various origins. for h. derbyanus, using a 253 bp fragment in the 3’ region (positions 768–1020 in the cytb alignment), we obtained six haplotypes (hd: 0.8030; pi: 0.01509; k: 3.818; n = 12; figure 4), organized in a star-like structure, with a central haplotype (h5) from borneo (and also zoo individuals); the siberut island (mentawai) individual (h6) was separated by 13 mutations from the two sumatra haplotypes (h2 and h4), which were separated from the borneo haplotype (h5) by two mutations; the individual from peninsula malaysia (h3) was separated by one mutation from the borneo haplotype; another haplotype (h1) comprising individuals from borneo and sumatra was three mutations distant from the borneo haplotype (h5). for c. owstoni, using another 253 bp fragment in the 5’ region (positions 123–375 in cytb alignment), we obtained six haplotypes (hd: 0.7000; pi: 0.00901; k: 2.280; n = 25; fig. 5); northern vietnam individuals formed a haplogroup of three haplotypes (h2, h5, h6), while central vietnam individuals formed a haplogroup of two haplotypes (h3, h4); the chinese individual (h1) was separated from these groups by a minimum of five and eight mutations, respectively; the northern and central vietnam groups were separated by a minimum of three mutations. we also computed haplotypes for the h. derbyanus fgb dataset (321bp), and we obtained three haplotypes (hd: 0.6000; pi: 0.00415; k: 1.333; n = 6); the siberut island (mentawai islands) individual was separated by three and four mutations from the other two haplotypes (not shown). fig. 4. median-joining haplotype network for hemigalus derbyanus (gray, 1837) cytb haplotypes (top: 837 bp, bottom: 253bp). the size of each circle is proportional to the haplotype frequency. the colours of the haplotypes correspond to those on the map: black = borneo; dark grey = siberut island (mentawai islands); light grey = penang island (peninsular malaysia); white = sumatra; hatched = zoo samples (on top network: singapore zoo (h2), negara zoo, kuala lumpur (h7) and batu secret zoo & maharani zoo, java (h8)). european journal of taxonomy 285: 1–20 (2017) 12 our results suggest that the h. derbyanus individuals from the singapore zoo (hdtc09), the negara zoo in kuala lumpur (tc-411), and the batu secret zoo & maharani zoo in java (tc-756) all come from borneo, despite the fact that the latter individual was supposed to come from sumatra. the individual from the cincinnati zoo (tc-20) was supposed to come from borneo and our results agree with this. discussion while wilting & fickel (2012) did not obtain a resolution for the banded, hose and owston’s civets relationships, our study found that the banded and owston’s civets are sister species, and also suggested that hose’s civet might be the sister species to these two. similar to wilting & fickel (2012), the sulawesi palm civet was found within the hemigalinae. it either clustered with the otter civet (although this was weakly supported) or its position was unresolved. these two species had missing data (nuclear genes) and long branches (in particular the otter civet), which could have affected their phylogenetic position; long branch attraction is a commonly recognized artefact in phylogenetic reconstruction that can produce misleading (but sometimes strongly supported) relationships (felsentein 1978; simmons 2012). also, the use of only mitochondrial data can lead to misleading relationships at ordinal and also lower taxonomic levels (e.g., for carnivores, sato et al. 2003). for example, prior to the addition of nuclear data, the small-toothed palm civet arctogalidia trivirgata was not retrieved within the paradoxurinae (patou et al. 2008). morphological features support the inclusion of the sulawesi palm civet in the paradoxurinae (gaubert et al. 2005), which either invalidates fig. 5. median-joining haplotype network for chrotogale owstoni thomas, 1912 cytb haplotypes (top: 837 bp, bottom: 235 bp). the size of each circle is proportional to the haplotype frequency. white = central vietnam clade; grey = northern vietnam clade, black = china. veron g. et al., systematics and biogeography of the hemigalinae civets 13 the molecular results, or indicates homoplasy in the morphological features. further genetic studies, with additional samples and data (nuclear genes) are needed to clarify further the phylogenetic position of the sulawesi palm civet. we advocate that further studies be conducted before moving macrogalidia to the hemigalinae. the sulawesi palm civet is a threatened species endemic to sulawesi (jennings & veron 2009; iucn 2015), and is the only native carnivore found east of wallace’s line (veron 2001). hall (2001) found evidence of a land bridge west of sulawesi ca 20 million years ago (mya), and suggested faunal exchanges between sulawesi and the rest of southeast asia occurred ca 10 mya. more recently, it has been suggested that colonisations of sulawesi started during the early miocene, and that speciation events happened as early as 20 mya (stelbrink et al. 2012). patou et al. (2008) proposed that the hemigalinae diverged 15.4–8.4 mya, which corresponds to the dates of origin of several sulawesi taxa (stelbrink et al. 2012). these findings suggest that the sulawesi palm civet diverged from the other species more than 10 mya, which is supported by its high cytb divergence (12.0–14.9%) from its relatives. this ancient speciation explains why the analysis of only mitochondrial sequences has not fully resolved its phylogenetic relationships. the northern distribution limit of the banded civet is close to the isthmus of kra, along the thai-malay peninsula (jennings et al. 2013), which is a transitional region between the indochinese and sundaic faunas (woodruff & turner 2009; meijaard 2009). the range of owston’s civet is restricted to southern china, laos, and vietnam (jennings &veron 2009). eustatic and climatic events in this region may have caused the speciation of these two civet species (with subsequent range restrictions due to habitat changes that occurred during the plio-pleistocene and more recently to human-induced modifications). hose’s civet is found only on borneo, where it is sympatric with the banded civet (although there is a tendency for these two species to separate spatially along an elevation gradient; jennings et al. 2013). however, we do not know if hose’s civet might have had a wider distribution in the past and was then restricted to forest refugia in the mountains of borneo during plio-pleistocene climatic variations, or if it speciated on borneo from an ancestor of hemigalus + chrotogale. the otter civet is a specialised viverrid that is adapted to aquatic life (with webbed feet, and nostrils and ears that can be closed by flaps) that ranges across the sundaic region (veron et al. 2006), and it could have diverged first within the subfamily. we found a strong geographical genetic structure in the banded and owston’s civets. within the banded civet, the individual from siberut island (mentawai islands) was very distant from the populations from both sumatra and borneo (for both cytb and fgb), and according to our haplotype networks it is closer to sumatra individuals. in contrast, the mentawai population of the common palm civet (paradoxurus sp.) was found to be closer to those from borneo and the philippines (paradoxurus philippinensis) than to the nearby sumatran populations (paradoxurus musangus) (patou et al. 2010; veron et al. 2015c). although the four mentawai islands (siberut, sipora, north and south pagai) are close to sumatra, they are separated by a deep sea channel, the mentawai strait, which has isolated populations and explains the high level of species endemism on these islands (wilting et al. 2012). their strong divergence from sumatran populations might also be explained by local extinctions during the pleistocene (patou et al. 2010; wilting et al. 2012) and the mentawai islands might have served as a forest refuge during glaciations. the divergence time of mentawai taxa has been estimated from 1.1 to 3.5 mya in mammals (patou et al. 2010; wilting et al. 2012). the mentawai islands have been separated from sumatra at least since the mid-pleistocene and might have remained separated despite the sea level fluctuations of the late pleistocene (voris 2000; wilting et al. 2012). the cytb distance of the siberut island individual to other banded civet populations ranged from 4.7 to 5.5%, which is close to the divergence found between other small carnivore species (veron et al. 2007, 2015a, 2015c); our nuclear data also confirmed this strong divergence. two subspecies have been described from the mentawai islands: hemigalus derbyanus minor miller, 1903 (south pagai island) and h. d. sipora chasen & kloss, 1927 (sipora island). very little is known about the population from siberut island, which has not been attributed to european journal of taxonomy 285: 1–20 (2017) 14 any subspecies (jennings et al. 2013). the four mentawai islands were likely connected during periods of lower sea levels (sathiamurthy & voris 2006), and there does not seem to be any endemism on each island, as several species occur on all or several of the four islands (sargis et al. 2014). it is therefore likely that banded civets from the islands of siberut, sipora and south pagai belong to the same taxon, which would be hemigalus derbyanus minor. banded civet individuals from sumatra were not strongly divergent from those from borneo, whereas a strong divergence between sumatran and borneo populations has been found in other forest species (veron et al. 2015a). borneo has been isolated from sumatra for ca 20 000 years, at the end of the last glacial maximum, and even when land connections existed between borneo and the rest of the sundaic region, open habitat might have restricted the movements of some forest animals (meijaard 2003; bird et al. 2005; cannon et al. 2009). our results suggest that there have been independent connections between borneo and sumatran populations, and between borneo and peninsular malaysian populations (in this study represented by an individual from penang island, which is very close and connected to the nw coast of peninsular malaysia). although peninsular malaysia and sumatra have been connected frequently during glacial periods, the banded civet individuals from these two areas did not form a monophyletic group in our analyses, whereas a close relationship and low differentiation of populations from sumatra and peninsular malaysia have been found in many other mammals, such as rodents, macaques, palm civets and mongooses (gorog et al. 2004; ziegler et al. 2007; patou et al. 2010; veron et al. 2015a). possible explanations for these findings would be independent dispersals during periods of lower sea levels (e.g., last glacial maximum), or that banded civets were transported from one area to another, as has been suggested for some other civet species (veron et al. 2014, 2015c), although there is no evidence that this has ever occurred for the banded civet. with regards to the structure obtained and the low divergences observed, we would suggest that the populations from sumatra, peninsular malaysia and borneo are placed in the same subspecies hemigalus derbyanus derbyanus (gray, 1837). within owston’s civet, we obtained two main groups in vietnam, which confirms the finding of veron et al. (2004), and we found a third group from china, quite distant from the northern vietnam group. this might have resulted from either geographic barriers or pleistocene climatic fluctuations (as suggested for bats, see tu et al. 2017) that shaped the population structure of this species. however, only one sequence was available for china (and it came from genbank so we cannot confirm its validity), and thus, additional data would be needed to confirm this finding. these three groups could have been maintained due to limited exchanges between isolated populations as a result of habitat destruction (see iucn 2015). to assess whether these populations should be considered subspecies would require further investigations, as the origin of several individuals was uncertain and samples from laos were missing in our dataset. this is a very poorly known, endangered civet species that is severely threatened by forest loss, hunting and the wildlife trade (willcox et al. 2011; coudrat et al. 2014). further studies are needed on the intraspecific diversity and genetic structure of the hemigalinae civets, and for the designation of conservation units, in particular, for island endemic species and subspecies. unfortunately, due to their elusive nature, these civet species have rarely been studied in the wild and are very rarely trapped, so obtaining fresh samples is extremely difficult. finally, ecological studies are urgently needed in order to better understand and assess the conservation status of these threatened species. acknowledgements we thank the following people and institutions for their contribution to this study: f. catzeflis (institut des sciences de l’evolution, université de montpellier), c.p. colon (fordham university), r. debruyne (ums cnrs 2700, mnhn, paris), a. erman & e. meijaard (people & nature consulting international), m.f. gorra (carnivore preservation trust), l. grassman (texas a&m university veron g. et al., systematics and biogeography of the hemigalinae civets 15 – kingsville), l.l. lee (taiwan university), b. long (world wildlife fund), p. martelli (singapore zoological gardens), j. patton (museum of vertebrate zoology, berkeley, ca), m.l. patou (biotope), s. roberton (ffi, owston’s palm civet conservation program), t.l. roth (cincinnati zoo), s. vellayan (zoo negara, kuala lumpur), siew te wong (borneo sun bear conservation center), servion zoo, d. boussarie, and s. heard. we thank the following curators and museums for access to specimens and samples: n. duncan, e. westwig, e. hoeger (american museum of natural history, new york), l.r. heaney, j.d. phelps, the late w.t. stanley (field museum of natural history, chicago), m. omura, b. zimkus, j. chupasko (harvard museum of comparative zoology, cambridge, ma), s. van der mije, p. kamminga (naturalis, leiden, the netherlands) and k.k.p. lim (raffles museum of biodiversity research, singapore). thanks are due to the forest protection department of vietnam’s ministry of agriculture & rural development for their permission to export samples. we thank the epu and perhilitan for permission to conduct research in malaysia, and s. ellagupillay (perhilitan) and z. akbar (university kebangsaan malaysia) for their support; funding for the malaysian carnivore project is acknowledged at www.smallcarnivores.org. molecular work was undertaken at the ‘service de systématique moléculaire’ (ums cnrs 2700, mnhn, paris), and we thank the staff of the lab for their help. the sequencing was supported by the network “bibliothèque du vivant” funded by cnrs, mnhn, inra and cea (genoscope, www.genoscope.fr). g.v. received financial support from mnhn, cnrs, and from the “action transversale du muséum, biodiversité actuelle et fossile” (mnhn, french ministry of research). we thank the associate editor and anonymous reviewers for their useful suggestions on an earlier version of this paper. references agnarsson i., kuntner m. & may-collado l.j. 2010. dogs, cats, and kin: a molecular specieslevel phylogeny of carnivora. molecular phylogenetics and evolution 54: 726–745. http://dx.doi. org/10.1016/j.ympev.2009.10.033 alfaro m.e., zoller s. & lutzoni f. 2003. bayes or bootstrap? 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botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands. http://dx.doi.org/10.1038/nature01303 http://dx.doi.org/10.1038/nature01303 http://dx.doi.org/10.1016/j.ympev.2005.01.017 http://dx.doi.org/10.1016/j.ympev.2004.08.001 european journal of taxonomy 391: 1–44 issn 2118-9773 https://doi.org/10.5852/ejt.2017.391 www.europeanjournaloftaxonomy.eu 2017 · prantoni a. et al. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:2e743067-031f-401e-aa7e-f85ec4d57e6d 1 global checklist of species of grania (clitellata: enchytraeidae) with remarks on their geographic distribution alessandro prantoni 1,*, paulo c. lana 2 & christer erséus 3 1,2 center for marine studies, federal university of paraná, av. beira mar, s/n, 83255-976 pontal do paraná, paraná, brazil. 3 department of biological and environmental sciences, university of gothenburg, box 463, se-405 30 göteborg, sweden. * corresponding author: aprantoni@gmail.com 2 email: lana@ufpr.br 3 email: christer.erseus@bioenv.gu.se 1 urn:lsid:zoobank.org:author:3fe88982-3151-43af-bc1c-5c8f863729b7 2 urn:lsid:zoobank.org:author:b5a74b14-a681-4d2a-8248-67de8fce9f7b 3 urn:lsid:zoobank.org:author:d98f606a-b273-4f50-95f5-c35f17b12c85 abstract. a checklist of all currently accepted species of grania southern, 1913 (annelida, clitellata, enchytraeidae) is presented. the genus is widespread over the world and comprises 81 species described to date. remarks on their geographical distribution, habitat, synonymies and museum catalogue numbers are provided. keywords. species list, annelida, marine clitellates, geographic distribution, interstitial fauna. prantoni a., lana p.c. & erséus c. 2017. global checklist of species of grania (clitellata: enchytraeidae) with remarks on their geographic distribution. european journal of taxonomy 391: 1–44. https://doi.org/10.5852/ ejt.2017.391 introduction grania southern, 1913 is a morphologically homogeneous and easily recognizable genus of marine enchytraeidae vejdovský, 1879 with a worldwide distribution. the worms are typically small, only a few mm long, with a filiform, nematode-like body and a characteristic pattern of few, stout chaetae (fig.1). most species live interstitially in intertidal or subtidal sands, but a few taxa are known from the deep sea. the genus was originally established for specimens of grania maricola southern, 1913, sampled from the west coast of ireland. later on, pierantoni (1915) regarded grania maricola and his own species, enchytraeus macrochaetus pierantoni, 1901, at the time referred to michaelsena ude, 1896 (pierantoni 1903), as synonyms. stephenson (1930) instead considered them as separate species within michaelsena. nielsen & christensen (1959) pointed out, however, that stephenson’s concept of michaelsena was artificial, and they transferred both species to the genus enchytraeus henle, 1837. https://doi.org/10.5852/ejt.2017.391 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:2e743067-031f-401e-aa7e-f85ec4d57e6d mailto:aprantoni%40gmail.com?subject= mailto:lana%40ufpr.br?subject= mailto:christer.erseus%40bioenv.gu.se?subject= http://zoobank.org/urn:lsid:zoobank.org:author:3fe88982-3151-43af-bc1c-5c8f863729b7 http://zoobank.org/urn:lsid:zoobank.org:author:b5a74b14-a681-4d2a-8248-67de8fce9f7b http://zoobank.org/urn:lsid:zoobank.org:author:d98f606a-b273-4f50-95f5-c35f17b12c85 https://doi.org/10.5852/ejt.2017.391 https://doi.org/10.5852/ejt.2017.391 european journal of taxonomy 391: 1–44 (2017) 2 two authors independently re-established the genus grania in 1966. kennedy (1966) re-described g. maricola and g. macrochaeta and described g. americana kennedy, 1966 from off the coast of florida, while lasserre (1966) transferred michaelsena postclitellochaeta knöllner, 1935 to grania. one year later, considering the similarities among g. maricola, g. macrochaeta and g. americana, lassserre (1967) proposed to divide g. macrochaeta into four subspecies, g. m. macrochaeta, g. m. americana, g. m. maricola and g. m. roscoffensis lasserre, 1967. this situation lasted for about a decade, after which many additional species of grania were described from different parts of the world (erséus & lasserre 1976; erséus 1977, 1980, 1990; jamieson 1977; coates & erséus 1985; coates 1990; rota & erséus 1996, 1997; locke & coates 1998, 1999; de wit & erséus 2007; rota et al. 2007; de wit et al. 2009; prantoni et al. 2016), bringing a better understanding of species-specific characters and raising the subspecies proposed by lasserre (1967) back to species status (coates 1984; de wit 2010). more recently, prantoni et al. (2016) updated the phylogeny of the genus together with the descriptions and genetic data of nine new species, for the first time including species from africa and the east coast of south america (brazil) (see discussion). the genus is morphologically and genetically well separated from other enchytraeid genera (erséus et al. 2010; de wit et al. 2011; prantoni et al. 2016). to provide a global taxonomic overview of grania, the present checklist summarizes historical and recently published data, including all valid species described to date. moreover, the general geographical distribution patterns of the various species are briefly discussed. material and methods the checklist is arranged in chronological order and based on a bibliographic survey. all records of grania from published papers and monographs were reviewed. when available, additional information on habitat, geographical distribution and museum catalogue numbers is included. fig. 1. specimen of grania chilensis prantoni, de wit & erséus, 2016. photograph by pierre de wit. prantoni a. et al., global checklist of grania 3 abbreviations ams = australian museum, sydney, australia bamz = bermuda aquarium, natural history museum and zoo bj = barrie g.m. jamieson collection, queensland, australia bnmh = natural history museum, london, united kingdom mczr = museo civico di zoologia di roma, rome, italy mimb = a.v. zhirmunsky institute of marine biology, far eastern branch of the russian academy of sciences, vladivostok, russia mnhm = muséum national d’histoire naturelle, paris, france mzut = museo e istituto de zoologia del’università de torino, turin, italy nminh = national museum of ireland (department of natural history), dublin, ireland ntm = museum and art gallery of the northern territory, darwin, australia qm = queensland museum, brisbane, australia qvm = queen victoria museum, launceston, tasmania, australia romiz = invertebrate zoology, royal ontario museum, toronto, canada samc = iziko south african museum, cape town, south africa smnh = swedish museum of natural history, stockholm, sweden usnm = national museum of natural history, washington dc, usa wam = western australian museum, perth, australia zma = zoological museum of amsterdam, the netherlands zmub = zoological museum of the university of bergen, norway zuec = museu de zoologia da universidade estadual de campinas, campinas, brazil results we report 81 currently accepted species of grania, of which 49 are known from the southern and 32 from the northern hemisphere. the only species found in both hemispheres (atlantic ocean) is the deep-sea grania atlantica coates & erséus, 1985. almost half of the known southern species (24) occur in australia. on the other hand, 15 of the 32 species described from the northern hemisphere to date are european (fig. 1). list of all described species of grania in the world phylum annelida lamarck, 1809 class clitellata michaelsen, 1919 order enchytraeida vejdovský, 1879 family enchytraeidae vejdovský, 1879 genus grania southern, 1913 1. grania monochaeta (michaelsen, 1888) enchytraeus monochaetus michaelsen, 1888: 66–68, figs 6a–c. enchytraeus monochaetus – beddard 1895: 339. — michaelsen 1900: 91. marionina monochaeta – nielsen & christensen 1959: 109. grania monochaeta – lasserre 1967: 279–280. — rota & erséus 1997: 29–34, fig. 2, tab. 1 (lectotype designation). hemigrania monochaeta – lasserre 1971: 454. european journal of taxonomy 391: 1–44 (2017) 4 non michaelsena monochaeta – michaelsen 1921: 3. — stephenson 1932: 263, fig. 14 (see rota & erséus 1997). non grania monochaeta – erséus & lasserre 1977: 299–300, figs 1a–d. type material lectotype united kingdom: south georgia (mzut 123.1). paralectotypes united kingdom: south georgia (mzut olig. coll. 123.2 –123.4 ). other material united kingdom: south georgia (smnh 362). type locality united kingdom: south georgia, sw atlantic ocean. habitat intertidal, subtidal to 20 m, shelly detritus, among the roots of seaweeds and in the canal system of sponges. distribution only known from the type locality. 2. grania macrochaeta (pierantoni, 1901) enchytraeus macrochaetus pierantoni, 1901: 201–202. michaelsena macrochaeta – pierantoni 1903: 409–444, figs 1–28; 1915: 48–50. — stephenson 1930: 776–777. enchytraeus macrochaetus – nielsen & christensen 1959: 89–91, tab. 9. grania macrochaeta – kennedy 1966: 403–404 (redescription). — lasserre 1966: 312–314 (redescription). — erséus 1974: 90–93, tab. 1. — rota 1995: tab. 2. grania macrochaeta macrochaeta – lasserre 1967: 280. — erséus & lasserre 1976: 122, fig. 1, tab. 1. type material not designated. other material italy: bay of naples (mnhm ah 61–63). this refers to three specimens collected in june, 1967 by j. renaud-mornant and examined by erséus & lasserre (1976). type locality italy: bay of naples. habitat subtidal, 4–13 m, coarse sand. distribution only known from the type locality. prantoni a. et al., global checklist of grania 5 3. grania paucispina (eisen, 1904) michaelsena paucispina eisen, 1904: 74, fig. 43. michaelsena paucispina – michaelsen 1907: 130. marionina paucispina – nielsen & christensen 1959: 109. — lasserre 1971: 454. grania paucispina – erséus & lasserre 1976: 127. — coates & erséus 1980: 1037–1038, fig. 1. — coates & ellis 1981: 2134. type material typus perditus — specimen dissolutus. according to coates & erséus (1980), the type material deposited in the california academy of science in san francisco was destroyed during the earthquake and fire in 1906. other material united states: california (usnm 58906–58907). canada: british columbia (usnm 58907). type locality united states: santa barbara, california. habitat in muddy sand, 2–6 m, with much organic material, brackish water. distribution california, usa and british columbia, canada. 4. grania principissae (michaelsen, 1907) michaelsena principissae michaelsen, 1907: 129–131, pl. 1, figs 1–2. hemigrania principissae – lasserre 1971: 454. grania principissae – erséus & lasserre 1976: 128. — coates 1990: 28–30, figs 2, 8. type material typus amissus. other material australia: western australia, princess royal harbour (wam 69–89, romiz 11277). type locality australia: princess royal harbour, albany area, western australia. habitat intertidal, sand with organic debris accumulated in mussel bed. distribution only known from the type locality. european journal of taxonomy 391: 1–44 (2017) 6 5. grania maricola southern, 1913 grania maricola southern, 1913: 14, figs 1–7. michaelsena macrochaeta – pierantoni 1915 (part): 48–50. michaelsena maricola – stephenson 1930: 776–777. enchytraeus maricolus – nielsen & christensen 1959: 89–91, tab. 9. grania maricola – kennedy 1966: 400–402, fig. 2 (redescription). — lasserre 1966: 312–314 (redescription). — erséus 1974: 90–93; 1976: 35, tab. 3. — erséus & lasserre 1976: 124–125, figs 4–5, tab. 1. — bonomi & erséus 1984: 209, tab. 1. — coates 1984: 40, fig. 7a, tab. 1. — rodriguez 1986: 82–83, fig. 2. — rota 1995: tab. 2. — locke & coates 1998: 1111–1112, tab. 1. — de wit 2006: 25. — van haaren 2016: 140–141, figs 20, 56. grania macrochaeta maricola – lasserre 1967: 280. type material holotype ireland: county kerry (nminh 1913.415). paratypes ireland: county kerry (nminh 1909.151, nminh 1914.313). type locality ireland: county kerry. habitat subtidal, 8–80 m, coarse shell sand and shell gravel. distribution ireland, iberian peninsula, italy, the netherlands, norway and sweden. 6. grania postclitellochaeta (knöllner, 1935) michaelsena postclitellochaeta knöllner, 1935: 449–455, figs 19–25, tab. 1. michaelsena postclitellochaeta – hagen 1954: 12–13. marionina postclitellochaeta – nielsen & christensen 1959: 109–110, tab. 11. grania postclitellochaeta postclitellochaeta – lasserre 1966: 299–300, 312–314, tabs 1–2. — erséus 1976: 35, tab. 3. — erséus & lasserre 1976: 124, tab. 1. hemigrania postclitellochaeta – lasserre 1971: 454–456, fig. 3c. grania postclitellochaeta – kossmagk-stephan 1983: 598. — rota & erséus 2003: 232–234, figs 10c–d. — van haaren 2016: 141–143, figs 21, 23, 25–26, 57. type material not designated. other material france: bassin d’arcachon (smnh 45646–45651, mczr oligochaeta 0116–0117); gulf of biscay (mnhm ah 66–68; see erséus & lasserre 1976). north sea: smnh 45652–45654, mczr oligochaeta 0118. iceland: hafnafjörðdur (smnh 45655– 45660, mczr oligochaeta 0119; see rota & erséus 2003). prantoni a. et al., global checklist of grania 7 norway: tromsø (smnh 45661–45665, mczr oligochaeta 0120; see rota & erséus 2003); bergen (smnh 107746–107749; see de wit & erséus 2010). sweden: koster islands (smnh 107730, 107736, 107738–107745, 108220; see de wit & erséus 2010). type locality germany: kiel bay. habitat intertidal, subtidal, 20–100 m in coarse shell sand, shell gravel, sometimes in brackish water (baltic sea), tolerating a wide range of salinity (11–35 ppt). distribution baltic sea, iceland (see rota & erséus 2003), north sea (germany, the netherlands and belgium), france, norway and sweden. remarks according to van haaren (2016), it is not possible to distinguish grania postclitellochaeta from g. occulta de wit & erséus, 2010 and g. ovitheca erséus, 1977 on morphological grounds only. 7. grania americana kennedy, 1966 grania americana kennedy, 1966: 404–405, fig. 3. grania macrochaeta americana — lasserre 1967: 78–280. grania americana – erséus 1974: 90–93, tab. 1. — healy & coates 1999: 111, 114, tab. 1. — locke & coates 1999: 598–623, figs 16–20 (redescription); 2000: 619–620, 625–626, figs 4a, 5. —locke 2000: 83–93, figs 1–14. grania americana – erséus & lasserre 1976: p. 123 (nomen dubium, see erséus & lasserre 1976; locke & coates 1999). type material holotype bahamas: north bimini (usnm 33005). paratype bahamas: north bimini (usnm 33039). other material bahamas: pearl island (bmaz 1999 180 009). bermuda: ferry reach (usnm 185957). united states: hutchinson island (usnm 185958). belize: carrie bow cay (usnm 185959–185960). type locality bahamas: north bimini. european journal of taxonomy 391: 1–44 (2017) 8 habitat intertidal, subtidal to 10 m, fine to coarse sand. distribution belize, bermuda, bahamas, florida. 8. grania roscoffensis lasserre, 1967 grania macrochaeta roscoffensis lasserre, 1967: 277–280. grania roscoffensis – erséus 1974: 90–93, tab. 1. — erséus & lasserre 1976: 125, fig. 6, tab. 1 (amended). — coates 1984: 49. — rota & erséus 2003: 218–221, figs 3a–c, 4. non grania roscoffensis — erséus 1977: 294, tab. 1 (see rota & erséus 2003). type material holotype france: roscoff, harbour (mnhm ah 64). paratype the originally designated paratype ah 65 belongs to another species (see rota & erséus 2003). other material france: roscoff, harbour (smnh 45614). spain: canary islands (smnh 45615, zmav.ol 9345). type locality france: roscoff, harbour. habitat intertidal, coarse sand and gravel. distribution france, sweden and canary islands. 9. grania pusilla erséus, 1974 grania pusilla erséus, 1974: 87–94, figs 1–6, tab. 1. grania pusilla – erséus 1976: 34, tab. 3. — de wit et al. 2011: 513, figs 1–5, tab. 1. grania macrochaeta pusilla – erséus & lasserre 1976: 122, fig. 2. — erséus 1977: 294, tab. 1. — coates 1984: 49. — kossmagk–stephan 1985: 77–78. non grania pusilla – locke & coates 1998: 1107–1112, figs 6–12 (see rota & erséus 2003). type material holotype norway: vågegrunnen (zmub 55050). paratype norway: vågegrunnen (zmub 55051). prantoni a. et al., global checklist of grania 9 other material sweden: koster islands (smnh 107775–107788). norway: bergen (smnh 107789–107796). type locality norway: vågegrunnen, hjeltefjorden. habitat subtidal, 35–500 m, fine to coarse shelly sand. distribution west coasts of norway and sweden, morocco. 10. grania bermudensis erséus & lasserre, 1976 grania macrochaeta bermudensis erséus & lasserre, 1976: 122–124, fig. 3 tab. 1. grania macrochaeta bermudensis – erséus & lasserre 1976: 453. — coates 1984: 49, fig. 8a. grania bermudensis – locke & coates 1999: 609–614, figs 6, 12–15, tab. 1; 2000: 619–621, 626, fig. 6c. type material holotype bermuda: castle island (usnm 53202). paratype bermuda: castle island (usnm 53203). type locality bermuda: castle island. habitat subtidal, 8–15 m, medium to coarse coral sand and gravel. distribution only known from bermuda. 11. grania longiducta erséus & lasserre, 1976 grania postclitellochaeta longiducta erséus & lasserre, 1976: 127, fig. 7, tab. 1. hemigrania postclitellochaeta – lasserre 1971: 454–456, fig 3a–b, d . grania postclitellochaeta longiducta – erséus 1977: 296–297. — coates 1984: 49. grania longiducta – coates & erséus 1985: 113–114, fig. 8. — diaz et al. 1987: tab. 1, 3. — locke & coates 2000: 619, 625. type material holotype united states: massachusetts, cape cod bay (usnm 43482). paratype united states: massachusetts, cape cod bay (usnm 53201). european journal of taxonomy 391: 1–44 (2017) 10 type locality united states: cape cod bay, massachusetts. habitat subtidal, 42–78 m, medium to coarse sand. distribution cape cod bay, massachusetts, georges bank (se of massachusetts), off the coast of new jersey, delaware and maryland, usa. 12. grania monospermatheca erséus & lasserre, 1976 grania monospermatheca erséus & lasserre, 1976: 127, fig. 9, tab. 1. grania monospermatheca – coates 1984: 49, fig. 8b. — coates & erséus 1985: 114–115, fig. 9. — diaz et al. 1987: tab. 1–3. — locke & coates 2000: 619, 626–628. type material holotype united states: massachusetts, cape cod bay (usnm 53204). paratype united states: massachusetts, cape cod bay (usnm 53205). type locality united states: cape cod bay, massachusetts. habitat subtidal, 3–48 m, fine to coarse well-sorted sands and sand mixed with shell or shell gravel. distribution along north american atlantic coast, from cape cod bay, massachusetts to biscayne bay, florida, usa. 13. grania variochaeta erséus & lasserre, 1976 grania variochaeta erséus & lasserre, 1976: 125–126, figs 10–11, tab. 1. grania variochaeta – erséus 1976: 35, tab. 3; 1977: 297–298, tab. 1. — coates 1984: 46, fig. 6. — rota & erséus 2003: 211, 234–235, fig. 11. — van haaren 2016: 143–144, fig. 58. type material holotype sweden: kosterfjorden (smnh 3132). paratypes sweden: kosterfjorden (smnh 3133–3136). type locality sweden: kosterfjorden, bohuslän. prantoni a. et al., global checklist of grania 11 habitat subtidal, 20–140 m, heterogeneous sand. distribution west coast of norway and sweden, the netherlands. 14. grania ovitheca erséus, 1977 grania ovitheca erséus, 1977: 125, figs 5–7. grania ovitheca – bonomi & erséus 1984: 209, tab. 1. — rota 1995: tab. 2. — rota & erséus 2003: 230–233, figs 10a–b. — de wit & erséus 2010: 286–291. type material holotype sweden: kosterfjorden (smnh 3071). paratypes sweden: kosterfjorden (smnh 3072–3073). other material sweden: off gullmar fjord (smnh 107753, 107755–107758, 107760–107768). type locality sweden: kosterfjorden, bohuslän. habitat subtidal, 15–30 m, shell sand with gravel and pebbles or coarse sand with stones, pebbles and shells. distribution west coasts of norway and sweden, italy. 15. grania trichaeta jamieson, 1977 grania macrochaeta trichaeta jamieson, 1977: 345–347, fig. 5, pl. 1g. grania macrochaeta trichaeta – coates 1984: 46, fig. 5a. grania trichaeta – de wit et al. 2009: 28–30, figs 8–9, 10a–e (redescription). type material holotype australia: queensland, wistari reef (qm 8863). paratypes australia: queensland, wistari reef (qm 8866, bnmh 1976.1.21–23, bj 1975.7.76–78); queensland, heron reef (bj 1975.7.74–75, bj 1975.7.84). other material australia: queensland, lizard island (ams w.35554–w.35559, smnh 105540–105559); queensland, heron island (smnh 105560–105584). european journal of taxonomy 391: 1–44 (2017) 12 type locality australia: wistari reef, great barrier reef, queensland. habitat intertidal, subtidal to 7 m, fine to medium heterogeneous sand. distribution lizard island, heron island and wistari reef, great barrier reef, queensland, australia. 16. grania pacifica shurova, 1979 grania pacifica shurova, 1979: 84–86, fig. 6. type material holotype russia: iturup island, sea of okhotsk (mimb 16017). type locality russia: sea of okhotsk, iturup island. habitat subtidal, 15–20 m, gravelly sediment. distribution only known from the type locality. 17. grania incerta coates & erséus, 1980 grania incerta coates & erséus, 1980: 1038–1040, fig. 2. grania incerta – coates & ellis 1981: 2134–2135. — coates 1984: 46, fig. 4. type material holotype united states: california, santa barbara (usnm 58908). paratypes united states: california, santa barbara (usnm 58909). canada: british columbia, rennison island (usnm 58910). type locality united states: santa barbara, california. habitat subtidal, 3–17 m, well-sorted fine sand. distribution california, usa and british columbia, canada. prantoni a. et al., global checklist of grania 13 18. grania parvitheca erséus, 1980 grania parvitheca erséus, 1980: 27–28, fig. 1. type material holotype united kingdom: ascension island (usnm 58738). paratype united kingdom: ascension island (usnm 58739). type locality united kingdom: ascension island, s atlantic ocean. habitat intertidal, among rocks and clumps of tubes of sabellariidae. distribution only known from the type locality. 19. grania atlantica coates & erséus, 1985 grania atlantica coates & erséus, 1985: 112–113, fig. 7. grania atlantica – diaz et al. 1987: 222–224, tabs 1, 3–4. — locke & coates 2000: 619, 626. — rota & erséus 1996: 182; 2003: 210–211, 235–237, fig. 10e. — erséus & rota 2003: 898, tab. 1. type material holotype united states: off massachusetts (usnm 96503). paratypes united states: off massachusetts (usnm 96504–96508). type locality united states: off massachusetts. habitat continental slope, 744–1796 m, fine ooze to silty deep-sea sediments. distribution widely distributed in the atlantic ocean from 48º35.4´ n to 09º05´ s in the east and from 39º51.2´ n to 70º40.8´ s in the west. 20. grania levis coates & erséus, 1985 grania levis coates & erséus, 1985: 111–112, fig. 6. grania levis – diaz et al. 1987: tabs 1, 4. — locke & coates 2000: 626. ?grania cf. levis – prantoni et al. 2016: 502. european journal of taxonomy 391: 1–44 (2017) 14 type material holotype united states: georges bank (usnm 96509). paratypes united states: georges bank (usnm 96510); off new jersey (usnm 96511). other material united states: off north carolina (usnm 1283176; immature specimen, see prantoni et al. 2016). type locality united states: georges bank, se of massachusetts, nw atlantic ocean. habitat intertidal, subtidal to 79 m (probably to 492 m, see prantoni et al. 2016), medium to coarse sand. distribution georges bank se of massachusetts, on the continental shelf (and slope?) off new jersey and north carolina, usa. 21. grania reducta coates & erséus, 1985 grania reducta coates & erséus, 1985: 110–111, fig. 5. grania reducta – diaz et al. 1987: tabs 3–4. — locke & coates 2000: 626, 628. type material holotype united states: off maryland (usnm 96512). paratype united states: off maryland (usnm 96513). type locality united states: off maryland. habitat intertidal, subtidal to 65 m, medium to coarse sand. distribution continental shelf off new jersey, maryland and delaware, usa. 22. grania ascophora coates, 1990 grania ascophora coates, 1990: 23–25, fig. 5. type material holotype australia: western australia, baker bay (wam 69.89). paratype australia: western australia, baker bay (romiz i2880). prantoni a. et al., global checklist of grania 15 type locality australia: baker bay, king george sound, western australia. habitat subtidal, 4 m, fine sand with shells and seagrass. distribution barker bay and princess royal harbour, albany area, western australia. 23. grania bykane coates, 1990 grania bykane coates, 1990: 21–23, figs 2, 4a–d. grania bykane – rota et al. 2007: 1001–1004, figs 1a–g, 2a. type material holotype australia: western australia, princess royal harbour (wam 55.8). paratype australia: western australia, princess royal harbour (wam 56.89). type locality australia: princess royal harbour, albany area, western australia. habitat intertidal, subtidal to at least 6 m, fine to coarse sand and in sediments under boulders and in seagrass beds. distribution southern coast of western australia (albany and esperance areas), australia. 24. grania crassiducta coates, 1990 grania crassiducta coates, 1990: 20–21, figs 2, 3a–d. grania crassiducta – coates & stacey 1993: 404–406, fig. 9a–f. — rota et al. 2007: 1004–1006, figs 2b, 3a–f. type material holotype australia: western australia, princess royal harbour (wam 51.89). paratypes australia: western australia, princess royal harbour (wam 53.89, romiz i1279). type locality australia: princess royal harbour, albany area, western australia. habitat intertidal, muddy coarse sand, gravel and mixed sand with pebbles and coral. european journal of taxonomy 391: 1–44 (2017) 16 distribution southern (albany and esperance areas) and western (rottnest island) coasts of western australia. 25. grania ersei coates, 1990 grania ersei coates, 1990: 17–20, figs 1a–d, 2. grania ersei – coates & stacey 1993: 406–408, figs 10a–f. — rota et al. 2007: 1008–1011, figs 4d–g, 5a. type material holotype australia: western australia, princess royal harbour (wam 61.89). paratypes australia: western australia, princess royal harbour (wam 62.68–68.89, romiz i1273-i1276). type locality australia: princess royal harbour, albany area, western australia. habitat intertidal, subtidal to 26 m, sand among boulders and pebbles, and with algal debris. distribution south (albany, esperance) and west (rottnest island) coasts of western australia. 26. grania hastula coates, 1990 grania hastula coates, 1990: 26–28, fig. 7. type material holotype australia: western australia, middleton beach (usnm 120714). type locality australia: middleton beach, albany area, western australia. habitat intertidal, sand among rocks in algal wash. distribution only known from the type locality. 27. grania hyperoadenia coates, 1990 grania hyperoadenia coates, 1990: 25–26, fig. 6. grania hyperoadenia – de wit et al. 2009: 30–31, fig. 9. type material holotype australia: western australia, baker bay (wam 54.85). prantoni a. et al., global checklist of grania 17 other material australia: queensland, lizard island (ams w.35560). type locality australia: baker bay, king george sound, western australia. habitat subtidal, 1.5–4 m in sand. distribution albany area, western australia and lizard island, great barrier reef, queensland, australia. 28. grania hongkongensis erséus, 1990 grania hongkongensis erséus, 1990: 311–12, fig. 22. type material holotype china: hong kong (new territories), crooked island (bmnh 1987.3.39). paratypes china: hong kong (new territories), crooked island (bmnh 1987.3.40, smnh 3717). type locality china: mirs bay, crooked island, hong kong (new territories). habitat intertidal, subtidal to 15 m, shelly sand. distribution only known from the type locality. 29. grania inermis erséus, 1990 grania inermis erséus, 1990: 314–315, fig. 24. type material holotype china: hong kong (new territories), crooked island (bmnh 1987.3.42). paratypes china: hong kong (new territories), crooked island (bmnh 1987.3.43, smnh 3719). type locality china: mirs bay, crooked island, hong kong (new territories). habitat shelly sand, 7–14 m. distribution only known from the type locality. european journal of taxonomy 391: 1–44 (2017) 18 30. grania stilifera erséus, 1990 grania stilifera erséus, 1990: 312–314, fig. 23. type material holotype china: hong kong (new territories), crooked island (bmnh 1987.3.41). paratype china: hong kong (new territories), crooked island (smnh 3718). type locality china: mirs bay, crooked island, hong kong (new territories). habitat subtidal, 5–8 m, shelly sand. distribution only known from the type locality. 31. grania alliata coates & stacey, 1993 grania alliata coates & stacey, 1993: 397–399, figs 3–4. type material holotype australia: western australia, rottnest island (wam 192-92). type locality australia: rottnest island, western australia. habitat intertidal, gravelly sand. distribution only known from the type locality. 32. grania conjuncta coates & stacey, 1993 grania conjuncta coates & stacey, 1993: 402–404, figs 7–8. type material holotype australia: western australia, rottnest island (wam 193-92). type locality australia: rottnest island, western australia. habitat subtidal, 2 m, medium to coarse sand. prantoni a. et al., global checklist of grania 19 distribution only known from the type locality. 33. grania longistyla coates & stacey, 1993 grania longistyla coates & stacey, 1993: 394–397, figs 1–2. type material holotype australia: western australia, rottnest island (wam 194-92). paratype australia: western australia, rottnest island (wam 195-92). type locality australia: rottnest island, western australia. habitat intertidal, gravelly sand. distribution only known from the type locality. 34. grania vacivasa coates & stacey, 1993 grania vacivasa coates & stacey, 1993: 400–402, figs 5–6. grania vacivasa – rota et al. 2007: 1018–1020, figs 8c, 9a–e. type material holotype australia: western australia, rottnest island (wam 196-92). paratype australia: western australia, rottnest island (wam 197-92). type locality australia: rottnest island, western australia. habitat subtidal, 1 m, in coarse sand. distribution south (esperance) and west (rottnest island) coasts of western australia. 35. grania acanthochaeta rota & erséus, 1996 grania acanthochaeta rota & erséus, 1996: 174–175, fig. 4, tab. 1. type material holotype antarctica: ross island (usnm 172142). european journal of taxonomy 391: 1–44 (2017) 20 paratype antarctica: ross island (usnm 172193). other material antarctica: ross island (usnm 172194–172397). type locality antarctica: ross island. habitat subtidal, 38 m, greyish brown gravelly mud with sponge spicules and valves of limatula. distribution only known from the type locality. 36. grania algida rota & erséus, 1996 grania algida rota & erséus, 1996: 179–181, fig. 8, tab. 1. type material holotype antarctica: ross island (usnm 172398). paratype antarctica: ross island (mzr oligochaeta 0065). type locality antarctica: ross island. habitat subtidal, 14–40 m, volcanic gravel and cobble. distribution only known from the type locality. 37. grania angustinasus rota & erséus, 1996 grania angustinasus rota & erséus, 1996: 177–178, figs 3b, 6, tab. 1. type material holotype antarctica: ross island (mczr oligochaeta 0059). paratypes antarctica: ross island (mczr 0060–0063, smnh 4759–4761). type locality antarctica: ross island. habitat subtidal, 35–126 m, fine sand. prantoni a. et al., global checklist of grania 21 distribution only known from the type locality. 38. grania antarctica rota & erséus, 1996 grania antarctica rota & erséus, 1996: 178–179, figs 3c, 7, tab. 1. type material holotype antarctica: ross island (usnm 172400). paratype antarctica: ross island (usnm 172402, mczr oligochaeta 0059). type locality antarctica: ross island. habitat subtidal, 14–31 m, volcanic gravel and cobble. distribution only known from the type locality. 39. grania carchinii rota & erséus, 1996 grania carchinii rota & erséus, 1996: 175–177, fig. 5, tab. 1. type material holotype antarctica: ross island (mczr oligochaeta 0057). paratype antarctica: ross island (mczr oligochaeta 0058). type locality antarctica: ross island. habitat subtidal, 35 m, fine sand with mica shale, shell debris and some pebbles. distribution only known from the type locality. 40. grania hirsuticauda rota & erséus, 1996 grania hirsuticauda rota & erséus, 1996: 175–177, fig. 5, tab. 1. type material holotype antarctica: ross island (usnm 172136). paratypes antarctica: ross island (usnm 172137–172138). european journal of taxonomy 391: 1–44 (2017) 22 type locality antarctica: ross island. habitat subtidal, 5–585 m, volcanic gravel, sandy mud, small rocks, ectoproct and sponge debris. distribution only known from the type locality. 41. grania darwinensis (coates & stacey, 1997) randidrilus darwinensis coates & stacey, 1997: 70–72, fig. 1. grania darwinensis – rota et al. 2003: 504–509, fig. 3. type material holotype australia: northern territory, darwin (ntm wo 0084). paratypes australia: northern territory, darwin (ntm wo 0085–0087, romiz i2457–i2458). type locality australia: darwin, northern territory. habitat intertidal, subtidal to 16 m, medium to coarse sand, clay or silty sediments. distribution northern territory and western australia. 42. grania eurystila coates & stacey, 1997 grania eurystila coates & stacey, 1997: 73–74, fig. 2. type material holotype australia: northern territory, darwin (ntm wo 0081). paratypes australia: northern territory, darwin (ntm wo 0082–0083, romiz i2479). type locality australia: darwin, northern territory. habitat intertidal rockpool. distribution only known from two locations in the inner part of darwin harbour, northern territory, australia. prantoni a. et al., global checklist of grania 23 43. grania integra coates & stacey, 1997 grania integra coates & stacey, 1997: 74–76, fig. 3. grania integra – rota et al. 2003: 499–501, fig. 1. type material holotype australia: northern territory, darwin (ntm wo 0079). paratype australia: northern territory, darwin (ntm wo 0080). type locality australia: darwin, northern territory. habitat intertidal crevices with sand gravel, pebbles and heterogeneous sediments. distribution darwin harbour, northern territory and nickol bay, dampier area, western australia. 44. grania lasserrei rota & erséus, 1997 grania lasserrei rota & erséus, 1997: 34–37, fig. 3, tab.1. enchytraeus monochaetus – michaelsen 1888: 66, figs 6a–c; part., pl. 2. michaelsena monochaeta – michaelsen 1921: 3, part. grania monochaeta – erséus & lasserre 1977: 299–300, figs 1a–d. type material holotype united kingdom: south georgia (smnh 4803). paratypes united kingdom: south georgia (smnh 48044806, mzut 1078, bmnh 1996:916). type locality united kingdom: south georgia, sw atlantic ocean. habitat intertidal, subtidal to 20 m. distribution only known from the type locality. 45. grania stephensoniana rota & erséus, 1997 grania stephensoniana rota & erséus, 1997: 37–39, figs 4–5, tab. 1. michaelsena monochaeta – stephenson 1932: 263, fig. 14. european journal of taxonomy 391: 1–44 (2017) 24 type material holotype united kingdom: south georgia (bmnh 1931.6.23.78). paratypes united kingdom: south georgia (bmnh 1933.2.23.946–1933.2.23.948). type locality united kingdom: south georgia, sw atlantic ocean. habitat 60 m, rocky bottom. distribution only known from the type locality. 46. grania mira locke & coates, 1998 grania mira locke & coates, 1998: 1103–1107, figs 1–5. grania sp. – healy 1996a: 53, 56–57, fig. 1, tabs 1–2; 1996b: 1287. type material holotype ireland: carnsore point (nmi 4.1998). paratypes ireland: carnsore point (nmi 5.1998–6.1998). type locality ireland: carnsore point, county wexford. habitat intertidal, sediments trapped in dense turf of corallina officinalis on horizontal or gently sloping rock. distribution only known from the type locality. 47. grania hylae locke & coates, 1999 grania hylae locke & coates, 1999: 605–609, figs 6–7, 11, tab. 1. grania hylae – locke & coates 2000: 619–621, 626, fig. 4b. type material holotype bermuda: paget island (usnm 185954). paratypes united states: florida, fowey rocks (usnm 185955). bermuda: castle island (usnm 185956, bamz 199 180 007). prantoni a. et al., global checklist of grania 25 type locality bermuda: paget island. habitat intertidal, subtidal to 17 m, medium to coarse sand with rocks. distribution rocky hill park, castle island and paget island, bermuda, and fowey rocks, miami, florida, usa. 48. grania laxartus locke & coates, 1999 grania laxartus locke & coates, 1999: 602–605, figs 2–6, 11, tab. 1. grania laxartus – locke & coates 2000: 619–621, 626–627, figs 4c, 6a. type material holotype bermuda: ferry point bridge (usnm 185951). paratypes bermuda: ferry reach (usnm 185952, bamz 199 180 006). belize: carrie bow cay (usnm 185953). type locality bermuda: ferry point bridge. habitat intertidal pools with accumulation of sand and fine to medium-coarse calcareous sand. distribution ferry point bridge, whalebone bay, pearl island, ferry reach and smith’s sound, bermuda, and carrie bow cay, belize. 49. grania dolichura rota & erséus, 2000 grania dolichura rota & erséus, 2000: 249–252, fig. 3. grania dolichura – rota et al. 2007: 1006–1008, figs 4a–c. type material holotype australia: tasmania, little musselroe (qvm 14: 3889). paratypes australia: tasmania, little musselroe (qvm 14:3890–14:3897, smnh 5203–5206, mczr oligochaeta 0085–0088). type locality australia: little musselroe, tasmania. european journal of taxonomy 391: 1–44 (2017) 26 habitat intertidal, silt-clay sediments. distribution widespread around tasmania, australia. 50. grania tasmaniae rota & erséus, 2000 grania tasmaniae rota & erséus, 2000: 247–249, fig. 2. type material holotype australia: tasmania, low head (qvm 14: 3887). paratype australia: tasmania, low head (qvm 14: 3888). type locality australia: low head, tamar estuary, tasmania. habitat intertidal, silt-clay sediments. distribution only known from the type locality. 51. grania aquitana rota & erséus, 2003 grania aquitana rota & erséus, 2003: 226–229, fig. 7, tab. 1. type material holotype france: bassin d’arcachon (smnh 5729). paratypes france: bassin d’arcachon (smnh 5730–5733). type locality france: bassin d’arcachon. habitat subtidal, 2–5 m, fine sand. distribution only known from the type locality. 52. grania canaria rota & erséus, 2003 grania canaria rota & erséus, 2003: 213–215, fig. 1, tab. 1. prantoni a. et al., global checklist of grania 27 type material holotype spain: tenerife (zma v.ol 9344). paratypes spain: tenerife (smnh 5710–5711). type locality spain: tenerife, canary islands. habitat intertidal, sand and gravel. distribution only known from the type locality. 53. grania fortunata rota & erséus, 2003 grania fortunata rota & erséus, 2003: 215–218, fig. 2, tab. 1. type material holotype spain: tenerife (smnh 5712). paratype spain: tenerife (smnh 5713, mczr oligochaeta 0103–0104). type locality spain: tenerife, canary islands. habitat subtidal, 12–17 m, fine and muddy sands associated with beds of the seagrass cymodocea nodosa. distribution only known from the type locality. 54. grania mauretanica rota & erséus, 2003 grania mauretanica rota & erséus, 2003: 224–226, fig. 6, tab. 1. type material holotype morocco: off casablanca (smnh 5718). paratypes morocco: off casablanca (smnh 5719–5720, mczr oligochaeta 0107). type locality morocco: off casablanca. european journal of taxonomy 391: 1–44 (2017) 28 habitat subtidal, 173 m, mud with shell debris. distribution only known from the type locality. 55. grania papillinasus rota & erséus, 2003 grania papillinasus rota & erséus, 2003: 239–240, fig. 13. type material holotype france: gulf of gascogne (smnh 5726). paratypes france: gulf of gascogne (smnh 5727–5728, mczr oligochaeta 0124–0126). type locality france: gulf of gascogne, lower continental slope. habitat deep sea, 2630–2885 m, most likely very fine sediments. distribution gulf of gascogne, off france and off the eastern usa (i.e., both sides of the north atlantic) (see erséus & rota 2003). 56. grania torosa rota & erséus, 2003 grania torosa rota & erséus, 2003: 237–239, fig. 12. type material holotype ne atlantic ocean: rockall trough (smnh 5721). paratypes ne atlantic ocean: rockall trough (smnh 5722–5725, mczr oligochaeta 0123). type locality ne atlantic ocean: rockall trough, off scotland. habitat continental slope, 1170–1800 m, fine sandy and hemi-pelagic ooze. distribution northern rockall trough, off the coast of scotland, to near the entrance to the english channel (ne atlantic ocean). 57. grania vikinga rota & erséus, 2003 grania vikinga rota & erséus, 2003: 222–224, fig. 5. prantoni a. et al., global checklist of grania 29 grania roscoffensis (part) – sensu erséus 1977: 294, tab. 1. grania vikinga – van haaren 2016: 144–145, figs 22, 24, 27–28, 59. type material holotype sweden: skagerrak (smnh 5714). paratypes sweden: skagerrak (smnh 5716–5717, mczr oligochaeta 0105–0106). type locality sweden: skagerrak, bohuslän. habitat subtidal, 10–46 m, sand. distribution west coast of sweden, the netherlands. 58. grania ocarina rota, erséus & wang, 2003 grania ocarina rota, erséus & wang, 2003: 502–504, fig. 2. type material holotype australia: western australia, withnell bay (wam v 4351). paratype australia: western australia, withnell bay (wam v 4352, smnh 5868, mczr oligochaeta 0128). type locality australia: withnell bay, dampier area, western australia. habitat barely subtidal, 0.5 m, medium to coarse sand. distribution only known from the type locality. 59. grania cinctura de wit & erséus, 2007 grania cinctura de wit & erséus, 2007: 33–36, fig. 3, tab. 1. type material holotype france: new caledonia, lifou (smnh 6559). paratypes france: new caledonia, lifou (smnh 6560–6564); new caledonia, touho (smnh 6565–6568, 6572). european journal of taxonomy 391: 1–44 (2017) 30 type locality france: lifou, loyalty islands, new caledonia. habitat intertidal, subtidal to 6 m, fine to coarse sand. distribution touho and nouméa areas, and loyalty islands, new caledonia. 60. grania curta de wit & erséus, 2007 grania curta de wit & erséus, 2007: 38–40, fig. 5, tab. 1. type material holotype france: new caledonia, lifou (smnh 6583). paratypes france: new caledonia, lifou (smnh 6584–6588). type locality france: lifou, loyalty islands, new caledonia. habitat barely subtidal, 0.5 m, heterogeneous sand. distribution only known from the type locality. 61. grania fiscellata de wit & erséus, 2007 grania fiscellata de wit & erséus, 2007: 45–47, fig. 9, tab. 1. type material holotype france: new caledonia, touho (smnh 6610). paratypes france: new caledonia, touho (smnh 6611–6613); new caledonia, lifou (smnh 6617). type locality france: touho, loyalty islands, new caledonia. habitat intertidal, subtidal to 3 m, heterogeneous sand. distribution touho area and lifou (loyalty islands), new caledonia. prantoni a. et al., global checklist of grania 31 62. grania fustata de wit & erséus, 2007 grania fustata de wit & erséus, 2007: 40–42, fig. 6, tab. 1. type material holotype france: new caledonia, touho (smnh 6589). paratypes france: new caledonia, touho (smnh 6590–6598). type locality france: touho, loyalty islands, new caledonia. habitat intertidal and barely subtidal, coarse sand and gravel. distribution only known from the type locality. 63. grania galbina de wit & erséus, 2007 grania galbina de wit & erséus, 2007: 36–38, fig. 4, tab. 1. type material holotype france: new caledonia, lifou (smnh 6573). paratypes france: new caledonia, lifou (smnh 6574–6582). type locality france: lifou, loyalty islands, new caledonia. habitat intertidal, subtidal to 13 m, heterogeneous sand. distribution nouméa area and lifou (loyalty islands), new caledonia. 64. grania novacaledonia de wit & erséus, 2007 grania novacaledonia de wit & erséus, 2007: 31–33, fig. 2, tab. 1. type material holotype france: new caledonia, touho (smnh 6549). paratypes france: new caledonia, touho (smnh 6550–6558). european journal of taxonomy 391: 1–44 (2017) 32 type locality france: touho, loyalty islands, new caledonia. habitat intertidal, subtidal to 21 m, heterogeneous sand. distribution touho and nouméa areas, new caledonia. 65. grania papillata de wit & erséus, 2007 grania papillata de wit & erséus, 2007: 42–45, figs 7–8, tab. 1. type material holotype france: new caledonia, lifou (smnh 6599). paratypes france: new caledonia, lifou (smnh 6600–6602); new caledonia, touho (smnh 6603–6609). type locality france: lifou, loyalty islands, new caledonia. habitat intertidal, subtidal to 22 m, heterogeneous sand. distribution touho area and lifou (loyalty islands), new caledonia. 66. grania quaerens rota, wang & erséus, 2007 grania quaerens rota, wang & erséus, 2007: 1011–1013, figs 5b–d, 6a–i. type material holotype australia: western australia, new island (wam v 7315). paratypes australia: western australia, new island (wam v 7316–7319, smnh 6803–6808, mczr oligochaeta 0146–0149). type locality australia: new island, western australia. habitat intertidal, medium to coarse sand. distribution south coast of western australia. prantoni a. et al., global checklist of grania 33 67. grania sperantia rota, wang & erséus, 2007 grania sperantia rota, wang & erséus, 2007: 1014–1017, figs 7a–h, 8a–b. type material holotype australia: western australia, lucky bay (wam v 7320). paratypes australia: western australia, lucky bay (wam v 7321–7326, smnh 6809–6817, mczr oligochaeta 0150–0155). type locality australia: lucky bay, western australia. habitat barely subtidal, 0.5–2 m, medium to coarse sand. distribution south coast of western australia. 68. grania breviductus de wit, rota & erséus, 2009 grania breviductus de wit, rota & erséus, 2009: 19–21, figs 2, 10a. type material holotype australia: queensland, heron island (ams w.35536). paratypes australia: queensland, heron island (ams w.35537–.35542, smnh 7761–7766). type locality australia: heron island, great barrier reef, queensland, australia. habitat intertidal, coarse sand. distribution only known from the type locality. 69. grania regina de wit, rota & erséus, 2009 grania regina de wit, rota & erséus, 2009: 21–23, figs 3–4, 10b. type material holotype australia: queensland, heron island (ams w.35543). type locality australia: heron island, great barrier reef, queensland, australia. european journal of taxonomy 391: 1–44 (2017) 34 habitat subtidal, 15 m, fine sand. distribution only known from the type locality. 70. grania homochaeta de wit, rota & erséus, 2009 grania homochaeta de wit, rota & erséus, 2009: 23–25, figs 5, 10c. type material holotype australia: queensland, heron island (ams w.35544). paratype australia: queensland, heron island (mnh 7767). type locality australia: heron island, great barrier reef, queensland, australia. habitat subtidal, 18 m, gravelly fine sand. distribution only known from the type locality. 71. grania colorata de wit, rota & erséus, 2009 grania colorata de wit, rota & erséus, 2009: 25–27, figs 6–7, 10d. type material holotype australia: queensland, heron island (ams w.35545). paratypes australia: queensland, heron island (ams w.35546–.35553, smnh 7768–7772). type locality australia: heron island, great barrier reef, queensland, australia. habitat subtidal, 7 m, in heterogeneous sand. distribution only known from the type locality. 72. grania occulta de wit & erséus, 2010 grania occulta de wit & erséus, 2010: 287–289, fig. 3. prantoni a. et al., global checklist of grania 35 type material holotype sweden: gullmar fjord (smnh 7844). type locality sweden: gullmar fjord, bohuslän. habitat subtidal, 10–25 m, shell sand with some mud. distribution only known from the type locality. 73. grania brasiliensis prantoni, de wit & erséus, 2016 grania brasiliensis prantoni, de wit & erséus, 2016: 489–491, fig. 1. type material holotype brazil: paraná state, paranaguá bay (zuec cli 04). paratypes brazil: paraná state, paranaguá bay (zuec cli 05); são paulo state, são paulo (zuec cli 06–07). type locality brazil: paranaguá bay, paraná state. habitat intertidal, subtidal to 7 m, medium to coarse sand with some mud and lots of mollusc and barnacle shells. distribution coasts of paraná and são paulo states, brazil. 74. grania bekkouchei prantoni, de wit & erséus, 2016 grania bekkouchei prantoni, de wit & erséus, 2016: 491–492, figs 2, 4a–c. type material holotype south africa: western cape (samc a82466). type locality south africa: western cape. habitat intertidal, coarse sand in rock crevice. distribution only known from the type locality. european journal of taxonomy 391: 1–44 (2017) 36 75. grania cryptica prantoni, de wit & erséus 2016 grania cryptica prantoni, de wit & erséus, 2016: 492–493, figs 3, 4d–f. type material holotype south africa: western cape (samc a82473). type locality south africa: western cape. habitat lower intertidal, rockpool. distribution only known from the type locality. 76. grania capensis prantoni, de wit & erséus, 2016 grania capensis prantoni, de wit & erséus, 2016: 493–495, fig. 5. type material holotype south africa: western cape (samc a82474). paratype south africa: western cape (samc a82475). type locality south africa: western cape. habitat lower intertidal, rockpool. distribution only known from the type locality. 77. grania simonae prantoni, de wit & erséus, 2016 grania simonae prantoni, de wit & erséus, 2016: 495–497, fig. 6. type material holotype south africa: western cape (samc a82476). paratypes south africa: western cape (samc 82477–82482). type locality south africa: western cape. prantoni a. et al., global checklist of grania 37 habitat intertidal, crevice between rocks. distribution only known from the type locality. 78. grania hinojosai prantoni, de wit & erséus, 2016 grania hinojosai prantoni, de wit & erséus, 2016: 497–498, fig. 7. grania sp. chile 1– de wit et al. 2011: 513. type material holotype chile: coquimbo (zuec cli 08). paratypes chile: coquimbo (zuec cli 09–12). type locality chile: coquimbo, elqui. habitat intertidal, sand among rocks. distribution puerto aldea to pampilla point, coquimbo, elqui, chile. 79. grania chilensis prantoni, de wit & erséus, 2016 grania chilensis prantoni, de wit & erséus, 2016: 498–500, fig. 8. grania sp. chile 2 – de wit et al. 2011: 513, 517. type material holotype chile: valdivia (zuec cli 13). paratypes chile: valdivia (zeuc cli 14–19). type locality chile: valdivia. habitat intertidal, sand among rocks and heterogeneous sand with organic material. distribution along coast of chile, from about 30.3° to 39.8° s. european journal of taxonomy 391: 1–44 (2017) 38 80. grania unitheca prantoni, de wit & erséus, 2016 grania unitheca prantoni, de wit & erséus, 2016: 500–501, fig. 9. type material holotype united states: off north carolina (usnm 1283175). type locality united states: off north carolina. habitat subtidal, 17 m, sand. distribution only known from the type locality. 81. grania carolinensis prantoni, de wit & erséus, 2016 grania carolinensis prantoni, de wit & erséus, 2016: 501–502, fig. 10. type material holotype united states: off north carolina (usnm 1283174). type locality united states: off north carolina. habitat continental shelf slope, 492 m, sand. distribution only known from the type locality. discussion despite the many species of grania described from the australian continent and a few other southern regions of the globe (e.g., new caledonia and antarctica, see fig. 2), taxa from the african and south american continents have been completely ignored until recently. this situation changed with the seven new species recently described from brazil (1), chile (2) and south africa (4 spp.) (prantoni et al. 2016). the geographical distribution of the various species of grania is strongly concordant with the phylogeny of the genus, as estimated by analyses of molecular data (de wit et al. 2011; prantoni et al. 2016). most of the individual species appear to be endemic to rather small geographical areas, which suggests a limited capability of dispersion. however, as a whole, the genus is divided into at least three distinct evolutionary lineages, each with a broad, but geographically coherent distribution in the world; this is based on a sample of 28 genetically analyzed species (prantoni et al. 2016). the first lineage (green numbers on fig. 2) comprises species from the atlantic ocean (including a sublineage of four species from south africa). the second (blue numbers) are species from australia and southern asia (hong prantoni a. et al., global checklist of grania 39 kong), and the third are species from the south pacific and atlantic regions (red numbers). an interesting aspect is the phylogenetic placement of g. americana in the north atlantic and g. brasiliensis in the south atlantic. these two species belong to an otherwise pacific group (the third lineage), and it is suggested that they share a common ancestor that migrated from the pacific region before the closing of the isthmus of panama 3 ma (de wit et al. 2011; prantoni et al. 2016). however, as most neotropical coasts remain unexplored, additional species of grania from both sides of south america are needed to corroborate (or refute) this hypothesis. in conclusion, more intense sampling efforts in many parts of the world are crucial to enable further studies of the evolutionary and biogeographical history of grania. the 81 species described to date is a high number for a marine genus of enchytraeidae, and yet, this number certainly does not represent the actual diversity of the genus grania. overall, our systematic knowledge of marine clitellates (oligochaetes and leeches) is poor, with perhaps only about 10% of the species diversity known (appeltans et al. 2012). the lack of specialists around the world is evidently one of the reasons for this (prantoni et al. 2014), but it is also a threat to any rapid improvement of the situation. as a partial solution, the combined efforts of taxonomists and ecologists may come as a first and necessary step towards a better understanding of the group as well as of marine clitellate species diversity as a whole. acknowledgements we thank pierre de wit for permission to publish his photo of a specimen of grania chilensis, and klara dozsa-farkas and one anonymous reviewer for good suggestions regarding the final version of the text. this study was supported by capes brazilian foundation (capes-process: bex 11676/13-2), which provided a phd fellowship for the first author. the second author was supported by the swedish taxonomy initiative (artdatabanken). fig. 2. world map showing the description localities, and the three evolutionary lineages (color-marked) of grania spp. genetically analyzed by prantoni et al. (2016: clades a, b and c). green numbers = atlantic species (clade a); red numbers = south pacific and atlantic species (clade b); blue numbers = australian and asian species (clade c); black numbers = species presently without molecular data, i.e., not yet allocated to any particular lineage. european journal of taxonomy 391: 1–44 (2017) 40 references appeltans w., ahyong s.t., anderson g., angel m.v., artois t., bailly n., bamber r., barber a., bartsch i., berta a., błażewicz-paszkowycz m., bock p., boxshall g., boyko c.b., brandão s.n., bray r.a., bruce n.l., cairns s.d., chan t.-y., cheng l., collins a.g., cribb t., curini-galletti m., dahdouh-guebas f., davie p.j.f., dawson m.n., de clerck o., decock w., de grave s., de voogd n.j., domning d.p., emig c.c., erséus c., eschmeyer w., fauchald k., fautin d.g., feist s.w., fransen c.h.j.m., furuya h., garcia-alvarez o., gerken s., gibson d., gittenberger a., gofas s., gómezdaglio l., gordon d.p., guiry m.d., hernandez f., hoeksema b.w., hopcroft r.r., jaume d., kirk p., koedam n., koenemann s., kolb j.b., kristensen r.m., kroh a., lambert g., lazarus d.b., lemaitre r., longshaw m., lowry j., macpherson e., madin l.p., mah c., mapstone g., mclaughlin p.a., mees j., meland k., messing c.g., mills c.e., molodtsova t.n., mooi r., neuhaus b., ng p.k.l., nielsen c., norenburg j., opresko d.m., osawa m., paulay g., perrin w., pilger j.f., poore g.c.b., pugh p., read g.b., reimer j.d., rius m., rocha r.m., saiz-salinas j.i., scarabino v., schierwater b., schmidt-rhaesa a., schnabel k.e., schotte m., schuchert p., schwabe e., segers h., self-sullivan c., shenkar n., siegel v., sterrer w., stöhr s., swalla b., tasker m.l., thuesen e.v., timm t., todaro m.a., turon x., tyler s., uetz p., van der land j., vanhoorne b., van ofwegen l.p., van soest r.w.m., vanaverbeke j., walker-smith g., walter t.c., warren a., williams g.c., wilson s.p., costello m.j. 2012 the magnitude of global marine species diversity. current biology 22: 2189–2202. https://doi.org/10.1016/j.cub.2012.09.036 beddard f.e. 1895. a monograph of the order oligochaeta. clarendon press, oxford. bonomi g. & erséus c. 1984. a taxonomic and faunistic survey of the marine tubificidae and enchytraeidae (oligochaeta) of italy. introduction and preliminary results. hydrobiologia 115: 207– 210. https://doi.org/10.1007/bf00027918 coates k.a. 1984. specific criteria in grania (oligochaeta, enchytraeidae). hydrobiologia 115: 45–50. https://doi.org/10.1007/bf00027891 coates k.a. 1990. marine enchytraeidae (oligochaeta, annelida) of the albany area, western australia. in: wells f., walker d., kirkman h. & lethbridge r. 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48: oligochaeta. proceedings of the royal irish academy 31: 1–15. stephenson j. 1930. the oligochaeta. clarendon press, oxford. stephenson j. 1932. oligochaeta. part i. microdrili. discovery reports 4: 233–264. van haaren t. 2016. oligochaeten van brakke en zoute wateren in nederland (annelida: oligochaeta). nederlandse faunistische mededelingen 46: 155–164. manuscript received: 25 may 2017 manuscript accepted: 5 july 2017 published on: 29 december 2017 topic editor: rudy jocqué desk editor: danny eibye-jacobsen printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain. european journal of taxonomy 283: 1–25 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2017.283 www.europeanjournaloftaxonomy.eu 2017 · philippe h. et al. this work is licensed under a creative commons attribution 3.0 license. d n a l i b r a r y o f l i f e , o p i n i o n p a p e r urn:lsid:zoobank.org:pub:4f1838fa-dd9d-402f-8d57-d5d38715d6e3 1 pitfalls in supermatrix phylogenomics hervé philippe 1,*, damien m. de vienne 2, vincent ranwez 3, béatrice roure 4, denis baurain 5 & frédéric delsuc 6 1,4 centre de théorisation et de modélisation de la biodiversité, station d’ecologie théorique et expérimentale, umr cnrs 5321, 09200 moulis, france. 1 département de biochimie, centre robert-cedergren, université de montréal, montréal, h3c 3j7 québec, canada. 2 laboratoire de biométrie et biologie evolutive, cnrs, umr 5558, université lyon 1, 69622 villeurbanne, france. 3 supagro, umr agap, 34398 montpellier, france. 5 inbios-phytosystems – eukaryotic phylogenomics, université de liège, liège, belgium. 6 institut des sciences de l’evolution, umr 5554, cnrs, ird, ephe, université de montpellier, montpellier, france. * corresponding author: herve.philippe@sete.cnrs.fr 2 email: damien.de-vienne@univ-lyon1.fr 3 email: vincent.ranwez@supagro.fr 4 email: bea.roure@gmail.com 5 email: denis.baurain@ulg.ac.be 6 email: frederic.delsuc@univ-montp2.fr 1 urn:lsid:zoobank.org:author:9a29f58c-d3e2-44da-9e38-7270e08b4264 2 urn:lsid:zoobank.org:author:1cf9e4f6-fc15-4e19-bf23-de2b132df2b1 3 urn:lsid:zoobank.org:author:48c8b062-e3e6-4d25-a12e-5f104bc071e3 4 urn:lsid:zoobank.org:author:db8adb04-d982-4a20-b559-c026c040962e 5 urn:lsid:zoobank.org:author:a6b5c24f-ea7b-4928-a89c-385054a02383 6 urn:lsid:zoobank.org:author:95d3fba1-cfd3-481d-8f38-a45bc3d7a7ad abstract. in the mid-2000s, molecular phylogenetics turned into phylogenomics, a development that improved the resolution of phylogenetic trees through a dramatic reduction in stochastic error. while some then predicted “the end of incongruence”, it soon appeared that analysing large amounts of sequence data without an adequate model of sequence evolution amplifies systematic error and leads to phylogenetic artefacts. with the increasing flood of (sometimes low-quality) genomic data resulting from the rise of high-throughput sequencing, a new type of error has emerged. termed here “data errors”, it lumps together several kinds of issues affecting the construction of phylogenomic supermatrices (e.g., sequencing and annotation errors, contaminant sequences). while easy to deal with at a single-gene scale, such errors become very difficult to avoid at the genomic scale, both because hand curating thousands of sequences is prohibitively time-consuming and because the suitable automated bioinformatics tools are still in their infancy. in this paper, we first review the pitfalls affecting the construction of supermatrices http://dx.doi.org/10.5852/ejt.2017.283 www.europeanjournaloftaxonomy.eu https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:4f1838fa-dd9d-402f-8d57-d5d38715d6e3 mailto:herve.philippe%40sete.cnrs.fr?subject= mailto:damien.de-vienne%40univ-lyon1.fr?subject= mailto:vincent.ranwez%40supagro.fr?subject= mailto:bea.roure%40gmail.com?subject= mailto:denis.baurain%40ulg.ac.be?subject= mailto:frederic.delsuc%40univ-montp2.fr?subject= http://zoobank.org/urn:lsid:zoobank.org:author:9a29f58c-d3e2-44da-9e38-7270e08b4264 http://zoobank.org/urn:lsid:zoobank.org:author:1cf9e4f6-fc15-4e19-bf23-de2b132df2b1 http://zoobank.org/urn:lsid:zoobank.org:author:48c8b062-e3e6-4d25-a12e-5f104bc071e3 http://zoobank.org/urn:lsid:zoobank.org:author:db8adb04-d982-4a20-b559-c026c040962e http://zoobank.org/urn:lsid:zoobank.org:author:a6b5c24f-ea7b-4928-a89c-385054a02383 http://zoobank.org/urn:lsid:zoobank.org:author:95d3fba1-cfd3-481d-8f38-a45bc3d7a7ad european journal of taxonomy 283: 1–25 (2017) 2 and the strategies to limit their adverse effects on phylogenomic inference. then, after discussing the relative non-issue of missing data in supermatrices, we briefly present the approaches commonly used to reduce systematic error. keywords. phylogenomics, supermatrix, systematic error, data quality, incongruence. philippe h., de vienne d.m., ranwez v., roure b., baurain d. & delsuc f. 2017. pitfalls in supermatrix phylogenomics. european journal of taxonomy 283: 1–25. http://dx.doi.org/10.5852/ejt.2017.283 from phylogenetics to phylogenomics the last two decades have seen significant changes taking place in the practice of phylogenetic inference from molecular data. these changes have mostly been triggered by the ever-increasing size of the datasets being assembled and analysed in the form of supermatrices of concatenated genes (driskell et al. 2004). the evolution of the size and shape of these datasets, thanks to new technological advances in dna amplification and sequencing, has been associated with different phases in the field of phylogenetic inference (fig. 1). in the early days of molecular phylogenetics based on pcr amplification and manual sanger sequencing of a handful of genes for a limited number of taxa, the field laid somewhat in the “uncertainty zone” dominated by stochastic error, even if some naïve over-interpretations did occur. then, the development of automated sanger sequencing typically led to a few standard genes (e.g., ssu rrna, elongation factors, rna polymerases) being sequenced for a large number of taxa, projecting the field into the “irresolution zone” with only limited information available to resolve a large number of nodes. yet, at that time the focus was not on resolving large-scale phylogenetic relationships, rather than on identifying species or strains by molecular means, which eventually gave rise to the field of dna barcoding (hebert et al. 2003). the first genome sequences from model organisms reverted the situation, with few taxa being sequenced for their entire set of genes causing a sort of “inconsistency zone” in which large multigene alignments combined with poor taxon sampling led to a number of phylogenetic reconstruction artefacts due to the prevalence of systematic error. finally, the development of genomics and transcriptomics (and more recently of high-throughput or next-generation sequencing technologies) allowed the field to enter an “optimal zone” by offering the opportunity to assemble phylogenomic datasets representing a good compromise between taxon and gene sampling and typically constituted by hundreds of genes for tens of taxa. however, the tempting up-scaling along these two axes resulted in an “overflow zone”, where missing data steadily increase and computing requirements explode at all stages of the analysis (fig. 1). in the mid-2000s, multigene phylogenetics therefore turned into phylogenomics (delsuc et al. 2005; philippe et al. 2005a). as with the beginnings of molecular phylogenetics, the early days of phylogenomics (i.e., the use of genome-scale data to infer phylogenetic relationships) started with a wave of optimism. it was first claimed that genomic data would allow “resolving incongruence in molecular phylogenies” (rokas et al. 2003), with some bidding even higher and promising “ending incongruence” (gee 2003). as a consequence, the number of newly sequenced nucleotides was given in the abstract of articles published in high-profile journals as a simple metric of quality (dunn et al. 2008). the assumption underlying this optimism was that the use of a large amount of characters would eliminate stochastic error, which had been plaguing morphology-based and single-gene phylogenies for too long. this view was clearly valid and numerous subsequent studies have yielded phylogenies with much better statistical support (e.g., delsuc et al. 2006; hampl et al. 2009; jarvis et al. 2014). however, a few discordant voices soon advocated caution (phillips et al. 2004; soltis et al. 2004; jeffroy et al. 2006). first, genuine topological incongruence, i.e., statistically supported contradictions, is in fact extremely rare when phylogenies are inferred from a limited set of characters, since stochastic error dominates in this case (this does not mean that weakly supported incongruences are not real, in particular these created by http://dx.doi.org/10.5852/ejt.2017.283 philippe h. et al., pitfalls in supermatrix phylogenomics 3 incomplete lineage sorting, and deserve consideration, see below). second, systematic error naturally becomes more apparent at the genomic scale, simply, as a typical example of statistical inconsistency, in which the inference method converges towards an incorrect solution as more and more data are analysed. as a result, phylogenomics rather was “the beginning of incongruence” among studies based on different datasets and (especially) inference methods (jeffroy et al. 2006). despite the fact that most articles do not study the same phylogenetic questions, incongruence appears to be common in phylogenomics. for instance, the first group of animals to branch out is proposed to be either ctenophora (dunn et al. 2008), bilateria (schierwater et al. 2009) or porifera (philippe et al. 2009); acoels are included within lophotrochozoa (dunn et al. 2008), as sister to all the remaining bilateria (hejnol et al. 2009) or within deuterostomia (philippe et al. 2011a); the sister-group of land plants is coleochaete (finet et al. 2010) or zygnematales (wodniok et al. 2011); scaphopods are sister to bivalvia + gastropoda (kocot et al. 2011), gastropoda (smith et al. 2011) or bivalvia (unpublished). the marked decrease in stochastic error is insufficient to avoid the generation of incompatible fig. 1. evolution of phylogenetic reconstruction over time. different zones can be delimited based on the number of genes (x axis) and number of taxa (y axis) composing the supermatrix at hand. generally speaking, the (upper) left part of the map has more to do with identifying species (as in barcoding studies), while the right part of the map corresponds to multigene and phylogenomic datasets assembled for recovering large-scale phylogenetic relationships. each of these zones suffers from its own combination of issues (stochastic error, systematic error, data errors, computational requirements and missing data). interestingly, the “optimal zone” in phylogenomics is not the one corresponding to the highest number of genes and taxa, because this computationally “intractable zone” is also the one where data errors and missing data are the most abundant. the latter aspect is due to the continuous shrinking of the number of orthologous genes when considering increasingly more species, owing to gene loss, gene duplication and gene transfer events. european journal of taxonomy 283: 1–25 (2017) 4 phylogenies and should have been accompanied, as expected by some, by a decrease in systematic error obtained through efforts in improving inference methods. unfortunately, it appears that genome-scale analyses also lead to the amplification of data errors, i.e., errors that occurred during the construction of the datasets (philippe et al. 2011b; laurin-lemay et al. 2012). this is due to the fact that the flood of genomic data (often of decreasing quality) has become intractable for manual controls, while few bioinformatics tools have yet been adapted to the complexity of the genomic scale. in this paper, we discuss major pitfalls of phylogenomics and ways to limit their impact on phylogenetic inference. first, we review the approaches, existing and to be developed, to assembling a phylogenomic dataset that is as free as possible of data errors, briefly reviewing the potential issue of missing data along the way. second, we describe efficient methods to reduce systematic error. construction of a phylogenomic dataset effects of data errors single-gene and small multigene datasets could easily be curated thoroughly by hand, allowing researchers to remove almost all major errors before analysis and publication. the few remaining errors were often due to contaminations in the case of organisms that were difficult to make axenic (pawlowski et al. 1996; bourlat et al. 2003). of course, due to the limitations of ancient sequencing technologies, sequencing errors were frequent and sequence quality was quite variable in these early datasets. highthroughput sequencing has greatly improved sequence quality, mainly through the large coverage of each nucleotide, but has simultaneously flooded researchers with an amount of data that is difficult, if not impossible, to handle by hand. while numerous custom scripts have been independently written to manage this deluge of data, there is currently no agreement in the community regarding the quality controls that should be implemented in phylogenomic dataset construction. more problematically, the practice of phylogenetic research has continued at the genomic scale with more or less the same approaches as those used at the single-gene and small multigene scales. in order to illustrate data errors and their consequences for phylogenomic inference, we will explore three example datasets (dunn et al. 2008; schierwater et al. 2009; finet et al. 2010) that were carefully verified after their original publication (philippe et al. 2011b; laurin-lemay et al. 2012). the most frequent errors observed in the protein-coding genes of these datasets are frameshifts and contaminant sequences. frameshifts are primarily due to sequencing errors (especially for transcriptomic data) and annotation errors (especially for genomic data). sequence contamination occurred at the sampling step (i.e., parasites or symbionts thriving inside, or in close association with, the organisms of interest) or at the laboratory or sequencing steps (cross-contamination). accordingly, many contaminants belong to evolutionary lineages that contain many parasites (e.g., apicomplexa, microsporidia, platyhelminthes), are widely distributed (e.g., fungi or ciliophora), or are extensively studied (e.g., mammalia or magnoliophyta). less frequent errors correspond to the use of an erroneous genetic code for translation, the inclusion of paralogous (or even non-homologous) genes. for instance, more than 4000 amino acids were incorrect and more than 30 sequences were contaminants in the supermatrix of dunn et al. (2008); 101 sequences were contaminants in the supermatrix of finet et al. (2010). one might be tempted to argue that these errors have a negligible impact on tree reconstruction. given the large size of phylogenomic alignments, non-phylogenetic signal should be overwhelmed by the wealth of genuine phylogenetic signal. to test this hypothesis, we introduced contaminant sequences into a large supermatrix (30 517 positions and 48 species) and observed that phylogenetic inference is indeed relatively robust to such errors (unpublished results): when contamination was below 4%, at most 4 partitions were incorrect (on average ~2 for 3%). as expected, all clades with a relatively long ancestral branch were accurately recovered. this demonstrates that obtaining a phylogenomic tree in philippe h. et al., pitfalls in supermatrix phylogenomics 5 good global agreement with previous knowledge does not imply that the underlying dataset is errorfree. in contrast, clades with a short ancestral branch appear to be very sensitive to computationally introduced contaminations, whereas such clades are the very target of phylogenomics, smaller-scale approaches having insufficient statistical power to resolve them. these simulations are nevertheless conservative, because we spiked contaminant sequences at random, which is usually not the case in practice. for instance, 55 out of 101 were cross-contaminations in the study of finet et al. (2010), with 29 sequences from the coleochaetalean chaetosphaeridium actually coming from the zygnematalean penium (laurin-lemay et al. 2012). these biased contaminations explain why two well-established clades (coleochaetales and zygnematales) were not recovered as monophyletic in the original study, in spite of relatively long ancestral branches in the true species tree. in the case of dunn et al. (2008), the impact of contamination has not been directly studied, but a supermatrix with the same set of genes and species was assembled not only to fix data errors in the former, but also to complete it with new sequences (cutting down missing data from 55.5% to 35.6%). moreover, we used less permissive gblocks parameters, reducing positions from 21 152 to 18 463 (philippe et al. 2011b). despite the use of the same genes, the same species and the same evolutionary model, trees inferred from the original and revised datasets differed by a robinson-foulds distance of 36, i.e., 18 clades among 74 were different. the distance was increased if a better fitting model (the cat model instead of the wag model) was used: 24 bipartitions among 74 (i.e., a robinson-foulds distance of 48) were different between trees inferred based on original (dunn et al. 2008) and revised (philippe et al. 2011b) datasets. for instance, acoels moved from within lophotrochozoa to sister of deuterostomia, ctenophora moved from sister to all the remaining animals to sister to all but porifera, or myzostoma moved from sister of gnathostomulida + acoela to within annelida, all these changes being of great biological significance. a large amount of data is therefore clearly insufficient to reconstruct the correct phylogeny, and data errors, as well as missing data and systematic error (see below), must be minimized as much as possible. data quality control no standard for the quality control of a phylogenomic supermatrix has been established yet. instead, only preliminary attempts for reporting data have been made (leebens-mack et al. 2006). obviously, manual and semi-automated controls that are in use for single-gene phylogenetics can still be applied, though they are very time-consuming (e.g., several weeks of tedious work in the case of our two previous studies, philippe et al. 2011b and laurin-lemay et al. 2012) and subjective (i.e., not easily reproducible nor easily assessable by referees). it is therefore of prime importance to develop bioinformatics tools to assemble high-quality datasets and necessary that the phylogenetics community establishes a standard for the quality of a supermatrix for phylogenetic analysis. note that errors can be very complex to track down, and even to define. for example, a gene horizontally transferred from a parasite to its host could easily (but erroneously) be viewed as a contaminant, and thus removed from the dataset; yet this would be the right thing to do only when the inference method assumes orthology, since such a xenologous gene might also provide valuable information (huang & gogarten 2006). similarly, the known case of a frameshift in the murf gene of buchnera, which is rescued by polymerase infidelity (tamas et al. 2008), looks very much like a sequencing error in the genomic sequence. while these exceptions certainly do exist, they are rare and can likely be safely ignored in the context of supermatrix construction. obvious ways of reducing data errors are to improve the sequencing technology, the genome assembly method, the gene annotation protocol and the identification of orthologous genes. these improvements, which are not specific to phylogenomics, are the object of active research and will not be discussed here. we will instead focus on the few tools that allow controlling the quality of a supermatrix after its assembly but before its use in phylogenetic tree reconstruction. the underlying assumption of these quality controls is that outliers (in terms of sequence similarity or phylogenetic position) are most often incorrect. granted, a very fast-evolving sequence is difficult to tell apart from a genuine outlier and might be discarded erroneously by such approaches. however, this unavoidable property is not european journal of taxonomy 283: 1–25 (2017) 6 really problematic for phylogenomic inference, since removal of fast-evolving characters/species is commonly carried out to reduce tree-reconstruction artefacts (e.g., brinkmann & philippe 1999; pisani 2004; brinkmann et al. 2005). reducing alignment errors in order to infer a phylogeny from amino-acid sequences, one has to chain up several analyses, first sequencing, assembling and identifying coding regions (annotation), then clustering orthologous sequences and, for each cluster, producing a multiple sequence alignment that will eventually be part of the phylogenomic supermatrix. since each step can introduce errors, the careful analysis of individual gene alignments is key to detecting and correcting those issues that would otherwise hamper the phylogenetic inference based on the final supermatrix. a crucial step is therefore to identify the homologous amino-acid residues (or nucleotides) at every position of the sequences. the importance of a correct alignment in phylogenetic inference has long been pointed out (morrison & ellis 1997; ogden & rosenberg 2006; talavera & castresana 2007; wong et al. 2008) and numerous efforts have been made to improve multiple sequence alignment software (higgins et al. 1992; notredame et al. 2000; edgar 2004; katoh et al. 2005; loytynoja & goldman 2005). yet, due to the lack of a tractable model of sequence evolution in the presence of insertion and deletion events (indels), the criteria optimized by alignment software are mostly ad hoc and based on the simplistic assumptions that homologous characters should be similar and that indels are rare events. automatic alignment programs have thus long been used as efficient tools to obtain a first draft of the alignment to then be manually curated by expert biologists. even if human curation greatly cripples reproducibility, this step may be strongly encouraged as a “way of introducing some biological (as opposed to bioinformatics) insight into the resulting alignment” (morrison 2006). reviewing the literature, morrison (2009) reported that manual curation is a common practice, with 78% of sequence alignment procedures described in 1280 papers published in 2007 involving human intervention. the landmark work on automatic alignment filtering is the famous gblocks program (castresana 2000), which (although based on a basic strategy) allows removal of the less reliable parts of an alignment and hence improving phylogenetic inference (talavera & castresana 2007). gblocks first considers each site independently; depending on the presence of gaps and on the percentage of identical residues, sites are partitioned in the three following categories: “non conserved”, “conserved” or “highly conserved”. then reliable blocks of sites are defined by taking into account the degree of conservation of sites inside and around those blocks. the fact that gblocks considers the neighbourhood of a site, when deciding whether to keep it or not, is a major reason for its efficiency. by default, the original gblocks program removed all sites containing gaps. this is generally too strict, especially when dealing with numerous sequences, and a threshold on the percentage of allowed gaps is preferable. similarly, the percentage of identical residues is a crude measure of site conservation. more recent alignment filtering methods, such as trimal (capella-gutierrez et al. 2009) or bmge (criscuolo & gribaldo 2010), refine this measure of site conservation through the use of, e.g., pam or blosum matrices (dayhoff et al. 1978; henikoff & henikoff 1992). moreover, unlike gblocks, these two filtering software programs do not require the user to provide fixed threshold values for determining sites to be trimmed. instead, bmge uses statistical tests to select the residues to keep while trimal dynamically adapts its thresholds based on the distribution of gaps and residue similarities observed in the alignment. another strategy to assert alignment reliability is to consider their stability. indeed, for a given set of sequences, there are often numerous (nearly) equally optimal alignments. the simple head or tail method that consists in aligning sequences from right to left in addition to from left to right (landan & graur 2007) and its variants (landan & graur 2008) are designed to measure the stability of residue philippe h. et al., pitfalls in supermatrix phylogenomics 7 pairing among such equally optimal alignments. other authors considered a broader view of alignment stability, trying to estimate stability by comparing alignments obtained with different parameter values (e.g., gap opening cost), different guide trees and even different alignment programs. among others, soap (loytynoja & milinkovitch 2001), t-coffee (poirot et al. 2003), mumsa (lassmann & sonnhammer 2007), trima1 (capella-gutierrez et al. 2009) or guidance (penn et al. 2010) propose to filter alignments based on stability criteria. since they require realigning the same set of sequences several times, these approaches are time consuming. moreover, their results strongly depend on the sampling of methods and parameters used to generate the alternative alignments. another option is to consider, for a given method and a given set of parameters, the stability of residue pairing among the sub-optimal alternative solutions. in this strategy, implemented in fsa (bradley et al. 2009), psar (kim & ma 2011) and zorro (wu et al. 2012), alternative alignments used to measure residue-pairing stability are obtained by a probabilistic sampling based on an approximation of the alignment posterior distribution (i.e., the better the alignment, the higher its probability of appearing in the sampling). this approach has the advantage of relying on a well-defined probabilistic framework, to allow for an efficient implementation (since alternative alignments are not built from scratch) and to provide filtered alignments leading to better phylogenies (wu et al. 2012). most of the above-mentioned methods attribute an elementary score to each residue pairing of the alignment, which can be turned into an elementary score per residue (e.g., using the average score of residue pairings containing a given residue). by considering the score of all residues of a given site, they can attribute a site score as done by gblocks, but they can also consider the score of all residues in a given sequence. this is particularly useful to identify rogue sequences, e.g., resulting from contaminations or erroneous homology predictions, before any further analysis. this can even be extended, as done for instance in bmge, guidance and fsa, to mask portions of sequences, which is particularly useful to detect annotation errors or frameshifts in coding sequences. the method statsigma (prakash & tompa 2005) uses an extension of the blast statistics (altschul & lipman 1990) to detect such nonorthologous portions of sequences (prakash & tompa 2005). alternatively, philippe et al. (unpublished) introduced hmmcleaner, a program that tackles the same problem by first building a hidden markov model profile of the alignment, and then measuring the score of the different sequence regions along this profile. the focus of those last two methods is clearly on evaluating a per (portion of) sequence quality rather than a per site quality as done by zorro or psar, for instance. although the rationale of removing part of the alignments because of potential errors is clear, it is also obvious that positions possibly hosting a correct phylogenetic signal are removed as well. hence, a recent study (tan et al. 2015) suggests that commonly used software for automated filtering alignments decreases single-gene phylogeny accuracy, in contradiction with a previous study (wu et al. 2012). although more work is needed to solve this point, the filtering is easier to justify in supermatrix phylogenomics, since the increase of stochastic error observed in single-gene phylogeny is naturally overcome when numerous genes are concatenated. finally, when dealing with protein-coding nucleotide sequences, it is highly recommended to align them based on their amino acid translations. three main reasons can be pointed out for doing so (morrison 2006): amino acid sequences are less variable and thus easier to align (due to the degeneracy of the genetic code), they are defined in a larger character-state space (which reduces the probability of random matches), and inserted gaps will preserve the reading frame (which is biologically more realistic). the alignment of protein-coding sequences is therefore often performed in three steps: 1) the nucleotide sequences are first translated into amino acids, 2) the amino-acid sequences are then aligned and 3) the resulting protein alignment is used for deriving the codon alignment by reporting inferred gaps. this strategy is implemented in transalign (bininda-emonds 2005), translatorx (abascal et al. 2010), european journal of taxonomy 283: 1–25 (2017) 8 and seaview 4 (gouy et al. 2010), for example. the main limitation of this approach is the inability to handle unexpected frameshifting indels (either real or due to sequencing errors) that result in (partially) incorrect translations. affected sequences hence look like highly divergent protein sequences and, in the best case, are removed by alignment filtering methods. the macse program handles such situations by simultaneously considering the nucleotide sequences and their amino-acid translations and by explicitly taking into account frameshift events during the alignment process (ranwez et al. 2011). this provides an automatic solution to detect undocumented frameshifts in public database sequences and highthroughput sequencing contigs. macse also allows aligning protein-coding gene datasets containing non-functional sequences (pseudogenes) without disrupting the underlying codon structure (fig. 2). sequences can thus be retained rather than removed from the alignment, provided that sites containing frameshifts are a posteriori filtered out. in addition, the accuracy of the alignments can be evaluated using protein structure (when available) as performed by tcs (chang et al. 2014). reducing orthology prediction errors although errors due to the inclusion of non-orthologous sequences in phylogenomics can have drastic consequences on the final results (philippe et al. 2011b; laurin-lemay et al. 2012), a hunt for such dubious sequences is rarely performed in phylogenomics and, when it is, fastidious manual approaches are most often used. an obvious type of non-orthology error that can be detected and corrected for is biological (or in silico) sequence contamination. blast similarity searches provide an easy way to detect contaminant sequences, as long as the reference database is taxonomically rich. hence, a much better hit on a sequence from a different clade than on a sequence from the expected clade (e.g., ciliate for a mollusc sequence) is an excellent indicator that the query sequence is “fishy”. however, especially when the difference between the two e-values is not that large, a confirmation through a phylogenetic tree is recommended (philippe et al. 2009; philippe et al. 2011b; laurin-lemay et al. 2012), owing to the limited power of blast (koski & golding 2001). disentangling in silico, laboratory and biological contamination may be difficult, especially when the biology of the organism at hand is poorly known. yet, when contaminant sequences belong to organisms also found in the laboratory having generated the data (e.g., escherichia coli or cross-contamination among the organisms of the study) or to clades rich in parasitic or symbiotic forms (e.g., microsporidia, fungi or dinoflagellates), laboratory or biological contamination, respectively, is the most straightforward interpretation. fig. 2. example of a frameshift-aware alignment produced by macse. trpc2(-like) sequences of bats were aligned at the nucleotide level (a) and amino acid level (b), unravelling several frameshifts (indicated by “!”) and stop codons (indicated by “*”) in the pseudogenes. hence, macse automatically provided an alignment that would otherwise require a lot of tedious manual work. it can also be used to replace these frameshifts and stop codons by standard codons (e.g., “nnn” or “---”) in order to obtain an alignment suitable for further analysis with standard tools (e.g., phylobayes or paml). philippe h. et al., pitfalls in supermatrix phylogenomics 9 cases of biological contamination are sometimes anticipated and addressed accordingly. for instance, kocot et al. (2011) analysed sequences from the order neomeniomorpha (the group containing solenogasters, small worm-like shell-less molluscs). the two neomenioid species included in the study feed on cnidarians and are known to harbour dna from their cnidarian prey (okusu & giribet 2003; kocot et al. 2011). to avoid contamination of the neomenioids by cnidarian sequences, the authors performed a blast search for every neomenioid sequence against the complete sets of proteins of a gastropod (lottia) and of a cnidarian (nematostella). if a sequence had a better hit against nemastostella than against lottia, it was considered as a contaminant and discarded. though sound in its principle, this approach is expected to be limited due to the use of a single cnidarian proteome. indeed, using multiple cnidarian species, we detected several cnidarian contaminants in the published dataset using a semiautomatic approach (data not shown). contamination is not always as simple to detect as in the ideal case of a very distantly related contaminant organism. further, the blast approach has some serious limitations. first, it requires knowledge of the phylogeny, which is often the question that needs to be answered in phylogenomic studies. second, if contamination occurs between closely related species and/or if taxon sampling is too sparse, the blast approach is unlikely to detect contaminants. finally, the absence of a blast hit on the genome used as a proxy for the contaminant organism does not ensure the inexistence of a closely related sequence. instead, it can simply mean that the sequence is missing from the reference genome. nevertheless, exactly as ncbi automatically removes vector sequences, this type of approach should be made systematic in phylogenomics, for it requires only limited resources (computing and human time) and becomes more and more powerful with the increasing taxonomic diversity of correctly annotated genomes. scientific journals should thus enforce this simple quality control, just as molecular biology and evolution once decided to require authors to report statistical support for publishing a phylogeny. contamination is not the only source of non-orthology, paralogy being a common underhand source of issues, given the high frequency of gene/genome duplication, gene conversion and gene loss. instead of looking for unexpected blast hits, it is thus preferable to build and check individual gene trees for unexpected bipartitions (i.e., aberrant placement of some species). again, this approach was also used by kocot et al. (2011). to this end, they inferred a maximum likelihood tree for each individual gene and “manually evaluated [...] any remaining cnidarian contamination in the neomenioid datasets”. these were neomenioid sequences that were forming a clade with nematostella or were part of a polytomy that included nematostella. their protocol was nonetheless focused on a single type of errors and more systematic strategies are certainly needed. one example of such an approach was developed to check the orthology in a supermatrix used to reconstruct deep animal relationships (philippe et al. 2009). for each gene tree, the presence of every bipartition with a bootstrap support value ≥ 70% was compared to the bipartition list of the tree obtained by concatenating all the genes (this tree is considered as a good approximation of the true, unknown, species tree). the authors found that the number of conflicting bipartitions was small and that incongruences almost always represented only minor rearrangements in the tree (often nearest neighbour interchanges). consequently, they concluded that paralogy was not playing a significantly negative role in their analysis. while this approach is reasonable for checking that a dataset is sufficiently free of nonorthology errors, it is seriously limited by stochastic error. indeed, in single-gene phylogenies, most of the nodes are poorly supported and, if the support threshold is reduced (e.g., down to 50%), too many nodes conflict with the tree inferred from the supermatrix, whatever the actual quality of the dataset. moreover, a manual, time-consuming step is needed to interpret each automatically detected conflict to determine whether it is due to a stochastic (e.g., partial sequence) or systematic (e.g., long-branch attraction (lba)) error, or caused by contamination or undetected hidden paralogy. european journal of taxonomy 283: 1–25 (2017) 10 an automatic approach for detecting outlier sequences in a phylogenomic dataset, named phylo-mcoa (de vienne et al. 2012) has been developed. it is based on a multivariate approach known as multiple co-inertia analysis (mcoa) (chessel & hanafi 1996). it takes as input a set of gene trees, converts all these trees into pairwise distance matrices (using either nodal or patristic distances between species) and compares all these matrices simultaneously. what is relevant here is the output given by the program and the possibilities it offers for detection and potential removal of problematic sequences. after mcoa has been applied to all pairwise distance matrices, the user is returned (1) a reference position of each species in a euclidean space, and (2) the position of each of these species in each individual gene tree in this same space. a so-called 2wr-matrix is then constructed by computing, for every species, the distance separating its position in each gene tree to its reference position. this matrix contains as many rows as the number of species and as many columns as the number of genes. a cell with a high value in this matrix represents a species whose position (or branch length) in a given gene tree is not concordant with its position in all other trees. an example of such a matrix for a hypothetical dataset of 40 genes and 40 species is presented in fig. 3. phylo-mcoa distinguishes two types of outliers, the “complete” outliers and the “cell-by-cell” outliers. complete outliers (black arrows in fig. 3) represent either species that have a position that is very variable with respect to other species “for all the genes” or genes for which “all the species” have a position that is not concordant with the other species in the other genes. this is clearly the case for two genes (g24 and g32) and three species (t23, t34 and t32) in the example matrix of fig. 3. phylo-mcoa provides a simple outlier detection method to identify these complete outliers (see de vienne et al. 2012 for details). cell-by-cell outliers (dashed circles in fig. 3) are specific species in specific genes whose position is not concordant with their position in the other gene trees. they can only be identified with confidence in the 2wr-matrix after complete outliers have been removed and the analysis performed anew. the final output of phylo-mcoa is a list of all complete outlier species, all complete outlier genes and all cell-by-cell outliers. these detected outliers may represent either problematic sequences due to contamination, frameshift events, incorrect annotations, erroneous chimerical sequences, wrong orthology assessment, or indicate interesting biological processes such as horizontal gene transfer, gene conversion, incomplete lineage sorting or hybridization. when the goal of the analysis is, as discussed here, to ensure having a dataset containing only orthologous sequences, one should focus on these detected outlier sequences to remove them. as an example, phylo-mcoa was used on the phylogenomic dataset of schierwater et al. (2009) known to contain problematic sequences that were manually identified by philippe et al. (2011b). by comparing the tree obtained after removing all the outliers identified by phylo-mcoa and the original tree of schierwater et al. (2009), it appeared that (1) the strong bootstrap support for the monophyly of diploblasts was reduced (from 100% to 63%), as expected from the fact that it is certainly not a monophyletic group, and (2) the monophyly of porifera disappeared (from 100% in the original tree), in agreement with the very low bootstrap support (36%) obtained by philippe et al. (2011b) on the same dataset after replacing the incorrect sequences they detected by better ones. even though the automatic approach performed with phylo-mcoa and the manual approach conducted by philippe et al. (2011b) only partly identified the same problematic sequences, the application of this tool to a complex dataset demonstrated its potential. nevertheless, there is certainly room for improvement and for the development of new tools dedicated to this important task. a first step is to be achieved soon with the development of an online version of phylo-mcoa that will facilitate its use and parameterization (unpublished). philippe h. et al., pitfalls in supermatrix phylogenomics 11 evaluating the impact of missing data assembling a large and complete dataset with hundreds of orthologous genes and tens to hundreds of species remains a challenge, since a gene may have not yet been sequenced, might have been lost in a lineage, or could have been replaced by a xenologous copy through lateral gene transfer. as a result, many supermatrices were published with a high amount of missing data (e.g., 55%, 74%, 79% and 81% for dunn et al. (2008), kocot et al. (2011), smith et al. (2011) and hejnol et al. (2009), respectively). the impact of missing data on the accuracy of phylogenomic inference has received a lot of attention, but studies focused on this issue have reached two opposite conclusions: (1) the effect of the incompleteness is deleterious and sparse supermatrices must be avoided (huelsenbeck 1991; lemmon et al. 2009) and fig. 3. typical output of phylo-mcoa. a matrix containing as many rows as the number of species and as many columns as the number of genes was computed, in which complete (black arrows) and cell-bycell (dashed circles) outliers can easily be detected. cells with a high value (dark grey) represent species whose position in a given gene is not concordant with their position in all the other genes. it is thus a measure of distance to the common signal present in the data. european journal of taxonomy 283: 1–25 (2017) 12 (2) missing data are not problematic per se as long as a sufficient number of characters are known for each taxon (wiens 2003; philippe et al. 2004; wiens & morrill 2011). because the impact of missing data has mainly been studied on medium-sized datasets and/or on simulated sequences, these conclusions are not easily transposable to phylogenomic datasets. the central question is whether the lack of data has detrimental effects beyond those of the associated reduction in information, i.e., a decrease of the resolution level when the amount of missing data increases. lemmon et al. (2009) suggested that “ambiguous characters [or missing data] produce misleading estimates of phylogeny through interaction with two other factors: bayesian priors and model misspecification”. their experiments using four species were nevertheless based on the incorrect assumption that “ambiguous sites provided no topological information because only 2 of the 4 characters had unambiguous states”. this assumption is not true within a probabilistic framework, as every ambiguous site, even with a single known character state over hundreds of species, provides information to estimate model parameters, such as stationary frequencies (roure et al. 2013). hence, the major topological changes observed by lemmon et al. (2009) are easily explained under the following theoretical framework: when more and more characters with known, but always different, states in two closely related species are added to the alignment (their fig. 7), these two species are more and more distantly related in the inferred phylogeny simply because the ambiguous sites provide a very strong signal in favour of a very long evolutionary path separating them (sum of branch lengths). in contrast, adding unbiased ambiguous sites does not modify the outcome of the phylogenetic inference (roure et al. 2013). since the distribution and nature of missing data are usually not biased, this supports the idea that missing data are not deleterious per se (wiens & morrill 2011). however, missing data and model violations (or misspecification) do interact. as a reminder, model violations lead to an incorrect interpretation of the multiple substitutions occurring at a given position, potentially yielding tree reconstruction artefacts. because using a rich taxon sampling facilitates the detection of multiple substitutions, it also helps reduce the impact of potential model violations. unfortunately, missing data reduce the number of taxa effectively present at a given position, so that the resulting decreased power in detecting multiple substitutions may increase the impact of model violations (roure et al. 2013). for instance, if a fast-evolving species is completely represented in the supermatrix but its close relatives are very incomplete, the detection of multiple substitutions along this long branch becomes inefficient, thus favouring potential long-branch attraction (lba) artefacts. to test whether missing data can generate such misleading effects in phylogenomics, roure et al. (2013) assembled an almost complete supermatrix (39 animals, 126 genes for 29 715 amino acids) and progressively increased the proportion of missing genes (20% to 80%) in some species (8 or 27), by introducing missing data following the patchy patterns observed in real transcriptomic-based datasets while controlling for information loss and decrease in taxon sampling. this empirical study demonstrated that a patchy distribution of missing data has an effect similar to the complete removal of species, but is much more detrimental than the complete removal of genes. missing data indeed exacerbate systematic error leading to lba artefacts by reducing the ability to detect multiple substitutions, as evidenced by tree length decreasing when the fraction of patchy missing data increases (roure et al. 2013). in this respect, it is worth noting that algorithmic approaches have been proposed to extract a subset of the original dataset (by removing genes and/or species) in order to reduce the proportion of missing data while preserving as much information as possible (sanderson et al. 2003; dutheil & figuet 2015). the concept of “effective number of species”, i.e., the real number of known character states at a typical position, is useful to understand the effect of missing data (fig. 4) and to assist in deciding when to stop adding increasingly incomplete genes to a supermatrix. though it is fundamentally a property of each column of the alignment, it can be considered globally by computing the product of the number of taxa by the frequency of unambiguous data (e.g., 94 × 0.19 = 18 for the hejnol et al. (2009) dataset). a low effective number of species provides an intuitive explanation for the exacerbation of systematic error philippe h. et al., pitfalls in supermatrix phylogenomics 13 fig. 4. distribution of missing data in phylogenomic datasets. the raw dimensions of a supermatrix (in numbers of genes and taxa) should always be accompanied by an estimate of its level (and ideally distribution) of missing data (a). indeed, some datasets were advertised as very large when published but required a lot of useless computing power due to a large proportion of missing data (b). more importantly, such datasets are more exposed to data errors, systematic error and phylogenetic artefacts, as their effective number of species is actually quite low for the largest part of their width. in other cases, a targeted completion (in the outgroups) was enough to improve the phylogenetic accuracy (d–e). in contrast, datasets assembled with the optimal phylogenomic zone in mind combine a sufficient number of genes and taxa while featuring a low proportion of missing data (c). these are the most appropriate datasets to produce accurate phylogenetic relationships. european journal of taxonomy 283: 1–25 (2017) 14 by missing data in a given dataset. the answer to the question “should we add a highly incomplete gene to a phylogenomic supermatrix?” is therefore rather no. it may be better to use a narrower, but almost complete dataset, than a wider and very incomplete one. in contrast, the answer to the question “should we add a very incomplete species to a phylogenomic supermatrix?” is more complex. in agreement with previous studies (philippe et al. 2005b; wiens 2005), we have shown that including incomplete species, especially slow-evolving ones, helped us to reduce the effects of systematic error (roure et al. 2013). yet, increasing the completeness of some species, in particular in outgroups, also reduced lba artefacts. therefore, the inclusion of incomplete taxa should not be prohibited, but the level of completeness of all species should be clearly indicated on the inferred phylogeny because it provides key information for identifying potential problems. for instance, when missing data are concentrated in one clade, the effects are more deleterious, questioning the reliability of the relationships within the clade, but virtually not affecting the rest of the tree. it is worth noting that the addition of incomplete species increases the computational time to a larger degree than the addition of complete species (roure et al. 2013). as a result, the decision to include incomplete species is a matter of balance between the improvement of phylogenetic accuracy and the inevitable increase in computational time. an open question is how the creation of chimerical sequences for incomplete, but closely related, species may help in solving this dilemma. finally, streicher et al. (2016) have recently shown that, in the case of ultraconserved elements of lizards, the optimal strategy depended on the approach. when dealing with supermatrices analysed by maximum likelihood, it was better to maximize the number of species while keeping the total amount of missing data below 50%, whereas for species trees inferred by neighbour joining, it was better to include all the markers but for a limited number of species. more empirical studies are therefore needed to establish consensual guidelines for finding the best level of missing data in phylogenomic inference. the last important lesson from empirical studies of missing data in phylogenomics is that the method used to infer the tree has a greater impact than the level of missing data itself (roure et al. 2013; hosner et al. 2016), especially when the latter is limited (between 20% and 40%). first, the alternative methods that have been suggested as being less sensitive to missing data, such as super-tree, maximum parsimony and gene-partitioning models, turned out to be more sensitive to missing data (albeit not significantly for the latter). second, the choice of the model of sequence evolution (e.g., the site-homogeneous wag + f + γ4 model versus the site-heterogeneous cat + γ4 mixture model) had a much larger impact than the level of missing data. in this particular case, the tree inferred from a dataset with 27 species having 80% of missing data using cat + γ4 was closer to the best phylogenetic estimate than the tree inferred from the complete dataset using wag + f + γ4, especially for the position of the fast-evolving taxa (roure et al. 2013). limiting systematic error once the issue of data errors in the supermatrix assembly is overcome, systematic error is the main, if not the sole, limitation to the power of phylogenomics. felsenstein (1978) described the lba artefact almost 40 years ago, demonstrating that maximum parsimony could converge towards an incorrect solution with additional characters. although the number of characters used in phylogenomics (typically from tens of thousands up to a few million) is far from infinite (i.e., the convergence of a method), the issue of statistical inconsistency is important and should not be neglected. the approaches used to eschew inconsistency have been reviewed several times (e.g., olsen 1987; felsenstein 1988; sanderson & shaffer 2002; soltis et al. 2004; philippe et al. 2005a, 2011b) and will only be succinctly discussed here. systematic error occurs when inference methods do not correctly handle multiple substitutions. in a probabilistic framework, systematic error stems from violations of the model of sequence evolution, philippe h. et al., pitfalls in supermatrix phylogenomics 15 which is designed in essence to predict the substitution history. it is generally considered that the problem is an underestimation of the true number of substitutions (related to the well-known issue of saturation), but it is quite possible that an incorrect design of the evolutionary model yields an overestimation (olsen 1987). three complementary approaches allow improving the correct estimation of multiple substitutions: (1) a rich taxon sampling, (2) realistic models of sequence evolution, and (3) removal of the most deviant data. a rich taxon sampling provides direct information to infer multiple substitutions, by breaking up branches into smaller pieces (hendy & penny 1989). from our experience, this strategy is the most efficient to resolve difficult phylogenetic questions, in particular for positioning fast-evolving lineages. for instance, the phylogenetic position of the very fast-evolving acoels is either sister to all the remaining bilaterians (baguna & riutort 2004), potentially resulting from an lba with the distant diploblast outgroup, or sister to the fast-evolving platyhelminthes (dunn et al. 2008), potentially resulting from another lba. the inclusion of sequences from xenoturbella, the likely sister-group of acoels, reduced the misleading signal for these two artefactual positions, but not enough to yield a resolved phylogeny (philippe et al. 2007). it was only the addition of sequences from several acoels, and also from their closest relatives, nematodermatids, that allowed positioning of this very fast-evolving lineage (philippe et al. 2011a), probably because nematodermatids evolve more slowly than the latter. even the best-fitting evolutionary model currently available is unable to accurately position acoels in the absence of xenoturbella and nematodermatids. two recent phylogenomic studies which focused on these organisms (cannon et al. 2016; rouse et al. 2016) illustrate that a larger number of genes (212 and 1178 in these studies vs 68 and 197 in older studies) exacerbates systematic error: when outgroups and fast-evolving acoelomorphs are included, xenacoelomorpha (xenoturbella + acoelomorpha) emerge far from ambulacraria, as sister of all other bilaterians, but when excluded, xenoturbella is robustly sister of ambulacraria in both studies (unpublished results), two obviously incompatible results. while very useful, the strategy of enriching the taxon sampling is limited in two different ways. the first one is inescapable: the history of speciation and extinction that has generated the tree of life has led to many taxon-poor clades (e.g., coelacanth or amborella), and ancient dna techniques are of very little help in filling in the gaps because they are restricted to recently extinct taxa. the second limitation is governed by time and money, given that the search for the optimal phylogenetic tree is an np-hard problem. in this respect, the development of more efficient heuristics (e.g., guindon & gascuel 2003; stamatakis & ott 2008) and of more efficient computers only partially alleviates the problem. the number of species included in a phylogenomic analysis will thus always remain a compromise between the resources available and the length of time one accepts to devote to it. the improvement of models of sequence evolution should strive for increased realism, not necessarily increased complexity (steel 2005). the use of inference-based comparison instead of data-based comparison in posterior predictive assessment, albeit very time-consuming, is the best way to evaluate any given improvement, since one assesses the ability of a model to correctly infer the topology, rather than to correctly simulate data (brown 2014). ideally, improvements should be focused on a better detection of the multiple substitutions that are the most likely to induce topological errors. this is the reason why the major examples of progress in the reduction of phylogenetic (lba) artefacts were obtained by including a more efficient detection of convergent substitutions (homoplasy): (1) the first probabilistic model, which included branch lengths and allowed the inference of more convergences along long branches (felsenstein 1981), (2) the modelling of the heterogeneity of the substitution rate among character states (e.g., the gtr model; lanave et al. 1984), which allowed the inference of more convergences for nucleotides of the same biochemical class or amino acids with similar properties, (3) the modelling of the rate variation across sites (yang 1993), which allowed the inference of more convergences at the fastest sites without inferring too many multiple substitutions at the slowest sites, file:///h:/a-wpdocs/ejt/library_of_life/11-philippe_16-83r1/l european journal of taxonomy 283: 1–25 (2017) 16 (4) non-stationary models accounting for the variation of nucleotide/amino-acid composition over time (galtier & gouy 1998; foster 2004), which allowed the inference of more convergences due to a similar bias among unrelated species, and (5) the modelling of the non-homogeneity of the substitution process across sites (mixture models such as cat; lartillot & philippe 2004; pagel & meade 2004) by locally decreasing the effective size of the alphabet, which allowed the inference of more convergences due to functional constraints (e.g., only a charged amino acid is acceptable). numerous other evolutionary model refinements have been made to take into account, for instance, the genetic code (through codon models), the secondary structure of rnas, heterotachy or gene-specific rate and/or composition (e.g., goldman & yang 1994; yang 1996; savill et al. 2001; huelsenbeck 2002). however, they all had little effect on the accuracy of phylogenetic inference, either because they do not allow the detection of the most problematic convergences or because the rest of the model is over-simplistic. for instance, taking into account the non-stationarity of the amino-acid composition is much more efficient when the heterogeneity of the substitution process across sites is also handled by the model (blanquart & lartillot 2008). future improvements will therefore be obtained by combining most, or even all, of these biologically reasonable properties into a single model, or by discovering which type of convergences are the most deleterious in order to specifically model them. clearly, the first approach will be limited by computational burden and a risk of over-parameterization. in contrast, it is unclear whether it is better to continue improving the current strategies based on a “static” supermatrix, or to further develop methods that simultaneously perform alignment and phylogenetic inference of a supermatrix (hein 1990; liu et al. 2012) or to develop methods that simultaneously infer single-gene trees and the species tree using reconciliation (to handle incomplete lineage sorting, duplication or horizontal gene transfer; reviewed in szollosi et al. 2015). careful (empirical) studies comparing the advantages of these different approaches are needed. in the very long run, a dream method that would simultaneously infer alignment, singlegene tree and the species tree using a realistic (complex) model for each of these three steps would probably be the most accurate, but perhaps not the most desirable because of its inevitable ecological footprint (see philippe 2011). the last widely used strategy to avoid systematic error consists in data removal. the underlying assumption, inspired by the parsimony approach (kluge & farris 1969), is that fastest-evolving sites and species should be removed because they are the most prone to homoplasy. it is therefore common practice in phylogenomics to discard fast-evolving sites and fast-evolving species, especially when slowly evolving close relatives are available. it is worth noting, however, that this is difficult to justify in a probabilistic framework, since the model is supposed to account for such rate variations through branch lengths or site-specific rates, respectively. in a probabilistic framework, the biggest issue is not the fastevolving objects per se, but the objects that most significantly violate the model of sequence evolution (philippe & roure 2011). for instance, a slowly evolving species, but with a marked compositional bias, will be more difficult to position than a fast-evolving but compositionally homogeneous species when using a model that assumes stationarity. similarly, a very slowly evolving site that independently, but unexpectedly according to the model, changes twice into the same character state (e.g., from aspartic acid to tryptophan) may be very detrimental to phylogenetic inference. recent studies have demonstrated that removing sites that violate the model of sequence evolution can overcome tree reconstruction artefacts in cases where removing fast-evolving sites alone cannot. as an illustration, roure & philippe (2011) obtained a sister-group relationship between cnidarians and poriferans in a phylogeny of animals based on proteins encoded in the mitochondrial genome using the cat + γ4 model. suspecting an lba artefact between the fast-evolving bilaterians and the outgroup, they identified heteropecillous positions, defined as positions with changing substitution profiles over time. interestingly, the progressive removal of these positions violating the assumption of homogeneity philippe h. et al., pitfalls in supermatrix phylogenomics 17 of the cat mixture model (lartillot & philippe 2004) led to a single topological change: the recovery of the more classical sister-group relationship between cnidarians and bilaterians. similarly, the gammaproteobacterial endosymbionts of insects have an at-rich genome and tend to be artificially clustered in trees based on nucleotide sequences husník et al. (2011). by progressively removing the positions that were the most compositionally biased (i.e., a position only containing a and t, or g and c, is unbiased), husník et al. (2011) were able to overcome this composition-based artefact and to recover a tree identical to the one obtained from amino-acid sequences. finally, recent studies of genome datasets in mammals (romiguier et al. 2013) and birds (jarvis et al. 2014) have revealed that at-rich genes hold a much clearer phylogenetic signal than gc-rich genes, which are subjects to gc-biased geneconversion (eyre-walker 1993) episodes possibly leading to an increased heterogeneity in both base composition and evolutionary rate among taxa. developing methods to detect and remove the sites and the species that violate the model of sequence evolution most significantly is a promising avenue of research. posterior predictive tests as implemented in programs such as phylobayes 3 (lartillot et al. 2013) and phycas (lewis et al. 2015) allow evaluation of model fit to the data or part of the data. sun et al. (2016) have recently used this kind of approach to identify sites in a whole chloroplast dataset that did not fit the model of sequence evolution. they showed that removing those sites from the analysis significantly improved phylogenetic inference of green algae relationships. even if data removal is intellectually less satisfying than developing a “perfect” model of evolution fully accounting for the complexity of the data, this kind of approach could be similarly efficient and certainly requires less computational resources, since these reduced datasets can be analysed with simpler models. conclusions with the democratization of high-throughput sequencing, acquiring new genome data has ceased to be the limiting factor in phylogenomics, except maybe for organisms that are difficult to sample from the environment. in contrast, phylogeneticists are now faced with a rise in data errors, which stems from a flood of increasingly bogus genomic data that has become intractable by hand. however, with more effort from the community, notably by investing thinking and time in the development of adequate bioinformatics tools, the issue of data errors should be solved in the next few years. given the size of current and upcoming phylogenomic datasets (thousands of genes for hundreds — and soon thousands — of species), computational requirements are emerging as the new limiting factor. as multiple possible improvements are theoretically able to limit systematic error (and artefacts) during phylogenetic inference, we will need both simulation and empirical studies to design the best compromise between using many species with a simple model and using few species with a complex model. similarly, more work is required to determine the set of properties that the ideal model of sequence of evolution should combine. in that respect, a sound strategy is to focus on model components that allow for a better detection of convergent multiple substitutions, without chasing biological realism. even if 1000s of genes may seem large for a biologist, it remains a modest sample size for a statistician. therefore, we should be ready to accept that the shortest branches of the tree of life may never be resolved. in that respect, it is somewhat paradoxical that the modern scientific tools developed to yield an accurate reconstruction of the tree of life are also very likely to contribute to the destruction of the species once thriving at its tips (see philippe 2011 for a more complete discussion). acknowledgments this work has benefited from support by the tulip laboratory of excellence (anr-10-labx-41) to hp and from an “investissements d’avenir” grant managed by agence nationale de la recherche european journal of taxonomy 283: 1–25 (2017) 18 (ceba, ref. anr-10-labx-25-01) to fd. this is contribution isem 2016-015 of the institut des sciences de l’evolution de montpellier. references abascal f., zardoya r. & telford m.j. 2010. translatorx: multiple alignment of nucleotide sequences guided by amino acid translations. nucleic acids research 38: w7–13. http://dx.doi.org/10.1093/nar/ gkq291 altschul s.f. & lipman d.j. 1990. protein database searches for multiple alignments. proceedings of the national academy of sciences 87: 5509–5513. http://dx.doi.org/10.1073/pnas.87.14.5509 baguna j. & riutort m. 2004. the dawn of bilaterian animals: the case of acoelomorph flatworms. bioessays 26: 1046–1057. http://dx.doi.org/10.1002/bies.20113 bininda-emonds o.r. 2005. transalign: using amino acids to facilitate the multiple alignment of proteincoding dna sequences. bmc bioinformatics 6: e156. http://dx.doi.org/10.1186/1471-2105-6-156 blanquart s. & lartillot n. 2008. a siteand time-heterogeneous model of amino acid replacement. molecular biology and evolution 25: 842–858. http://dx.doi.org/10.1093/molbev/msn018 bourlat s.j., nielsen c., lockyer a.e., littlewood d.t. & telford m.j. 2003. xenoturbella is a deuterostome that eats molluscs. nature 424: 925–928. http://dx.doi.org/10.1038/nature01851 bradley r.k., roberts a., smoot m., juvekar s., do j., dewey c., holmes i. & pachter l. 2009. fast statistical alignment. plos computational biology 5: e1000392. http://dx.doi.org/10.1371/journal. pcbi.1000392 brinkmann h. & philippe h. 1999. archaea sister group of bacteria? 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editors: line le gall, frédéric delsuc, stéphane hourdez, guillaume lecointre and jean-yves rasplus desk editor: danny eibye-jacobsen printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands. http://dx.doi.org/10.1080/10635150390218330 http://dx.doi.org/10.1080/10635150500234583 http://dx.doi.org/10.1093/sysbio/syr025 http://dx.doi.org/10.1093/sysbio/syr025 http://dx.doi.org/10.1186/1471-2148-11-104 http://dx.doi.org/10.1126/science.1151532 http://dx.doi.org/10.1371/journal.pone.0030288 http://dx.doi.org/10.1007/bf02352289 european journal of taxonomy 13: 1-62 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2012.13 www.europeanjournaloftaxonomy.eu 2012 · alexandr b. ryvkin this work is licensed under a creative commons attribution 3.0 license. m o n o g r a p h 1 new species and records of stenus (nestus) of the canaliculatus group, with the erection of a new species group (insecta: coleoptera: staphylinidae: steninae) alexandr b. ryvkin laboratory of soil zoology & general entomology, severtsov institute of problems of ecology & evolution, russian academy of sciences, leninskiy prospect, 33, moscow, 119071 russia. bureinskiy nature reserve, zelyonaya 3, chegdomyn, khabarovsk territory, 682030 russia. leninskiy prospekt, 79, 15, moscow, 119261 russia. email: vr.staph@gmail.com abstract. the canaliculatus species group of stenus (nestus) is redefined. four new palaearctic species of the group are described and illustrated: s. (n.) alopex sp. nov. from the putorana highland and taymyr peninsula, russia; s. (n.) canalis sp. nov. from se siberia and the russian far east; s. (n.) canosus sp. nov. from the narat mt ridge, chinese tien shan; s. (n.) delitor sp. nov. from c & se siberia. new distributional data as well as brief analyses of old records for fourteen species described earlier are provided from both palaearctic and nearctic material. s. (n.) milleporus casey, 1884 (= sectilifer casey, 1884) is revalidated as a species propria. s. (n.) sphaerops casey, 1884 is redescribed; its aedeagus is figured for the first time; the aedeagus of s. (n.) caseyi puthz, 1972 as well as aedeagi of eight previously described palaearctic species are illustrated anew. a key for the identification of all the known palaearctic species of the group is given. a morphology and ecology based analysis of the main evolutionary trends within the group is provided. a lectotype is designated for s. (n.) melanopus marsham, 1802; its siberian and ne european records are supposed to be erroneous; the monotypic melanopus species group is erected. key words. holarctic, palaearctic, nearctic, fauna, bionomics. ryvkin a.b. 2012. new species and records of stenus (nestus) of the canaliculatus group, with the erection of a new species group (insecta: coleoptera: staphylinidae: steninae). european journal of taxonomy 13: 1-62. http://dx.doi.org/10.5852/ejt.2012.13 introduction the canaliculatus group was erected by l. benick (1925) in his keys for palaearctic stenus species groups. in the short preface to the named article, benick indicated that the keys provided by him were meant for both practical identification and reflecting phylogenetic relationships between different species. it is clear that a harmonious combination of these two principles is an unrealizable ideal, especially in the case of steninae, which are a group with extensive parallelism in many lineages. in addition, this circumstance makes all attempts to apply formal cladistics in morphology-based phylogenetic studies of the subfamily ineffective. benick’s definitions of the species groups could not avoid the http://creativecommons.org/licenses/by/3.0/ mailto:vr.staph%40gmail.com?subject= http://dx.doi.org/10.5852/ejt.2012.13 european journal of taxonomy 13: 1-62 (2012) 2 problems connected with the form of the key in his article. the canaliculatus group within the subgenus nestus rey, 1884 was defined by the following characters (in the translation below i try to reformulate most hazy items): 1) abdominal segments [3 to 7] distinctly uninterruptedly margined on each side; 2) [anterior visible] abdominal tergites with four short longitudinal basal keels each; 3) pubescence not greyish-silvery; 4) maxillar palpi with basal segment yellow; 5) head not broader than elytra between humeri; 6) body with ground-sculpture developed, otherwise head unusually narrow; 7) body larger; legs usually dark; 8) median longitudinal furrow of pronotum complete, rarely shortened. one can see that the characters have been of unequal diagnostic value. the structure of abdominal tergites as well as their lateral margination and even coloration of maxillar palpi may be regarded as features at the species group level, whereas the body size, colour of legs, proportions of the forebody parts, etc. are fit for the discrimination of particular species only. the character of puncturation, macroand microsculpture vary widely within each of the species groups in nestus. the long and sharp median longitudinal furrow of the pronotum seems to be a detail typical of any species of the group being discussed; but a similar feature has also arisen independently in other complexes (e.g. in s. (n.) melanopus (marsham, 1802), see below), and careful phylogenetic analysis is necessary in every case to prove a real relationship; on the other hand, the furrow may be very feeble and vague in some species of the canaliculatus group (s. (n.) raddei ryvkin, 1987, some samples of s. (n.) canaliculatus gyllenhal, 1827 and s. (n.) nitens stephens, 1833, etc). it is for the reasons aforementioned that a species group definition based on external morphological criteria solely or primarily has proved to be unsatisfactory. fortunately, the male genitalia of steninae, being extremely multiform at the species level, can nevertheless be used in their phylogenetically conservative general structure to establish and confirm the natural groups. therefore, i have used these characters in the new diagnosis of the canaliculatus group (see below). i do not consider here the synonymization of the subgenus nestus rey, 1884 with stenus s.str. undertaken by puthz (2001), as the matter is discussed by me in a separate paper (ryvkin 2011). since a key for identification with brief distributional characteristics was given by me for continental palaearctic species of the canaliculatus group (ryvkin 1987), some new taxa have been described (puthz 1987, 2006), and vast material of the group, from both the palaearctic and the nearctic, became available to me for study. the material from the former ussr and china is of particular interest since both countries together cover the greater part of the palaearctic region and are still the least investigated in the faunal aspect. steninae of the usa and canada seem also to be studied insufficiently: numerous papers of puthz (see the list of references below) dealt only with part of the species and with part of the territory; the nearctic material of many american museums looks rather scanty in comparison with their neotropical collections. the list of staphylinid beetles provided by campbell & davies (1991) for canada & alaska contains no references to material and sources published; as a result, it is impossible to verify the data cited, and many erroneous data of old authors remain in the list concurrently with recent records. unfortunately, the same problem arises with the palaearctic catalogue by smetana (2004) which, owing to the absence of references to the sources of information, turned into a simple check-list, untestable and unusable, in spite of the herculean labour undertaken by the author. the annotated list given below contains the results of processing the material on the canaliculatus group, including the descriptions of four new species as well as numerous new data on distribution with brief analyses of old records for fourteen representatives previously described from both the palaearctic and the nearctic (other species, for which no new data have been provided, are listed in the ‘taxa included’ section only, each marked with an asterisk and supplied with short distributional data and a reference to the source). a key for the identification of all the known palaearctic species of the group is given. ryvkin a.b., stenus species of the canaliculatus group 3 material and methods the original material, which the present paper is based on, is represented by 591 specimens (288 ♂♂, 303 ♀♀). a large proportion of these has been captured during my numerous field trips in 1976–2009; others have been either donated or loaned to me for study by many colleagues and institutions. the names of the respective collections in which the material is deposited now are abbreviated and listed below. in the annotated list below, both type and unique specimen labels are cited in single quotes completely (the type and historic labels with vertical strokes to separate different lines of a label); square brackets are used to complete label data; if necessary, the labels are supplied with remarks in angled brackets; the labels of other specimens are given in english without indicating the original language. the material identified questionably is shown with a question-mark in angled brackets. catalogue references are given only for the sources concerning essential aspects of taxonomy or distribution; for other references see herman (2001). catalogues and compilations have been used mainly if they include any original data and/or interpretations (j. sahlberg 1899, 1900; jakobson 1909; bernhauer & schubert 1911; tichomirova 1973, campbell & davies 1991, ryabukhin 1999), while others (heyden 1881, 1898, etc) are mentioned, as needed, either ad notam or for correcting appreciable errors. the numerous latest regional check-lists, where sources of individual records have been concealed, are mostly not cited below. in cases of doubtful citing, the question-mark in angled brackets was placed either behind the species name (for faunistic records) or before it (for doubtful synonymy) in the respective lines of catalogue sections. if a quite evident misidentification has not been proved by a revision of the material till now, this fact is indicated in the respective catalogue record as ‘?!’ in angled brackets behind the species name. in all cases, when the material identified before by other authors is used, the references to their identifications are given. the line drawings were made with the drawing apparatus ra-7 (lomo); the photomicrographs were produced with a webbers myscope 130-m digital microscope camera (1.5 mpix); the images were processed using helicon focus 3.20.3 free software (www.helicon.com.ua). the measured proportions of body parts are given in points of an eyepiece linear micrometer in a binocular microscope at 56x magnification; prementum of some stenus species was measured dorsally from its base to the base of the glossae, in glycerine preparations; the head length is measured along the midline from the neck restriction to the base of the labrum. abbreviations ♂, ♂♂: male, males. ♀, ♀♀: female, females. ex: specimen, specimens. ht: holotype. pt: paratype. ar: collection of a.b. ryvkin, moscow, russia. ash: collection of a.v. shavrin, daugavpils, latvia. avs: collection of a.v. sokolov, moscow, russia. ayus: collection of a.yu. solodovnikov, copenhagen, denmark. bm: natural history museum (=british museum (natural history)), london, uk (m. barclay, m. brendell). dul: systematic biological institute, daugavpils university, latvia (a.v. shavrin). fmnh: field museum of natural history, chicago, usa (m. thayer, a. newton). gmr: collection of g.m. de rougemont, london, uk. european journal of taxonomy 13: 1-62 (2012) 4 ibpm: institute of biological problems of the north, magadan, russia (d.i. berman, e.g. matis, a.s. ryabukhin). ig: collection of i.n. goreslavets, samara, russia. irsn: institut royal des sciences naturelles de belgique, bruxelles (d. drugmand, m. peeters) kg: collection of k.a. grebennikov, st. petersburg, russia. lh: collection of l. hromádka, praha, czech republic. mhng: muséum d’histoire naturelle, genève, suisse (g. cuccodoro). mtd: museum für tierkunde, dresden, germany (k.-d. klass). onk: collection of o.n. kabakov, st. petersburg, russia. shin: collection of sh.-i. naomi, chiba, japan. sstc: smolensk state teacher’s college, smolensk, russia (m.yu. gildenkov). ti: collection of tateo ito, kyoto, japan. uasm: e.h. strickland entomological museum, university of alberta, edmonton, canada (d. shpeley). vp: collection of v. puthz, schlitz, germany. vs: collection of v.b. semenov, moscow, russia. zin: zoological institute, russian academy of sciences, st. petersburg, russia (†g.s. medvedev). zmh: zoological museum helsinki, finland (h. silfverberg). zmmu: zoological museum of moscow state university, moscow, russia (a.a. gusakov). zmru: zoological museum of rostov state university, rostov-on-don, russia (e.a. khachikov). results: systematic part redefinition of the canaliculatus group diagnosis body size moderate; body length of the known species: 2.8 to 4.7 mm. head fairly small to fairly broad in comparison to elytra and pronotum. upper surface between eyes feebly evenly convex, nearly flat, to slightly concave as a whole; with longitudinal elevations and impressions nearly absent to well developed. antennae rather short to long, with the club segments more or less elongated. internal tooth of each mandible placed a little before the middle, directed only slightly dorsally of the main plane. maxillar palpi with yellow and not shortened basal segment. a considerable reduction of the adhesion capture apparatus may be supposed for all the members of the group (for more details see the remarks section below). pronotum with long and sharp longitudinal median furrow more or less deep, less often very feeble, stroke-shaped. elytra nearly rectangular, large, with short rounded humeri, to shortened, slightly trapezoid with humeri less developed. legs moderately long; the segment 4 of tarsi without emargination; segment 1 of metatarsi about the same length as to a bit longer than segment 5 but distinctly shorter than segments 2 to 4 together. abdomen moderately convex, with paratergites evident on abdominal segments 3 to 7; four anterior visible tergites each with four short but evident longitudinal keels at basal part; posterior margin of the tergite 7 with well-developed fine membranous fringe. puncturation of pronotum and elytra diverse, but always non-rugose. ryvkin a.b., stenus species of the canaliculatus group 5 ground sculpture cellular or net-shaped to entirely absent. pubescence moderate, contiguous to fairly long, outstanding. male. legs without specific features; posterior margin of abdominal sternites 6–7 shallowly emarginated to about straight; abdominal sternite 8 with a broad but not deep, rounded or angularly rounded emargination of posterior margin; abdominal sternite 9 with large posterolateral teeth not incurved inwards; ventral depressions of sternites 6–8, if present, flat and feeble, never flanked by sharp keels. aedeagus relatively large; median lobe archetypically lanceolate, with apical sclerotized part to a variable extent angular to angularly rounded; endophallus with paired medial bands (optionally, an unpaired bar may also be distinct), small but visible lateral parts, and h-shaped expulsion clasp producing lateral portions fairly broad; basal tube of diverse structure, more or less sclerotized. parameres cylindrical, with apical broadening, if present, very feeble, rather vague; apico-internal setiferous surface differentiated poorly to moderately, setae fairly uniform. female. posterior margin of abdominal sternite 8 broadly rounded to angularly rounded; each valvifer with posterolateral tooth directed backwards. spermatheca sclerotized to a greater or lesser extent. in the structure of abdomen, the male genitalia, the abdominal sternite 9 of both males and females (valvifera), the group under consideration is most closely related to the boops and palposus groups of the same subgenus nestus. it differs from these by the structure of the mandibles, by the adhesion capture apparatus more or less shortened, by the pronotum with long and sharp, line-shaped, longitudinal median furrow, by the evidently narrower aedeagus, by the endophallus with the expulsion clasp’s transversal bridge adjoining to the lateral pieces near their middle, by the abdominal sternite 9 with posterolateral teeth not being curved inwards, by the male legs without peculiar features (not all the species of the named groups have such features well developed); from the palposus group it can be easily distinguished by the yellow basal segment of the maxillar palpi. the differences from s. (n.) melanopus marsham, 1802, similar externally to some species of the canaliculatus group, are given in the diagnosis of the melanopus group and in the key (see below). in many characters of the structure of abdomen and male genitalia, the canaliculatus group also resembles the aggregate including the large and diverse atratulus-fuscipes-cautus-crassus complex and some isolated american members, as s. (n.) mendosus puthz, 1971 (1971d) and s. (n.) sordidus puthz, 1988, but phylogenetic relations within the named aggregate remain poorly understood; therefore a thorough analysis seems to be premature. in external morphology, the canaliculatus group can easily be distinguished from all the species of the mentioned complex by the presence of the long and sharp, line-shaped, longitudinal median furrow of the pronotum. taxa included s. (n.) alopex sp. nov., s. (n.) brivioi puthz, 1972, s. (n.) canaliculatus gyllenhal, s. (n.) canalis sp. nov., 1827, s. (n.) canosus sp. nov., s. (n.) caseyi puthz, 1972, s. (n.) confusus j. sahlberg, 1876, s. (n.) delitor sp. nov., s. (n.) dolosus casey, 1884, s. (n.) geminorum puthz, 1973* (usa: washington; canada: bc, see puthz 1973b: 208), s. (n.) idoneus sharp, 1887* (mexico, see sharp 1887: 799; puthz 1968: 11 <as laniger puthz, 1968>), s. (n.) illotulus puthz, 1972, s. (n.) illusor ryvkin, 1987, s. (n.) immigratus puthz, 2006* (china: taiwan: ilan hsien: shen mi lake, 1100 m, see puthz, 2006: 187), s. (n.) labilis erichson, 1840, s. (n.) latipennis j. sahlberg, 1880, s. (n.) milleporus casey, 1884, s. (n.) nitens stephens, 1833, s. (n.) raddei ryvkin, 1987, s. (n.) shogun puthz, 1987* (japan: nishigo uzan, see puthz 1987: 45), s. (n.) sphaerops casey, 1884, s. (n.) vinnulus casey, 1884. european journal of taxonomy 13: 1-62 (2012) 6 notes on comparative morphology and evolution as most other polytypic groups in steninae, the canaliculatus group reveals diverse evolutionary trends quite evident at a comparison between species of the same lineage. a concise review of the mentioned trends is given below. a. upper surface of head between eyes tends either to become flattened, at most slightly convex (s. shogun, s. nitens, s. illotulus, s. raddei, s. caseyi, s. canaliculatus, s. canalis, s. canosus, s. alopex) or to produce distinct longitudinal impressions with median elevation in between (s. labilis, s. idoneus, s. vinnulus, s. brivioi, s. confusus, s. delitor, s. illusor, s. latipennis, s. immigratus, s. geminorum, s. sphaerops, s. dolosus, s. milleporus). the latter tendency seems to arise and develop independently, at least within the complexes labilis-confusus-delitor-illusor-latipennis-sphaerops and dolosus-milleporus, in spite of the fact that the latter is closely related in the shape of the male genitalia to the ‘canaliculatus s.str.’ complex characterised by a flat front. b. for three species studied by him (s. canaliculatus, s. nitens, s. labilis), betz (1996) has shown a considerable reduction of the adhesion capture apparatus. it is expressed both in the degeneration of sticky cushions of paraglossae (decreasing of their surface and number of adhesive setae etc.) and in a shortening of the labium. no data on labium length have been provided by the named author but, as it follows from the text, s. milleporus (as sectilifer casey, 1884), s. caseyi, s. vinnulus, as well as s. melanopus, which is regarded as a member of the same group, have also been studied in this respect. i have measured the relative length of the prementum (eulabium after weinreich 1968) for six species of the canaliculatus group: s. canaliculatus (i: 17, 0.53; ii: 16, 0.53), s. nitens (33, 0.87), s. labilis (37, 1.12), s. illotulus (26, 0.84), s. raddei (35, 1.09), s. latipennis (31, 0.79); and for three species of other groups of the subgenus nestus rey, 1884: s. ruralis erichson, 1840 (47, 1.52) from the palposus group, s. argus gravenhorst, 1806 (47, 1.74) from the fuscipes group, and s. melanopus marsham, 1802 (41, 1.46) presumably representing a monotypic group. in each case above, the first number in brackets means length of prementum whereas the second one is a rounded ratio between the former and the length of the head (two specimens have been measured for s. canaliculatus). one can see that the data split the examined species into three clusters: a) s. canaliculatus; b) the remaining species of the canaliculatus group; c) the species of other groups; both total and relative length of prementum increases from category a) to c). it can be supposed that the reduction of the eulabium is a characteristic feature of all the members of the canaliculatus group, but it must be verified by a specific study. the shortening of the adhesion capture apparatus seems to correlate with some other modifications of the mouth parts, namely the more proximal position of the internal tooth of mandibles as well as its lesser deflection off the main plane of the mandible. the small median notch of the labrum is a specific character of s. canaliculatus and the most closely related palaearctic species (s. canalis, s. canosus, s. alopex). though the functional explanation of this feature is obscure as yet, one can suppose it to correlate with the extreme reduction of the adhesion capture apparatus in this species complex. the nearctic members of the same complex (s. milleporus and s. dolosus) have only a broad and shallow emargination on the anterior margin of the labrum; the margin is slightly sinuate at the middle in s. nitens, s. illotulus, s. raddei, s. caseyi, s. brivioi, s. vinnulus; lastly, the species of the labilis-confusus-delitor-illusor-latipennis-sphaerops line have the margin nearly straight. betz (1996) supposed the reduction of the adhesion capture apparatus in the canaliculatus group to be a result of miniaturisation, but this explanation does not stand up to any criticism. indeed, the smallest specimens of the species within the group may be considered among the medium-sized representatives of the genus, while even much smaller members of both the pusillus and the pumilio groups have the adhesion capture apparatus well developed (s. (n.) pusillus stephens, 1833 has been studied by betz (1996) himself). ryvkin a.b., stenus species of the canaliculatus group 7 i believe that the modification of the mouth parts should be connected more likely to a particular suite of potential prey species. unfortunately, betz used only springtails and (to a lesser extent) winged aphids in his interesting prey-capture experiments, but those may not exhaust the list of possible prey objects for steninae. i did not experiment with species of the canaliculatus group; however other stenus species were capturing nymphs of miridae (insecta: hemiptera: heteroptera) and cicadellidae (insecta: hemiptera: auchenorrhyncha), juvenile spiders of the family linyphiidae (arachnida: aranei), small mites (arachnida: acari), small pyralid caterpillars (insecta: lepidoptera: pyralidae), small flies (diptera), soil enchytraeidae (oligochaeta), in addition to the aforementioned objects, in my unpublished experiments of 1985–1986. i consider it necessary to undertake a more detailed study of prey for the species of the group being discussed. c. the tendency of legs to become longer has most likely taken place independently in two species complexes: delitor-illusor-latipennis-confusus-sphaerops and nitens-illotulus-raddei (only the species with the 1st segment of the metatarsus longer than the 5th are listed). s. canaliculatus, s. canalis, s. canosus, s. alopex, s. milleporus, s. caseyi have metatarsi with the 1st segment about as long as 5th; in s. dolosus, s. brivioi, s. vinnulus, and s. labilis, metatarsi remain short (with the 1st segment evidently shorter than the 5th). d. the plesiomorphic aedeagus with a lanceolate median lobe and unmodified cylindrical parameres (as in s. labilis) can be transformed by different ways. the distal sclerotized part of the median lobe may be shortened, either angularly (as in s. sphaerops) or rounded-angularly (in the delitor-illusor-latipennis line); in only one case with a small apical denticle (in s. vinnulus); this modification usually correlates to a lengthening of the whole aedeagus (see figs. 6, 7a–i) or only the median lobe (as in s. confusus, see fig. 2e), although this correlation has not been observed in s. raddei, belonging to another lineage (fig. 2a, b). the tendency of the median lobe to become narrower (as in s. shogun, s. brivioi, s. alopex) is another way of transformation, being peculiar to the lineages of nitens and ‘canaliculatus s.str.’; one can suppose that the aedeagus of mexican s. idoneus has evolved in this way. all the modifications mentioned may be followed by forming a more or less developed medial impression or/and keel at the distal sclerotized part of the median lobe. the main trend in the evolution of the parameres is a differentiation of their setiferous surface. it is expressed more frequently in a flattening of the apicointernal side of each paramere or, on the contrary, in forming a one-sided swell which setae are attached to. based on all the characters and tendencies briefly discussed above, we can subdivide the canaliculatus group into four subgroups: the monotypic labilis subgroup (s. labilis), the nitens subgroup (s. immigratus, s. nitens, s. raddei, s. illotulus, s. shogun, s. caseyi, s. geminorum, s. brivioi, s. idoneus), the canaliculatus s.str. subgroup (s. dolosus, s. milleporus, s. canaliculatus, s. canalis, s. canosus, s. alopex), and the confusus subgroup (s. confusus, s. delitor, s. illusor, s. latipennis, s. sphaerops, s. vinnulus). bionomics the majority of the species for which enough data have been presented can be grouped into swamp inhabitants and alluvial ones. the former category, comprising the species dwelling in moss and litter of eutrophic and mesotrophic wet habitats, definitely includes s. nitens, s. raddei, s. illotulus, s. caseyi, s. brivioi and s. sphaerops(?), whereas the latter category consists of the members inhabiting open alluvia, like shingle beds, sandy, clayey or silty banks, sometimes sparsely overgrown with grass but without any more or less evident peat-moss or litter layer: s. labilis, s. delitor, s. latipennis, s. confusus, s. dolosus, s. canaliculatus, s. canosus, s. canalis. there are also some species (s. alopex, s. vinnulus, and supposedly s. illusor) living in biotopes of both categories. the peculiar swamp species, excluding european journal of taxonomy 13: 1-62 (2012) 8 s. sphaerops for which too little data are available, are represented only by the nitens subgroup, which is supposed to be closest to the common ancestor of the canaliculatus group. general distribution palaearctic except subtropical regions (spotty-relict in the southernmost temperate territories); the whole nearctic; the caribbean islands. many species of the group are represented in the northernmost territories of the holarctic.thus, five of the fifteen palaearctic species have been recorded for the taimyr peninsula and putorana highland, which is more than 1/4 of the regional stenine fauna. species list stenus (nestus) labilis erichson, 1840 (fig. 1e) stenus labilis erichson, 1840: 697. stenus labilis – thomson 1857: 223. — j. sahlberg 1871: 415. — j. sahlberg 1900: 29. — l. benick 1921a: 146. — puthz 1971b: 34. — silfverberg 1988: 21 <types of lapponicus j. sahlberg, 1876>. — sokolov 2003: 1272. — shavrin & puthz 2007: 122. stenus labilis <?!> – fauvel 1865: 306. — iljin 1926: 225. stenus (nestus) labilis – poppius 1909b: 19. — jakobson 1909: 481. — renkonen 1935: 29. — palm 1961: 90, 98. — puthz 1967a: 49. — puthz1967d: 293. — tichomirova 1973: 173. — shilov 1975: 58. — ryvkin 1987: 159. stenus (nestus) labilis <?!> – poppius 1909a: 17. stenus (s. str.) labilis – campbell & davies 1991: 111. stenus canaliculatus – c. sahlberg 1832: 428. stenus lapponicus j. sahlberg, 1876: 56. stenus lapponicus – j. sahlberg 1880: 78. — heyden 1881: 78. — j. sahlberg 1899: 340. stenus (nestus) lapponicus – jakobson 1909: 481. — hansen et al. 1939: 32 (pars). stenus latipennis – renkonen 1941: 105. material examined finland: 1 ♂, ‘lac. kemi’ <printed on yellowed white rectangle>, ‘j. sahlb.[erg]’ <printed on yellowed white rectangle>, ‘77’ <hand-written with indian ink on grey square>, ‘st. labilis er. typ (=lapponicus j. sahlb.) <black indian ink> j. sahlberg det. <printed in italics>’ <common determinative label by j. sahlberg> (zin); 1 ♀, ‘turtola’ <printed on yellowed white rectangle>, ‘j. sahlb.[erg]’ <printed on yellowed white rectangle>, ‘stenus lapponicus’ <black indian ink> (zin); 1 ♂, ‘tenojoki’ <printed on yellowed white rectangle>, ‘j. sahlb.[erg]’ <printed on yellowed white rectangle>, ‘st. labilis er. (=lapponicus j. sahlb.)’ <black indian ink>, ‘c.[ollection] of a. jakovlev’ <printed label in russian> (zin). russia: 1 ♂, arkhangelsk area, 10 km n of naryan-mar, near iskateley, sandy bank of pechora river, 14 jul. 1988, p.k. yeryomin leg. (ar); 1 ♂, arkhangelsk area, 5 km s of naryan-mar, bank of right confluent of pechora river, on silt, 19 jul. 1988, p.k. yeryomin leg. (ar); 1 ♀, arkhangelsk area, naryanmar, swamp, 29 jul. 1988, p.k. yeryomin leg. (ar); 1 ♂, arkhangelsk area, nenetzkiy autonomous region, pechora river delta, sedge meadow at lake shore, 7 sep. 1998, n.s. mazura leg. (ar); 1 ♀, ‘yugorskiy shar, near velikaya river, tundra, 27.viii.1921, e. abakumova leg.’ <in russian>, ‘labilis er. l. benick det.’ (zin); 1 ♂, arkhangelsk area, near amderma, flood-plain of amderminka river, ryvkin a.b., stenus species of the canaliculatus group 9 fig. 1. – a–d: stenus (nestus) nitens stephens, 1833: ♂ genitalia, ventral view. a–b: (russia: moscow area: gorenki). c–d: (russia: evenkia: stolbovaya river). a, c. aedeagus. b, d. apical part of median lobe. – e. s. (n.) labilis erichson, 1840: ♂ (russia: s yamal: nr. salekhard). aedeagus, ventral view. – scales = 0.1 mm. european journal of taxonomy 13: 1-62 (2012) 10 on silt, 9 jul. 1988, p.k. yeryomin leg. <the specimen destroyed by dermestids, remained on cotton> (ar); 1 ♂, south of yamal peninsula, 110th km by railway n off obskaya station, puddle, 6 aug. 2001, p. petrov leg. (ar); 1 ♂, w siberia, salekhard, poluy river bank, 27 jun. 1954, i. telishev leg. <det. v. puthz, 1971> (zmmu); 1 ♂, yamalo-nenetskiy autonomous region, nr. salekhard, 5 jul. 1940 (ar); 1 ♀, muzhi-na-obi, 23 jun. 1952, i. telishev leg. (zmmu); 1♂, beryozov, 13 jul. 1932, telishev leg. <det. v. puthz, 1971, with earlier determinative label by unknown author: ‘nestus melanarius steph.’> (zmmu); 7 ♂♂, 12 ♀♀, khanty-mansi autonomous region, near shapsha, 61.085°n 69.458°e, 42 m a.s.l., 1–4 aug. 2010, k. tomkovich leg. (ar, shin); 1 ♂, khanty-mansi autonomous region, surgutskiy district, surgutskiy refuge, ob’ river basin, materikovyi pasl channel, bank of rill flowing into channel: near water (clay), carex sp., poaceae gen. sp., veronica ? longifolia, asteraceae, small mosses, 8 aug. 2000, a.b. ryvkin leg. (ar); 1 ♂, khanty-mansi autonomous region, surgutskiy district, yuganskiy nature reserve, malyi yugan river, near cordon below lyarykni river mouth, on sand and clay and in grass litter among carex spp. at river bank, 15 aug. 2003, a.b. ryvkin leg. (ar); 1 ♂, 1 ♀, tyumen area, uvatskiy district, 10 km s of gornoslinkino, near tobolsk field research station of severtsov institute, among carex spp. on flood-plain of varpak river, 25 sep. 2003, a.b. ryvkin leg. (ar); 2 ♂♂, 4 ♀♀, sw taimyr, upper reaches of nizhnyaya agapa river, nyapan ridge, 70°04´n 87°36´e, detritus at shore of ladannakh lake, 17 jul. 2001, a.v. sokolov leg. (avs); 1 ♀, same locality and collector, shingle bed, 2 jul. 2001 (avs); 2 ♂♂, 3 ♀♀, same locality, soil traps on swamp, 8 jul.–15 aug. 1999, o. makarova leg. (avs); 1 ♂, 1 ♀, taymyr, tammot lake, 10 km ene of lukunskoye cordon, on silt under flood-plain, salix sp., 15 jul. 1989, p.k. yeryomin leg. (ar); 1 ♂, same locality, slightly silty sandy lake shore with small shingles, 24 jul. 1989, p.k. yeryomin leg. (ar); 1 ♀, taymyr, island on khatanga river, 10 km ne of oboynaya village, slightly silty sandy river bank, 19 jul. 1989, p.k. yeryomin leg. (ar); 1 ♂, taymyr, maymecha river mouth, yantardakh mount, 7 jul. 1971 a.p. rasnitsyn, a.g. ponomarenko, i.d. sukatcheva, v.v. zherikhin leg. (ar); 1 ♂, 1 ♀, taymyr, khatangskiy district, taymyr nature reserve, ary-mas: right bank of novaya river, lake ese of ulakhan-yuryakh river mouth, mosses, litter and sweeping at lake shore: carex spp., comarum palustre etc., 20 jul. 1992, a.b. ryvkin leg. (ar); 1 ♂, taymyr, khatangskiy district, taymyr nature reserve, ary-mas: novaya river basin, bogatyr’-yuryakh river near mouth, steep river bank and high river plain, clay, drift, poaceae gen. sp., carex spp., salix spp., alnus fruticosa, ribes sp., 21 jul. 1992, a.b. ryvkin leg. (ar); 5 ♂♂, taymyr, khatangskiy district, taymyrskiy nature reserve, ary-mas: left bank of novaya river near field research station, river bank, sand with thin layer of silt and drift near water, 22 jul. 1992, a.b. ryvkin leg. (ar). canada: 2 ♀♀, ‘nwt, loc. 06-77, anderson river delta, boat island, salix alexensis zone: traps, 26 jun.–15 jul 1977, d. shpeley, g.e. ball leg. (anderson river exp. – 1977)’ (uasm). remarks this species was named by erichson (1840) from ‘fennia’, having been misidentified earlier by c. sahlberg (1832) as s. canaliculatus gyllenhal, 1827; redescribed as s. lapponicus j. sahlberg, 1876 from finland (up to 68°30’n), but cited as labilis (only!) by j. sahlberg (1900) in his catalogue of the beetles of the fauna of finland. only the norwegian, swedish, finnish, and n russian records (thomson 1857; j. sahlberg 1871, 1899, 1900; poppius 1909b; jakobson 1909; palm 1961; puthz 1971b) seem to be reliable for the european range of the species. the records for france/italy (fauvel 1865), as well as for the yekaterinoslav government (=dnepropetrovsk area) of the ukraine (iljin 1926), are to be regarded as results of misidentification. in western and central siberia, s. labilis has been known from polar ural to w taymyr (j. sahlberg 1880, l. benick 1921a, puthz 1967a, shilov 1975, sokolov 2003 <the latter is based on my identifications>). puthz (shavrin & puthz 2007), without new material, considers the species to be distributed in e siberia, but the only record by poppius (1909a), for the lena basin (‘mehrere exemplare auf feuchten, moos-bewachsenen wiesen auf der insel agrafena, 1.viii!.’), has been confirmed by neither subsequent authors nor my vast material from this spacious territory. ryvkin a.b., stenus species of the canaliculatus group 11 thus, s. labilis is unlikely to be represented in siberia eastwards of e taymyr. the new material above confirms the distribution of the species in n america (campbell & davies 1991: alaska, yukon territory, northwest territories). most captures known to me are confined to alluvial habitats. stenus (nestus) nitens stephens, 1833 (fig. 1a–d) stenus nitens stephens, 1833: 300. stenus nitens – waterhouse 1858: 28. — j. sahlberg 1876: 58. — poppius 1899: 39. — j. sahlberg 1899: 340. — j. sahlberg 1900: 29. — szujecki 1960: 297. — puthz 1971b: 34. —tichomirova 1982: 213. — pisanenko & puthz 1991: 170. — puthz & zanetti 1995: 18. — shavrin & puthz 2007: 125. stenus (nestus) nitens – jakobson 1909: 481. — reitter 1909: 158. — renkonen 1935: 29. — szujecki 1961: 34. — palm 1961: 90. — puthz 1967d: 293. — puthz 1971a: 82. — tichomirova 1973: 173. — shilov 1975: 58. — ryvkin 1987: 159. — dauphin 1993: 188. — semenov 2004: 12. stenus (nestus) nitens <?!> – poppius 1909a: 16. — bordoni 2004: 120. stenus aemulus erichson, 1839: 541. stenus aemulus – hochhuth 1872: 154 (as s. aemulus gyllenhal). material examined finland: 1 ♂, ‘fennia. reitter’, ‘stenus nitens steph.’, ‘c.[ollection] of a. jakovlev’ <printed label in russian> (zin); 1 ♂, 1 ♀, ‘fennia’<printed label on red paper>, ‘j. sahlb.[erg]’ <printed on yellowed white rectangle>, ‘stenus nitens stph.’ <black indian ink> (zin: ex coll. semenov-tian-shanskiy); 1 ♂, ‘helsingfors. 4126[=stenus nitens steph.] – 3’ (zin). russia: 1 ♀, ‘petropolis. 4126[=stenus nitens steph.] – 5’, ‘nitens steph.’ (zin); 1 ♀, tyumen area, nefteyuganskiy district, 90 km s of nefteyugansk, near bolshoye kayukovo lake, geobotanical field research station of moscow university, floating island on sedge eutrophic swamp: in hummock base, 20 aug. 1985, r.a. rakitov leg. (ar); 1 ♂, 2 ♀♀, khanty-mansi autonomous region, surgutskiy district, yuganskiy nature reserve, nyogus’yakh river, bisarkina cordon, moss and litter on swamp amid flood plain forest near foot of slope: menyanthes trifoliata, equisetum spp., sphagnum squarrosum (sparse), sph. ? centrale, sph. spp., nardosmia frigida, carex spp., poaceae gen. sp., salix sp., comarum palustre, filipendula ulmaria, ? tomenthypnum sp., plagiomnium sp., etc., 27 jul. 1998, a.b. ryvkin leg. (ar); 1 ♂, khanty-mansi autonomous region, surgutskiy district, yuganskiy nature reserve, nyogus’yakh river basin near pechpan’yakh river mouth, mosses and litter on large swamp: carex spp., comarum palustre, equisetum sp., nardosmia frigida, eriophorum sp., poaceae gen. spp., rumex sp., chamaenerion angustifolium, salix sp., ? aulacomnium sp., plagiomnium sp., ptilium cristacastrensis, sphagnum ?russovii, sph. sp. etc., 15 sep. 2000, a.b. ryvkin leg. (ar); 1 ♂, 1 ♀, khantymansi autonomous region, surgutskiy district, yuganskiy nature reserve, ay-magromsy river basin, medvezhyi ugol cordon, mosses and litter at southern border of sedge-mossy swamp: carex rostrata, c. spp., comarum palustre, equisetum spp., sphagnum girgensohnii, sph. spp., plagiomnium sp., aulacomnium sp., calamagrostis sp., menyanthes trifoliata, etc. among sparse young birches (t=+10°– +5°c), 4 oct. 2002, a.b. ryvkin leg. (ar); 1 ♂, khanty-mansi autonomous region, surgutskiy district, yuganskiy nature reserve, malyi yugan river basin, 3–5 km sw of cordon at kol-kochen-yagun river, mosses and litter by swampy margin at n side of lake: carex spp., sphagnum spp., poaceae gen. spp., 26 sep. 1999, a.b. ryvkin leg. (ar); 1 ♀, khanty-mansi autonomous region, surgutskiy district, yuganskiy nature reserve, malyi yugan river, cordon below lyarykni river mouth, open swamp with tussocks of carex spp. and calamagrostis sp. with comarum palustre, salix sp., sphagnum squarrosum, sph. spp., ? aulacomnium sp. etc. among sparse young betula sp., 17 aug. 2003, a.b. ryvkin leg. (ar); european journal of taxonomy 13: 1-62 (2012) 12 1 ♂, krasnoyarsk territory, turukhanskiy district, bakhta river basin, near keteollo lake, noya river source, 323 m a.s.l., mosses and litter among carex spp., filipendula ulmaria, poaceae on tussocks and among those at open river bank, 6 aug. 1992, a.b. ryvkin leg. (ar); 1 ♂, evenkia, baykitskiy district, central siberian biosphere reserve, stolbovaya river 8 km up-stream of river mouth, 60 m a.s.l., mosses and litter on open swamp with carex spp., comarum palustre, sparse menyanthes trifoliata, true mosses, sphagnum spp. etc., 20 sep. 1991, a.b. ryvkin leg. (ar); 1 ♂, evenkia, baykitskiy district, podkamennaya tunguska river, 175 km up-stream of river mouth, near belaya kosa island, 55 m a.s.l., swamp with comarum palustre, sphagnum spp., equisetum sp., plagiomnium sp., carex spp., etc., 23 aug. 1990, a.b. ryvkin leg. (ar). remarks terra typica: ‘london; suffolk’; for aemulus: ‘brandenburg’. the species is known from c & n europe and w siberia (tichomirova 1973). hochhuth (1872) reported it (as aemulus gyllh. <sic!>) for the kiev government (‘ziemlich selten’); j. sahlberg cited it for n karelia (1876: ‘nordligast har jag funnit den vid soukelo i ryska lappmarken (66°40’)’), ‘peninsula kola, petschora’ (1899); jakobson (1909) summarized records for ‘arkhangelsk government (lapponia, pechora), <…> olonets government, st.-petersburg government, kiev government’. puthz (pisanenko & puthz 1991) mentioned that ‘aus der europäischen sowjetunion war diese art horion 1963 unbekannt’, but, besides the records named above, puthz (1971a) himself cited an additional find for the leningrad area (‘viipuri’=vyborg); tichomirova (1982) recorded the species for the moscow area. the sketchy figure of an aedeagus provided by bordoni (2004) for s. nitens from italy (‘veneto: verona’) seems to belong to a specimen of another species. unfortunately the specimen which has been figured by bordoni is not found in museo civico di storia naturale, verona (a. zanetti pers. comm.); thus, the italian records of the species require a confirmation. the new material proves that the species is widely distributed in w & c siberia. the e siberian records by poppius (1909a: ‘an der mittleren lena weit verbreitet, aber einzeln. <...> olekminsk, ytyk-haja, önkyr-yrjä, ust-aldan, zwischen ust-aldan und batylym, shigansk’) have been confirmed by neither subsequent writers nor the new material. the aforecited material from w evenkia seems to provide the easternmost verified records of the species until now. the shape of the aedeagus of the male from evenkia corresponds well to that of the specimen from european russia (see fig. 1a–d). stenus (nestus) raddei ryvkin, 1987 (fig. 2a, b) stenus (nestus) raddei ryvkin, 1987: 156. stenus raddei – herman 2001: 2365. stenus (nestus) raddei – naomi & puthz 1994: 219. stenus (s. str.) raddei – smetana 2004: 561. material examined russia: 1 ♀, maritime province, lazovskiy nature reserve, 10 km w of preobrazheniye, swampy meadow, t=20°, ph=5.4, 17 aug. 1986, a. shatrovskiy leg. (ar); 1 ♂, amur province, selemdzhinskiy district, near selemdzhinsk, 270–280 m a.s.l., 12 aug. 1976, e.m. veselova & a.b. ryvkin leg. (ar); 1 ♀, amur area, selemdzhinskiy district, norskiy reserve, near fevral’sk, 268th km of belogorskfevral’sk road, tikhiy rill, 275 m a.s.l., mosses and plant debris between sedge & gramineous tussocks among alnus sp., salix sp., spiraea sp. with sphagnum squarrosum, sph. spp., etc., 8 oct. 2008, a.b. ryvkin leg. <?> (ar); 3 ♂♂, 4 ♀♀, amur area, selemdzhinskiy district, norskiy nature reserve, selemdzha river basin near dvadtsatikha cordon, open swamp: carex sp., spiraea spp., geranium sp., vicia sp., poaceae gen. sp., salix spp., filipendula palmata, sparse plagiomnium sp., sphagnum sp., polytrichum sp., etc., 4 aug. 2004, a.b. ryvkin leg. (ar); 1 ♀, same locality, mosses and plant debris ryvkin a.b., stenus species of the canaliculatus group 13 among tussocks at small open swamp with carex spp. and poaceae gen. sp., 10 aug. 2004, a.b. ryvkin leg. (ar); 3 ♂♂, 3 ♀♀, amur area, selemdzhinskiy district, norskiy nature reserve, selemdzha river basin, 1.5 km ne of dvadtsatikha cordon, banks and burnt flood-plain of a rill inflowing to lake, 222 m a.s.l., mosses and leaf litter among carex spp., alnus sp., salix spp., padus sp., undergrowth of betula spp. and populus tremula, etc., 18 aug. 2006, e.m. veselova & a.b. ryvkin leg. (ar); 6 ♂♂, 4 ♀♀, amur area, selemdzhinskiy district, norskiy nature reserve, selemdzha river basin, 2 km ne of dvadtsatikha cordon, open swamp near lakeside: plant debris and sparse mosses among tussocks with carex spp., poaceae gen. sp., salix spp., etc., 9 aug. 2004, a.b. ryvkin leg. (ar, shin); 4 ♂♂, 3 ♀♀, amur area, selemdzhinskiy district, selemdzha river basin, flood-plain of aldikon river near norsk, mosses and leaf litter under salix sp. and along the edge of open sedge-gramineous tussock swamp, 5 sep. 2004, a.b. ryvkin leg. (ar, lh); 11 ♂♂, 21 ♀♀, amur area, selemdzhinskiy district, norskiy nature reserve, nora river basin near maltsevskiy cordon, mosses and plant debris on swampy floodplain lakeside: carex spp., calamagrostis sp., filipendula palmata, geranium sp., comarum palustre, spiraea sp., salix sp., sphagnum ? girgensohnii, sph. squarrosum, polytrichum commune, p. sp., climacium sp., iris sp., etc., 28 aug. 2004, a.b. ryvkin leg. (ar); 1 ♂, amur area, selemdzhinskiy district, norskiy nature reserve, rill (left confluent of nora river) up-stream of gryashchinskaya mt., true mosses and plant debris among carex spp., poaceae gen. sp., alnus sp., padus sp., salix sp., etc., 26 aug. 2004, a.b. ryvkin leg. (ar); 7 ♂♂, 3 ♀♀, amur area, selemdzhinskiy district, norskiy nature reserve, nora river basin, 1.5 km up-stream of gryashchinskaya mt., mosses and plant debris on open swamp along lake side: tussocks of carex spp. and poaceae with sphagnum ? girgensohnii, sph. ? angustifolium, sph. ? magellanicum, sph. squarrosum, sph. spp., chamaedaphne calyculata, vaccinium uliginosum, sparse ledum palustre, salix sp., betula ? fruticosa, etc., 24 aug. 2004, a.b. ryvkin leg. (ar); 1 ♂, 3 ♀♀, same locality, moss and plant debris on carex-eriophorum swamp with chamaedaphne calyculata, gramineous admixture, comarum palustre, salix sp., vaccinium uliginosum, sphagnum spp., etc., 25 aug. 2004, a.b. ryvkin leg. (ar); 3 ♂♂, 9 ♀♀, amur area, selemdzhinskiy district, norskiy nature reserve, nora river basin, 2 km up-stream of gryashchinskaya mt., mosses and plant debris on small open swamp on high flood-plain: tussocks of calamagrostis sp. and carex spp. with sphagnum ? girgensohnii, sph. squarrosum, sph. centrale, sph. spp., rubus arcticus, convallaria keiskei, salix sp., etc., 22 aug. 2004, a.b. ryvkin leg. (ar); 3 ♂♂, 1 ♀, amur area, selemdzhinskiy district, norskiy nature reserve, nora river basin near meunskiy cordon, mosses and plant debris among carex spp. & poaceae with comarum palustre, sphagnum squarrosum, sph. spp., aulacomnium sp. and other true mosses near side of flood-plain lake and swampy road, 13 aug. 2004, a.b. ryvkin leg. (ar); 1 ♂, same locality, swamp with sedge tussocks, poaceae gen. sp., sphagnum squarrosum, sph. spp., etc., 14 jul. 2005, a.b. ryvkin leg. (ar); 3 ♂♂, amur area, selemdzhinskiy district, norskiy nature reserve, nora river basin near meun river mouth, small swamp with carex spp., poaceae & sphagnum squarrosum, 19 aug. 2004, a.b.ryvkin leg. (ar); 2 ♂♂, 5 ♀♀, same locality & biotope, 20 aug. 2004, a.b. ryvkin leg. (ar); 1 ♀, amur area, selemdzhinskiy district, norskiy nature reserve, meun river mouth, plant debris among carex spp. & equisetum sp. and leaf litter under salix spp. & alnus sp. on low bank of creek, 21 aug. 2004, a.b. ryvkin leg. (ar); 1 ♂, amur area, selemdzhinskiy district, norskiy nature reserve, burunda river bank 3 km up-stream of mouth, 215 m a.s.l., mosses and plant debris among sedges etc. and on shingle bed, 3 aug. 2006, e.m.veselova & a.b. ryvkin leg. (ar); 1 ♀, amur area, selemdzhinskiy district, norskiy nature reserve, burunda river near ozyornyi rill mouth, mosses and plant debris among sedge-gramineous tussocks on very gentle slope near intermittent channel: carex spp., calamagrostis sp., polytrichum sp. and other true mosses, spiraea sp., etc., 27 sep. 2004, a.b. ryvkin leg. (ar); 2 ♂♂, 3 ♀♀, same locality & biotope. 28 sep. 2004, a.b. ryvkin leg. (ar); 5 ♂♂, 1 ♀, same locality, plant debris and mosses among tussocks of carex spp. and poaceae with spiraea sp., vicia sp., small true mosses, etc. on gently sloping side of small flood-plain lake, 2 oct. 2004, a.b. ryvkin leg. (ar); 22 ♂♂, 2 ♀♀, amur area, selemdzhinskiy district, norskiy nature reserve (buffer zone), burunda river basin, 1.5–2 km nw of burunda cordon, mosses and plant debris on swamp and among sedge-gramineous tussocks in european journal of taxonomy 13: 1-62 (2012) 14 fig. 2. – a–b: stenus (nestus) raddei ryvkin, 1987: ♂ (russia: magadan area: sibit-tyhellakh). a. aedeagus, ventral view. b. apical part of aedeagus, ventral view. – c–d: s. (n.) illotulus puthz, 1972: ♂ (russia: amur area: zeyskiy nature reserve). c. aedeagus, ventral view. d. apical part of aedeagus, ventral view. right paramere removed. – e. s. (n.) confusus j. sahlberg, 1876: ♂ (russia: kalinin area: putilovo). aedeagus, ventral view. – scales = 0.1 mm. ryvkin a.b., stenus species of the canaliculatus group 15 sparse growth of burnt betula platyphylla and larix gmelinii near kamyshovoye lake with chamaedaphne calyculata, comarum palustre, vaccinium uliginosum, ledum palustre, sphagnum spp., polytrichum sp., hypnum sp., etc., 19 sep. 2004, a.b. ryvkin leg. (ar); 1 ♀, amur area, selemdzhinskiy district, norskiy nature reserve (buffer zone), burunda river basin, 1.5 km nw of burunda cordon, sedge-gramineous swamp near rill, and also mosses and leaf litter among carex spp. and under alnus sp. on rill bank, 10 sep. 2004, a.b. ryvkin leg. (ar); 1 ♀, same locality, plant debris among carex sp. under sparse alnus sp. with young growth of larix gmelinii along rill bank near road, 18 sep. 2004, a.b. ryvkin leg. (ar); 1 ♀, amur area, selemdzhinskiy district, norskiy nature reserve (buffer zone), burunda river basin, 0.5–1 km nw of burunda cordon, plant debris and small true mosses among carex spp. & poaceae (+sweeping) along rill bank, 12 sep. 2004, a.b. ryvkin leg. (ar); 3 ♀♀, amur area, selemdzhinskiy district, norskiy nature reserve (buffer zone), burunda river basin near burunda cordon, plant debris and mosses (+sweeping) on swampy sides of flood-plain lake: carex spp., poaceae gen. sp., spiraea sp., sparse climacium sp., hypnum sp., sphagnum sp., polytrichum sp., etc., 9 sep. 2004, a.b. ryvkin leg. (ar); 1 ♂, 3 ♀♀, amur area, selemdzhinskiy district, norskiy nature reserve (buffer zone), burunda river basin, 1 km se of burunda cordon, mosses and plant debris among sedge tussocks on wide carex-eriophorum-gramineous swamp with small true mosses, sphagnum squarrosum, sph. sp., ledum palustre, vaccinium uliginosum, sparse salix sp., young growth of betula sp., etc., 21 sep. 2004, a.b. ryvkin leg. (ar); 1 ♀, amur area, selemdzhinskiy district, norskiy nature reserve (buffer zone), burunda river basin, 2.5 km se of burunda cordon, small true mosses and plant debris among carex spp. on sand, small shingles and clay along dried ephemeral stream near border of complex swamp, 21 sep. 2004, a.b. ryvkin leg. (ar); 1 ♀, khabarovsk territory, okhotsk, airport, litter in larch forest, 15 jul. 1987, v.v.zherikhin & i.d.sukacheva leg. (ar); 1 ♀, khabarovsk territory, okhotskiy district, khetana river (confluent of amka river, ulya river basin), 7 km up-stream of river mouth, 31 jul. 1985, v. zherikhin, a. rasnitsyn & d. shcherbakov leg. (ar); 1 ♀, same locality & collectors, 19 aug. 1985 (ar); 1 ♀, khabarovsk territory, verkhnebureinskiy district, 20th km of chegdomyn-urgal road, mosses and plant debris on swampy sides of wayside lake with carex spp., poaceae gen. sp., ledum palustre, chamaedaphne calyculata, vaccinium uliginosum, spiraea sp., sphagnum spp., undergrowth of betula platyphylla and larix gmelinii, 6 oct. 2006, a.b. ryvkin leg. (ar); 1 ♂, khabarovsk territory, verkhnebureinskiy district, bureinskiy nature reserve (buffer zone) near ‘strelka’ cordon, 550 m a.s.l., mosses, leaf litter and drift on banks of old channel of umal’ta-makit river: sphagnum spp., polytrichum sp., carex spp., poaceae gen. spp., etc., 30 sep. 2006, a.b. ryvkin leg. (ar); 1 ♀, khabarovsk territory, verkhnebureinskiy district, bureinskiy nature reserve, ‘strelka’ cordon, mari (bog), 28 aug. 2006, a. barševskis & u. valainis leg. (dul); 3 ♂♂, 4 ♀♀, khabarovsk territory, verkhnebureinskiy district, bureinskiy nature reserve, pravaya bureya river basin near medvezhye winter hut, 1050 m a.s.l., mosses and litter in slope sparse larch forest with sphagnum spp., rubus chamaemorus, ledum palustre, ptilium crista-castrensis, pleurozium schreberi, polytrichum spp., carex ? globularis, etc., 29 jul. 2007, a.b. ryvkin leg. (ar); 1 ♂, same locality, 1100–1240 m a.s.l., mosses, lichens and litter near rill in upper part of slope and on bald mountain crest: picea ajanensis, pinus pumila, rhododendron aureum, duschekia sp., larix gmelinii, sphagnum spp., ptilium crista-castrensis, pleurozium schreberi, hylocomium splendens, rubus chamaemorus, carex ? globularis, etc., 29 jul. 2007, a.b. ryvkin leg. (ar); 1 ♀, khabarovsk territory, verkhnebureinskiy district, left side of olga river valley 3 km up-stream of sofiysk, 880 m, mosses, leaf litter & plant debris on gentle slope near rill amid mari with hummocks of sphagnum spp., carex ? globularis, c. spp., poaceae gen. spp., ledum sp., vaccinium uliginosum, v. vitis-idaea, polytrichum sp., hypnum sp., hylocomium splendens, pleurozium schreberi, ptilium crista-castrensis, betula spp., etc., 19 aug. 2008, a.b. ryvkin leg. (ar); 1 ♂, khabarovsk territory, verkhnebureinskiy district, ust’urgal nature park (project), near semicha river mouth, se side of old channel, 280 m a.s.l., moss and plant debris in larch-ledum sparse forest with betula divaricata, vaccinium uliginosum, chamaedaphne calyculata, sphagnum spp., polytrichum spp., dicranum spp., cladonia spp., eriophorum sp., carex sp., etc., 29 aug. 2009, a.b. ryvkin leg. (ar); 2 ♀♀, khabarovsk territory, verkhnebureinskiy district, european journal of taxonomy 13: 1-62 (2012) 16 dublikanskiy nature refuge, right side of dublikan river valley, 1 km up-stream of cordon, nnw slope of bald mountain, 400 m a.s.l., mosses and litter on stones with pika burrows (permafrost!): pleurozium schreberi, hylocomium splendens, ptilium crista-castrensis, sphagnum spp., ledum sp., vaccinium vitis-idaea, carex sp., poaceae gen. sp., etc. among larix gmelinii, betula platyphylla, alnus sp., duschekia sp., undergrowth of abies nephrolepis and picea ajanensis, 27 aug. 2008, a.b. ryvkin leg. (ar); 2 ♂♂, 1 ♀, same locality & biotope, 30 aug. 2008, a.b. ryvkin leg. (ar); 1 ♂, magadan area, ten’kinskiy district, near sibit-tyhellakh, peat-moss stripe, 28 may–07 jun. 1982, d.i. berman leg. (ar); 2 ♂♂, 1 ♀, magadan area, ten’kinskiy district, near sibit-tyhellakh, jack london lake, sphagnum swamp on alluvial cone, 18 aug. 1984, k.yu. eskov leg. (ar); 1 ♀, magadan area, ten’kinskiy district, near sibit-tyhellakh, ‘aborigen’ field research station, alder shrubs near rill: in sphagnum hillock, 24 aug. 1984, k.yu. eskov leg. (ar); 1 ♀, magadan area, ten’kinskiy district, detrin river basin, vakkhanka rill, northern swamp, soil traps, 30 may–9 jun. 1981, s.p. bukhkalo leg. (ar); 1 ♂, same locality, swamp near landing place, soil traps, 7–17 jun. 1982, s.p. bukhkalo leg. (ar); 1 ♂, same locality, swamp on glacial drift, 6–16 jun. 1983, s.p. bukhkalo leg. (ar); 1 ♀, same locality, peat-moss stripe, 7–17 jun. 1983, s.p. bukhkalo leg. (ar). remarks this species was originally described based on two males (ht: [china:] island in sungari river; pt: [russia: s maritime province:] kedrovaya pad’ [nature reserve]); it was later cited for hokkaido, japan, and ‘amur region [without more precise locality]’ (naomi & puthz 1994). it should be noted that neither herman (2001) nor smetana (2004) mentioned the chinese occurrence of this species in their respective catalogues. both authors seem to have misinterpreted the terra typica as a russian territory. based on the new material above, s. raddei proves to be a very common species distributed widely in the russian far east. stenus (nestus) illotulus puthz, 1972 (fig. 2c, d) stenus (nestus) illotulus puthz, 1972b: 169. stenus illotulus – puthz 1972d: 107. — shavrin & puthz 2007: 120. stenus (nestus) illotulus – ryvkin 1987: 159. material examined russia: 1 ♀, tuva, todjenskiy district, azas nature reserve, near cordon at azas lake, 940 m a.s.l., left bank of channel: sedge-gramineous bog (+ trollius sp. etc.) with tussocks, 15 jun. 1991, a.b. ryvkin leg. (ar); 3 ♂♂, 3 ♀♀, buryatia, okinskiy district, 1900 m, il’chir lake, 21–23 aug 2007, a.v. shavrin leg., <det. v. puthz, 2008> (ar, ash); 2 ♂♂, amur area, zeyskiy nature reserve, ‘34th km’ cordon, gulik river valley, in litter, 23 jun. 1978, v.v. belov & s.a. kurbatov leg. (ar); 1 ♂, amur area, selemdzhinskiy district, byssa river 10 km below ‘tyoplyi klyuch’ spa, 290 m a.s.l., leaf litter among tussocks with spiraea sp., calamagrostis sp., carex spp., etc. in high flood-plain, 3 jun. 2007, e.m. veselova & a.b. ryvkin leg. (ar); 1 ♂, 1 ♀, amur area, selemdzhinskiy district, byssa river basin, bottom part of mountainside nw of ‘tyoplyi klyuch’ spa, 370 m a.s.l., litter under alnus sp. with spiraea sp., undergrowth of betula platyphylla, trientalis europaea, poaceae gen. spp., rubus arcticus, etc., 22 jun. 2007, e.m. veselova & a.b. ryvkin leg. (ar); 1 ♂, 1 ♀, amur area, selemdzhinskiy district, norskiy nature reserve, selemdzha river basin, 2 km ne of dvadtsatikha cordon, open swamp near lakeside: plant debris and sparse mosses among tussocks with carex spp., poaceae gen. sp., salix spp., etc., 9 aug. 2004, a.b. ryvkin leg. (ar); 1 ♂, amur area, mazanovskiy district, nora river mouth, 210 m a.s.l., mosses and leaf litter on natural levee and in flood-plain forest with alnus ryvkin a.b., stenus species of the canaliculatus group 17 sp., salix spp., padus sp., ferns, poaceae gen. spp., carex spp., smilacina davurica, etc., 6 aug. 2006, e.m. veselova & a.b. ryvkin leg. (ar); 2 ♀♀, amur area, selemdzhinskiy district, norskiy nature reserve, nora river basin near maltsevskiy cordon, mosses and plant debris on swampy flood-plain lakeside: carex spp., calamagrostis sp., filipendula palmata, geranium sp., comarum palustre, spiraea sp., salix sp., sphagnum ? girgensohnii, sph. squarrosum, polytrichum commune, p. sp., climacium sp., iris sp., etc., 28 aug. 2004, a.b. ryvkin leg. (ar); 5 ♀♀, amur area, mazanovskiy district, nora river basin, sorokavyorstnaya channel, upper reaches of zolotoy rill, 270 m a.s.l., mosses and plant debris along edge of slope swamp near mountain crest: eriophorum sp., chamaedaphne calyculata, vaccinium uliginosum, ledum palustre, rhododendron sp., betula ? fruticosa, b. platyphylla, alnus sp., oxycoccus sp., etc., 28 jul. 2006, e.m. veselova & a.b. ryvkin leg. (ar); 1 ♂, amur area, mazanovskiy district, nora river basin, sorokavyorstnaya channel, foot of s slope of sosnovaya mountain, 215 m a.s.l., flood-plain birch forest with betula platyphylla, alnus sp., padus sp., tussocks of carex spp. and poaceae gen. spp., pteridium aquilinum, convallaria keiskei, thalictrum sp., equisetum sylvaticum, etc., 29 jul. 2006, e.m. veselova & a.b. ryvkin leg. (lh); 3 ♂♂, amur area, selemdzhinskiy district, norskiy nature reserve, nora river, 0.5 km up-stream of gryashchinskaya mt., mosses and leaf litter under betula platyphylla and larix gmelinii with calamagrostis sp., equisetum sylvaticum, e. pratense, maianthemum bifolium, polygonatum sp., etc. in burnt forest along river bank, 27 aug. 2004, a.b. ryvkin leg. (shin); 2 ♂♂, 1 ♀, amur area, selemdzhinskiy district, norskiy nature reserve, nora river basin, 1.5 km up-stream of gryashchinskaya mt., mosses and plant debris on open swamp along lake side: tussocks of carex spp. and poaceae with sphagnum ? girgensohnii, sph. ? angustifolium, sph. ? magellanicum, sph. squarrosum, sph. spp., chamaedaphne calyculata, vaccinium uliginosum, sparse ledum palustre, salix sp., betula ? fruticosa, etc., 24 aug. 2004, a.b. ryvkin leg. (ar); 1 ♂, amur area, selemdzhinskiy district, norskiy nature reserve, nora river basin near meunskiy cordon, mosses and plant debris among carex spp. & poaceae with comarum palustre, sphagnum squarrosum, sph. spp., aulacomnium sp. and other true mosses near side of flood-plain lake and swampy road, 13 aug. 2004, a.b. ryvkin leg. (ar); 1 ♂, amur area, selemdzhinskiy district, norskiy nature reserve, nora river basin near meunskiy cordon, mosses and plant debris at flood-plain of a rill: alnus sp., carex spp., poaceae gen. sp., trientalis europaea, convallaria keiskei, sphagnum squarrosum, sph. spp., hypnum sp., etc., 15 aug. 2004, a.b. ryvkin leg. (ar); 1 ♀, amur area, selemdzhinskiy district, norskiy nature reserve (buffer zone), burunda river basin, 1.5 km nw of burunda cordon, plant debris among tussocks of carex spp. under sparse alnus sp. with young growth of larix gmelinii along rill bank near road, 17 sep. 2004, a.b. ryvkin leg. (ar); 2 ♀♀, same locality and biotope, 18 sep. 2004, a.b.ryvkin leg. (ar); 1 ♂, amur area, selemdzhinskiy district, norskiy nature reserve (buffer zone), burunda river basin, 0.5–1 km nw of burunda cordon, plant debris and small true mosses among carex spp. & poaceae (+sweeping) along rill bank, 12 sep. 2004, a.b. ryvkin leg. (ar); 1 ♂, same locality, mosses and plant debris among sedge tussocks with spiraea sp., ? aulacomnium sp., sphagnum squarrosum, sph. ? girgensohnii, small true mosses, etc., 5 oct. 2004, a.b. ryvkin leg. (ar); 1 ♂, amur area, selemdzhinskiy district, norskiy nature reserve (buffer zone), burunda river basin, 1.5–2 km se of burunda cordon, mosses and leaf litter under betula platyphylla with ledum palustre, chamaedaphne calyculata, salix sp., sphagnum spp., hypnum sp., poaceae gen. sp., etc., 6 oct. 2004, a.b. ryvkin leg. (ar); 1 ♀, amur area, selemdzhinskiy district, norskiy reserve, near fevral’sk, 268th km of belogorsk–fevral’sk road, tikhiy rill, 275 m a.s.l., mosses and plant debris between sedge & gramineous tussocks among alnus sp., salix sp., spiraea sp. with sphagnum squarrosum, sph. spp., etc., 8 oct. 2008, a.b. ryvkin leg. (ar); 1 ♀, khabarovsk territory, verkhnebureinskiy district, lower reaches of verkhniy mel’gin river near 1st rapid, 300–350 m a.s.l., moss and litter on steep rocky se slope with betula platyphylla, abies nephrolepis, larix gmelinii, acer ukurunduense, rhododendron dauricum, poaceae gen. spp., carex sp., vaccinium vitis-idaea, linnaea borealis, maianthemum bifolium, trientalis europaea, ledum sp., alnus sp., ferns, equisetum pratense, hylocomium splendens, sphagnum girgensohnii, dicranum sp., ptilium crista-castrensis, polytrichum commune, etc., 20 aug. 2009, a.b. ryvkin leg. (ar); 1 ♂, 1 ♀, khabarovsk territory, verkhnebureinskiy district, ust’-urgal european journal of taxonomy 13: 1-62 (2012) 18 nature park (project), near semicha river mouth, se side of old channel, 280 m a.s.l., leaf litter and moss in birch forest with larix gmelinii, ledum sp., vaccinium vitis-idaea, poaceae gen. spp., carex sp. by edge of larch-ledum sparse forest, 29 aug. 2009, a.b. ryvkin leg. (ar). remarks this very nice and distinctive species was described based only on the male holotype, from southern cis-baikalia (‘südsibirien: tunkun-sajan’). in the same year, puthz (1972c), based on two specimens deposited in the helsinki museum, reported s. illotulus for the n lena basin: ‘shigansk [=zhigansk] und <…> ust-aldan’. the latter specimen is a male, and the sketch of the aedeagus that i recently received from dr. puthz proves that it is unlikely that it belongs to s. illotulus. i think it necessary to accumulate additional material from the n lena basin to give an adequate interpretation of these specimens. shavrin & puthz (2007) provided records for the irkutsk area (i have seen the specimen) and se buryatia; the mention of the krasnoyarsk territory and n mongolia in distributional remarks by the named authors was not accompanied by a reference to any material. the remark concerning the krasnoyarsk territory was a misprint (a. shavrin pers. comm.). the remark for n mongolia is based on 2 ♂♂, 6 ♀♀, mongolia, terelji, 8–12 jul. 1996, t. ito leg. (gmr, ti, vp) (v. puthz pers. comm.). the new data cited above from the khabarovsk territory, amur area, and tuva republic widen the known range of s. illotulus as regards both its sw and its e limits. stenus illotulus, as well as s. raddei, inhabits moss and leaf litter in habitats moist to moderately wet, but is as a rule less abundant than the latter in those communities. stenus (nestus) caseyi puthz, 1972 (fig. 5f) stenus (nestus) caseyi puthz, 1972b: 172. stenus caseyi – puthz 1975b: 125. stenus (s. str.) caseyi – campbell & davies 1991: 110. material examined holotype ♂, ‘[usa] michigan’, ‘♂ | holotype’<printed red label with hand-written male symbol>, ‘stenus | caseyi | n. sp. | det. v. puthz 1971’<standard puthz’s determinative label> (mnhg). usa: 1 ♂, 4 ♀♀, mich. berrien co., 2 mi. n. new buffalo, gallien r. w bank, 2 nov. 1980, litter in marsh, l.e. watrous leg. (fmnh); 4 ♂♂, 1 ♀, wisconsin, kenosha co., carol beach, chiwaukee prairie, 24 oct. 1980, fmhd#80-35 litter in wet grassy pockets, l. watrous & w. suter leg. (fmnh); 2 ♂♂, 1 ♀, same locality, 1 may 1980, fmhd#80-3162, rhizomes, grasses at ditch, w.s. suter leg. (fmnh); 2 ♀♀, same locality, 19 mar. 1974, w. suter leg., fm(hd)#74-49, ber.: litter in semi-dry ditch, ws#74-22b (fmnh); 1 ♀, same locality, 4 jun. 1974, w. suter leg., fm(hd)#74-163, ber.: marsh floor at grass hummocks, ws#74-44a (fmnh); 2 ♂♂, 1 ♀, same locality, 10 may 1975, w. suter leg., fm(hd)#75-189, ber.: grassy moss clumps marsh area (fmnh); 1 ♂, same locality, 26 apr. 1974, w. suter leg., fm(hd)#74-66, ber.: floor at cattails, ws#74-30d (fmnh); 1 ♂, same locality, 26 apr. 1974, w. suter leg., fm(hd)#74-65, ber.: sandy soil at water table, ws#74-30c (fmnh); 1 ♀, same locality, 11 sep. 1976, w. suter leg., fm(hd)#76-160, ws#76-65b, temporary pond, floor peat (fmnh). remarks the species was originally described from michigan, usa, has subsequently been reported from quebec, canada (puthz 1975b) and also cited for the canadian provinces ontario, quebec, new brunswick and nova scotia (campbell & davies 1991) without citing material. the new data above widen the range to ryvkin a.b., stenus species of the canaliculatus group 19 include wisconsin. since the genital preparation of the holotype had been somewhat deformed owing to excessively quick dehydration, the figure of the aedeagus in the original description was not quite adequate. based on the available material, i herein provide a new figure for this structure (see fig. 5f). stenus (nestus) brivioi puthz, 1972 stenus (nestus) brivioi puthz, 1972c: 12. stenus brivioi – puthz 1972b: 171 (note). — puthz 1975a: 124. stenus (s. str.) brivioi – campbell & davies: 110. stenus vinnulus – casey 1884: 112 (pars). material examined canada: 1 ♂, alberta, 5 km w. ashmont rte. 28a, camas-lily marsh, 12 jun. 1977, g.e. ball leg. (uasm); 1 ♂, yukon territory, proctor’s saw mill, on stewart river, 63°33’30”n 137°25’w, collected by treading grass tussocks 2’–8’ from shore of pond, 3 jul. 1977, r.e. morlan, j.v. matthews, r.e. roughley leg. (yukon refugium project) (uasm). remarks terra typica: usa: michigan, maryglade college, e. of memphis, macomb co; paratype: eagle har., l.[ake] sup.[erior]. puthz (1972a) attributed paralectotypes of s. vinnulus casey, 1884 (see below) from white fish point to s. brivioi as well, and also reported the latter for quebec, canada (1975). campbell & davies (1991), without citing material, reported s. brivioi for alaska and most provinces of canada (excluding british columbia and prince edward island). stenus (nestus) dolosus casey, 1884 stenus dolosus casey, 1884: 108. stenus dolosus – puthz 1975a: 124. stenus (nestus) dolosus – puthz 1972b: 171. stenus (s. str.) dolosus – bernhauer & schubert 1911: 157. — campbell & davies 1991: 111. stenus jejunus casey, 1884: 112. stenus villosus casey, 1884: 109. stenus villosus – casey 1892: 711. stenus (nestus) villosus – l. benick 1938: 266. stenus (s. str.) villosus – bernhauer & schubert 1911: 168. material examined canada: 1 ♀, yukon territory, 24 mi w whitehorse, alaska hwy., 2000’, 60°51’n 136°36’w, 21 jun. 1977, r.e. morlan, j.v. matthews, r.e. roughley leg. (yukon refugium project) (uasm). usa: 1 ♀, ‘wisconsin. kenosha co.; chiwaukee prairie, carol beach. 26:iv:1974. leg: w. suter.’, ‘fm(hd)#74-65. ber.: sandy soil at water table. ws#74-30c’ <without top abdominal segments> (fmnh); 1 ♀, ithaca, cayuga lake, 17 aug. 1928, dr. rambousek leg., l. hromadka det., 1979 (ar); 1 ♂, ottawa co., ohio, south bath is., squaw harbor, put-in-bay, washed-up eel-grass on sand beach, 2 aug. 1979, s.w. nichols leg. (fmnh); 1 ♂, michigan, clinton co., rose l. exp. sta. ex mud edge, european journal of taxonomy 13: 1-62 (2012) 20 typha marsh, 3 sep. 1965, thomas hlavac leg. (fmnh); 1 ♂, ‘volga s.d. edge of pond. 5 may 1966. v.m. kirk’, ‘n.m. downie colln. 1992 acc. z-18,343 field museum’ (fmnh); 1 ♂, ‘rossie, n.y. c-15 1965. n.m. downie leg.’, ‘n.m. downie colln. 1992 acc. z-18,343 field museum’ (fmnh); 1 ♀, ‘tompkins co, n.y. f-15 1967. n.m. downie leg.’, ‘n.m. downie colln. 1992 acc. z-18,343 field museum’ (fmnh). remarks terra typica: usa: ‘fort garland, colorado’; for s. villosus: ‘california’; for s. jejunus: ‘british columbia’. casey (1892) himself synonymized s. jejunus with s. villosus. revising all three holotypes, puthz (1972a) found them to be conspecific and placed s. villosus in synonymy with s. dolosus. campbell & davies (1991) reported this species for alaska and most provinces of canada: yukon territory, northwest territories, alberta, saskatchewan, manitoba, ontario, quebec and new brunswick, without citing material. stenus (nestus) milleporus casey, 1884 sp. propria stenus milleporus casey, 1884: 111. stenus milleporus – casey 1892: 711. stenus illustris l. benick, 1926: 263. stenus (nestus) illustris – l. benick 1938: 266. stenus odius blackwelder, 1943: 213. stenus sectilifer casey, 1884: 110. stenus sectilifer – herman 2001: 2384. stenus (nestus) sectilifer – puthz 1972b: 171. stenus (s. str.) sectilifer – bernhauer & schubert 1911: 167. stenus teter notman, 1920: 699. material examined usa: 2 ♀♀, florida, marion co., 3 mi n blichton, 26 mar. 1976, c.w.o’brien & g.b.marshall (fmnh). remarks originally described from the usa: ‘new york, 3; st. catharine island, georgia, 3; florida, 7’ <the digits refer to the number of specimens in the original type series>; the holotype of s. sectilifer: ‘district of columbia’. stenus teter was described from florida. terra typica of s. illustris: mexico; of s. odius: cuba, soledad near cienfuegos. acting as first reviser, casey (1892) himself synonymized s. sectilifer with s. milleporus. however, bernhauer & schubert (1911) mistakenly regarded s. milleporus as a junior synonym of s. sectilifer, and this treatment was repeated by puthz (1972a) when placing s. teter, s. illustris, and s. odius in the synonymy of s. sectilifer. under articles 24.2.1 and 24.2.2 of iczn, the valid name for the species under consideration is stenus milleporus casey, 1884. ryvkin a.b., stenus species of the canaliculatus group 21 stenus (nestus) canaliculatus gyllenhal, 1827 (fig. 3d, e) stenus canaliculatus gyllenhal, 1827: 501. stenus canaliculatus – fauvel 1869: 492. — j. sahlberg 1871: 415. — hochhuth 1872: 154. — fauvel 1873: 255. —j. sahlberg 1876: 55, 323. — eppelsheim 1880: 94. — j. sahlberg 1880: 77. — heyden 1881: 78. — casey 1884: 115. — fauvel 1889: 119. — hamilton 1889: 114. — eppelsheim 1893: 56. — hamilton 1894a: 20. — hamilton 1894b: 371. — poppius 1899: 38. — j. sahlberg 1900: 28. — fauvel 1902: 75. — jakovlev 1902: 112. — l. benick 1921a: 148. — l. benick 1924: 254. — lebedev 1925: 134. — iljin 1926: 225. — szujecki 1958: 118. — puthz 1970b: 174, 178. — korge 1971: 6. — puthz 1971c: 139. — puthz 1973a: 27. — uhlig & vogel 1981: 98. — tichomirova 1982: 213. — dubeshko 1984: 52. — silfverberg 1988: 20 <‘holotype’ of congener maeklin, 1853>. — pisanenko & puthz 1991: 170, 174. — puthz & zanetti 1995: 17. — shavrin 2007: 141. — shavrin & puthz 2007: 114 (pars). stenus (nestus) canaliculatus – rey 1884: 273. — jakobson 1909: 481. — poppius 1909a: 15. — reitter 1909: 158. — renkonen 1935: 29. — palm 1961: 88. — szujecki 1961: 34. — puthz 1967b: 78. — puthz 1971a: 82. — puthz 1972a: 259. — puthz 1972c: 12.— tichomirova 1973: 173. — shilov 1975: 58. — ryvkin 1987: 159. — ryvkin 1990: 168. — kuznetsova 1991: 171. — dauphin 1993: 188. — ryabukhin 1999: 46. — bordoni 2004: 120. — semenov 2004: 11. stenus (s. str.) canaliculatus – heyden 1898: 34. — bernhauer & schubert 1911: 156. — scheerpeltz 1933: 1150. — campbell & davies 1991: 110. — puthz 2001: 36. stenus affinis stephens, 1833: 298. <?>stenus congener maeklin, 1853: 192. <?>stenus congener – casey 1884: 114. — hamilton 1889: 114. — l. benick 1921b: 114. <?>stenus (s. str.) congener – scheerpeltz 1933: 1152. stenus cordaticollis leinberg, 1900: 187. stenus cordaticollis – j. sahlberg 1900: 28. stenus (nestus) cordaticollis – jakobson 1909: 481. stenus (nestus) foveifrons rey, 1884: 272. material examined france: 1 ♀, ‘caen (ga) raffray’ <white rectangle>, ‘fvl[?fauvel] caen 67.’ <green circle>, ‘fvl[?fauvel] stenus canaliculatus gyll.’ <white circle>, ‘c.[ollection] of a. jakovlev’ <printed label> (zin). poland: 1 ♂, ‘silesia. 4106[=stenus canaliculatus gyll.] – 3’ (zin); 1 ♀, ‘danzig’, <underside> ‘kraatz.’, ‘st. canaliculatus gyll.’, ‘c.[ollection] of a. jakovlev’ <printed label> (zin). hungary: 3 ♂♂, 1 ♀, nagycovaczy, 22.v.1982, v.g. shilenkov leg. (ar); 1 ♀, budapest, 10 apr. 1982, v.g. shilenkov leg. (ash). ukraine: 1 ♀, podol’sk government, kamenets-podol’sk, 12 apr. 1911, v. & i. yakubovskiy leg. (zin); 1 ♂, ‘odessa, krivaya balka. 4.iii.1920. d. znojko leg.’, ‘stenus (nestus) morio grav.?’ (zin); 1 ♀, odessa, fontanka, slope, lakeside, 4 jul. 1970, s.ya. blinshtein leg. (ar); 1 ♀, nikolayev area, rybakovka, sandy seashore, 28 jul. 1970, s.ya. blinshtein leg. (ar). european journal of taxonomy 13: 1-62 (2012) 22 russia: 1 ♂, kaliningrad area, rybachiy, 6 jun. 1959, o.l. kryzhanovskiy leg. (zin); 1 ♀, same locality and collector, 28 jun. 1959 (zin); 1 ♂, petropol.[is]. (zin); 1 ♀, same locality, a. moravitz collection (zin); 1 ♂, st. petersburg, 30 mar. 1897, bianki leg., l. benick det. (zin); 1 ♂, 1 ♀, [st. petersburg] smolenskoye cemetery, 18 aug. 1890 (zin); 1 ♀, same locality, 5 apr. 1891 (zin); 1 ♀, same locality, 9 may 1896, g.g. jakobson leg. (zin); 2 ♂♂, 1 ♀, petrograd [=st. petersburg], smolenskoye cemetery, 11 apr. 1920, a.a. stackelberg leg., l. benick det. (zin); 2 ♀♀, [st. petersburg] ekaterinhof, sifting, leaves, 2 apr. 1921 (zin); 1 ♀, ‘small islands on neva river, shlisselburg uyezd [=district], 1 jun. 1906. g.g. jakobson leg.’, ‘canaliculatus’ (zin); 1 ♀, pargolovo, finlandskaya railway, 8 aug. 1905, somina leg., l. benick det. (zin); 1 ♂, st. petersburg government, tsarskosel’skiy uyezd [=district], orlinskoye lake, 26 apr. 1905, d. glazunov leg. (zin); 1 ♀, [st. petersburg government], luzhskiy district, ploskoye, 7 apr. 1906, yu.i. bekman leg. (zin); 1 ♂, [? st. petersburg government], lesnoy, 12 oct. 1897 (zin); 1 ♂, st. petersburg government, luzhskiy uyezd [=district], preobrazhenskaya, 2 may 1905, d. glazunov leg. (zin); 1 ♀, ‘[st.-petersburg government,] yamburg. [skiy] u.[yezd] [= yamburgskiy district]. 12.1912.–01.1913. a.p. rimskiy-korsakov leg.’ (zin); 1 ♀, smolensk, bank of dnepr river, on moist sand, 10 may 1993, m.yu. gildenkov leg. (sstc); 1 ♀, moscow government, klinskiy uyezd [=district], boblovo, 16 apr. 1906, d. smirnov leg. (zin); 1 ♀, vladimir area, petushki, 1 may 1983, v.b. semenov leg. (ar); 1 ♀, same locality and collector, in litter, 21 apr. 1984 (ar); 1 ♀, kostroma area, manturovskiy district, near shilovo, andromeda polifolia – sphagnum magellanicum, jun.1981, a.l. tichomirova leg. (ar); 1 ♀, ‘urzhum, drift of urzhumka river, 21.iii.1901, [l. krulikovskiy leg.]’, ‘canaliculatus d. bernh.[auer]’, ‘c.[ollection] of a. jakovlev’ <printed label> (zin); 1 ♀, ‘urzhum, drift of urzhumka river. 23.iii.1901. [l. krulikovskiy leg.]’, ‘canaliculatus [?bernhauer det.]’, ‘c.[ollection] of a. jakovlev’ <printed label> (zin); 1 ♀, ‘urzhum, drift of urzhumka river. 24–25.iii.1901. l. krulikovskiy leg.’, ‘c.[ollection] of a. jakovlev’ <printed label> (zin); 1 ♂, 2 ♀♀, ‘[urzhum,] drift of urzhumka river. 09.iv.1906. l. krulikovskiy leg.’, ‘canaliculatus det. shakhonin’, ‘c.[ollection] of a. jakovlev’ <printed label> (zin); 2 ♀♀, ‘malmyzh, drift of shoshma river. 03–04.iv.1899. l. krulikovskiy leg.’, ‘canaliculatus’, ‘c.[ollection] of a. jakovlev’ <printed label> (zin); 2 ♂♂, 3 ♀♀, ‘a. jakowlew| jaroslaw[l’ government]’, ‘c.[ollection] of a. jakovlev’ , ‘stenus canaliculatus gyll. d. fauvel’, ‘c.[ollection] of a. jakovlev’ <printed label> (zin); 1 ♂, 1 ♀, ‘yaroslav[l’]’, <underside> ‘eppelsh[eim]’, ‘c.[ollection] of a. jakovlev’ <printed label> (zin); 1 ♂, ‘[yaroslavl’ government,] manikha, sifting from floating litter. 15.iv.1898.’, ‘yaroslavskiy u.[yezd] [=district]. c.[ollection] of a. jakowlew’, ‘stenus carbonarius gyll. [det. ?a.jakowlew]’ (zin); 1 ♀, yaroslavl’ area, nekouzskiy district, borok, shore of reservoir, 6 jun. 1979, e.m. veselova leg. (ar); 1 ♀, ‘ryaz[an]’<white rectangle with thin black frame and species name> ‘stenus aterrimus er.’, ‘c.[ollection] of a. jakowlew’ (zin); 1 ♂, 1 ♀, ‘riasan.’ <printed label on red paper>, ‘stenus canaliculatus gyll.’ <black indian ink in dubble box> (zin: ex coll. semenov-tian-shanskiy); 1 ♀, gremyachka, dankovskiy uyezd [=district], ryazan’ government, 13 aug. 1899, a. semenov leg. (zin: ex coll. semenov-tian-shanskiy); 1 ♀, same locality and collector, 19 sep. 1901 (zin); 1 ♀, tataria, stolbishchevskiy district, collective farm ‘13 let oktyabrya’, perennial grass, clover: in trap buckets, 1 jun. 1956, utrobina leg. (ar); 1 ♀, voronezh area, khopyorskiy nature reserve, near water, 5 jun.? 1976. (ar); 1 ♂, samara area, kinel’skiy district, krasnosamarskoye forestry, flood-plain of samara river, 2 jul. 1995, i.n.goreslavets leg., ‘stenus sp-6 det. goreslavetz i.n.’ (ig); 1 ♀, saratov area, khvalynsk, sterkh river, 30 jul. 1974, v.i. lomakin leg. (ar); 1 ♀, [rostov area] env. novocherkassk, razliv, 17 apr. 1913, v. kiseritzky leg. (zin); 1 ♀, rostov area, nizshne-kundryuchenskoye hunting farm, 2 jun. 1990, v. grebennikov leg. (zmru); 1 ♀, ‘sarepta. [18]95. becker.’, ‘canaliculatus d. bernh.[auer]’, ‘c.[ollection] of a. jakovlev’ <printed label> (zin); 3 ♂♂, astrakhan’ area, near dosang, 16 may 1997, k.a. grebennikov leg., ‘stenus (nestus) ? canaliculatus gyll. k.a. grebennikov det.1997’ (kg); 1 ♀, kalmyk assr, lysyi liman, 26 jun. 1977, arzanov leg. (zmru); 1 ♂, kalmykiya, arshanya zelmen’, 3 jun. 1980, n.s. kalyuzhnaya leg. (kg); 1 ♀, nw caucasus [krasnodar territory], malyi tkhach mt., 2000 m a.s.l., subalpine belt, clayey bank of puddle, 4 jun. 1994, a.yu. solodovnikov leg. (ayus); 1 ♀, murmansk area, poyakonda, ryvkin a.b., stenus species of the canaliculatus group 23 biological field research station of moscow university, ejected fucus, 14 jun. 1979, e.m. veselova leg. (ar); 1 ♀, same locality and collector, alopecurus ventricosus, 13–17 jul. 1979 (ar); 1 ♀, near solovetskiy monastery, bol’shoy solovetskiy island, 1893, g.g. jakobson leg. (zin); 1 ♂, komi assr, gibyu river, near ukhta, fluvial terrace above flood-plain, in sod, 26 jun. 1974, v.f. shilov leg. (zin); 1 ♂, komi assr, near ukhta, in drift after flood, 23 jun. 1974, v.f. shilov leg. (zin); 1 ♂, same locality and collector, 5 may 1974 (zin); 1 ♂, sob’ river basin, bol’shoy ural, obdorsk [=salekhard], 8 jul. 1925, fridolin leg. (zin); 1 ♀, tyumen area, uvatskiy district, 10–11 km s of gornoslinkino, steep right bank of irtysh river up-stream of tobolsk field research station of severtsov institute, small mosses, plant debris, denuded loam among tussilago farfara, carex sp., poaceae gen. sp., fabaceae etc. near small rill draining by slope, 11 jun. 2004, a.b. ryvkin leg. (ar); 1 ♀, tyumen area, uvatskiy district, 11 km s of gornoslinkino, clayish right bank of irtysh river (narrow fold at rill mouth) upstream of tobolsk field research station of severtsov institute, in leaf litter and sweeping under salix sp. with small true mosses, marchantia sp., equisetum sp., carex sp., calamagrostis sp. etc., 11 jun. 2004, a.b. ryvkin leg. (ar); 1 ♂, 2 ♀♀, tomsk government, ob’ river between barnaul and chesnokovka (postal station), 13 jun. 1897, silant’yev leg. (zin); 1 ♂, 1 ♀, tuva, east shore of chagytay lake, river bank, 18 jul. 1980, b.a. korotyayev leg. (ar); 1 ♂, krasnoyarsk territory, beryozovskiy district, ‘stolby’ nature reserve, mana river near berly cordon, shingles at river bank, 22 jun. 1990, a.b. ryvkin leg. (ar); 1 ♀, evenkia, baykitskiy district, central siberian biosphere reserve, stolbovaya river basin: dulkuma river (iv), 12 km below topkaya river mouth, 165 m a.s.l., river banks: leaf litter among poaceae gen. sp. and salix sp., shingles, sand, silt, drift, 1 jul. 1990, a.b.ryvkin leg. (ar); 1 ♂, evenkia, baykitskiy district, central siberian state reserve, stolbovaya river basin: lower reaches of birapchana river near mouth of kruten’kiy stream, 110 m a.s.l., river bank: shingles, silt, sand, clay, poaceae gen. sp., mosses, carex spp., etc., 21 jul. 1991, a.b. ryvkin leg. (ar); 2 ♂♂, krasnoyarsk territory, turukhanskiy district, nizhnyaya sarchikha river near kamenka river mouth, 150 m a.s.l., under mosses on river bank, 6–7 jul. 1992, v.b. semenov leg. (ar); 1 ♀, same locality and collector, river bank: sand, 10 jul. 1992 (ar); 1 ♂, krasnoyarsk territory, turukhanskiy district, yeloguy river, 7–10 km below tyna river mouth, 70 m a.s.l., sandy river bank with small shingles, 19 jul. 1989, a.b. ryvkin leg. (ar); 1 ♂, 1 ♀, same locality and collector, river bank: carex spp., poaceae gen. sp., mosses, shingles, 24 jul. 1989 (ar); 1 ♂, same locality, sphagnum bog with eriophorum sp., ledum palustre, oxycoccus sp., rubus chamaemorus, vaccinium uliginosum and sparse pinus sylvestris, 1 jul. 1989, r.a. rakitov leg. (ar); 1 ♂, 3 ♀♀, krasnoyarsk territory, turukhanskiy district, yeloguy refuge, yeloguy river 6 km below tyna river mouth, river bank: under drift, 22–23 jul. 1992, v.b. semenov leg. (ar); 1 ♂, krasnoyarsk territory, turukhanskiy district, yeloguy refuge, tyna river 12 km up-stream of river mouth, under mosses on stony bank of river, 12 aug. 1992, v.b. semenov leg. (ar); 1 ♂, putorana highland, ayan lake near ayan river source, in stony stream channel, 9 jul. 1983, k.yu. eskov leg. (ar); 1 ♂, taymyr, maymecha river mouth, 5 jul. 1971, a.p. rasnitsyn, a.g. ponomarenko, i.d. sukatcheva, v.v. zherikhin leg. (ar); 1 ♂, taymyr, khatangskiy district, kotuy river, 6 km up-stream of kresty, salix sp., 13 jul. 1976, a.p. rasnitsyn & i.d. sukatcheva leg. (ar); 1 ♂, 5 ♀♀, ‘irkutsk. v.ye. yakovlev leg.’, ‘canaliculatus gyll. l. benick det.’ (zin); 1 ♀, ‘irkutsk. v.ye. yakovlev leg.’, ‘canal. v. cordaticollis leinbg. l. benick det.’ <immature specimen> (zin); 1 ♂, ‘irkutsk. v.ye. yakovlev leg.’, ‘coll. semenov-tian-shansky’ (zin); 1 ♂, buryatia, selenga river, sedge community, 3 jul. 1982, dubeshko leg. (ar); 1 ♂, buryatia, middle reaches of vitim river, baysa, 3 aug. 1969, v.v. zherikhin, ye. shnitnikova leg. (ar); 1 ♂, [yakutia], ‘5 verst off yakutsk, suburban settlement. 18.vi.1925. bianki leg.’, ‘yakutian research expedition of acad. of sci.’ (zin); 1 ♀, ‘yakutsk. 21.ix.1927. moskvin leg.’, ‘168’, ‘canaliculatus gyll. l. benick det.’ (zin); 1 ♂, ‘2nd tyhylyminskiy nasleg, yakutsk region. 02.vii.1925. bianki leg.’, ‘yakutian research expedition of acad. of sci.’, ‘canaliculatus gyll. l. benick det.’ (zin); 1 ♀, central yakutia, nizhniy bestyakh (ash); 1 ♂, w yakutia, 11 km sw of toybokhoy, uchugey syhylakh, sedge-gramineous hygrophytous meadow, 25 aug. 1997, stepanov leg., ‘stenus carbonarius gyll.’ (ash); 1 ♀, khabarovsk territory, okhotsk, airport, wall of timber house, 4 jul. 1985, v. zherikhin, a. rasnitsyn, d. shcherbakov leg. european journal of taxonomy 13: 1-62 (2012) 24 fig. 3. – a–c: stenus (nestus) alopex sp. nov.: ♂. a. aedeagus, ventral view (holotype). b. apical part of aedeagus, ventral view (holotype). c. abdominal sternite 9 (paratype). – d–e: s. (n.) canaliculatus gyllenhal, 1827: ♂. d. aedeagus, ventral view (russia: rostov area: nedvigovka). e. abdominal sternite 9 (russia: taimyr: maymecha river). – scales = 0.1 mm. ryvkin a.b., stenus species of the canaliculatus group 25 (ar); 1 ♀, maritime prov., khasan, under boards on littoral, 3 jun. 1967, a.l. tichomirova leg. <?> (ar). mongolia: 1 ♂, seleng.[a] aimak, shamar, 18 jul. 1984, s.l. kuz’min leg. (ar). usa: 1 ♀, ‘wisconsin: walworth co.; troy center. (sw) girl scout camp area. w. suter leg. 27:vi:1976.’, ‘fm(hd)#76-92. ws#76-55. mosses ex boggy field.’ (fmnh); 1 ♂, ‘usa: ill., kendall co., silver springs st. pk. 26-iv-1981.’, ‘fmhd#81-164, ex rotting wood chips, l. watrous leg.’ (fmnh); 1 ♀, ‘usa: mich. berrien co., 2 mi. n. new buffalo, gallien r. w bank 2-xi-80.’, ‘litter in marsh. l.e. watrous leg.’ (fmnh); 1 ♀, ‘19’ <indian ink on white rectangle>, ‘kenai’ <indian ink on green stripe>, ‘stenus| congener| mäkl.| am. b. occ.’ <indian ink on green rectangle> (zmmu: motschulsky collection). remarks the species is variable in both external characters and, to a lesser extent, in the shape of the aedeagus. terra typica: ‘suecia’ [= sweden]. widespread in the holarctic. in an earlier article devoted to the caucasian stenine fauna, i summarized the range as follows: ‘palearctic and alaska <…>, michigan <…>, but missing evidently in subtropics and in the southernmost districts of the pontic biogeographical province (turkestanian and kurdian). unknown to me also from the crimean, derbentian, caucasian, lenkoranian districts. the records nearest to the caucasus are in the rostov area (nedvigovka, azhinov, manychskaya, yerokhin, podtyolkovo, liventsovka). one cannot deny a possibility of finding it in ciscaucasia’ (ryvkin 1990; the biogeographical units follow those of razumovskiy 1980). in this excerpt i missed the important record from ne turkey: ‘benliahmed südwestl. kars’ (korge 1971); though the locality does not belong to the caucasus in a strict sense, it lies right at the border between the caucasian and kurdian biogeographical districts. this species is missing in the lists for the nw caucasus (solodovnikov 1998) and the samara area (goreslavets et al. 2002), but my restudy of the collections of the named authors makes it possible to report s. canaliculatus for both the regions (see above: the material section). both records of s. canaliculatus in shavrin (1998) (for “alar', irkutsk area”, and “bol'shoye alginskoye lake, buryatia”) are to be referred to s. incrassatus erichson, 1839 (i have seen the material in shavrin’s collection). among the material listed under the name ‘canaliculatus’ by shavrin & puthz (2007), at least the specimen from ‘barun-torej, myrgen’, chita area should be referred to the new species s. canalis sp. nov. described below; some other transbaikalian specimens unknown to me may also prove to belong to this new species. in that article, all the records marked as relating to the material deposited in ‘coll. a.shavrin’ and ‘irkutsk state university’, except ‘sarma’, ‘shelekhov’, and ‘selenga delta’ (the latter locality, based on my identifications, has been published by dubeshko (1984)), should be verified because the respective specimens were not present in the complete collection that dr. shavrin has sent me for restudy. ryabukhin (1999), based on my identifications in the ibpm collection, listed three localities from kamchatka. campbell & davies (1991) reported this species for alaska and all the provinces of canada, excluding saskatchewan, labrador, and prince edward island, without citing material. the american specimens i know from the northernmost localities are very similar to those from ne asia; but the material from the usa main territory as well as from s canada demonstrates definite differences both in body proportions (e.g. the head somewhat more wide) and in the shape of the male genitalia; i believe the variability mentioned above requires further analysis based on more representative sampling. european journal of taxonomy 13: 1-62 (2012) 26 s. congener maeklin, 1853, according to the original description, was described for a single specimen captured on kadjak island (‘in insula kadjak a d.holmberg semel captus’). no specimens from the kenai peninsula were mentioned in maeklin’s text; these two localities were distinguished well by both mäklin and mannerheim (a good map was provided in mannerheim 1853, which incorporated maeklin’s descriptions); therefore, assumptions by silfverberg (1988) and puthz (2001) concerning the type status of kenai specimens in different museums seem to be as premature as unfounded (‘semel captus’<!> means that the type series cannot be distributed widely over all the main european museums). casey (1884), l. benick (1921b), and scheerpeltz (1933) treated s. congener as a species propria; nevertheless, fauvel (1869, 1873, 1889), rey (1884), bernhauer & shubert (1911), and others regarded this name as a synonym of s. canaliculatus. i think it important that fauvel (1889), when citing the synonymy, mentioned maeklin’s type as known to him and cited the terra typica as ‘kadjak’. consequently, the synonymy of congener and canaliculatus should be regarded as hypothetical before a reliable type specimen can be found. stenus (nestus) alopex sp. nov. (fig. 3a–c) material examined russia: holotype: ♂, ‘putorana highland, ayan lake near | source of ayan river. | lakeshore osier | tundroid. 22.08.1983. | k.yu. eskov. #74.’ <printed label in russian>, ‘holotypus’ <my standard printed red label>, ‘stenus ht | alopex sp. n. | a.b. ryvkin det., 1994’ <my standard determinative label> (zmmu). – paratypes: 1 ♂, ‘putorana highland, ayan lake near | source of ayan river. | lakeshore tundroid. | 23.08.1983. | k.yu. eskov. #75.’ <printed label in russian> (ar); 1 ♀, ‘putorana highland, ayan lake. | alpine belt. | 28.07.1983. | k.yu. eskov. #54.’ <printed label in russian> (ar); 2 ♂♂, ‘taymyr, khatangskiy district, | taymyrskiy nature reserve, | ary-mas: right bank of novaya river | near field research station. [mosses and litter under dwarf salix sp. and betula ?exilis] near small rill. | 17.07.1992. a. ryvkin. #93.’ <printed label in russian> (irsn, mtd); 1 ♂, 1 ♀, ‘taymyr, khatangskiy district, | taymyrskiy nature reserve, | ary-mas: right side of novaya river | near ulakhan-yuryakh river mouth. bank | of [novaya] river[: shingles, sand]. 19.07.1992. a. ryvkin. #98.’ <printed label in russian> (ar); 2 ♀♀, ‘taymyr, khatangskiy district, | taymyrskiy nature reserve, | ary-mas: right side of novaya river | near ulakhan-yuryakh river mouth. | swampy floodplain of [small] rill[: mosses and litter among carex spp., salix spp., betula ?exilis etc.]. | 19.07.1992, a. ryvkin. #99.’ <printed label in russian> (ar); 1 ♀, ‘taymyr, khatangskiy district, | taymyrskiy nature reserve, | ary-mas: right bank of novaya river | near field research station. shore of ‘small’ | lake [ese of ulakhan-yuryakh river mouth. mosses, litter and sweeping at lake shore: carex spp., comarum palustre]. 20.07.1992. a. ryvkin. #104.’ <printed label in russian> (ar); 2 ♂♂, ‘taymyr, khatangskiy district, | taymyrskiy nature reserve, | ary-mas: right bank | of novaya river, near lake between | small rill and ulakhan| yuryakh river.’, ‘mosses and litter under larix [gmelinii] with | cassiope tetragona, pyrola | sp., vaccinium vitis-idaea etc. | on s slope. 23.07.1992. | a.b. ryvkin. #115.’ <printed labels in russian> <one specimen without head and pronotum> (ar); 1 ♂, 1 ♀, ‘taymyr, khatangskiy district, | taymyr nature reserve, | ary-mas: right bank | of novaya river near mouth of ulakhan| yuryakh river (watershed of novaya river and | small rill).’, ‘mosses and litter on glade | in sparse larch forest | with betula ?exilis, salix spp., | vaccinium vitis-idaea, v. | uliginosum, pyrola sp., | cassiope tetragona, cladonia | spp., carex spp., etc. | 24.07.1992. a.b. ryvkin. #118.’ <printed labels in russian> (ar). — the paratypes listed above are also provided with both “paratypus” printed red label and my standard determinative label (‘stenus | alopex sp. nov. | a.b.ryvkin det., 1994’) for each lot. — 1 ♀, chukot peninsula, 78-84, ayopechan island, ust’-chaun, 19 jul. 1978, e.g. matis leg. <?> (ar). ryvkin a.b., stenus species of the canaliculatus group 27 differential diagnosis this species resembles s. (n.) canaliculatus gyllenhal, 1827 and can be distinguished from it by the narrower and shorter elytra, by the much elongated pronotum, by the character of puncturation, and by the shape of the aedeagus. external differences from all the known species of the canaliculatus-group are given in the key below. etymology the specific name is derived from the latin generic name of the arctic fox alopex lagopus (linnaeus, 1758). description length. 3.2–3.9 mm (the last value for the specimen with abdomen extended). coloration. black, moderately shining, with moderately long and dense greyishor yellowish-silvery pubes cence. legs pitchy black, tibial apices and tarsi somewhat lighter. antennae pitchy brown, with segments 1–2 black and club vaguely infuscate; 1st segment of maxillar palpi yellow, 2nd brown with yellowish base, 3rd dark brown, with base a bit lighter. head. about 1/5 to 1/6 broader than pronotum (47:39), exactly as broad as elytra between humeri (47:47), distinctly narrower than those in broadest part (47:54). front feebly evenly convex, with very vague and shallow lateral impressions distinguishable only between antennal tubercles and eyes. puncturation moderately dense and regular, nonconfluent, as a rule, except of periocular areas, without smooth spots and strips; average diameter of punctures about as large as basal cross–section of antennal segment 3. frontal slope before antennal fossae rather flat. anterior margin of labrum with a small median notch. antennae moderately long, reaching the middle 1/3 of pronotum. length proportions of anten nal segments 2–11 = 6:5:4.5:5:4:4:3:3.5:4.5:4; segments of club a little longer than broad (3.5:3, 4.5:4, 4:3.5). pronotum. evidently longer than broad (45:39), broadest near middle, narrowed feebly convexly towards anterior margin and feebly concavely towards posterior one. longitudinal median furrow not deep but evident over almost entire length of disk. laterobasal depressions very shallow, nearly vanishing. puncturation dense and fairly deep, sometimes nonrugosely confluent near midline; punctures somewhat larger than those of head. elytra. a bit broader than long (54:52), distinctly longer than pronotum (52:45 in holotype), at suture nearly as long as the latter (46:45 in holotype). humeral angles short but evident, lateral sides uniformly moderately divergent posteriorly (47:54). humeral and sutural depressions nearly absent. puncturation deep and fairly regular, evidently larger in diameter than that of pronotal disk, sometimes confluent, but not forming long furrows. legs. fairly short; segment 1 of metatarsi about as long as segment 5. abdomen. moderately convex, with well developed paratergites and two pairs of keels at bases of anterior visible tergites. lateral sides nearly parallel, feebly convergent in apical part. posterior margin of tergite 7 with fine light fringe. puncturation of tergites much smaller than that of head and pronotum, on anterior visible tergites denser laterally, medioposteriorly with some interstices comparable to diameter of punctures. microsculpture. the whole dorsal side with fine and dense ground sculpture visible throughout. european journal of taxonomy 13: 1-62 (2012) 28 male. mesoand metatibiae without specific characters; posterior margin of 8th abdominal sternite with a fairly broad but not deep angularly-rounded emargination, without deep impressions and keels, slightly depressed in posterior half; abdominal sternite 9 as in fig. 3c; aedeagus as in fig. 3a, b. female. posterior margin of 8th abdominal sternite angularly rounded. remarks this new species seems to be a late derivative of s. (n.) canaliculatus gyllenhal, 1827; the latter also inhabits both the taymyr peninsula and the putorana highland (see above). additional material including male specimens is necessary to prove the distribution of the new species up to the chukot peninsula. stenus (nestus) canosus sp. nov. (figs. 4a–b, 5d) material examined china: holotype: ♂, ‘nw china, xinjiang uygur | zizhiqu, tien | shan, southern slope of narat mt ridge: | koktau mts, right confluent | of koksu river, 3200 m, stone-| sandy bank of rill with sedge tussocks and | small willows. 25.07.[20]01. [s.v.]saluk. no11’ <printed label in russian>, ‘holotypus’ <my standard printed red label>, ‘stenus ht | canosus sp. n. | a.b.ryvkin det., 2010’ <my standard determinative label> (zmmu). differential diagnosis this new species is externally very similar to s. (n.) canaliculatus gyllenhal, 1827 and differs from it by the character of the puncturation, which is much coarser, more sparse and irregular, by the less developed microsculpture, by the median furrow of pronotum, which is deeper and broader, and by the shape of the aedeagus. external differences from all the known species of the canaliculatus-group are given in the key below. etymology the name of this species is the latin adjective “canosus” (grey-haired). description length. 4.0 mm (with abdomen extended). coloration. black, distinctly shining, with moderately long and dense silvery pubes cence. legs dark brown with knees somewhat darkened, brownish-black. antennae dark brown, with segments 1–2 pitchy-black and club vaguely infuscate; 1st segment of maxillar palpi yellow, 2nd brown with yellow base, 3rd dark brown with very base yellowish. head. distinctly broader than pronotum (50:44), somewhat narrower than elytra between humeri (50:53), much narrower than those in their broadest part (47:54). front feebly evenly convex, with very vague and shallow lateral impressions distinguishable mainly between antennal tubercles and eyes. puncturation moderately dense, somewhat irregular, evidently smaller and denser between antennal tubercles, partly nonrugosely confluent both there and at periocular areas, without distinct smooth spots and strips, but obviously sparser and greater in the middle; average diameter of punctures in the middle distinctly larger than the greatest cross-section of antennal segment 3. frontal slope before antennal fossae rather flat. anterior margin of labrum with small median notch. antennae moderately long, scarcely reaching middle of pronotum. length proportions of anten nal segments 2–11 = 6:5:6:4:4:4:3:4:4.5:6; segments of club longer than broad (4:3, 4.5:4, 6:4). ryvkin a.b., stenus species of the canaliculatus group 29 fig. 4. – a–b: stenus (nestus) canosus sp. nov.: ♂ (holotype). a. aedeagus, ventral view. b. apical part of aedeagus, ventral view. – c–d: s. (n.) canalis sp. nov.: ♂ (holotype). c. aedeagus, ventral view. d. apical part of aedeagus, ventral view. [note that the difference in shape of the apical lobe compared to that in c is due to the level of magnification.] – scales = 0.1 mm. european journal of taxonomy 13: 1-62 (2012) 30 pronotum. a bit longer than broad (45:44), broadest near middle of length, narrowed feebly convex ly towards anterior margin and concavely towards posterior one. longitudinal median furrow deep and sharp over entire length except for very anterior piece. laterobasal depressions very shallow, nearly vanishing. puncturation deep and irregular, partly nonrugosely confluent; punctures distinctly larger than those of head. elytra. a bit broader than long (60:58), much longer than pronotum (58:45), at suture distinctly longer than the latter (48:45). humeri anglarly rounded, lateral sides uniformly moderately divergent posteriorly (53:60). humeral and sutural depressions feeble but evident in anterior half of disk. puncturation deep, much more regular, evidently larger in diameter than that of head and pronotum, sometimes confluent, but not forming long furrows. legs. fairly short; metatibia about 1/3 longer than metatarsus (40:29); segment 1 of metatarsi about as long as segment 5 (8:8). abdomen. moderately convex, with well developed paratergites and two pairs of keels at bases of anterior visible tergites. lateral sides feebly uniformly convergent. posterior margin of tergite 7 with fine light fringe. puncturation of tergites evidently smaller than that of head and pronotum; on anterior visible tergites much denser and smaller laterally, medioposteriorly with some smooth interstices distinctly wider than average diameter of punctures. microsculpture. very fine mesh-like ground sculpture visible between punctures, mainly on elytra and abdominal tergites 8–9, as well as on lateral parts of head, pronotum and anterior abdominal tergites, median parts of abdominal tergites quite smooth or slightly irregularly netted only on edges of punctures. male. mesoand metatibiae without specific characters; 8th abdominal sternite without conspicuous impressions and keels, its posterior margin with very broad and not deep angularly-rounded emargination; abdominal sternite 9 as in fig. 5d; aedeagus as in fig. 4a, b. female. unknown. remarks no records of s. (n.) canaliculatus were previously known from china or from adjacent countries of central asia. the capture of s. (n.) canosus sp. nov., which is closely related to s. (n.) canaliculatus, proves that their common ancestor was recently widely distributed throughout these spacious territories. stenus (nestus) canalis sp. nov. (figs. 4c–d, 5e) stenus canaliculatus – shavrin & puthz 2007: 114 (pars). material examined russia: holotype: ♂, ‘magadan area, ten’kinskiy | district, detrin river basin, | vakkhanka rill. plateau: open | plots, soil traps. | 01–11.06.1983. s.p. bukhkalo [leg.]’ <printed label in russian>, ‘holotypus’ <my standard printed red label>, ‘stenus ht | canalis sp. n. | a.b. ryvkin det., 2010’ <my standard determinative label> (zmmu). paratypes: 1 ♂, ‘magadan area, ten’kinskiy | district, near sibit-tyhellakh. | dry birch forest. 14–24.07.1982. | d.i. berman [leg.]’ <printed label in russian> (ar); 1 ♂, ‘khabarovsk territory, jewish | autonomous area, obluchenskiy | district, near pashkovo. puddle | bank under steep. | 05.07.1977. a.b. ryvkin [leg.]’ <printed label in russian>, ‘stenus | ryvkin a.b., stenus species of the canaliculatus group 31 canaliculatus gyll. | a.b. ryvkin det. 1978’ (ar); 1 ♂, ‘maritime province, khasan, | golubinyi utyos. | 4.viii. | 1991 s.a. kurbatov [leg.]’ <photo label in russian> (ar); 1 ♂, ‘maritime prov. | flood-plain of poyma river | 30.07.–06.08.[19]99 | shavrin a. [leg.]’ <printed label in russian>, ‘stenus gyll. | canaliculatus | det. v. puthz 2005’ (ar); 1 ♂, ‘chita area | ne side of barun-torey lake | myrgen cape | 18–19.06.1994 | v.[g.] shilenkov leg.’ <printed label in russian>, ‘stenus gyll. | canaliculatus | det. v. puthz 1999’ (ar); 1 ♂, ‘5.08.[19]86 <transverse> | buryatya | zaigraevo[=zaigrayevo] 1695 [=5695th km] st[ation] | voincov[=voinkov] a. [leg.]’ <printed label>, ‘3-23 | canal.’, ‘stenus | canaliculatus | shavrin a. det. 2006’ (ash). — the paratypes listed above are also provided with both ‘paratypus’ printed red label and my standard determinative label (‘stenus | canalis sp. n. | a.b. ryvkin det., 2010’) for each lot. — 1 ♀, magadan area, upper reaches of kolyma river, 7 km nne of omsukchan, 10 jul. 1980, o. mashukova leg. <?> (ar); 1 ♀, magadan area, upper reaches of kolyma river, 5 km ne of spornoye, 21 jun. 1981, a. ryabukhin leg. <?> (ar). differential diagnosis this species is closely related to s. (n.) canaliculatus gyllenhal, 1827 and differs from it by the smaller body size, by the much more irregular puncturation, by the less developed ground sculpture, by the posterior margin of 8th abdominal sternite, which is much more shallowly emarginated, and by the shape of the aedeagus. external differences from all the known species of the canaliculatus-group are given in the key below. etymology the specific name is the latin adjective ‘canalis’ (canine). description length. 2.8–3.7 mm (the latter value for the paratype specimen with abdomen extended). coloration. pitchy black, distinctly bronze-coloured shining, with moderately long and dense yellowish-silvery pubescence. legs pitchy black to brownish-black, tibial apices and tarsi somewhat lighter. antennae dark brown, with 1st segment black and 2nd brownish-black; 1st segment of maxillar palpi yellow, 2nd brown, 3rd dark brown, with very base a bit lighter. head. about 1/8 to 1/6 broader than pronotum (46:41, all measurements, except where specifically indicated, are from the holotype), a little narrower than elytra between humeri (46:48), distinctly narrower than elytra in broadest part (46:55). front feebly evenly convex to nearly flat, with very vague and shallow lateral depressions, distinguishable only between antennal tubercles and eyes and behind the tubercles. puncturation dense and irregular, partly nonrugosely confluent, sometimes with small but evident smooth spots and strips situated both medially and laterally; average diameter of punctures in middle about as large as the greatest cross-section of antennal segment 3. frontal slope before antennal fossae rather flat. anterior margin of labrum with a small median notch. antennae rather short, scarcely reaching middle 1/3 of pronotum. length proportions of anten nal segments 2–11 = 6:5:5:4:3:3:2:3:4:6; segments 9–10 slightly transverse (3:3.5, 4:4.5), the segment 11 distinctly elongate (6:4). pronotum. somewhat longer than broad (45:41), broadest just before middle, narrowed feebly convex ly towards anterior margin and concavely towards posterior one. longitudinal median furrow sharp and deep, well-developed over entire length of disk as a rule, but less evident in some paratypes in anterior 1/3 as well as near both anterior and posterior margins. laterobasal depressions very shallow, nearly vanishing. puncturation rather deep and irregular, partly nonrugosely confluent but not forming long furrows; punctures somewhat larger on average than those of head; some bulging smooth interstices between punctures evidently smaller in diameter than surrounding punctures. european journal of taxonomy 13: 1-62 (2012) 32 elytra. about as broad as long (55:55) to a bit broader, much longer than pronotum (55:45), by suture a bit longer than pronotum (46:45). humeral angles short, angularly rounded, lateral sides uniformly moderately divergent posteriorly (48:55). humeral and sutural depressions very feeble to nearly absent. puncturation larger in diameter and sparser, more regular than that of pronotal disk, as a rule nonconfluent, but individual furrows of merged punctures visible in some paratypes; interstices between punctures distinctly smaller than half average diameter of puncture. legs. fairly short; metatibia much longer than metatarsus (38:27 in paratype from magadan area); segment 1 of metatarsi about as long as segment 5 (7:7 in the same paratype). abdomen. moderately convex, with well developed paratergites and two pairs of keels at bases of anterior visible tergites. lateral sides of abdomen nearly parallel, feebly convergent in apical part. posterior margin of tergite 7 with fine light fringe. puncturation of anterior visible tergites comparable in average diameter of punctures to that of head but not so coarse and much more regular; denser laterally, medioposteriorly with some interstices comparable to diameter of punctures or larger. microsculpture. variable in different specimens. very fine mesh-like ground sculpture visible, as a rule, between punctures or by edges of punctures on elytra in humeral impressions and near suture, as well as on periocular parts of head; abdominal tergites quite smooth and glossy at least in their median parts. male. mesoand metatibiae without specific characters; 8th abdominal sternite without conspicuous impressions and keels, its posterior margin with a very broad and shallow rounded emargination; abdominal sternite 9 as in fig. 5e; aedeagus as in fig. 4c, d. female. no reliable female specimens included in type series. remarks both the females listed were captured at localities from which no male material is known. therefore, they are not included in the type series and their attribution to this new species remains questionable. this new species seems to replace s. (n.) canaliculatus gyllenhal, 1827 in the southernmost and, in part, easternmost territories of e siberia and the far east. additional material is necessary to ascertain the precise distribution of both species in the e palaearctic. stenus (nestus) delitor sp. nov. (figs. 5c, 6a, 7a) material examined russia: holotype: ♂, ‘evenkia, [baykitskiy district,] basin of podka| mennaya tunguska river: ko| chumdek river 8–10 km up-stream of mouth |. 70 m a.s.l. [rill bank, among stones (mosses and litter). (he+hm+ho)]. 22–24.vii.1990. a.b. ryvkin.’ <photo label in russian>, ‘262’, ‘holotypus’ <my standard printed red label>, ‘stenus ht | delitor sp. n. | a.b. ryvkin det., 2008’ <my standard determinative label> (zmmu). – paratypes: 1♂, together with holotype, ‘262’ (ar); 2 ♂♂, 1 ♀, ‘evenkia, [baykitskiy district, ] central sibe| rian [biosphere] reserve, basin of | stolbovaya river: [upper reaches of ]raskol rill (confluent of dulkuma river), s | slope of dulkuma | residual mountain. 250–330 [280] m a.s.l. [bank of rill (left confluent of raskol rill): litter under alnus sp., abies sibirica, picea obovata and mosses near water. ]16.vii. | 1991. a.b. ryvkin’ <photo label in russian>, ‘132’ (ar, irsn); 1 ♂, ‘tuva, [todjenskiy district, ]azas nature reserve, basin | of azas river: upper reaches of kara | -tesh river [(environs of the 3rd hut)]. 1150 m a.s.l. [carex spp., mosses and poaceae with sparse salix spp. etc. on river bank and near small flood-plain lake.] 06.vi. | 1990. a.b. ryvkin’ ryvkin a.b., stenus species of the canaliculatus group 33 fig. 5. ♂ copulative structures. a–e: abdominal sternite 9. a. stenus (nestus) latipennis j. sahlberg, 1880 (russia: krasnoyarsk territory: mirnoye). b. s. (n.) illusor ryvkin, 1987 (russia: magadan area: 2 km w of splavnaya). c. s. (n.) delitor sp. nov. (holotype). d. s. (n.) canosus sp. nov. (holotype). e. s. (n.) canalis sp. nov. (holotype). f. s. (n.) caseyi puthz, 1972 (u.s.a.: michigan: berrien co.). aedeagus, ventral view. – scales = 0.1 mm. european journal of taxonomy 13: 1-62 (2012) 34 <photo label in russian>, ‘64’, ‘stenus | illusor ryv. | a.b. ryvkin det., 1993’ <my standard determinative label> (ar); 1 ♂, ‘khamar-daban mt. ridge | pass in upper reaches | of l.[evaya] mishikha river, 1800 m a.s.l. | foot of glacier | 1.vii.[19]75. shilenkov’ <photo label in russian>, ‘stenus | latipennis | j. sahlbg. | det. v.puthz 1979’ <standard determinative label by puthz>, ‘stenus | illusor a. ryv. | a.b. ryvkin det. 1984’ <my standard determinative label> <specimen with endophallus dissected by puthz> (ar). — the paratypes listed above are also provided with both ‘paratypus’ printed red label and my standard determinative label (‘stenus | delitor sp. n. | a.b. ryvkin det., 2008’) for each lot. differential diagnosis stenus delitor sp. nov. is closely related to both s. (n.) latipennis j. sahlberg, 1880 and s. (n.) illusor ryvkin, 1987; it can be distinguished from these species by its smaller elytra, by the character of the puncturation and ground sculpture, and by the shape of the aedeagus. external differences from all the known species of the canaliculatus-group are given in the key below. etymology the name of this species is the latin noun ‘delitor’ (destroyer). description length. 4.0–4.6 mm (the latter value for specimens with abdomen extended). coloration. black, shining, with moderately long and dense greyish-silvery pubes cence. legs black, tibial apices and tarsi sometimes a bit lighter. anterior margin of labrum nearly sraight, without median notch. antennae brown to dark brown, with segments 1–2 black to brownish-black; 1st segment of maxillar palpi yellow, 2nd brown with yellowish base, 3rd dark brown, with base a bit lighter. head. much broader than pronotum (56:46), as broad as to a bit broader than elytra between humeri (56:56 in holotype), somewhat narrower than elytra in broadest part (56:62). front with broad, moderately prominent longitudinal median ridge, obtuse-angled in cross-section, and flanked with two well developed lateral impressions; median elevation more than twice as broad as each of lateral portions in basal part (20:9). puncturation fairly coarse and dense, irregular, partly nonrugosely confluent in twos to threes; basal part of median elevation punctured somewhat sparser but without conspicuous spots or strips impunctate; average diameter of punctures somewhat larger than basal cross-section of antennal segment 3. frontal slope before antennal fossae fairly gentle. antennae moderately long, scarsely reaching middle of pronotum. length proportions of anten nal segments 2–11 = 7:9:7:7:6:6:4:5:5:5; segments of club a little longer than broad (5:3.5, 5:4.5, 5:4). pronotum. about 1/5 to 1/6 longer than broad (54:46 in holotype), broadest near middle, narrowed convex ly towards anterior margin and concavely towards posterior one. deep median longitudinal furrow well developed over almost entire length of disk. laterobasal depressions very shallow, though laterobasal prominences evident. puncturation fairly coarse, irregular, somewhat greater in middle of lengh, partly transversely confluent; punctures much greater than those of head. elytra. as broad as to a bit broader than long (62:61 in holotype), distinctly longer than pronotum (61:54 in holotype), although a bit shorter than pronotum (52:54 in holotype) by suture. humeral angles short but evident, lateral sides uniformly moderately divergent posteriorly (56:62). humeral and sutural depressions very feeble, visible near base only. puncturation deep, more regular and evidently larger in average diameter than that of pronotum, partly confluent, but not forming long furrows. legs. moderately long; metatibia nearly ½ longer than metatarsus (54:37); 1st segment of metatarsi a bit longer than 5th segment (11:10.5). ryvkin a.b., stenus species of the canaliculatus group 35 abdomen. moderately convex, with well developed paratergites and two pairs of keels at bases of anterior visible tergites. lateral sides of abdomen nearly parallel, feebly convergent in apical part. tergite 7 with well developed light fringe at posterior margin. puncturation of tergites fairly dense, on anterior visible tergites irregular, distinctly sparser medioposteriorly, on preapical tergites more scattered medially; average diameter of punctures on anterior visible tergite about equal to that of median elevation of head. microsculpture. very fine and dense mesh-like ground sculpture well developed throughout but entirely absent in some areas both on head and pronotum and, mainly, on varnish shining interstices between punctures of elytra and along midline of anterior visible abdominal tergites; abdominal tergites 7–10 with evident reticulation. this character seems to be fairly variable in different populations. male. mesoand metatibiae without specific characters; abdominal sternites 6–7 slightly depressed along midline and cut medioposteriorly; 8th abdominal sternite with fairly broad but not deep, angularlyrounded emargination of posterior margin and with flat depression in posterior half; abdominal sternite 9 as in fig. 5c; aedeagus as in figs. 6a, 7a. female. abdominal sternite 8 with broad, rounded apical margin. remarks until recently i have interpreted this species as being a western form of s. illusor ryvkin, 1987. the specimen from the khamar-daban was not included in the type series of the latter owing to evident differences in both external appearance and the shape of the aedeagus. study of the material from middle siberia and tuva demonstrates that s. delitor is a valid new species. stenus (nestus) illusor ryvkin, 1987 (figs. 5b, 7b) stenus (nestus) illusor ryvkin, 1987: 155. material examined russia: 1 ♀, amur area, selemdzhinskiy district, selemdzha river basin, angelokit river, 52°53.497’n 132°24.543’e, 430 m a.s.l., mosses and litter under salix spp. and alnus sp. with poaceae gen. sp., sphagnum squarrosum, sph. sp., plagiomnium sp., etc. in flood-plain, 3 jul. 2007, e.m. veselova & a.b. ryvkin leg. (ar). – 1 ♂, amur area, selemdzhinskiy district, selemdzha river basin, angelokit river near mouth, 52°53.49’n 132°24.5’e, 420 m a.s.l., mosses and litter under populus sp., padus sp., alnus sp., abies nephrolepis, picea ajanensis, etc. with spiraea spp., swida alba, rosa sp., rhododendron sp., sparse betula platyphylla, poaceae gen. spp., carex spp., smilacina davurica, pyrola daurica, trientalis europaea, etc. in flood-plain forest, 4 jul. 2007, e.m. veselova & a.b. ryvkin leg. (ar); 1 ♀, amur area, selemdzhinskiy district, left side of selemdzha river, 3 km below butov kamen’ rapid, gorge with waterfall, 52°50.94’n 132°11.44’e, 400 m a.s.l., mosses and litter on stones and rocks near, above and below the waterfall, 8 jul. 2007, e.m. veselova & a.b. ryvkin leg. (ar). – 1 ♂, khabarovsk territory, verkhnebureinskiy district, island on niman river 700 m upstream of niman cordon of bureinskiy nature reserve, 1040 m a.s.l., mosses and leaf litter in floodplain forest with populus sp., salix sp., spiraea spp., sorbaria sorbifolia, poaceae gen. spp., carex spp., sphagnum sp., polytrichum sp., hylocomium splendens, etc., 11 aug. 2008, a.b. ryvkin leg. (ar); 1 ♂, 1 ♀, magadan area, 2 km w of splavnaya, №236, 2 sep. 1981, a. ryabukhin leg. (ar); 1 ♀, magadan area, 35 km n of magadan, №184, 16 aug. 1981, a. ryabukhin leg. (ar); 1 ♀, same locality, №194, 20 aug. 1981, a. ryabukhin leg. (ar); 1 ♀, kamchatka, esso, flood-plain of bystraya river, 27 jun. european journal of taxonomy 13: 1-62 (2012) 36 1975, b.a. korotyayev leg. (ar); 1 ♀, chukot peninsula, anadyr river basin, novyi yeropol, middle flood-plain, osier-bed with poaceae on silty clay sand: soil trap, 15 jul. 1986, e.g. matis leg. (ar). remarks the species was previously known only for the male holotype: ‘khabarovsk territory, amur basin, lyanchli river, confluent of gorin river, 5.ix.1975, o.n. kabakov leg.’ (zin). based on the new material, i regard it as being widely distributed across the russian far east, except the southernmost territories. stenus (nestus) latipennis j. sahlberg, 1880 (figs. 5a, 7c–h) stenus latipennis j. sahlberg, 1880: 78. stenus latipennis – heyden 1881: 78. — puthz 1972d: 107. — puthz 1974: 112. — shavrin & puthz 2007: 123 (pars). stenus latipennis<?> – j. sahlberg 1899: 340. stenus (nestus) latipennis – jakobson 1909: 481. — palm 1961: 98. — puthz 1965: 27 (pars). — puthz 1967a: 49. — tichomirova 1973: 173. — shilov 1975: 58. — ryvkin 1987: 159. stenus (nestus) latipennis<?> – poppius 1909b: 19. stenus (s. str.) latipennis – campbell & davies 1991: 111. material examined russia: 1 ♂, south of the yamal peninsula, priuralskiy district, tarcheda-yakha river near mouth, 23 jul. 1980, e.m. veselova leg. (ar); 1 ♀, 70°04´n 87°36´e, sw taimyr, upper reaches of nizhnyaya agapa river, nyapan ridge, at shore of ladannakh lake, 11–16 jul. 2001, a.b. babenko leg. (ar); 1 ♂, same locality and collector, snowfield, 5–9 jul. 2001 (ar); 7 ♂♂, 7 ♀♀, taymyr, maymecha river mouth, yantardakh mt, 7 jul. 1971, a.p. rasnitsyn, a.g. ponomarenko, i.d. sukatcheva, v.v. zherikhin leg. (ar, mtd); 1 ♂, 2 ♀♀, same locality and collectors, 3 aug. 1971 (ar); 1 ♂, taymyr, khatangskiy district, kotuy river, 6 km up-stream of kresty, salix sp., 11 jul. 1976, a.p. rasnitsyn & i.d. sukatcheva leg. (ar); 1 ♂, taymyr, kotuy river 6 km up-stream of mouth, 18 aug. 1971, a.p. rasnitsyn, a.g. ponomarenko, i.d. sukatcheva, v.v. zherikhin leg. (ar); 1 ♂, taymyr, khatangskiy district, bol’shaya romanikha river mouth, swampy old channel, 18 jul. 1977, i.d. sukatcheva leg. (ar); 1 ♀, taymyr, near khatanga town, flood-plain salix bushes with larix, soil traps, 16–30 aug. 1989, p.k. yeryomin leg. (ar); 1 ♂, putorana highland, ayan lake near ayan river source, mossy coastal osier-bed, 7 jul. 1983, k.yu. eskov leg. (ar); 1 ♂, same locality and collector, in stony stream channel, 9 jul. 1983 (ar); 1 ♂, krasnoyarsk territory, turukhanskiy district, bol’shaya varlamovka river 2–6 km up-stream of mouth, 40 m a.s.l., river banks, 12 jul. 1988, a.b. ryvkin leg. (ar); 1 ♂, 6 ♀♀, krasnoyarsk territory, turukhanskiy district, mirnoye, traps, plate 11, 6–16 jul. 1991, l.b. rybalov leg. (ar, mhng, shin); 1 ♀, same locality and collector, trap 10, 2–12 aug. 1990 (ar); 1 ♂, 1 ♀, same locality and collector, plate 7, 9 traps, 8–18 jun. 1989 (ar); 4 ♂♂, evenkia, central siberian biosphere reserve, stolbovaya river basin: lower reaches of birapchana river near kruten’kiy stream, 110 m a.s.l., river bank: shingles (limestone), 23 jun. 1993, v.b. semenov leg. (ar); 1 ♂, east siberia, magadan area, ten’kinskiy distr., ‘aborigen’ field research station, 500 m, pebble river banks, 28 jul.–5 aug. 1990, l. penev leg. (ar); 1 ♀, magadan area, 15 km sse of atka, 29 jun. 1981, a.s. ryabukhin leg. (ar); 1 ♂, magadan area, near palatka, khasyn, 27 aug. 1978, v.v. zherikhin leg. (ar). ryvkin a.b., stenus species of the canaliculatus group 37 remarks the type series was collected by nordenskjöld’s yenisei expedition at five localities from chyornyi ostrov (‘tschornaya ostrov’ in the original description, present-day chernoostrovsk village) to dudinka (‘habitat ad ripas rivulorum torrentium inter lapillos in territorio frigido et arctico et parte boreali territorii silvosi rarius’). this species was reported for the first time for ne european russia (‘petschora’, without more precise locality) by j. sahlberg (1899), based on the collection made by the botanist o. kihlman; a decade later it was recorded from the kanin peninsula (‘zwei exemplare unter moos am rande von schneefeldern auf dem bergrücken (paë) bei bugranitza am 4.vii. und ein drittes auf gleichartigen stellen bei madoha am 16.vii gefunden’, poppius 1909b); dr. puthz has seen the two females from kanin in zmh, but has not seen the specimens from pechora (pers. comm). jakobson (1909), not being acquainted with the latter kanin findings, summarized the range as ‘arkhangelsk government (pechora); yenisei government’. puthz (1965), based on earlier misidentifications by hellén (hansen et al. 1939; renkonen 1941), believed the species to be represented in n finland, notwithstanding the fact that palm (1961) had revised all fennoscandian records a few years earlier and noticed them to be erroneous (see s. (n.) labilis above and s. (n.) confusus below); nevertheless, finland (without citing new material or references) was recently mentioned in the distribution section for this species by shavrin & puthz (2007); just before the present manuscript was completed, dr. puthz informed me that two unpublished specimens (male and female) from ‘lapponia inariensis’ had been identified by him as latipennis in 1974, but that the material should be revised again (e-mail of 2 dec. 2010). the figure of the aedeagus of the male from dudinka (puthz 1965) is more or less adequate, except the missing longitudinal middle keel at the apical part of the median lobe. all further mentions of this species from the palaearctic are confined to siberia and polar ural: ‘polarer ural, gouv.[ernment] tobolsk’ (puthz 1967a); ‘im museum helsinki je ein stück von irkutsk und ytyk-haja’ (puthz 1972d; i believe the specimen from the former of these two localities may be related to s. delitor sp. nov.; the female specimen is repeatedly cited by shavrin & puthz 2007); records for komi assr (puthz 1974; shilov 1975) do not go beyond the northernmost part of the territory (sob’ river, vorkuta, yeletskiy, ‘polyarnyi ural’ railway station, etc). tichomirova (1973) indicated the range as siberian only. this species has also been reported for the nearctic (puthz 1974: ‘belege aus dem north west territory, dem yukon territory und alaska im museum ottawa und in waschington’; campbell & davies 1991: ‘ak, yk, nt’), but i do not know of any reliable american specimens and cannot state whether the range of s. latipennis is really subcircumpolar or whether the nearctic findings concern a related species. in an earlier paper (ryvkin 1987), i regarded s. latipennis warily as a n-siberian species; after some closely related taxa have been described, it would probably be useful to revise all the material from other territories where the species has been recorded again. stenus (nestus) vinnulus casey, 1884 stenus vinnulus casey, 1884: 112. stenus vinnulus – fall 1926: 61. — puthz 1972d: 107. stenus (nestus) vinnulus – puthz 1972b: 171. — ryvkin 1987: 159. — silfverberg 1988: 20 <holotype of confusoides renkonen, 1935>. — ryabukhin 1999: 46. stenus (s. str.) vinnulus – campbell & davies 1991: 112. stenus (nestus) confusoides renkonen, 1935: 27. stenus confusoides – strand 1954: 66. — puthz 1970a: 39. stenus (nestus) confusoides – renkonen 1936: 179. — palm 1961: 90. — puthz 1965: 27. — puthz 1967a: 49. — tichomirova 1973: 173. european journal of taxonomy 13: 1-62 (2012) 38 material examined russia: 1 ♂, 1 ♀, karelia, white sea, bay of kandalaksha, n shore of chupa inlet, 500 m ne of nizhnyaya polunga: blizhneye lake, 18–19 jul. 2005, p. petrov leg. (ar); 1 ♀, tuva, todjenskiy district, azas nature reserve, environs of azas lake, zelyonoye lake, 980 m a.s.l., mossy swamps with ledum palustre, eriophorum sp., carex spp., rubus chamaemorus, rhododendron sp. etc. near banks of rill (rhadon !) – in moss, 3 jun. 1990, a.b. ryvkin leg. (ar); 1 ♀, evenkia, baykitskiy district, central siberian biosphere reserve, stolbovaya river 8 km up-stream of river mouth, 60 m a.s.l., mosses and litter on open swamp with carex spp., comarum palustre, sparse menyanthes trifoliata, true mosses, sphagnum spp. etc., 20 sep. 1991, a.b. ryvkin leg. (ar); 1 ♀, evenkia, central siberian biosphere reserve, stolbovaya river basin: lower flow of birapchana river near kruten’kiy stream, 110 m a.s.l., backwashing of limestone shingles at river bank, 29 jun. 1993, v.b. semenov leg. (ar); 1 ♂, 1 ♀, putorana highland, nr. ayan lake, kapchug river, riverside ‘tundroid’, bog with mosses and eriophorum sp., 11 jun. 1983, k.yu. eskov leg. (ar); 1 ♂, yakutia, vilyui river basin, kempendyayi river upstream of kempendyay village, 5 aug. 1988, v. blagoderov & v. zherikhin leg. (ar); 1 ♂, magadan area, 3 km n of shirokoye, 7 jul. 1974, b.a. korotyayev leg. (ar+1ex: ibpm); 1 ♀, amur area, selemdzhinskiy district, near fevral’sk, 268th km of belogorsk–fevral’sk road, tikhiy rill, 275 m a.s.l., mosses and plant debris between sedge & gramineous tussocks among alnus sp., salix sp., spiraea sp. with sphagnum squarrosum, sph. spp., etc., 8 oct. 2008, a.b. ryvkin leg. (ar); 2 ♂♂, amur area, near zeya town, 4 jun. 1978, v.v. belov & s.a. kurbatov leg. (ar); 1 ♀, amur area, near zeya town, gulik, 19 oct. 1979, s.a. serbenyuk leg. (ar); 1 ♀, amur area, selemdzhinskiy district, norskiy nature reserve, nora river basin, 2 km up-stream of gryashchinskaya mt., mosses and plant debris on small open swamp on high flood-plain: tussocks of calamagrostis sp. and carex spp. with sphagnum ? girgensohnii, sph. squarrosum, sph. centrale, sph. spp., rubus arcticus, convallaria keiskei, salix sp., etc., 22 aug. 2004, a.b. ryvkin leg. (ar); 1 ♀, amur area, selemdzhinskiy district, norskiy nature reserve, nora river basin near maltsevskiy cordon, e side of maltsevskoye lake, 210 m a.s.l., sweeping on carex spp., poaceae gen. spp. & motley grass, 1 oct. 2008, e.m. veselova & a.b. ryvkin leg. (ar); 1 ♂, khabarovsk territory, jewish autonomous area, obluchenskiy district, se of radde, dichun river, about 2 km off river mouth, near water, 7 aug. 1977, a.b. ryvkin leg. (ar); 2ex, [khabarovsk territory,] ussuri river basin, bikinskiy district, birskoye, 27 jun. 1958, o.n. kabakov leg. (onk); 1 ex, same locality, 1 jul. 1958, o.n. kabakov leg. (onk); 1 ♀, maritime province, spasskiy district, nr. novoselskoye, rice field, t=27°, ph=5.8, 11 aug. 1986, a. shatrovskiy leg. (ar); 1 ♂, maritime province, spasskiy district, yevseyevka, in stream, 17 jul. 1976, e. berlov leg., ‘stenus sp.’, ‘stenus vinnulus cas., det. v. puthz 2006’ (ash). canada: 1 ♀, yukon territory, klokut archeol. site, 6 mi. n of old crow, 67°54’n 136°36’w, ex. shallow margin of small lake, 19 jul. 1977, r.e. morlan, j.v. matthews, r.e. roughley leg. (yukon refugium project) (uasm); 1 ♀, ‘n.w.t. -21 m.e. tuktoyaktuk. 17–21.vii.[19]71. d.m. wood’, ‘stenus illotulus puthz det. v. puthz 1978’, ‘ob abw. vinnulus?’, ‘eigentum cnc!’ <?> (cnc). remarks originally described from the usa. when revising casey’s heterogeneous type series, puthz (1972a) designated the specimen from cambridge, massachusetts as the lectotype and placed s. confusoides renkonen, 1935, that had been known until then from fennoscandia (renkonen 1936; strand 1954; palm 1961; puthz 1965, 1970) and e siberia (puthz 1967a: chita area: ‘dorf udotschnoje am ingodazufluss’), in the synonymy of s. vinnulus. the specimens from isle royale, lake superior and marquette, michigan, were also attributed to the latter species, but the paralectotypes from white fish point were identified as s. brivioi puthz, which was described in the same year (puthz 1972c). the same author cited e siberian ‘poppius-funde von ytyk-haja, ust aldan und olekminsk’ for s. vinnulus in the same year (puthz 1972d). campbell & davies (1991), without providing material, reported the species for alaska and most provinces of canada, excluding british columbia, saskatchewan, prince edward ryvkin a.b., stenus species of the canaliculatus group 39 fig. 6. aedeagus, ventral view. a. stenus (nestus) delitor sp. nov. (holotype). b. s. (n.) sphaerops casey, 1884 (u.s.a.: new york: albany co.). – scales = 0.1 mm. european journal of taxonomy 13: 1-62 (2012) 40 island, and newfoundland (the first records for alaska and the yukon territory had been provided by fall 1926). ryabukhin (1999) mentioned a single specimen of this species from the magadan area, russia, based on my identifications from the ibpm collection (see material examined above). stenus (nestus) sphaerops casey, 1884 (figs. 6b, 7i–j) stenus sphaerops casey, 1884: 68. stenus (s. str.) sphaerops – campbell & davies 1991: 112. material examined usa: 1 ♂, ‘u.s.a.: new york. s. westerlo: bear swamp. 12viii:1974. leg: w. suter. albany co.’, ‘fm(hd)#74-195. berlese: sphagnum. ws#74–61d.’ (fmnh). differential diagnosis based on many external characters, this species might be placed separately within the canaliculatus group. nevertheless, from the shape and structure of the aedeagus, one can suppose it to be most closely related to the latipennis-illusor-delitor complex. s. sphaerops differs from all the known species of this complex by the broader head with eyes much more convex, by the greater and more bulging pronotum, by the shorter elytra, by the much coarser forebody puncturation, by the less developed ground sculpture, and by the shape of aedeagus. redescription length. 4.0–4.1 mm. coloration. the single specimen on hand is somewhat immature; therefore, the genuine coloration is probably darker than described here. body pitchy black, moderately shining, with fairly short silvery pubescence. legs dark brown with middle parts of femora somewhat lighter, reddish; antennae reddishbrown with 1st segment pitchy black and 2nd segment dark brown; palpi brown, with segment 1 and base of segment 2 yellow. head. evidently broader than elytra between humeri (69:62), nearly as broad as elytra in posterior quarter (69:70). front with a pair of fairly broad and deep longitudinal impressions and a broad keelshaped median elevation that is not very prominent but well developed; median elevation 1.8 times as broad as each of lateral pieces. puncturation coarse and dense, irregular, partly rugose along internal margins of eyes, with small smooth spot in middle of median longitudinal ridge; diameter of coarsest punctures evidently larger than basal cross of antennal segment 3. frontal slope before antennal fossae fairly gentle. antennae long, reaching basal 1/4 of pronotum. length proportions of antennal segments 2–11 = 8:11.5:10:8:7:7:5:6:6:7.5; segments of club distinctly elongate (6:3.5, 6:4.5, 7.5:4). pronotum. distinctively large, bulging, uneven, a bit longer than broad (55:52), broadest near middle of length, narrowed convex ly anteriorly and concavely posteriorly. median longitudinal groove deep over almost entire length of disk, vanishing just behind slightly elevated anterior margin. laterobasal depressions shallow though laterobasal prominences developed. puncturation coarse, in part nonrugosely obliquely confluent, distinctly greater in middle of length; punctures much larger than those of head. elytra. somewhat broader than long (70:63), nearly as broad between humeri as long (62:63), about 1/7 longer than pronotum (63:55) although a bit shorter than pronotum (53:55) by suture. humeral angles short but prominent, lateral sides slightly convexly and moderately divergent posteriorly (62:70). ryvkin a.b., stenus species of the canaliculatus group 41 humeral and sutural depressions rather feeble, vanished posteriorly. puncturation deep, distinctively larger in diameter than that of pronotum, more regular, partly nonrugosely confluent but not forming long furrows. legs. rather long; segment 1 of metatarsi obviously longer than segment 5 (15:12) though shorter than segments 2–4 together. abdomen. moderately convex, with well developed paratergites and two pairs of keels at bases of anterior visible tergites. lateral sides uniformly convergent posteriorly. tergite 7 with very fine light fringe at posterior margin. puncturation of tergites fairly dense, on anterior visible tergites distinctly sparser medioposteriorly; average diameter of punctures on anterior visible tergites about equal to that of median elevation of head. microsculpture. very fine but regular mesh-like ground sculpture evident between punctures on abdominal tergites 6–10; other surfaces smooth or with infrequent, extremely vague netting mainly by margins of punctures. male. mesoand metatibiae without specific characters; abdominal sternites 6–7 slightly depressed along midline and slightly emarginated medioposteriorly; abdominal sternite 8 with broad but not deep, rounded emargination of posterior margin and with flat depression in posterior half; lateral pieces uniformly rounded; abdominal sternite 9 as in fig. 7j; aedeagus as in figs. 6b, 7i. remarks originally described for the male holotype only: ‘massachusetts’. campbell & davies (1991), without citing material, recorded it for the canadian provinces ontario, quebec, and nova scotia, as well as the northwest territories. the original description, though fairly long and detailed, did not contain some necessary data comparable to those of present-day publications. inasmuch as neither descriptions nor figures have been provided for s. sphaerops since casey (1884), i thought it useful to give such a description here. stenus (nestus) confusus j. sahlberg, 1876 (fig. 2e) stenus confusus j. sahlberg, 1876: 58. stenus confusus – poppius 1899: 39. — j. sahlberg 1900: 29. — munster 1921: 119. — l. benick 1924: 254. — haberman 1983: 101, 108. — silfverberg 1988: 20. — puthz 1998: 149. — gollkowsky 2001: 194. — shavrin & puthz 2007: 118. stenus (nestus) confusus – jakobson 1909: 481. — poppius 1909a: 17. — renkonen 1935: 29. — hansen et al. 1939: 32. — palm 1961: 90, 98. — ryvkin 1987: 159. — semenov 2004: 12. stenus (s. str. + nestus) confusus – puthz 1973c: 50. <?>stenus aemulus – thomson 1857: 224. stenus (nestus) latipennis – hansen et al. 1939: 32 (pars). stenus (nestus) protensicollis krása, 1941: 166. material examined russia: 1 ♂, ‘carelia bor.[ealis] 4122 [=stenus confusus sahlb.] – 1.’, ‘4122. confusus sahlb.’ (zin); 2 ♀♀, ‘petrograd, smolenskoye cemetery. 11.iv.1920. a.a. stackelberg leg.’, ‘68’ <round>, ‘confusus european journal of taxonomy 13: 1-62 (2012) 42 sahlb. l. benick det.’ (zin); 1 ♀, ‘yaroslavl’. 08.vi. n. filippov.’, ‘stenus confusus sahlb. det. v. puthz, 1976’ (zmmu); 3 ♂♂, 1 ♀, kalinin [=tver’] area, 18 km sw of kalinin, near putilovo, volga river bank, 22 jul. 1982, i.a. ushakov leg. (ar); 1 ♀, ‘[moscow,] on snow in petrovsko-razoumovskiy park, 28.x.1910, v. boldyrev leg.’, ‘stenus nitens steph.’ (zin); 1 ♀, south of yamal peninsula, 110th km by railway n off obskaya station, puddle, 6 aug. 2001, p. petrov leg. (ar); 1 ♀, krasnoyarsk territory, achinsk, 10 jul. 1973, v. zolotikhin leg. (ar); 1 ♀, ‘5 verst off yakutsk, sergelyakh suburban settlement. 24.viii.1926. l. bianki’, ‘yakutian research expedition of acad. of sci.’, ‘47’ <round>, ‘confusus sahlb. l. benick det.’ <?> (zin); 2♀♀, ‘yakutsk. 22.vi.1925. bianki’, ‘yakutian research expedition of acad. of sci.’, ‘63’ <rectangle>, ‘confusus sahlb. l. benick det.’ <?> (zin). remarks terra typica: russian karelia and finland (‘jag har funnit den vid vigsjön [= ‘lac. wig’ (silfverberg, 1988)] (63°50’) och svir i ryska karelen, i kihtelysvaara, eno och nurmis socknar i norra karelen äfvensom vid jyväskylä och helsingfors’). in the original description, sahlberg supposed that thomson’s (1857) record of s. aemulus erichson, 1839 (see s. (n.) nitens above) for sweden should be referred to s. confusus, but this seems to be doubtful since the species description is quite adequate to s. nitens stephens, 1833, for which aemulus is in fact a synonym. poppius (1909a) reported the species from the middle reaches of the lena river: ‘an der mittleren lena an lehmigen ufern, ein exemplar bei jakutsk, 1.vii!, ein anderes auf einer insel nördlich von önkyr-yrjä, 8.vii!.’ (the east siberian latipennis specimens, identified by poppius as confusus and found by puthz (1972c) in zmh, were collected at other localities and cannot fig. 7. ♂ copulative structures. a–i, k: aedeagus, ventral view. j: abdominal sternite 9. a. stenus (nestus) delitor sp. nov. (holotype). b. s. (n.) illusor ryvkin, 1987 (russia: amur area: angelokit river). c–h: s. (n.) latipennis j. sahlberg, 1880. c. (russia: sw taimyr: ladannakh lake). d. (russia: taimyr: kotuy river). e. (russia: putorana: ayan lake). f–g. (russia: evenkia: kruten’kiy stream). h. (russia: magadan area: ‘aborigen’). i–j. s. (n.) sphaerops casey, 1884 (u.s.a.: new york: albany co.). k. s. (n.) melanopus marsham, 1802 (ukraine: odessa area: krinichnoye). – scales = 0.1 mm. ryvkin a.b., stenus species of the canaliculatus group 43 be correlated to the material cited in poppius 1909a). jakobson (1909) summarised the range as follows: ‘sweden <the record is most likely based on thomson’s misidentification of confusus as aemulus: see above>; lapponia, finland, olonets government’. munster (1921) provided some distributional data for northernmost norway: ‘kaafjord i alten, sirma i tanen samt neiden, langfjordbunden, vaggatim og graense-jakobselv i syd-varanger !’; l. benick (1924), a bit later, cited the only female from se siberia (‘tschita’); palm (1961) confirmed n swedish records of this species (norrbotten, jämtland); haberman (1983) provided three mapped localities from estonia; a pair of specimens (1 ♂, 1 ♀) have been published recently from the moscow area (semenov, 2004). the terra typica of s. protensicollis krása, 1941, which was placed in synonymy with confusus (puthz, 1973c, 1998): ‘simbirsk, rossia m.[ed]’. the new material above confirms that s. confusus is rather widespread in the northern palaearctic, but the species has proved to be rare and sporadic in its distribution. it is essential that all the siberian records known to me are represented by females only. i expect the range to have a relict pattern. what is stenus melanopus (marsham, 1802)? preliminary remarks stenus melanopus has been included in the present paper for the following reasons. firstly, some authors consider the species to be closely related to the canaliculatus group (e.g. betz l.c.); i believe such an interpretation to be not perfectly correct. secondly, owing to its external resemblance to several other species, including those of the canaliculatus group, s. melanopus has been recorded repeatedly for territories where it is in fact not represented. in addition, inasmuch as my study of the material from stephens’ collection (bm) revealed heterogeneity in the series that probably contains the type of s. melanopus, a lectotype designation appears to be called for in order to provide stability in name usage. lastly, a preliminary analysis of the literature on the species under consideration forces me to call into question some synonyms which have been established earlier and, at least, to raise a question regarding the thorough revision of these synonyms. a new monotypic species group, the melanopus group, is defined and diagnosed below; its differences from the canaliculatus group, as well as from other groups of the subgenus nestus, are provided; a lectotype of s. melanopus is designated; a brief analysis of the main published records makes it possible to define the species distribution more exactly and to evaluate the reliability of the synonymy provided. definition of the melanopus group diagnosis body size moderate; body length of only known species: 2.8 to 3.7 mm. head rather small in comparison to elytra and pronotum. upper surface between eyes bisulcate, with longitudinal elevations and impressions well developed. labrum with anterior margin straight, neither sinuate nor notched. internal tooth of each mandible placed much more distally than middle, turned evidently dorsally of the main plane. maxillar palpi with basal segment yellow and not shortened. prementum of normal length, not reduced. antennae fairly short, with penultimate club segments globular to slightly transverse, the last segment a bit longer than broad. pronotum with sharp longitudinal median furrow, rather broad and deep in basal half, vague or vanishing before the middle. european journal of taxonomy 13: 1-62 (2012) 44 elytra moderately long, slightly trapezoid with humeri developed. legs moderately long; segment 4 of tarsi without emargination; segment 1 of metatarsi a bit shorter than segment 5, distinctly shorter than segments 2 to 4 together. abdomen moderately convex, with paratergites evident on abdominal segments 3 to 7; four anterior visible tergites with four short but evident longitudinal keels at basal part each; posterior margin of tergite 7 with well developed, fine membranous fringe. puncturation of pronotum and elytra moderate, non-rugose, with interstices fairly flat. ground sculpture entirely absent, upper surface varnish shining. pubescence moderate, contiguous. male. legs without specific features; posterior margin of abdominal sternites 6–7 about straight; abdominal sternite 8 with broad and very shallow, rounded emargination of posterior margin, without evident depression; abdominal sternite 9 with posterolateral teeth very short, not incurved inwards. aedeagus relatively small; median lobe archetypically lanceolate, with apical sclerotized part angularly rounded; endophallus with paired medial bands and unpaired bar distinct in middle, with very small but visible lateral parts, and with h-shaped expulsion clasp producing fairly broad lateral portions; basal tube evidently sclerotized. parameres cylindrical, without apical broadening; apico-internal setiferous surface poorly differentiated, setae rather uniform. female. posterior margin of abdominal sternite 8 obtuse-angularly rounded; each valvifer with very short posterolateral tooth, backwards directed. spermatheca sclerotized. in the structure of the mandibles, the abdominal tergites, the male genitalia and abdominal sternite 9 of both males and females (valvifera), the melanopus group is most closely related to the atratulusfuscipes-cautus-crassus complex. it differs from all the species of this complex by the presence of a long and sharp, longitudinal median furrow on the pronotum, as well as by the posterolateral teeth of abdominal sternite 9 of both males and females, which are very short and not incurved inwards. taxa included s. (n.) melanopus (marsham, 1802). bionomics see below. general distribution see below. stenus (nestus) melanopus (marsham, 1802) (fig. 7k) staphylinus melanopus marsham, 1802: 528. stenus melanopus – stephens 1829: 290. — stephens 1833: 299. — stephens 1839: 411. — hardy 1851: 45. — waterhouse & janson 1855: 138, 151. — rye 1864: 41. — fauvel 1869: 492 (pars). — rye 1870: 84. — fauvel 1873: 256. — j. sahlberg 1876: 59 (pars; original record: false). — fauvel 1878: ryvkin a.b., stenus species of the canaliculatus group 45 102 (pars). — fauvel 1886: 29. — fowler 1888: 336 (pars). — j. sahlberg 1900: 29 (false). — fauvel 1902: 75 (pars). — sainte-claire deville 1906: 82. — l. benick 1921a: 144. — wüsthoff 1934: taf. ii. — normand 1935: 364. — l. benick 1947: 90. — focarile 1964: 63. — puthz 1966: 146. — puthz 1970b: 174, 178. — puthz 1972e: 49. — puthz 1973a: 26. — anderson 1984: 247. — lucht 1987: 96. — puthz & zanetti 1995: 18. — herman 2001: 2278. — monsevičius & pankevičius 2001: 41. — puthz 2008: 150. stenus melanopus<?!> – j. sahlberg 1880: 78. — heyden 1881: 78. — poppius 1899: 39. — iljin 1926: 225. stenus (nestus) melanopus – ganglbauer 1895: 576 (pars). — jakobson 1909: 481 (pars). — reitter 1909: 158 (pars). — johansen 1914: 505. — l. benick 1929: 44 (pars). — tottenham 1954: 62. — smetana 1959: 202. — palm 1961: 88. — szujecki 1961: 33. — horion 1963: 339. — scheerpeltz 1963: 417. — lohse 1964: 116. — puthz 1965: 27. — puthz 1967c: 8. — puthz 1967d: 305. — puthz 1971a: 83, 97. — puthz 1972a: 259. — tichomirova 1973: 173. — bordoni 1974: 13. — dauphin 1993: 187 (pars). — outerelo et al. 1995: 80. — bordoni 2004: 119. stenus (s. str.) melanopus – bernhauer & schubert 1911: 162 (pars). — scheerpeltz 1933: 1160. — campbell & davies 1991: 111. <?>stenus (nestus) cribrellus rey, 1884: 261. <?>stenus nitidus lacordaire, 1835: 450. <?>stenus nitidus – erichson 1840: 703. — kiesenwetter 1845: 224. — lucas 1846: 122. — fairmaire & laboulbène 1856: 576. — kraatz 1857: 756. — thomson 1857: 224. — kraatz 1858: 123. — thomson 1860: 225. — seidlitz 1872–1875(1874): 255. — seidlitz 1887–1891(1889): 365. <?>stenus (nestus) nitidus – rey 1884: 256. <?>stenus sulcicollis stephens, 1833: 295 (nomen dubium). stenus tythus schaufuss, 1882: 621. material examined uk: lectotype (designated herein): ♀, <small square of red paper>, ‘53’ <hand-written with black indian ink on small white oval>, ‘british isles. j. stephens coll. bm 1853–46’ <printed rectangular label>, ‘standing in stephens coll. as stenus melanopus marsham’ <printed rectangular label>, ‘lectotypus’ <my standard printed red label>, ‘lectotypus | stenus | melanopus marsham | a.b. ryvkin des. 2003’ <my standard printed determinative label> (bm). france: 1 ♀, ‘806’, ‘stenus | melanopus | marsh. | gallia’ [ex coll. semenov-tian-shanskiy] (zin); 1 ♀, ‘corsica. 4121 [=stenus melanopus marsh.] – 2.’, ‘4121. melanopus marsh.’ (zin). ukraine: 1 ♂, 3 ♀♀, odessa area, bolgradskiy district, near krinichnoye, fresh-water lake shore, in reed drift, 5 may 2003, a.v. gontarenko leg. (ar). algeria: 1 ♂, ‘186’, ‘? algeria | ? deyrolle.’ (zin). measurements of the lectotype entire body length: 3.1 mm; head width: 48; pronotum length: 45; pronotum width: 45; elytral length: 59; elytral length by the suture: 50; elytral width between humeri: 53; maximum elytral width (in posterior ¼): 64. european journal of taxonomy 13: 1-62 (2012) 46 remarks the lectotype of s. melanopus is designated from among the four specimens under this name in stephens’ collection and originating likely, at least in part, from marsham’s collection. only this female belongs to melanopus in the commonly accepted interpretation. only this specimen has been marked with a small patch of red paper and only for this specimen is the number ‘53’ indicated, which corresponds to the number of the species in stephens’ ‘illustrations of british entomology’; it is believed to have been placed by waterhouse on a small white oval label, without additional daggers. i have transferred it to a new rectangular board; the old, smaller board remains on the same pin. similar problems that have arisen during the revision of the collections of marsham, kirby, and stephens have been discussed many times by many authors (e.g. tottenham 1937; hammond 1972). an excellent historical reference was kindly compiled by mrs. sharon shute (department of entomology, bm), in addition to the material received by me on loan in 2002. below, i give a slightly abbreviated version of the text of this reference with the permission of the author and management of the department [e-mail of february 20 2004]: “stenus melanopus marsham this species was described by marsham 1802. stephens redescribes this species, in illustrations of british entomology […]. stephens acquired specimens from marsham’s collection when it was sold after his death and normally indicates in his text if he had a marsham specimen. the species name in the illustrations is attributed to marsham, however, he does not mention that he had marsham’s specimen. the staphylinidae in the stephens collection normally bear small, white, round labels if they come from the marsham collection, (see hammond 1972[…]). it is thought these were put on by stephens, although they do not occur on the specimens of the species of many other families in his collection although we know from stephens sale list of the marsham collection that he did indeed purchase specimens of many of these species. the larger oval white labels with black ink numbers are ( ? waterhouse) attached to all four specimens of this species in the stephens collection. the numbers on these labels refers to the species number in the illustrations vol. v. there is no indication what the asterisks mean. none of these specimens have a marsham round, white label, whereas other species within this genus do. this is a mystery because waterhouse in his paper with janson, on some of the british stenus species, […], says that the ‘type’ is in the stephens collection. this may have been an assumption on his part and he may not have known what the round, white labels meant. the fact that stephens does not mention having marsham’s specimen at the time of the redescription i think can be taken to assume he did not have it in his collection. although the series label is not in his hand and could be marsham’s ???. […] it may be advisable to select a neotype from the stephens collection as without a label on the specimen it will not be possible to identify which if any of the four specimens came from the marsham collection. please note that as the specimens in the stephens and kirby collections do not have any locality data it is not possible to tell if the specimens are those he had in front of him at the time of his descriptions or later additions. it is known that he exchanged material and often disposed of specimens if he collected ‘better’ ones which is why in a number of cases it has been found that the specimens standing as a particular species in stephens collections do not fit his description. care must be taken when using these specimens to ‘fix’ names, to take account of common usage of names and the common interpretations of such species before changing names as a result of examining stephens specimens, which should in every case be carefully checked against stephen’s description. […]” [e-mail of may 16 2002] unfortunately, marsham's original description cannot help sufficiently with identification of the original type specimen: ‘st. niger obscurus scaber, pedibus concoloribus. long. corp. 1½ lin. habitat -----. descr. simillimus st. immuni, at pedes omnino nigri.’ every third stenus species appears to conform to the diagnosis cited. even the reference to s. immunis, described by marsham in the same monograph, can add little clarity to solving this problem. the latter name, having been synonymized with no evident ryvkin a.b., stenus species of the canaliculatus group 47 proof with either s. (nestus) circularis gravenhorst, 1802 or s. (hemistenus) pallipes gravenhorst, 1806, at different times, requires a separate revision in itself. the redescription by stephens (1833), on the contrary, is much more detailed and corresponds well, in main features, to the melanopus in commonly accepted interpretation. the only inconsistency may be found in the coloration of the maxillar palpi: ‘antennae and palpi black, immaculate’ whereas the basal segment of the palpi is in fact yellow. the following information provides an explanation for this inconsistency: ‘it is said of stephens that he declined to use a microscope regularly believing that what could not be seen with the naked eye was not worth studying’ (darby 2010). i can confirm that the colour of the basal segment of the palpi may be poorly visible in a total specimen even at a high magnification. the redescription given by waterhouse & janson (1855) ‘according to the type specimen in stephens’ coll.’ relates undoubtedly to s. melanopus in commonly accepted interpretation and conforms to the unique female mentioned above. it is necessary to remember that the paper by waterhouse & janson (1855) was published just about three years after the death of stephens, by authors who had been close to him and had known his collection for many years. thus, both the redescriptions obviously indicate that stephens had at least a specimen of the ‘true’ melanopus in front of him at the time of his redescription. i suppose ‘the label problem’ to be confused in this case not so far as the author of the aforementioned historical reference finds. waterhouse & janson (1855) reported that ‘the specimens which mr. stephens used to call his ‘type specimens’ are almost always marked by some kind of ticket attached to the pin holding the insect. the marshamian specimens are thus marked by a round yellow ticket; and when the species is described in the ‘entomologia britannica’, a number will be found on the under side of the ticket, corresponding to the number of the species in that work. other type specimens are either marked by a round white ticket without a number, or by a small square ticket with a number. mr. stephens’ own species are not marked’. i should like to stress the words ‘almost always’ in the text cited. in my opinion, they may prove that not all the ‘type specimens’ in the collection were actually marked in the way they described. about the small round labels hammond (1972) mentions: ‘all internal evidence from the collection suggests that the colour of these labels is of no great significance but that they all indicate specimens originating from marsham’s collection’; however, he did not present valid arguments against the possibility that specimens without such round tickets or having labels of other shapes may also have belonged to the named collection. the same author further stated: ‘the significance of certain other labels attached to various specimens, including white rectangles bearing a number and small red squares of paper, has not been discovered’; however, these ‘white rectangles’ may have conformed to the ‘small square tickets with a number’ in the aforecited fragment by waterhouse & janson (1855). apropos, the latter authors did not indicate a colour of the ‘tickets’, and the small red squares of paper may be analogs of the white rectangles: as we can see in the text, waterhouse & janson did not regard the numbers as indispensable attributes of the type labels. stephens never stated that all the material from marsham’s collection would be specified separately in his ‘illustrations’. indeed, he mentioned such material directly in the stenus section in the following three cases: 1. under s. marshami stephens, 1833 (=s. (tesnus) brunnipes stephens, 1833; =s. immunis marsham, 1802, pars); it was, at least in part, based on material in the series of immunis from marsham’s collection (see the ‘systematic catalogue’: stephens 1929: 289); 2. under s. immunis marsham, 1802 (the name requires a revision, see above); since the series proved to be heterogeneous, it was reasonable to show that a part of the original series remained under the name of immunis; besides, since marsham’s descriptions did not contain precise references to localities, stephens gave additional data (london & norfolk), having mentioned that his own specimens came ‘from the marshamian collection’; european journal of taxonomy 13: 1-62 (2012) 48 3. under s. laevis stephens, 1833 (=s. (metatesnus) pubescens stephens, 1833), which he attributed to marsham and therefore, to explain the posthumous attribution, found it appropriate to mention that his specimen had been ‘obtained from the marshamian cabinet’. however, i can find no reasons for particularly citing material from marsham's cabinet under the name melanopus: the species was thoroughly described and published by marsham. it was a well known fact that stephens had purchased marsham’s collection. in this case, stephens merely gave the redescription more detailed based on the ‘type specimen’ and then added: ‘also found, not uncommonly, within the metropolitan district and in norfolk’, with a reference to the habitat data provided by l. w. dillwyn, esq. the word ‘also’ here means, in my opinion, that the above redescription is based on marsham’s original material. based on the statement by waterhouse & janson (1855), as well as on all the facts aforesaid, i consider the specimen under discussion to be an original type specimen that can be designated as the lectotype. the remaining three specimens (1 ♀ of stenus (tesnus) brunnipes stephens, 1833; 1 ♂, 1 ♀ of stenus (nestus) boops ljungh, 1810) have not been marked as paralectotypes inasmuch as their belonging to the original type series seems improbable; all those have been supplied with my common determinative labels. each of these specimens has also been labelled by waterhouse: ‘the species follow in succession, in the cabinet, in accordance with the descriptions in the “illustrations,” but in one or two instances there have un doubtedly been some accidental transpositions; and to prevent further changes of this nature, all the species, and indeed nearly all the specimens, have now been numbered to correspond with the numbers of the species as given in the “manual.” [=stephens 1839]. the numbers here alluded to are on small oval tickets attached to the specimens […]’ (waterhouse & janson 1855). besides the number ‘53’, the oval individual labels bear additional markings: one dagger under s. brunnipes; two daggers (one under another) under each of the specimens of s. boops. the daggers may have marked extraneous species within each series. in different times different species have been erroneously synonymized with s. melanopus: s. (n.) capitatus eppelsheim, 1878 (fauvel 1878, 1902; fowler 1888; ganglbauer 1895; reitter 1909), s. (n.) discretus rey, 1884 (now a synonym of s. (n.) crassus stephens, 1833) (ganglbauer 1895; fauvel 1902; reitter 1909; bernhauer & schubert 1911; l. benick 1929; dauphin 1993), s. (n.) ignotus eppelsheim, 1890 (fauvel 1902), s. (n.) piscator saulcy, 1864 (fauvel 1902), s. (n.) arctulus hochhuth, 1849 (now a synonym of s. (n.) incanus erichson, 1839) (fauvel 1902), s. (n.) morulus baudi, 1870 (now a synonym of s. (n.) piscator saulcy, 1864) (fauvel 1902) and s. (n.) sulcifrons eppelsheim, 1878 (now a synonym of s. (n.) piscator saulcy, 1864) (fauvel 1902). among the names having been regarded as synonyms of s. melanopus till now, only s. tythus schaufuss, 1882 does not raise doubts: its female type was studied and identified as melanopus by puthz (1967c). none of the names s. cribrellus rey, 1884, s. nitidus lacordaire, 1835 or s. sulcicollis stephens, 1833 have been revised based on the type material. the last of these seems to be a nomen dubium. waterhouse & janson (1855: 148-149) indicated: ‘the detailed description in this work is pos sibly from a small specimen of st. [(hemistenus)] subaeneus, er. the insect in mr. stephens’ cabinet which stands for sulcicollis evidently is misplaced by a black-legged species, i.e. st. melanopus, of marsham’. the result of this mess was marked well by rye (1870): ‘this [sulcicollis] and the preceding [assimilis] are given as valid species in gemm.[inger] & v. harold’s cat.[alogue], ii, though (? because) they are not re cognizable or known to british entomologists. the former is represented by brunnipes, and the latter by melanopus, in stephens’ collection. see wat. & janson ([…] 1855), from whose account of the con fusion with regard to these insects it is evidently impossible to do anything but ignore them altogether.’ fauvel (1869), without any explanation, put the ryvkin a.b., stenus species of the canaliculatus group 49 name sulcicollis in the synonymy of melanopus. the assumption by puthz (2008) that fauvel had seen the ‘type’ in stephens’ collection before synonymizing sulcicollis (‘fauvel hat das 1869 wohl ebenso gesehen, sonst hätte er seine synonymie nicht ausgesprochen’) is contrary to fact: unlike many other names, which are supplied with remarks ‘ex typ.’ in the paper cited, the pair ‘s. sulcicollis steph. = s. melanopus marsh.’ has no such remark. in addition, the reference by puthz (2008) to the ‘indisputable authority’ of fauvel, who actually had the name of ‘master of false synonymy’ (lohse 1985; see, for example, the list of erroneous synonyms above) seems not to be the best way to confirm ‘ausgesprochene synonymie.’ by the way, in the later work (1873), fauvel himself discussed the problem much more deliberately, with a reference to rye (1870): ‘le sulcicollis steph. (ill. brit., v, 295) , représenté dans la collection stephens par le melanopus, est une espèce méconnaissable à rayer des catalogues (v, rye, ent. annual, 1870, 84)’. bernhauer & schubert (1911), based on the original description, placed s. sulcicollis within the subgenus hemistenus motschulsky, 1860 (s.l.) as a valid species. thus, i find the synonymy of s. sulcicollis with s. melanopus to be ill-founded. the original descriptions of both s. cribrellus and s. nitidus correspond rather to species of the atratulus complex than to s. melanopus: ‘un exemplaire [...] à prothorax paraissant un peu plus court, subtransverse, avec une fossette poncti-forme, obsolète, seulement visible suivant un certain jour et située près de la base [...]’ (rey 1884); ‘prothorax un peu plus long que large, [...] avec une fossette oblongue peu mar quée sur le disque [...]’ (lacordaire 1835). it is obvious that the small ‘fossette ponctiforme, obsolète’ as well as ‘fossette oblongue peu mar quée’ cannot conform with the long and deep furrow on the pronotum in s. melanopus. therefore, most subsequent records of s. nitidus may have been the result of misidentification of s. melanopus. thus, s. tythus may be regarded as the only reliably established synonym of s. melanopus. this species, widely distributed over the mediterranean, w & c europe, and also introduced into the nearctic (see puthz 1966; campbell & davies 1991), is rather variable and similar to many other representatives of different groups in many characters; therefore the range: ‘mittel-südeuropa, südliches nordeuropa, nordafrika, kanaren, kleinasien, westsibirien, östliches nordamerika’ (puthz 1971a, 1972), seems to be greatly exaggerated. j. sahlberg (1876) first recorded s. melanopus for s finland (‘sällsynt; funnen några gånger invid åbo af o.reuter, och förf.[attare]’). the same data (‘regio aboënsis’) was cited by him in the subsequent catalogue (j. sahlberg 1900). although l. benick (1921a) ascertained that this record had resulted from misidentification (‘ein von sahlberg […] genanntes stück (reuter, åbo) in der sammlung des mus. hels. (‘reuter, pargas’) ist atratulus er.’), many subsequent authors continued to cite finland in the range description of melanopus (palm 1961; horion 1963; etc). benick (l.c.) soundly questioned also the record by poppius (1899: ‘statsradet a. günther [...] i trakten kring petrosawodsk’) for n european russia (‘die angabe von poppius, nach der die art in russ. karelien […] vorkommen soll, ist nachzuprüfen’). in his well-known work on siberian coleoptera, j. sahlberg (1880) found s. melanopus to be distributed over the ob’ basin (‘in territorio silvoso prope oppidum tobolsk 3/6 unicum specimen invenit bergroth’). this misleading record was repeated by heyden (1881) and became a source of erroneous citation of this species for w siberia by many authors up to present time (jakobson 1909; l. benick 1929; scheerpeltz 1933; smetana 1959; szujecki 1961; horion 1963; puthz 1971a, 1972; tichomirova 1973; dauphin 1993). neither puthz (pers. comm.) nor i have seen melanopus specimens from territories eastwards of denmark and southernmost sweden. one can confidently suppose the n russian and siberian records to relate to another species; most likely, it may be a species of the canaliculatus group, either s. labilis or s. confusus. the record for yekaterinoslav [=dnepropetrovsk] of the ukraine (iljin 1926) may also be a result of misidentification. the only material i have seen from the territory of the former ussr is given in the material section (see above: odessa area). the recent record of this species for lithuania (monsevičius & pankevičius 2001: 1 specimen of undisclosed sex) requires confirmation. european journal of taxonomy 13: 1-62 (2012) 50 the bionomics of this species seem to be rather diverse. along with different open alluvia: seashore, riverside, and lakeside (hardy 1851; rye 1864; fauvel 1873; fowler 1888; johansen 1914; l. benick 1929, 1947; palm 1961; horion 1963; anderson 1984; dauphin 1993), many other types of habitats have been reported: hay-stack rubbish (rye 1864), manure heaps (tottenham, 1954), dry manure (dauphin 1993), compost heaps (anderson 1984), detritus (smetana 1959; dauphin 1993), humus (bordoni 1974) and bogs, including peat and sphagnum (horion 1963). the suppositions of halophily for s. melanopus (l. benick 1947; puthz 1965, 1971, 1972) seem to be ill-founded, inasmuch as inhabiting seashore alluvia and lakesides of brackish basins is not a sufficient ground to consider a species to be halophilous. the idea was argued and rejected by horion (l.c.: ‘von einer allgemeinen halophilie dieser art kann keine rede sein’). i believe that the fact that s. melanopus inhabits sphagnum bogs is quite enough for the assumption of halophily to be denied. as to the records of the occurrence of this species in manure and compost heaps, i should like to note that those habitats are common shelters which many stenus species use during adverse weather or in adverse seasons. key to the palaearctic species of the canaliculatus group 1. front with deep longitudinal impressions and high median elevation in between. seventh abdominal tergite varnish shining, without ground-sculpture between punctures. puncturation of pronotum and elytra fairly deep and coarse, average diameter of punctures on pronotum larger than basal cross of antennal segment 3. disk of pronotum with deep longitudinal furrow rather broad and not lineshaped throughout. internal tooth of each mandible shifted distally, directed mediodorsally of the main plane. abdominal sternite 9 of both males and females (valvifera) with posterolateral teeth very short. aedeagus small (see fig. 7k). 2.8–3.7 mm ……………………………………………… ……………………………………………………[melanopus group] melanopus (marsham, 1802) – if front with deep longitudinal impressions and 7th abdominal tergite between punctures without regular ground sculpture, puncturation of pronotum very fine and dense with average diameter of punctures distinctly smaller than basal cross of antennal segment 3. longitudinal median furrow of pronotum more or less deep but always long, line-shaped. internal tooth of each mandible placed a little before the middle, directed only slightly dorsally of the main plane. abdominal sternite 9 of both males and females (valvifera) with posterolateral teeth normally developed (see fig. 5a–e). aedeagus large ………………………………………………………………[canaliculatus group] 2 2. front with distinct lateral impressions and median elevation in between …………………………3 – front feebly convex or feebly concave, with median elevation and lateral depressions rather vague or entirely absent ……………………………………………………………………………8 3. median elevation of front broad and knob-like. puncturation of pronotum very fine and dense, average diameter of punctures distinctly smaller than basal cross of antennal segment 3. disk of abdominal tergite 7 without regular ground sculpture or with reticulation very feeble and vague. aedeagus as in fig. 1e. 3.3–4.5 mm ………………………………………………………labilis erichson, 1840 – median elevation of front ridge-shaped, angular in its cross section, flanked by broader and less sharp lateral impressions. puncturation of pronotum coarser, average diameter of punctures about as large as or larger than basal cross of antennal segment 3 ………………………………………4 4. apical abdominal tergites without ground sculpture. for aedeagus, see puthz 2006 (fig. 4). 2.9–4.0 mm …………………………………………………………………………immigratus puthz, 2006 ryvkin a.b., stenus species of the canaliculatus group 51 – at least tergites 7–10 distinctly reticulated …………………………………………………………5 5. elytral suture about 1/4–1/5 longer than pronotum. aedeagus as in fig. 7c–h. 4.0–4.7 mm …………………………………………………………………………latipennis j. sahlberg, 1880 – elytral suture not more than l/7 longer than pronotum ……………………………………………6 6. elytral suture a bit shorter than pronotum. aedeagus as in figs. 6a, 7a. 4.0–4.6 mm …delitor sp. nov. – elytral suture distinctly longer than pronotum ……………………………………………………7 7. elytra more than 1/4 broader than head. puncturation less coarse, much more regular; interspaces between punctures along the midline of anterior visible tergites distinctly wider than average diameter of punctures. aedeagus as in fig. 2e. 3.4–4.0 mm ……………confusus j. sahlberg, 1876 – elytra by 1/4–1/5 broader than head. puncturation much coarser, distinctly irregular; interspaces between punctures along the midline of anterior visible tergites smaller than average diameter of punctures. aedeagus as in fig. 7b. 3.9–4.4 mm ………………………………illusor ryvkin, 1987 8. elytra not longer than pronotum, nearly as broad as head. abdominal tergites with very dense and deep ground-sculpture. aedeagus as in fig. 2c, d. 3.5–4.7 mm (the latter value for specimens with the abdomen extended) ……………………………………illotulus puthz, 1972 – elytra longer than pronotum, broader than head …………………………………………………9 9. front as a whole feebly concave, with median elevation very vague, nearly flat, situated below internal margin level of eye; the upper surface except abdominal tergite 10 without ground sculpture. for aedeagus, see puthz, 1987 (abb. 1). 3.3–3.7 mm …………………………shogun puthz, 1987 – if front as a whole concave, at least abdominal tergites 7–8 with evident ground sculpture ……10 10. front with median elevation and lateral depressions rather feeble but evident. median parts of anterior abdominal tergites with puncturation very small, scattered, and shallow. for aedeagus, see renkonen, 1935 (fig. 1). 3.0–3.8 mm ……………vinnulus casey, 1884 – front with median elevation entirely absent or, occasionally (in s. nitens stephens, 1833), represented by narrow, smooth longitudinal strip ………………………………………………11 11. elytra with fairly small and dense puncturation. anterior margin of labrum with a small median notch ………………………………………………………………………………………………12 – elytra with puncturation larger and more or less sparse. anterior margin of labrum without distinct median notch, at most feebly sinuate …………………………………………………15 12. pronotum 1/7–1/8 longer than broad. aedea gus as in fig. 3a, b. 3.2–3.9 mm (the latter value for specimens with abdomen extended) …………………………………………alopex sp. nov. – pronotum not, or only a little, longer than broad …………………………………………………13 13. entire body with ground sculpture more or less visible. aedea gus as in fig. 3d. 3.0–4.5 mm (the latter value for the american specimen with abdomen extended, the maximum value for palaearctic specimens known to me: 4.3 mm) ………………………canaliculatus gyllenhal, 1827 – at least median part of abdominal tergites 3–7 without traces of microsculpture ………………14 14. elytra between punctures with ground sculpture throughout. aedea gus as in fig. 4a, b. 4.0 mm (with abdomen extended) …………………………………………………canosus sp. nov. – elytra between punctures with ground sculpture mainly in humeral impressions and near suture. aedea gus as in fig. 4c, d. 2.8–3.7 mm (the latter value for specimen with abdomen extended) …………………………………………………………………………canalis sp. nov. european journal of taxonomy 13: 1-62 (2012) 52 15. middle of front punctured much sparser than lateral parts, occasionally with a small longitudinal strip or spot broader than the average diameter of punctures. elytra only a bit (l/14th to l/15th) broader than head, being punctured larger and sparser. aedeagus as in fig. 1a–d. 3.4–4.0 mm (the latter value for specimens with the abdomen extended) …………………nitens stephens, 1833 – front with puncturation more regular, without evident smooth strips or spots. elytra l/7 to l/10 broader than head, being smaller and denser punctured. aedeagus as in fig. 2a, b. 3.1–4.1 mm (the latter value for specimens with the abdomen extended) ………………raddei ryvkin, 1987 ackowledgements i wish to thank v.v. belov, k.yu. eskov, s.a. kurbatov, s.l. kuz'min, p.n. petrov, r.a. rakitov, l.b. rybalov, v.b. semenov, s.a. serbenyuk, a.v. sokolov, i.d. sukacheva, k.p. tomkovich, i.a. ushakov (moscow), o.n. kabakov, b.a. korotyayev (st. petersburg), e.a. khachikov, v.i. lomakin (rostov-on-don), l.n. dubeshko, v.g. shilenkov (irkutsk), e.g. matis, d.i. berman, a.s. ryabukhin (magadan), a.v. shavrin (daugavpils), s.v. saluk (minsk), a.v. gontarenko (odessa), a.g. shatrovskiy (kharkov), s.ya. blinshtein (dortmund), a.yu. solodovnikov (copenhagen), l. penev (sofia), the late a.l. tichomirova, p.k. yeryomin, v.v. zherikhin, for the material donated. i am indebted to the curators and holders of the collections named in the material and methods section, who gave me an opportunity to study respective material. i express my profound gratitude to mrs. sharon shute (bm) for the exhaustive historical reference on the series of s. melanopus marsham, 1802, as well as to martin brendell (bm) for useful advice concerning the specimens. i am thankful to alexey solodovnikov (copenhagen), who has supplied me with a copy of marsham’s original description of s. melanopus, and to oliver betz for the reprints of his splendid works. i am indebted also to the administration and staff of the aforenamed nature reserves who have assisted in my own field work in these protected areas. my best thanks are due to my dear colleagues arnaldo bordoni (florence), adriano zanetti (verona), alexey shavrin (daugavpils) for information concerning material; and also to volker puthz (schlitz) for various useful data including the photo of the aedeagus of s. sphaerops casey, 1884 from his collection. and finally, i am perpetually indebted to my wife and colleague elena veselova for the help i have received both in our common collecting trips and in our joint office study. references anderson r. 1984. staphylinidae (coleoptera) in ireland. 3: steninae. the irish naturalist’s journal 21(6): 242–251. benick l. 1921a. über nord-palaearktische steninen, vorwiegend aus dem zoologischen museum in helsingfors (col., staphyl.). mit 4 fig. meddelanden af societas pro fauna et flora fennica 46(1919– 1920): 135–156. benick l. 1921b. über stenus-typen von mäklin und motschulsky aus dem zoologsichen museum in helsingfors, nebst 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[in russian]. in: pochvennyie bespozvonochnyie moskovskoy oblasti: 201–222. nauka, moscow. tottenham c.e. 1937. note on species of philonthus in the stephens collection. the entomologist’s monthly magazine 73: 190–191. tottenham c.e. 1954. coleoptera, staphylinidae. section (a) piestinae to euaesthetinae. handbooks for the identification of british insects 4(8a): 1–79. uhlig m. & vogel j. 1981. zur staphylinidenfauna der umgebung von wahren/müritz (mecklenburg). − mitteilungen aus dem zoologischen museum in berlin 57(1): 75–168. http://dx.doi.org/10.1002/ mmnz.19810570107 waterhouse g.r. 1858. catalog of british coleoptera: i–iv+1–117. taylor and francis, london. http://dx.doi.org/10.5962/bhl.title.8987 http://dx.doi.org/10.5962/bhl.title.8987 http://dx.doi.org/10.5962/bhl.title.8133 http://dx.doi.org/10.5962/bhl.title.59227 http://dx.doi.org/10.1002/mmnz.19810570107 http://dx.doi.org/10.1002/mmnz.19810570107 european journal of taxonomy 13: 1-62 (2012) 62 waterhouse g.r. & janson e.w. 1855. british species of the genus stenus. transactions of the entomological society of london (n. ser.) 3: 136–156. http://dx.doi.org/10.1111/j.1365-2311.1855. tb02667.x weinreich e. 1968. über den klebfangapparat der imagines von stenus latr. (coleopt., staphylinidae) mit einem beitrag zur kenntnis der jugendstadien dieser gattung. zeitschrift für morphologie der tiere 62: 162–210. wüsthoff w. 1934. beitrag zur kenntnis der mitteleuropäischen stenusarten. entomologische blätter 30(2): 62–64+taf. i–iv. manuscript received: 9 october 2011 manuscript accepted: 9 december 2011 published on: 24 may 2012 topic editor: malcolm scoble in compliance with the iczn, printed versions of all papers are deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; national botanic garden of belgium, meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgium institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark. http://dx.doi.org/10.1111/j.1365-2311.1855.tb02667.x http://dx.doi.org/10.1111/j.1365-2311.1855.tb02667.x european journal of taxonomy 310: 1–32 issn 2118-9773 https://doi.org/10.5852/ejt.2017.310 www.europeanjournaloftaxonomy.eu 2017 · van ginneken m. et al. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e urn:lsid:zoobank.org:pub:e07a58ba-7d3b-4231-9c77-97a031128e62 1 morphometry and dna barcoding reveal cryptic diversity in the genus enteromius (cypriniformes: cyprinidae) from the congo basin, africa marjolein van ginneken 1,†, eva decru 2,*,†, erik verheyen 3 & jos snoeks 4 1 department of biology, systemic physiological and ecotoxicological research, university of antwerp, groenenborgerlaan 171, 2020 antwerpen, belgium. 2,4 royal museum for central africa, section vertebrates, ichthyology, leuvensesteenweg 13, 3080 tervuren, belgium. 2,4 department of biology, laboratory of biodiversity and evolutionary genomics, ku leuven, charles deberiotstraat 32, 3000 leuven, belgium. 3 royal belgian institute of natural sciences, od taxonomy and phylogeny, vautierstraat 29, 1000 brussels, belgium; department of biology, evolutionary ecology group, university of antwerpen, campus drie eiken, building d, room d.150 universiteitsplein 1, 2610 antwerpen, belgium. † equally contributing authors * corresponding author: eva.decru@africamuseum.be 1 email: marjolein.vanginneken@uantwerpen.be 3 email: everheyen@naturalsciences.be 4 email: jos.snoeks@africamuseum.be 1 urn:lsid:zoobank.org:author:fdadb435-8b18-49c9-b803-f925abf21a22 2 urn:lsid:zoobank.org:author:1aeb7eed-c939-4702-8590-b3fca7076324 3 urn:lsid:zoobank.org:author:86b40463-e3d9-4147-9ed3-d7302e0d64b6 4 urn:lsid:zoobank.org:author:13a8ab26-ff46-437c-9806-d49e11c5e15d abstract. one of the main challenges to adequately conserve the african fish fauna is to improve our so far unsatisfactory taxonomic knowledge of important portions of the ichthyofauna. in the present study, we attempted to unravel the taxonomic diversity of some species of enteromius cope, 1867, a problematic african fish genus, recently collected in the north-eastern part of the congo basin. we used an integrative approach, combining dna barcodes and morphological analyses. for one of the species complexes found, the e. miolepis/eutaenia species complex, we evaluated taxonomic diversity over a larger geographic scale within the congo drainage system. although initial literature-based species identifications allowed us to assign all examined specimens to four tentative species, dna barcodes indicated the presence of 23 distinct mitochondrial lineages. the majority of these lineages appeared endemic to particular rivers, and in most rivers multiple lineages occur in sympatry. subsequent exploratory morphometric analyses indicated that almost all these lineages are morphologically distinguishable and that they may therefore represent undescribed species. as only a part of the congo basin and a subset of the species diversity within enteromius were examined, it appears that the species richness of enteromius in the congo basin is severely underestimated. keywords. ‘barbus’, coi, integrative taxonomy, species richness. https://doi.org/10.5852/ejt.2017.310 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:e07a58ba-7d3b-4231-9c77-97a031128e62 mailto:eva.decru%40africamuseum.be?subject= mailto:marjolein.vanginneken%40uantwerpen.be?subject= mailto:everheyen%40naturalsciences.be?subject= mailto:jos.snoeks%40africamuseum.be?subject= http://zoobank.org/urn:lsid:zoobank.org:author:fdadb435-8b18-49c9-b803-f925abf21a22 http://zoobank.org/urn:lsid:zoobank.org:author:1aeb7eed-c939-4702-8590-b3fca7076324 http://zoobank.org/urn:lsid:zoobank.org:author:86b40463-e3d9-4147-9ed3-d7302e0d64b6 http://zoobank.org/urn:lsid:zoobank.org:author:13a8ab26-ff46-437c-9806-d49e11c5e15d european journal of taxonomy 310: 1–32 (2017) 2 van ginneken m., decru e., verheyen e. & snoeks j. 2017. morphometry and dna barcoding reveal cryptic diversity in the genus enteromius (cypriniformes: cyprinidae) from the congo basin, africa. european journal of taxonomy 310: 1–32. https://doi.org/10.5852/ejt.2017.310 introduction with some 3060 species in about 375 genera already described (froese & pauly 2017), cyprinidae is the largest family of freshwater fishes in the world. within this family, the former genus barbus sensu lato (s.l.) cuvier & cloquet, 1816, was a very extensive paraphyletic aggregation of over 800 species occurring in europe, africa and asia. based on ploidy level, barbus s.l. has been divided into three groups: diploids (2n = 48 or 50), tetraploids (2n = 100) and hexaploids (2n = 148–150) (agnèse et al. 1990; oellerman & skelton 1990; rab et al. 1995; berrebi & valiushok 1998). the phylogeny of some of these groups has been examined (e.g., agnèse et al. 1990; tsigenopoulos et al. 2010), and a number of studies suggested to split barbus s.l. into several genera at least based on the level of ploidy (berrebi et al. 1996). due to their uncertain taxonomic position, the african small-sized diploid species were, until recently, referred to as ‘barbus’ (between quotation marks) as proposed by berrebi et al. (1996). a recent study explored the classification of the subfamily cyprininae that includes barbus s.l., using mitochondrial and nuclear genes (yang et al. 2015). the results of this study supported a close relationship within the diploid species between the asian puntius hamilton, 1822 species and the african small ‘barbus’. as such, they confirmed earlier results based on karyology (golbutsov & krysanov 1993; rab et al. 1995). yang et al. (2015) proposed the revalidation of the genus enteromius cope, 1867 to accommodate all african diploid ‘barbus’ species, as it is the oldest available genus group name for these fishes. recently, this revalidation has been criticized (schmidt & bart jr. 2015); we prefer, however, the use of enteromius over ‘barbus’. the taxonomy of species of enteromius is to date insufficiently known, resulting in difficulties in identifications and incomplete inventories of the species. it also hampers further studies on phylogenetic relationships, which are still obscure among species of enteromius (berrebi et al. 2014; yang et al. 2015; ren & mayden 2016). while these problems have been tackled in various asian taxa (berrebi & tsigenopoulos 2003; pethiyagoda et al. 2012), the need for such revisions remains for the african species. this study focuses on some species of enteromius from the congo basin. the congo basin comprises the second largest catchment area in the world after the amazon (snoeks et al. 2011). it harbours a very high diversity of fishes with at least 957 valid species currently listed, excluding lakes kivu and tanganyika as well as the malagarazi system (froese & pauly 2017; musschoot, rmca, pers. comm.). the congo basin is generally divided into three main sections: the upper congo (or lualaba), which runs from the basin’s source in zambia down to the wagenia falls near the city of kisangani in north-eastern dr congo; the middle congo, which starts at the wagenia falls, and flows in a large arc, first northwestwards, then southwards, before widening into pool malebo; and the lower congo, which starts at the outlet of pool malebo and runs down to the mouth of the basin in the atlantic ocean (runge 2007). despite the fact that the congo basin is recognized as a hotspot for fish diversity (snoeks et al. 2011), the ichthyofauna in large parts of the basin remains poorly studied. to date, 90 morphologically often very similar species of enteromius are recognized from the congo basin (vreven, rmca, pers. comm.). identification keys for the region are lacking and the designation of so-called species complexes is the result of the identification difficulties. one of the specific taxonomic problems within the congolese species of enteromius concerns the e. miolepis/eutaenia complex, which comprises species characterized by a strongly ossified first dorsal ray that is serrated along its posterior margin and a blackish mid-lateral band that extends from the tip of the snout to the caudal fin base or onto the mid-central part of the caudal fin (tweddle & skelton 2008; banyankimbona et al. 2012). due to the similarity of the species within https://doi.org/10.5852/ejt.2017.310 van ginneken m. et al., unexpected cryptic diversity in enteromius 3 this complex (poll 1976), the available information on morphological characteristics is often insufficient to arrive at a correct species identification. in addition, e. miolepis (boulenger, 1902) and e. eutaenia (boulenger, 1904) are both characterized by high intraspecific morphological variation, which suggests that these taxa could be polyspecific. some congolese specimens were identified as e. holotaenia (boulenger, 1904) or e. kerstenii (peters, 1868), although both species were described from outside the congo basin, from the ogowe river in gabon and from a coastal river in zanzibar respectively. with their ossified, serrated dorsal spine and blackish mid-lateral band, they can be considered members of the e. miolepis/eutaenia complex. enteromius holotaenia can only be distinguished from other members of the e. miolepis/eutaenia complex by the presence of a black dorsal fin tip, which is not always clearly visible, and e. kerstenii by the presence of a red opercular spot and the absence of large scales on the dorsal fin base. however, in the congolese specimens identified as e. kerstenii, some enlarged scales seem to be present at the dorsal fin base, although not very pronounced. in order to unravel the taxonomy of species of enteromius from the congo basin and delineate the various species, a morphologic approach alone may be inadequate. the present study ascertains whether a multidisciplinary approach might facilitate the differentiation of taxa within the genus enteromius from the congo basin. for this, some species of the north-eastern part of the basin, a particularly poorly explored region, were selected as a case study. additionally, because of the taxonomic problems in the e. miolepis/eutaenia complex discussed above, we have examined specimens from this complex from other regions within the congo basin as well. we explored whether this multidisciplinary approach would result in the discovery of cryptic species, and the detection of (additional) morphological characteristics to discriminate between these species. to attain these objectives, we used dna barcoding (hebert et al. 2003) together with traditional morphometrical techniques. molecular techniques like dna barcoding have already proven to be powerful for the identification of both marine and freshwater fish taxa (becker et al. 2011; pereira et al. 2013), and to detect cryptic species diversity or taxonomic inconsistencies (goodier et al. 2011; collins & cruickshank 2012; decru et al. 2016). however, the uncritical use of molecular tools to identify species, especially using only a single molecular locus (mt genome), is unwarranted (desalle et al. 2011), and traditional morphology has proven its value as a suitable technique to assess the diversity in many taxa. therefore, we combined both methodologies in the present study, an approach that has already proven to be successful in several taxonomic studies (e.g., olayemi et al. 2012; stiassny et al. 2013; lavoué & sullivan 2014). material & methods specimen selection (fig. 2b, table 1) in total, we used 181 specimens of enteromius from the congo basin for morphological and/or genetic analyses. to investigate the taxonomic diversity of enteromius in the north-eastern part of the congo basin (middle congo), we examined specimens of all species of enteromius for which tissue samples had been recently collected from rivers in this region, i.e., the ituri, its tributary the epulu, the lobilo, and the lomami/lobaye system. to unravel the e. miolepis/eutaenia complex, we also examined specimens from various sampling sites in the upper, middle and lower congo, identified as e. miolepis, e. eutaenia, e. holotaenia or e. kerstenii. for the middle congo, we included specimens from the léfini, the itimbiri, the ituri/epulu, the lobilo, and the lomami/lobaye. for the upper congo we included samples from the loboya (a tributary of the maiko), and the luapula. finally, for the lower congo we used samples from the rivers luki and inkisi. these specimens were collected during ten expeditions that were carried out between 2005 and 2012. european journal of taxonomy 310: 1–32 (2017) 4 relevant types were selected based on the literature-based identifications and include the holotype of e. miolepis described from the yembe river at banzyville (ubangi river system); 15 syntypes of e. holotaenia described from the ogowe river in gaboon; one lectotype and five paralectotypes of e. eutaenia described from huila (mossamedes) in angola; the lectotype of e. kerstenii described from the coast opposite zanzibar; the holotype of e. brazzai (pellegrin, 1901) described from the sangha river in mobaka; the holotype and five paratypes of e. tshopoensis (de vos, 1991), a junior synonym of e. brazzai, which was described from the tshopo river; the lectotype and three paralectotypes of e. pellegrini (poll, 1939) described from lake kivu; and two paratypes of e. atromaculatus (nichols & griscom, 1917) described from the yakuluku river (lévêque & daget 1984) (see also appendix 1). we attempted to use the same specimens for the morphological and genetic analyses, but this was not always possible. as some specimens were lost or too damaged, 40 specimens that were genetically examined could not be used for morphological analyses. in 19 of these cases, literature-based identifications could not even be done, and the specimens were identified as enteromius sp. as such, we morphologically examined 177 specimens (table 1, appendix 1), including several type specimens. similarly, not all specimens that we examined morphologically could be successfully dna barcoded, and genetic samples are unavailable for the type specimens. morphological analyses specimen identifications based on published characteristics, we assigned the non-type specimens to four ‘a priori’ groups: 60 to e. cf. miolepis (which represents the e. miolepis/eutaenia complex); 15 to e. cf. brazzai; 26 to e. cf. pellegrini; and 40 to e. cf. atromaculatus. we used ‘cf.’ to indicate the uncertainty connected to their identifications. the four groups mainly differ in the morphology of their dorsal spine (ossified or flexible; serrated or not) and colour pattern (fig. 1). enteromius cf. miolepis has an ossified serrated dorsal spine and a mid-lateral band on the flank and through the eye; e. cf. brazzai has a flexible unserrated dorsal spine and no melanin markings; e. cf. pellegrini has an ossified serrated dorsal spine and large dark fig. 1. schematic representation of the four ‘a priori’ enteromius groups with their characteristic morphological features (dorsal spine morphology and melanin pattern). a. e. cf. miolepis (boulenger, 1902) (38.1–111.0 mm). b. e. cf. pellegrini (poll, 1939) (40.7–81.3 mm). c. e. cf. brazzai (pellegrin, 1901) (44.8–82.8 mm). d. e. cf. atromaculatus (nichols & griscom, 1917) (28.3–55.8 mm). drawings modified from bamba et al. (2011). van ginneken m. et al., unexpected cryptic diversity in enteromius 5 spots that can fuse into a mid-lateral band; and e. cf. atromaculatus has a flexible unserrated dorsal spine and small spots that can fuse into a mid-lateral band. morphometry for the 177 examined specimens, we obtained 17 measurements with a vernier caliper (helios, 0.05 mm), as well as 10 meristics. counts and measurements were based on bamba et al. (2011), with some small modifications: the pre-occipital distance, post-dorsal distance i, post-anal distance i, and body-depth i were not included; the standard length, post-dorsal distance ii, and post-anal distance ii were measured up to the insertion of the first caudal fin ray of the upper lobe instead of to the middle of the caudal peduncle base. barbel lengths were also coded according to bamba et al. (2011). head measurements were expressed as percentages of head length (hl) and body measurements as percentages of standard length (sl). abbreviations of the measurements and counts: afr = anal fin rays bd = body depth cp sc = caudal peduncle scales dfr = dorsal fin rays d-l sc = scales dorsal fin-lateral line a region river spec. examined (morphological/ genetical) spec. examined morphological & genetical middle congo ituri 36/28 16 epulu (tributary ituri) 32/7 7 lobilo 10/11 10 lobaye (tributary lomami) 3/3 3 b upper congo luapula 6/7 6 loboya (tributary maiko) 0/1 0 middle congo léfini 5/5 5 itimbiri 2/2 2 ituri 6/13 4 epulu (tributary ituri) 19/7 5 lobilo 4/7 4 lomami 1/1 1 lobaye (tributary lomami) 4/4 3 lower congo inkisi 3/8 3 luki 10/10 5 table 1. number of examined non-type specimens of enteromius cope, 1867 per river (system) studied. a. specimens other than those belonging to the e. miolepis/eutaniea complex. b. specimens belonging to the e. miolepis/eutaenia complex. numbers represent morphometrically analysed specimens vs genetic samples, then the number of specimens analysed morphologically as well as genetically. european journal of taxonomy 310: 1–32 (2017) 6 dofbl = dorsal finbase length dofl = dorsal fin length ed = eye diameter hl = head length iow = interorbital width l-b sc = scales lateral line-belly ll sc = lateral line scales l-p sc = scales lateral line-pelvic fin mncpd = minimum caudal peduncle depth mxcpd = maximum caudal peduncle depth pd sc = pre-dorsal scales pecfr = pectoral fin rays pelfr = pelvic fin rays poad = post-anal distance podd = post-dorsal distance prad = pre-anal distance prdd = pre-dorsal distance propd = pre-occipital distance prpecd = pre-pectoral distance prpeld = pre-pelvic distance sl = standard length snl = snout length data analyses we used principal component analysis to explore the multivariate data matrix and to reduce the large number of variables into a few meaningful axes (snoeks 2004; decru et al. 2012). pcas were executed in past 3.15c (hammer et al. 2001). raw data were used for meristics, log-transformations for the measurements. by using log-transformations, the first component can be interpreted as a proxy for size, since it is characterized by loadings with the same sign and same order of magnitude for all variables (bookstein et al. 1985). because the barbel lengths were coded (bamba et al. 2011) as categorical variables, they were not included in the pcas. pc loadings of the illustrated pcas are given in the appendices 2–8. genetic analyses dna extraction, pcr amplification and sequencing we successfully barcoded 114 specimens using dna extracted from finclips (table 1). the method used is based on the fish-bol (fish barcode of life) protocol (steinke & hanner 2011). dna was extracted using a ‘nucleospin® tissue kit’ following the instructions of the manufacturer (macherey-nagel, germany). subsequently, polymerase chain reactions (pcr) were used to amplify the mitochondrial cytochrome c oxidase i (coi) gene. a part of the samples was amplified with ‘fish cocktail’, an m13 tailed primer combination (ivanova et al. 2007) (table 2). a standard 25 µl pcr mix consisted of 2.5 µl pcr buffer (10x); 2.5 μl dntp (2 mm); 1.25 μl ‘fish cocktail’ (2 μm); 0.2 taq dna polymerase (5 units per μl); 16.75 μl mq-h2o and 2.0 μl of the extracted dna sample. the pcr profile was 3 min at 94°c, followed by 35 cycles of 40 s at 94°c, 40 s at 52°c, and 1 min at 72°c, plus a final extension of 10 min at 72°c. the success rate of this pcr-mix was relatively low, which led to the development of specific primers, bbus f and bbus r, for the amplification of samples that weren’t successfully sequenced with ‘fish cocktail’ (table 2). in this pcr mix, instead of ‘fish cocktail’, 2.5 µl of each specific primer was used, and an annealing temperature of 54°c instead of 52°c. amplified products were verified on 1.2% agarose gels. afterwards the pcr products were purified using a nucleofast® van ginneken m. et al., unexpected cryptic diversity in enteromius 7 96 pcr kit (macherey-nagel, germany), and sequenced bidirectionally using bigdye terminator v.3.1 and an abi 3130 capillary sequencer. dna analyses the dna sequences were assembled and visually checked in codoncode aligner 5.1.4 (codoncode corporation) and aligned in mega 5.2 using muscle alignment (edgar 2004; tamura et al. 2011). the longer m13-sequences were shortened to the length of the bbus-sequences (558 bp). next, mega 5.2 was used to execute a model test and to create a phylogenetic tree. the model test indicated the gtr+g+i as the optimal model using the aic criterion. this model was used to construct a maximumlikelihood (ml) tree with 1000 bootstrap replications. as both the ml tree and nj (neighbour joining) tree gave similar results, statistical node support of both methods are visualised on the ml tree. results class actinopterygii klein, 1885 subclass neopterygii regan, 1923 division teleostei müller, 1846 order cypriniformes rafinesque, 1810 family cyprinidae rafinesque 1815 genus enteromius cope, 1867 evaluation of the literature-based identifications in the ml tree obtained (fig. 2), lineages with less than 2% sequence divergence were collapsed and named after the river system from which the specimens were collected. specimens from the lomami/ lobaye system and the lobilo were grouped under the label ‘kisangani region’, since there appeared to be little or no genetic difference between samples of these nearby affluents. the ml tree shows 23 lineages within the assayed enteromius samples, representing the four ‘a priori’ morphospecies in the following quantities/properties: 13 of which belong to the e. cf. miolepis group; three to the e. cf. brazzai group; one to the e. cf. pellegrini group; and six to the e. cf. atromaculatus name cocktail name / primer sequence 5’–3’ ‘fish cocktail’ (m13-tail): c_fishf1t1-c_fishr1t1 (ratio 1:1:1:1) vf2_t1 tgtaaaacgacggccagtcaaccaaccacaaagacattggcac fishf2_t1 tgtaaaacgacggccagtcgactaatcataaagatatcggcac fishr2_t1 caggaaacagctatgacacttcagggtgaccgaagaatcagaa fr1d_t1 caggaaacagctatgacacctcagggtgtccgaaraaycaraa reference: ivanova et al. (2007) primer sequences for m13-tails m13f tgtaaaacgacggccagt m13r caggaaacagctatgac reference: messing (1983) primer sequences for enteromius specific primers bbus f tgagccggaatagtgggaac bbus r cctgcrgggtcraagaatgt table 2. sequences 5’–3’ of the primers used for the pcr reactions. european journal of taxonomy 310: 1–32 (2017) 8 fig. 2. a. ml tree based on 558-bp-long enteromius coi sequences with 1000 bootstrap replications, with node support shown as nj/ml bootstrap (bootstrap values > 95% are shown; lineages < 2% sequence divergence were collapsed), the label ‘kisangani region’ contains samples from the lomami/ lobaye system and the lobilo. b. map of the congo basin with the sampled river stretches indicated according to the phylogenetic lineages. van ginneken m. et al., unexpected cryptic diversity in enteromius 9 group. enteromius cf. miolepis, e. cf. atromaculatus and e. cf. pellegrini each form clearly supported clades, while this is not the case for e. cf. brazzai. there is often a substantial genetic distance between lineages occurring in different rivers, but also between some lineages detected in the same river. for example, within the e. cf. miolepis group, we observed a considerable difference between specimens of the luapula (three lineages; 14.6% sequence divergence between luapula 1 and 2; 15.3% between luapula 1 and 3; 19.0% between luapula 2 and 3), and the luki (two lineages; 7.7% sequence divergence). interestingly, the sequences obtained for the upper, middle and lower congo populations did not group according to these zoogeographic regions. however, two out of the three lineages of the luapula, a river system geographically remote from the other rivers included in this study, also show the largest genetic distances within the e. cf. miolepis clade. enteromius cf. brazzai is not resolved as a single clade, and consists of three lineages, two of which were detected in the ‘kisangani region’ (19.8% sequence divergence). in contrast to the other groups, e. cf. pellegrini consists of a single lineage that is composed of specimens from a single river, the ituri. the e. cf. atromaculatus group contains an important amount of genetic variation, with five of its six lineages occurring in the ituri river. remarkably, one of those lineages also contains one sample from the ‘kisangani region’. morphometric groupings firstly, pcas were executed on all the specimens examined, after which more detailed pcas were performed on each literature-based ‘a priori’ group separately. overall analyses the highest loadings on pc2 for a pca on 17 log-transformed measurements (n = 177) are for the eye diameter (ed), the post-anal distance (poad) and the dorsal fin length (dofl); pc1 is a proxy for size (see above). on a scatterplot of pc2 against pc1, the four ‘a priori’ groups cannot all be distinguished from each other (fig. 3). however, specimens of e. cf. pellegrini and e. cf. brazzai are completely separated from each other on pc2. this is mainly because e. cf. pellegrini has a smaller ed, a smaller dorsal fin base length (dofbl) and a smaller poad than e. cf. brazzai. clearly e. cf. miolepis occupies the largest morphospace on pc2, which comprises most of the type specimens included, except for two paralectotypes of e. eutaenia and one syntype of e. holotaenia. the e. cf. atromaculatus polygon comprises the paratypes of e. atromaculatus. the specimens of e. cf. pellegrini, however, only overlap slightly with the type specimens of e. pellegrini and the specimens of e. cf. brazzai are separated from the holotype of e. brazzai and types of e. tshopoensis. furthermore, the type specimens of e. tshopoensis are separated from the other groups, mainly on pc2, due to a smaller poad and a larger dofl. the highest loadings on pc1 for a pca on 10 meristics (n = 177) are for the number of scales between the lateral line and the belly (l-b sc), the number of scales between the dorsal fin and the lateral line (d-l sc) and the number of scales between the lateral line and the pelvic fin (l-p sc); on pc2 for the number of scales on the lateral line (ll sc), the number of scales between the occiput and the base of the first dorsal fin ray (pd sc) and the number of pelvic fin rays (pelfr). similar to the analysis for the measurements, a scatterplot of pc2 against pc1 does not allow the separation of the four ‘a priori’ groups (fig. 4). again, e. cf. miolepis is the group showing the largest variation. also, e. cf. pellegrini and e. cf. brazzai are again completely separated on pc1, mainly due to e. cf. pellegrini having a higher d-l sc (4.5–5.5 vs 3.5) and a higher l-b sc (5–6 vs 4–5). furthermore, the four ‘a priori’ groups all overlap with their respective type specimens. to examine whether morphological differences could be detected between the different genetic lineages within each group, we performed pcas on e. cf. miolepis, e. cf. brazzai and e. cf. atromaculatus separately. to investigate whether one or more lineages represent a currently valid species, relevant type european journal of taxonomy 310: 1–32 (2017) 10 specimens were also included in these analyses. this resulted in a multitude of analyses for which the most important outcomes are presented below. morphometric comparisons among e. cf. miolepis lineages because of the high number of lineages and specimens for e. cf. miolepis, we analysed the major geographical regions, the upper, the middle and the lower congo, separately (see also table 1), but still included the relevant type specimens to check whether some groups could be allocated to these species. only the results of the latter are discussed as an example. for the lower congo, we detected three genetic lineages within e. cf. miolepis (fig. 2), i.e., one group containing specimens from the inkisi and two groups with specimens from the luki (luki 1 and luki 2). the highest loadings on pc1 for a pca on 10 meristics (n = 36) are for ll sc, cp sc and pecfr; on pc2 again for pecfr, afr and pd sc. the specimens from the inkisi are well separated from the luki lineages on both pc1 and pc2; and there is only one specimen overlap between luki 1 and luki 2 (fig. 5). in addition, specimens from these latter two lineages noticeably differ in the length of their barbels. lineages inkisi and luki 1 can be clearly distinguished from all type specimens (fig. 5). luki 2 overlaps with the type specimens of e. eutaenia and e. holotaenia, but luki 2 differs from the e. eutaenia types in barbel length. a pca of the log-transformed measurements did not separate the genetic lineages from each other or from the type specimens (not illustrated). fig. 3. scatterplot of pc2 against pc1 for a pca on 17 log-transformed measurements (n = 177) of enteromius cope, 1867: e. cf. miolepis (boulenger, 1902) (◊), e. cf. brazzai (pellegrin, 1901) (♦), e. cf. pellegrini (poll, 1939) (∆), and e. cf. atromaculatus (nichols & griscom, 1917) (▲). also shown are the type specimens examined of: e. miolepis (boulenger, 1902) (○), e. holotaenia (boulenger, 1904) (●), e. eutaenia (boulenger, 1904) (□), e. kerstenii (peters, 1868) (■), e. brazzai (pellegrin, 1901) ( ), e. tshopoensis (de vos, 1991) (▼), e. pellegrini (poll, 1939) (+), and e. atromaculatus (nichols & griscom, 1917) ( ). van ginneken m. et al., unexpected cryptic diversity in enteromius 11 for the middle congo and upper congo, we detected respectively six and four genetic lineages of e. cf. miolepis (fig. 2). similar exploratory morphometric analyses of these genetic lineages, as for the lower congo discussed above, indicated that almost all could be separated from each other as well as from the included type specimens. however, specimens from ‘kisangani region’ 1, itimbiri and the syntypes of e. holotaenia clustered on the pcas. yet, the itimbiri specimens had a different colour pattern (a black anal fin tip), which is absent in the ‘kisangani region’ 1 specimens as well as in the syntypes of e. holotaenia. when comparing all lineages of e. cf. miolepis (lower, middle and upper congo), each could be morphologically distinguished from the others based on meristics (fig. 6), measurements and/or barbel length (not illustrated). only specimens from kisangani region’ 1 and itimbiri could not be separated from each other, but differed in colour pattern (see above). morphometric comparisons among e. cf. brazzai lineages we observed three genetic lineages within e. cf. brazzai (fig. 2). the highest loadings on pc1 for a pca on 10 meristics (n = 22) are for ll sc, pecfr and pelfr; on pc2 for cp sc, l-b sc and again pecfr. specimens from the ‘kisangani region’ 2 are well separated from the ituri 3 lineage on pc2 and both genetic lineages differ from the ‘kisangani region’ 3 lineage and type specimens on pc1 (fig. 7). the polygon of specimens from the ‘kisangani region’ 3 lineage comprises the holotype of e. brazzai and overlaps with the types of e. tshopoensis. however, the eight specimens from the ‘kisangani region’ 3 fig. 4. scatterplot of pc2 against pc1 for a pca on 10 meristics (n = 177) of enteromius: e. cf. miolepis (boulenger, 1902) (◊), e. cf. brazzai (pellegrin, 1901) (♦), e. cf. pellegrini (poll, 1939) (∆), and e. cf. atromaculatus (nichols & griscom, 1917) (▲). also shown are the type specimens examined of: e. miolepis (boulenger, 1902) (○), e. holotaenia (boulenger, 1904) (●), e. eutaenia (boulenger, 1904) (□), e. kerstenii (peters, 1868) (■), e. brazzai (pellegrin, 1901) ( ), e. tshopoensis (de vos, 1991) (▼), e. pellegrini (poll, 1939) (+), and e. atromaculatus (nichols & griscom, 1917) ( ). european journal of taxonomy 310: 1–32 (2017) 12 lineage as well as the holotype of e. brazzai differ from the types of e. tshopoensis by the absence of barbels. morphometric comparisons among e. cf. atromaculatus lineages we detected six genetic lineages within e. cf. atromaculatus (fig. 2). although there was only 1.75% sequence divergence between the lineages of epulu 2 and ituri 8, we interpreted these lineages as separate operational taxonomic units (otus, i.e., clusters of similar dna sequences) because of the observed differences in colour pattern (specimens from the epulu 2 lineage have mid-lateral dots, while specimens from ituri 8 display a vague mid-lateral band). the single specimen from ituri 7 was lost and could not be measured. the highest loadings on pc1 for a pca of 10 meristics (n = 42) are for ll sc, pecfr and d-l sc; on pc2 for cp sc, again pecfr and d-l sc. a plot of pc2 vs pc1, separates the two specimens from the ituri 6 lineage from all other lineages based on pc2 (fig. 8). the specimens from the ituri/‘kisangani region’ lineage are separated from all other lineages mainly on pc1. the ituri 5 lineage is separated from the ituri 6 and epulu 2 lineages along pc2, from the e. atromaculatus type specimens on pc1, and from the ituri/’kisangani region’ and ituri 8 lineages on a combination of pc1 and pc2. as the initial pca resulted in a great overlap between the epulu 2 and ituri 8 lineages, we carried out a second pca on 10 meristics, only including specimens from these lineages and the types of e. atromaculatus. the highest loadings on pc1 are for ll sc, d-l sc and cp sc; on pc2 for pecfr, pd sc and the number of pelvic fin rays (pelfr). on a plot of pc2 versus pc1, specimens from epulu 2 and ituri 8 still overlap and the types of e. atromaculatus overlap with specimens from epulu 2 (fig. 9). these groups also overlap on a pca of 17 log-transformed measurements (not illustrated). fig. 5. scatterplot of pc2 against pc1 for a pca on 10 meristics (n = 36) of e. cf. miolepis specimens from the lower congo: inkisi (◊), luki 1 (♦) and luki 2 (∆). also shown are the type specimens examined of: e. miolepis (boulenger, 1902) (○), e. holotaenia (boulenger, 1904) (●), e. eutaenia (boulenger, 1904) (□) and e. kerstenii (peters, 1868) (■). van ginneken m. et al., unexpected cryptic diversity in enteromius 13 discussion multidisciplinary approach to detect cryptic species specimens from the genus enteromius are notoriously difficult to identify. as, apart from some regional reviews (e.g., bamba 2012), no extensive continent-wide reviews have been carried out for this genus, the available literature for species identifications is often limited to the accounts of the original species descriptions. recent molecular studies already revealed enteromius to be non-monophyletic (yang et al. 2015; ren & mayden 2016). our study examined whether a multidisciplinary approach could provide more insight into the taxonomy of some species of enteromius from the congo basin and whether it might reveal cryptic diversity. for the examined species in this study, a literature-based identification led to the delineation of four so-called ‘a priori’ species. the reliability of these ‘a priori’ identifications was tested using a combined approach that consisted of dna barcoding and a detailed morphometric approach that allowed the comparison of the specimens from our collection with relevant type specimens. this approach revealed a high number of potentially new species of enteromius within the examined samples/specimens. dna barcoding uncovered the existence of 23 genetic lineages within the four ‘a priori’ species obtained by literature-based identifications. the genetic distances between many of the lineages, even within the ‘a priori’ species, were substantial. sometimes they even reached almost 20%, which is considerably larger compared to most other african freshwater fish taxa, that usually have fig. 6. scatterplot of pc2 against pc1 for a pca on 8 meristics (n = 60) of e. cf. miolepis (boulenger, 1902) specimens from the congo basin (excluding types): ‘kisangani region’ 1 (◊), ituri 1 (♦), itimbiri (∆), léfini (▲), epulu 1 (○), inkisi (●), luapula 1 (□), luki 1 (■), luapula 2 ( ), luapula 3 (▼), ituri 2 (+), and luki 2 ( ). specimens from luapula 1 and luapula 2 can be separated from each other based on a pca on the log-transformed measurements; specimens of luki 2 fall separated when barbel lengths are included; specimens from kisangani region’ 1 and itimbiri can be distinguished based on colour pattern. european journal of taxonomy 310: 1–32 (2017) 14 interspecific divergences lower than or around 10% for the coi gene (see, e.g., lowenstein et al. 2011; decru et al. 2016). only three of the four ‘a priori’ species agreed with supported clades in our ml tree, i.e., e. cf. miolepis, e. cf. pellegrini and e. cf. atromaculatus, while the lineages of e. cf. brazzai do not form a supported clade. subsequent exploratory morphometric analyses revealed that, despite the small sample sizes, most lineages could be separated on the basis of their morphology. the fact that specimens from separate genetic lineages are morphologically distinguishable, suggests that they may represent different species. therefore, our hypothesis that at least the e. miolepis/eutaenia complex represents a polyspecific complex appears to be confirmed by our results, as within e. cf. miolepis we detected no less than 13 putative species. also in the ml tree of ren & mayden (2016), samples from the e. miolepis/ eutaenia complex formed five distinct lineages, three identified as e. kerstenii, one as e. miolepis, and one as e. holotaenia. furthermore, respectively three and six putative species were detected in the ‘a priori’ species e. cf. brazzai and e. cf. atromaculatus. interestingly, one of the important differences between the e. cf. brazzai lineages was the number of branched pelvic fin rays (7 vs 8). as far as we are aware, variation in the number of pelvic fin rays is uncommon in enteromius. enteromius cf. pellegrini is the only ‘a priori’ species that appears to be monospecific within the study area. providing formal descriptions with diagnoses of the undescribed species of enteromius lies outside the scope of the present study, as it requires a detailed morphological examination of more specimens of each otu and from other areas to unequivocally delineate species-specific diagnostic characteristics. in some analyses, type specimens ended up within or very close to the polygon of a particular otu: the types of e. holotaenia partially overlapped with two otus of e. cf. miolepis, i.e., luki 2 and ‘kisangani region’ 1; the holotype of e. brazzai fell within an otu of e. cf. brazzai from the kisangani region (‘kisangani region’ 3); and the types of e. atromaculatus overlapped with an otu of e. cf. atromaculatus from the epulu (epulu 2). however, this does not necessarily imply that the otus are conspecific with the respective types. our results indicated that in almost every studied river stretch at least one distinct species of enteromius occurs, which makes it very unlikely that one of our studied otus from the congo basin indeed represents e. holotaenia, a species originally described from the ogowe river in the lower guinea ichthyofaunal province. most of the specimens examined lacked the fig. 7. scatterplot of pc2 against pc1 for a pca on 10 meristics (n = 22) of e. cf. brazzai (pellegrin, 1901): ‘kisangani region’ 2 (◊), ituri 3 (♦) and ‘kisangani region’ 3 (∆). also shown are the type specimens examined of e. brazzai (pellegrin, 1901) (○) and e. tshopoensis (de vos, 1991) (●). van ginneken m. et al., unexpected cryptic diversity in enteromius 15 black dorsal tip present in e. holotaenia. however, it was present in most of the specimens from luki 2, in two of the nine specimens examined from ‘kisangani region 1’, in both specimens from itimbiri 1 and in one of the five specimens from the léfini. this demonstrates that this feature is not consistently present in the lineages. hence, it does not seem a good criterion for species to distinguish these taxa, also because this characteristic is paraphyletic in the ml tree (fig. 2). for e. atromaculatus and e. brazzai the situation appears to be different. enteromius atromaculatus is described from the yakuluku river (uele river), which is in the vicinity of the epulu river; and enteromius brazzai is described from the sangha river, which is also part of the middle congo. it is therefore plausible that the epulu 2 population is conspecific with e. atromaculatus, and that ‘kisangani region’ 3 is conspecific with e. brazzai. this implies that at least 21 of our 23 otus may represent new species that will need to be formally described in the future. distribution patterns and speciation processes all species to which the specimens were originally assigned are presumed to have large distribution ranges that, except for e. atromaculatus, extend beyond the congo basin. these wide distribution ranges of the four ‘a priori’ species on the one hand, and the often much narrower distribution ranges of the putative new species detected through dna barcoding on the other, may be a reason why up to now, these otus were not detected as different species. indeed, as the majority of earlier studies on ichthyofaunal diversity concern collections from small areas and even individual rivers, the studied specimens are often identified as one of the morphologically similar species presumed to occur in the studied region. unfortunately, these identifications rarely imply detailed comparisons with specimens from other regions, let alone type specimens. in our study, most of the sympatric lineages were morphologically clearly distinct, and were as such already assigned to different ‘a priori’ species. in addition, the allopatric lineages within each of these ‘a priori’ species were morphologically sufficiently similar to remain grouped for as long as they were not subjected to comparisons based on multivariate morphometric methods. although minor morphological differences can be interpreted as intraspecific geographic fig. 8. scatterplot of pc2 against pc1 for a pca on 10 meristics (n = 42) of e. cf. atromaculatus (nichols & griscom, 1917): ituri 5 (◊), ituri 6 (♦), ituri/‘kisangani region’ (∆), epulu 2 (▲), and ituri 8 ( ). also shown are the type specimens of e. atromaculatus (nichols & griscom, 1917) (○). european journal of taxonomy 310: 1–32 (2017) 16 variation among different populations, the detected morphological and genetic differences among the examined specimens of enteromius, suggests that these populations probably represent separate, undescribed species. although dna barcodes allowed for the clustering of putative conspecifics, the obtained tree does not necessarily reflect the phylogenetic relationships within the genus enteromius. a review of the phylogeny would not only require a comprehensive sampling of all species of enteromius (some 200 species), but also the inclusion of outgroups. nevertheless, some scenarios can be proposed concerning their evolutionary history. as in each of the geographically isolated river stretches studied distinct putative species appear to occur, allopatric speciation may well have been the main mode of speciation in this genus. however, the phylogenetic relationships among the species remain largely unresolved, as there is a lack of resolution in several parts of the ml tree. while it is possible that the lack of resolution is due to incomplete taxon sampling and/or to the small number of characters, we suggest a plausible cause to be rapid radiation, which often results in difficulties in resolving phylogenetic patterns (fernández & vrba 2005; koblmüller et al. 2010). the unresolved phylogenies would in that case be the result of an almost concurrent differentiation between populations from different river stretches on a short evolutionary timescale. the fact that in this scenario adaptation to different niches has not been necessary, could thus be the cause that only minor morphological differences exist, which were only detected after multivariate exploratory techniques. for e. cf. atromaculatus and e. cf. pellegrini, the populations from the ituri river and its most important right-bank affluent, the epulu river, have a slightly different colour pattern, which seems to support the scenario of allopatric speciation. however, for e. cf. atromaculatus, there was only a genetic divergence of 1.75% between these populations (epulu 2 and ituri 8 on fig. 2a), and even no divergence for e. cf. pellegrini (ituri/epulu on fig. 2a). also based on counts and measurements, both populations of e. cf. atromaculatus (e.g., figs 8–9) and e. cf. pellegrini (not illustrated) overlap. these two instances could be examples of intraspecific geographical colour variation rather than of two separate species in each of the two groups. this conclusion is unexpected, since even morphologically very similar populations were found to represent separate putative species in almost every individual river stretch. the observation that the populations from the ituri and the epulu cannot be distinguished based on the fig. 9. scatterplot of pc2 against pc1 for a pca on 10 meristics (n = 36) of e. cf. atromaculatus (nichols & griscom, 1917): epulu 2 (▲), and ituri 8 ( ). also shown are the type specimens of e. atromaculatus (nichols & griscom, 1917) (○). van ginneken m. et al., unexpected cryptic diversity in enteromius 17 multivariate analyses and do not appear to be genetically distinct, is even more surprising since these two river stretches are isolated. indeed, the discussed populations are separated from each other by the presence of two waterfalls, one on the epulu, just upstream of its confluence with the ituri, and one on the ituri itself (vreven, rmca, pers. comm.). these falls could form important barriers for fish species and populations, at least for upstream dispersal. however, this hypothesis does not appear to be supported by our results. interestingly, in some rivers multiple lineages of the same ‘a priori’ species appear to co-exist. this implies that in some instances morphologically similar putative (a posteriori) species may occur in sympatry; this is, e.g., the case for three lineages of e. cf. miolepis in the luapula and two in the luki river. why morphologically very similar but separate species have evolved in the same rivers cannot be inferred from the present data set, and would require a phylogenetic analysis on multiple genetic markers and, e.g., additional information on ecology. the distribution patterns, in combination with morphological (dis)similarities and the unresolved phylogenetic relationships, indicate that multiple allopatric speciation events on a short evolutionary timescale is a plausible mode of speciation for the examined species of enteromius. such allopatric divergences can occur in river systems when hydrological changes cause (simultaneous) disconnections of river stretches. remarkably, some morphologically similar putative species occur however in sympatry. impact on documented species richness by applying dna barcoding in combination with multivariate analyses of morphometric data to several enteromius populations from the congo basin and the relevant type specimens, the number of species identified has increased from four based on literature to putatively 23. considering the fact that only a part of the congo basin and only some species have been studied, our results could imply a considerable increase of the number of species within this genus when extrapolated to the entire congo basin or even africa as a whole. a similar approach applied to other tropical vertebrates has yielded similar results. for instance, in a study on african giant pouched rats (cricetomys, nesomyidae), the combined use of dna barcoding and cranial measurements lead to the discovery of at least three new species (olayemi et al. 2012). also for the fish genus pseudobarbus (cyprinidae) occurring in southern africa, 15 separate lineages were identified, using cytb and 16s data, within the (at that time) seven valid species within the genus, most of which were confirmed by morphological results (swartz et al. 2009). also in neotropical frogs, a study based on 16s rdna indicated a huge underestimation of the current species richness, as up to 115% additional species were discovered (fouquet et al. 2007). in the latter study however, no morphological analyses were performed to support the validity of the putative new species. these different studies indicate that not only for african fishes, but for several other vertebrate taxa in different regions on earth, the current species richness could be severely underestimated. however, every case has its specific patterns, which indicates that a taxon-specific approach is needed for species detection and delineation. systematically using a multidisciplinary approach could therefore result in an enormous increase of the overall documented species richness. correct insight in the number of species within a certain taxon is urgently needed to address conservation issues. when the number of species is highly underestimated, the decline of species may also be far worse than initially detected (fouquet et al. 2007). therefore, using molecular techniques can be of key importance for traditional taxonomy in accelerating the pace of species detection and description. acknowledgements we would like to thank koen de gelas for his help in the lab as well as james maclaine (nhm), patrice pruvost (mnhn) and barbara brown (amnh) for the loan of type specimens under their care. we european journal of taxonomy 310: 1–32 (2017) 18 are grateful to emmanuel vreven (rmca) for providing literature-based compilations of the species of enteromius, and for sampling the “réserve de faunes à okapi”, epulu region and in the ituri river; these expeditions were funded by the ‘leopold iii-fonds voor natuuronderzoek en natuurbehoud’, ‘directorate-general for development cooperation and humanitarian aid’ and the ‘stichting tot bevordering van het wetenschappelijk onderzoek in afrika’. the expedition to the kisangani area in 2013 was funded by the belspo science for a sustainable development (ssd) project entitled ‘congo basin: from carbon to fishes’ (cobafish). this study is part of the phd research of e.d., funded by a belspo action 2 programme. references agnèse j.f., berrebi p., lévêque c. & guégan j.f. 1990. two lineages, diploid and tetraploid, demonstrated in african species barbus (osteichthyes, cyprinidae). aquatic living resources 3: 305– 311. https://doi.org/10.1051/alr:1990031 bamba m. 2012. the ‘barbus’ species (cypriniformes, cyprinidae) from côte d’ivoire: a systematic revision of some west african species complexes. phd thesis, ku leuven, belgium. bamba m., vreven e.j. & snoeks j. 2011. description of barbus teugelsi sp. nov. 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members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands. https://doi.org/10.1016/j.ympev.2015.01.014 https://doi.org/10.1016/j.ympev.2015.01.014 european journal of taxonomy 310: 1–32 (2017) 22 a g en et ic li ne ag e l ab el c ol le ct io n l oc at io n sl (m m ) e . c f. m io le pi s ‘k is an ga ni re gi on ’ 1 _1 03 23 b 009 -p -0 09 6 r iv . l om am i, le ft b an k, d ja bi r, si te 3 1 (0 0° 28 ′3 0. 4″ n , 2 4° 09 ′4 7. 8″ e ) 52 .8 5 ‘k is an ga ni re gi on ’ 1 _2 29 12 b 009 -p -0 08 800 89 r iv . l ob ay e, ri gh t b an k, d ja bi r, si te 6 (0 0° 29 ′2 8. 1″ n , 2 4° 10 ′4 2. 8″ e ) 46 .4 0 ‘k is an ga ni re gi on ’ 1 _3 29 13 b 009 -p -0 08 800 89 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79 6 r iv . l él o, c lo se t o th e ag ri cu ltu ra l ar ea o f th e ce nt re o f e pu lu , tr ib ut ar y of t he e pu lu , ba ss in i tu ri (r fo ), in p ri m ar y fo re st 75 .8 0 e pu lu 1 _1 9 / a 902 9p07 80 -0 79 6 r iv . l él o, c lo se t o th e ag ri cu ltu ra l ar ea o f th e ce nt re o f e pu lu , tr ib ut ar y of t he e pu lu , ba ss in i tu ri (r fo ), in p ri m ar y fo re st 72 .8 0 in ki si _1 61 1 a 607 -p -0 48 604 89 r iv . n gu fu , n ea r t he b ri dg e of th e vi lla ge l ua ng u, k av ua ya , t ri bu ta ry o f t he in ki si , l ow er c on go 98 .7 5 in ki si _2 61 2 a 607 -p -0 48 604 89 r iv . n gu fu , n ea r t he b ri dg e of th e vi lla ge l ua ng u, k av ua ya , t ri bu ta ry o f t he in ki si , l ow er c on go 83 .8 0 in ki si _3 61 3 a 607 -p -0 48 604 89 r iv . n gu fu , n ea r t he b ri dg e of th e vi lla ge l ua ng u, k av ua ya , t ri bu ta ry o f t he in ki si , l ow er c on go 80 .7 0 l ua pu la 1 _1 15 25 a 821 -p -0 37 403 76 r iv . m al au sh i, ri ve rs id e 47 .2 5 l ua pu la 1 _2 15 31 a 821 -p -0 38 5 r iv . m al au sh i, br id ge 69 .5 5 l ua pu la 1 _3 16 65 a 821 -p -0 42 9 c ha nt et e, ri v. m ul em bo 11 0. 95 l uk i 1 _1 54 5 a 731 -p -0 91 209 19 r iv . l uk i, +/ 2 .5 k m o f k iw él éw él é 48 .5 5 l uk i 1 _2 94 3 a 731 -p -0 91 009 11 r iv . l uk i, ne ar th e br id ge o f t he b io sp he re re se rv e of l uk i 68 .6 5 l uk i 1 _3 10 62 a 820 -p -1 38 4 r iv . l uk i, ne ar th e br id ge o f t he v ill ag e k iz a 38 .1 0 l ua pu la 2 _1 15 24 a 821 -p -0 37 403 76 r iv . m al au sh i, ri ve rs id e 43 .8 0 l ua pu la 3 _1 16 35 a 821 -p -0 47 2 po nt oo n, ri v. k as an ka , l ef t b an k 56 .0 0 l ua pu la 3 _2 16 60 a 821 -p -0 53 0 k at w a, ri v. k as an ka 48 .2 5 it ur i 2 _1 / b 111 -g -0 05 43 r iv . e la ke , v ill ag e e la ke u ps tr ea m o f t he c on flu en ce w ith it ur i p n 57 .0 5 european journal of taxonomy 310: 1–32 (2017) 24 l uk i 2 _1 04 67 a 607 -p -0 69 106 92 r iv . l uk i, up st re am o f t he b ri dg e in th e l uk i f or es t r es er ve , l ow er c on go 66 .8 0 l uk i 2 _2 04 68 a 607 -p -0 68 9 r iv . l uk i, up st re am o f t he b ri dg e in th e l uk i f or es t r es er ve , l ow er c on go 77 .1 5 l uk i 2 _3 10 37 a 820 -p -1 36 8 r iv . l uk i, co nfl ue nc e w ith th e l ov o, s ev er al k m s do w ns tr ea m o f k iw él éw él é 51 .9 5 l uk i 2 _4 10 65 a 820 -p -1 36 9 r iv . l uk i, up st re am o f t he b ri dg e at th e vi ll. k iz a 70 .0 0 l uk i 2 _5 / a 820 -p -1 37 113 72 r iv . l uk i, on th e ro ut e of th e ca m p up st re am o f t he b ri dg e 71 .9 5 l uk i 2 _6 / a 820 -p -1 37 113 72 r iv . l uk i, on th e ro ut e of th e ca m p up st re am o f t he b ri dg e 67 .0 5 l uk i 2 _7 / a 820 -p -1 37 0 r iv . l uk i, up st re am o f t he b ri dg e at th e vi ll. k iz a 53 .8 0 e . c f. br az za i k is an ga ni re gi on ’ 2 _1 12 31 7 b 330 v ill . b us uk ul u, ri v. l ob ilo , r ig ht b an k up st re am o f t he k ib an gu is t m is si on 76 .9 5 k is an ga ni re gi on ’ 2 _2 12 38 6 b 330 v ill . b us uk ul u, ri v. l ob ilo , l ef t b an k up st re am o f t he k ib an gu is t m is si on 82 .8 0 k is an ga ni re gi on ’ 2 _3 12 41 1 b 330 v ill . b us uk ul u, ri v. l ob ilo , l ef t b an k up st re am o f t he k ib an gu is t m is si on 68 .1 0 k is an ga ni re gi on ’ 2 _4 12 43 0 b 330 v ill . b us uk ul u, ri v. l ob ilo , r ig ht b an k up st re am o f t he b ri dg e 66 .2 0 it ur i 3 _1 31 36 a 929 v ill . m ol ok ay , r iv . e pe ne , d ow ns tr ea m o f t he b ri dg e 53 .9 5 it ur i 3 _2 31 37 a 929 v ill . m ol ok ay , r iv . e pe ne , d ow ns tr ea m o f t he b ri dg e 48 .5 0 it ur i 3 _3 31 55 a 929 v ill . m ol ok ay , r iv . e pe ne , d ow ns tr ea m o f t he b ri dg e 48 .9 0 ‘k is an ga ni re gi on ’ 3 _1 89 69 b 330 v ill . d ja tiz i, ri v. l ob ay e, u ps tr ea m o f t he fi sh in g ca m p l ik un gu 55 .0 0 ‘k is an ga ni re gi on ’ 3 _2 89 70 b 330 v ill . d ja tiz i, ri v. l ob ay e, u ps tr ea m o f t he fi sh in g ca m p l ik un gu 53 .0 0 ‘k is an ga ni re gi on ’ 3 _3 12 41 7 b 330 v ill . b us uk ul u, ri v. l ob ilo , r ig ht b an k up st re am o f t he k ib an gu is t m is si on 49 .8 0 ‘k is an ga ni re gi on ’ 3 _4 12 41 8 b 330 v ill . b us uk ul u, ri v. l ob ilo , r ig ht b an k up st re am o f t he k ib an gu is t m is si on 45 .2 0 ‘k is an ga ni re gi on ’ 3 _5 12 41 9 b 330 v ill . b us uk ul u, ri v. l ob ilo , r ig ht b an k up st re am o f t he k ib an gu is t m is si on 48 .0 0 ‘k is an ga ni re gi on ’ 3 _6 12 42 0 b 330 v ill . b us uk ul u, ri v. l ob ilo , r ig ht b an k up st re am o f t he k ib an gu is t m is si on 44 .7 5 ‘k is an ga ni re gi on ’ 3 _7 12 47 2 b 330 r iv . c on go , d ow ns tr ea m o f t he c on flu en ce w ith th e l ob ilo , r ig ht b an k di ag on al ly o f l ot ok ila 49 .3 5 ‘k is an ga ni re gi on ’ 3 _8 12 47 7 b 330 r iv . c on go , d ow ns tr ea m o f t he c on flu en ce w ith th e l ob ilo , r ig ht b an k di ag on al ly o f l ot ok ila 47 .5 5 e . c f. pe lle gr in i it ur i/e pu lu _1 36 81 a 902 9p08 84 -0 88 5 r iv . a ko ko ra , w at er fa lls o f k ou ko u, tr ib ut ar y of th e e pu lu , b as si n it ur i ( r fo ) 55 .6 0 it ur i/e pu lu _2 36 82 a 902 9p08 84 -0 88 5 r iv . a ko ko ra , w at er fa lls o f k ou ko u, tr ib ut ar y of th e e pu lu , b as si n it ur i ( r fo ) 57 .8 5 it ur i/e pu lu _3 / a 902 9p08 86 -0 88 9 r iv . a ko ko ra , w at er fa lls o f k ou ko u, tr ib ut ar y of th e e pu lu , b as si n it ur i ( r fo ) 58 .7 5 it ur i/e pu lu _4 / a 902 9p08 86 -0 88 9 r iv . a ko ko ra , w at er fa lls o f k ou ko u, tr ib ut ar y of th e e pu lu , b as si n it ur i ( r fo ) 55 .5 0 it ur i/e pu lu _5 / a 902 9p08 86 -0 88 9 r iv . a ko ko ra , w at er fa lls o f k ou ko u, tr ib ut ar y of th e e pu lu , b as si n it ur i ( r fo ) 62 .2 0 it ur i/e pu lu _6 / a 902 9p08 86 -0 88 9 r iv . a ko ko ra , w at er fa lls o f k ou ko u, tr ib ut ar y of th e e pu lu , b as si n it ur i ( r fo ) 54 .6 5 it ur i/e pu lu _7 31 63 a 902 9p08 82 -0 88 3 r iv . n ep us e, s m al l t ri bu ta ry o f t he e pu lu in e pu lu , b as si n it ur i ( r fo ) 49 .7 0 it ur i/e pu lu _8 31 64 a 902 9p08 82 -0 88 3 r iv . n ep us e, s m al l t ri bu ta ry o f t he e pu lu in e pu lu , b as si n it ur i ( r fo ) 50 .7 5 van ginneken m. et al., unexpected cryptic diversity in enteromius 25 it ur i/e pu lu _9 / a 902 9p08 81 r iv . n ep us e, s m al l t ri bu ta ry o f t he e pu lu in e pu lu , b as si n it ur i ( r fo ) 40 .7 0 it ur i/e pu lu _1 0 32 14 a 929 -p -0 96 609 68 r iv . n em bo ng o, n ea r t he b ri dg e on th e ro ut e to e pu lu , t ri bu ta ry o f t he e pu lu , b as si n it ur i ( r fo ) 57 .9 0 it ur i/e pu lu _1 1 32 15 a 929 -p -0 96 609 68 r iv . n em bo ng o, n ea r t he b ri dg e on th e ro ut e to e pu lu , t ri bu ta ry o f t he e pu lu , b as si n it ur i ( r fo ) 59 .5 0 it ur i/e pu lu _1 2 32 16 a 929 -p -0 96 609 68 r iv . n em bo ng o, n ea r t he b ri dg e on th e ro ut e to e pu lu , t ri bu ta ry o f t he e pu lu , b as si n it ur i ( r fo ) 54 .8 0 it ur i/e pu lu _1 3 / a 902 9p09 63 -0 96 5 r iv . n em bo ng o, n ea r t he b ri dg e on th e ro ut e to e pu lu , t ri bu ta ry o f t he e pu lu , b as si n it ur i ( r fo ) 51 .0 0 it ur i/e pu lu _1 4 / a 902 9p09 63 -0 96 5 r iv . n em bo ng o, n ea r t he b ri dg e on th e ro ut e to e pu lu , t ri bu ta ry o f t he e pu lu , b as si n it ur i ( r fo ) 43 .9 5 it ur i/e pu lu _1 5 / a 902 9p09 63 -0 96 5 r iv . n em bo ng o, n ea r t he b ri dg e on th e ro ut e to e pu lu , t ri bu ta ry o f t he e pu lu , b as si n it ur i ( r fo ) 47 .0 5 it ur i/e pu lu _1 6 / a 902 9p08 90 c on flu en ce n du ye -b ia sa , d ow ns tre am o f th e br id ge o f n du ye , t rib ut ar y of th e ep ul u, b as si n itu ri (r fo ) 55 .8 5 it ur i/e pu lu _1 7 36 03 a 929 -p -0 89 1 r iv . l ol w a in l ol w a, u ps tr ea m o f t he b ri dg e, tr ib ut ar y of th e e pu lu 63 .4 5 it ur i/e pu lu _1 8 36 13 a 929 -p -0 89 208 93 r iv . l ol w a in l ol w a, u ps tr ea m o f t he b ri dg e, tr ib ut ar y of th e e pu lu 81 .3 5 it ur i/e pu lu _1 9 36 14 a 929 -p -0 89 208 93 r iv . l ol w a in l ol w a, u ps tr ea m o f t he b ri dg e, tr ib ut ar y of th e e pu lu 73 .2 0 it ur i/e pu lu _2 0 / a 902 9p08 94 -0 90 0 r iv . l ol w a in l ol w a, u ps tr ea m o f t he b ri dg e, tr ib ut ar y of th e e pu lu 71 .9 5 it ur i/e pu lu _2 1 / a 902 9p08 94 -0 90 0 r iv . l ol w a in l ol w a, u ps tr ea m o f t he b ri dg e, tr ib ut ar y of th e e pu lu 55 .5 0 it ur i/e pu lu _2 2 / a 902 9p08 94 -0 90 0 r iv . l ol w a in l ol w a, u ps tr ea m o f t he b ri dg e, tr ib ut ar y of th e e pu lu 69 .7 0 it ur i/e pu lu _2 3 / a 902 9p08 94 -0 90 0 r iv . l ol w a in l ol w a, u ps tr ea m o f t he b ri dg e, tr ib ut ar y of th e e pu lu 70 .1 0 it ur i/e pu lu _2 4 / a 902 9p08 94 -0 90 0 r iv . l ol w a in l ol w a, u ps tr ea m o f t he b ri dg e, tr ib ut ar y of th e e pu lu 68 .0 0 it ur i/e pu lu _2 5 / a 902 9p08 94 -0 90 0 r iv . l ol w a in l ol w a, u ps tr ea m o f t he b ri dg e, tr ib ut ar y of th e e pu lu 62 .5 0 it ur i/e pu lu _2 6 / a 902 9p08 94 -0 90 0 r iv . l ol w a in l ol w a, u ps tr ea m o f t he b ri dg e, tr ib ut ar y of th e e pu lu 69 .5 0 e . c f. at ro m ac ul at us it ur i 5 _1 30 94 a 902 9p09 03 -0 90 4 r iv . a ng ol om u in b ab ek e, c on flu en ce w ith th e is ay e, tr ib ut ar y of th e it ur i ( r fo ) 51 .5 5 it ur i 5 _2 30 95 a 902 9p09 03 -0 90 4 r iv . a ng ol om u in b ab ek e, c on flu en ce w ith th e is ay e, tr ib ut ar y of th e it ur i ( r fo ) 45 .0 0 it ur i 5 _3 39 18 a 902 9p09 37 -0 94 0 r iv . m ak ul um ba in a va ku bi , s m al l t ri bu ta ry o f t he it ur i, up st re am o f t he b ri dg e 44 .8 0 it ur i 5 _4 39 19 a 902 9p09 37 -0 94 0 r iv . m ak ul um ba in a va ku bi , s m al l t ri bu ta ry o f t he it ur i, up st re am o f t he b ri dg e 36 .9 5 it ur i 5 _5 39 20 a 902 9p09 37 -0 94 0 r iv . m ak ul um ba in a va ku bi , s m al l t ri bu ta ry o f t he it ur i, up st re am o f t he b ri dg e 37 .6 5 it ur i 5 _6 39 21 a 902 9p09 37 -0 94 0 r iv . m ak ul um ba in a va ku bi , s m al l t ri bu ta ry o f t he it ur i, up st re am o f t he b ri dg e 37 .7 0 it ur i 6 _1 37 51 a 902 9p09 32 r iv . i tu ri , a cr os s fr om th e vi ll. p en ge , d ow ns tr ea m o f t he p en ge ra pi ds (r fo ) 48 .6 0 it ur i 6 _2 38 87 a 902 9p09 34 -0 93 5 r iv . i tu ri in a va ku bi , u ps tr ea m o f t he b ri dg e an d th e la st ra pi ds 48 .4 5 it ur i 6 _3 38 88 a 902 9p09 34 -0 93 5 r iv . i tu ri in a va ku bi , u ps tr ea m o f t he b ri dg e an d th e la st ra pi ds 47 .5 0 it ur i 6 _4 / a 902 9p09 36 r iv . i tu ri in a va ku bi , u ps tr ea m o f t he b ri dg e an d th e la st ra pi ds 50 .6 0 it ur i/’ k is an ga ni re gi on ’_ 1 37 76 a 902 9p09 33 r iv . i tu ri , u ps tr ea m o f t he p en ge ra pi ds (r fo ) 55 .6 5 it ur i/’ k is an ga ni re gi on ’_ 2 73 25 b 002 1p08 83 r iv . l ob ay e, ri gh t b an k, b on ba nd ji ii s ite 1 14 (0 0° 11 ′3 3. 6″ n , 2 5° 31 ′5 1. 1″ e ) 44 .3 0 european journal of taxonomy 310: 1–32 (2017) 26 e pu lu 2 _1 30 37 / r iv . e pu lu in e pu lu , r ig ht b an k do w ns tr ea m o f t he b ri dg e 47 .1 5 e pu lu 2 _2 30 38 / r iv . e pu lu in e pu lu , r ig ht b an k do w ns tr ea m o f t he b ri dg e 48 .5 0 e pu lu 2 _3 30 55 a 929 -p -0 90 109 02 r iv . m ay i m bi li, s m al l t ri bu ta ry o f t he e pu lu , t ri bu ta ry o f t he it ur i ( r fo ) 35 .7 5 e pu lu 2 _4 30 56 a 929 -p -0 90 109 02 r iv . m ay i m bi li, s m al l t ri bu ta ry o f t he e pu lu , t ri bu ta ry o f t he it ur i ( r fo ) 34 .0 0 e pu lu 2 _5 32 05 / r iv . e pu lu , u ps tr ea m o f t he b ri dg e an d ac ro ss fr om c hi m pa nz ee is la nd 41 .0 0 e pu lu 2 _6 32 06 / r iv . e pu lu , u ps tr ea m o f t he b ri dg e an d ac ro ss fr om c hi m pa nz ee is la nd 44 .2 0 e pu lu 2 _7 35 55 a 902 9p09 08 r iv . k au be li, tr ib ut ar y of th e e pu lu in b an di se nd e, b as si n it ur i 40 .4 0 e pu lu 2 _8 / a 902 9p09 06 -0 90 7 r iv . k au be li, tr ib ut ar y of th e e pu lu in b an di se nd e, b as si n it ur i 28 .2 5 e pu lu 2 _9 36 67 a 902 9p09 09 -0 91 0 r iv . n du ye i n n du ye , up st re am o f th e la st b ri dg e be hi nd t he p ol ic e ca m p, t ri bu ta ry o f th e e pu lu , ba ss in it ur i ( r fo ) 51 .4 5 e pu lu 2 _1 0 36 68 a 902 9p09 09 -0 91 0 r iv . n du ye i n n du ye , up st re am o f th e la st b ri dg e be hi nd t he p ol ic e ca m p, t ri bu ta ry o f th e e pu lu , ba ss in it ur i ( r fo ) 51 .5 0 e pu lu 2 _1 1 / a 902 9p09 11 -0 91 3 r iv . n du ye i n n du ye , up st re am o f th e la st b ri dg e be hi nd t he p ol ic e ca m p, t ri bu ta ry o f th e e pu lu , ba ss in it ur i ( r fo ) 53 .6 0 e pu lu 2 _1 2 / a 902 9p09 11 -0 91 3 r iv . n du ye i n n du ye , up st re am o f th e la st b ri dg e be hi nd t he p ol ic e ca m p, t ri bu ta ry o f th e e pu lu , ba ss in it ur i ( r fo ) 50 .8 0 e pu lu 2 _1 3 / a 902 9p09 11 -0 91 3 r iv . n du ye i n n du ye , up st re am o f th e la st b ri dg e be hi nd t he p ol ic e ca m p, t ri bu ta ry o f th e e pu lu , ba ss in it ur i ( r fo ) 50 .7 0 e pu lu 2 _1 4 36 71 / r iv . u et ef e in n du ye , r ig ht b an k 36 .2 5 e pu lu 2 _1 5 36 85 a 902 9p09 19 -0 92 0 r iv . a ko ko ra , w at er fa lls o f k ou ko u, tr ib ut ar y of th e e pu lu , b as si n it ur i ( r fo ) 54 .8 5 e pu lu 2 _1 6 36 86 a 902 9p09 19 -0 92 0 r iv . a ko ko ra , w at er fa lls o f k ou ko u, tr ib ut ar y of th e e pu lu , b as si n it ur i ( r fo ) 55 .7 5 it ur i 8 _1 33 17 a 929 -p -0 95 409 55 r iv . b an go in b an go , t ri bu ta ry o f t he it ur i 51 .6 0 it ur i 8 _2 33 18 a 929 -p -0 95 409 55 r iv . b an go in b an go , t ri bu ta ry o f t he it ur i 45 .9 5 it ur i 8 _3 / a 902 9p09 72 -0 97 3 r iv . b an go in b an go , u ps tr ea m o f t he b ri dg e al on g th e m in in g op er at io n, b as si n it ur i 40 .5 5 it ur i 8 _4 / a 902 9p09 72 -0 97 3 r iv . b an go in b an go , u ps tr ea m o f t he b ri dg e al on g th e m in in g op er at io n, b as si n it ur i 45 .7 0 it ur i 8 _5 / a 902 9p09 56 -0 96 2 r iv . b an go in b an go , t ri bu ta ry o f t he it ur i 46 .0 5 it ur i 8 _6 / a 902 9p09 56 -0 96 2 r iv . b an go in b an go , t ri bu ta ry o f t he it ur i 47 .1 5 it ur i 8 _7 / a 902 9p09 56 -0 96 2 r iv . b an go in b an go , t ri bu ta ry o f t he it ur i 51 .3 5 it ur i 8 _8 / a 902 9p09 56 -0 96 2 r iv . b an go in b an go , t ri bu ta ry o f t he it ur i 49 .9 0 it ur i 8 _9 / a 902 9p09 56 -0 96 2 r iv . b an go in b an go , t ri bu ta ry o f t he it ur i 45 .0 0 it ur i 8 _1 0 / a 902 9p09 56 -0 96 2 r iv . b an go in b an go , t ri bu ta ry o f t he it ur i 48 .6 0 it ur i 8 _1 1 / a 902 9p09 56 -0 96 2 r iv . b an go in b an go , t ri bu ta ry o f t he it ur i 48 .8 5 it ur i 8 _1 2 33 47 a 929 r iv . b an go in b an go , u ps tr ea m o f t he b ri dg e al on g th e m in in g op er at io n, b as si n it ur i 45 .7 0 van ginneken m. et al., unexpected cryptic diversity in enteromius 27 b s pe ci es l ab el c ol le ct io n l oc at io n sl (m m ) e . m io le pi s_ 1 ho lo ty pe r iv . y em be in b an zy vi lle (u ba ng i r iv er s ys te m ) 63 .9 e . h ol ot ae ni a_ 1 sy nt yp e 18 82 .2 .1 3. 7 r iv . o go w e in g ab oo n 65 .4 0 e . h ol ot ae ni a_ 2 sy nt yp e 18 88 .1 2. 13 .3 1 an d 18 89 .3 .2 .1 0 r iv . o go w e in g ab oo n 84 .9 5 e . h ol ot ae ni a_ 3 sy nt yp e 18 88 .1 2. 13 .3 1 an d 18 89 .3 .2 .1 0 r iv . o go w e in g ab oo n 88 .5 0 e . h ol ot ae ni a_ 4 sy nt yp e 18 69 .5 .3 .2 324 r iv . o go w e in g ab oo n 58 .6 0 e . h ol ot ae ni a_ 5 sy nt yp e 18 69 .5 .3 .2 324 r iv . o go w e in g ab oo n 64 .5 5 e . h ol ot ae ni a_ 6 sy nt yp e 18 96 .5 .5 .1 07 .1 12 r iv . o go w e in g ab oo n 66 .9 0 e . h ol ot ae ni a_ 7 sy nt yp e 18 96 .5 .5 .1 07 .1 12 r iv . o go w e in g ab oo n 79 .7 5 e . h ol ot ae ni a_ 8 sy nt yp e 18 96 .5 .5 .1 07 .1 12 r iv . o go w e in g ab oo n 62 .9 5 e . h ol ot ae ni a_ 9 sy nt yp e 18 96 .5 .5 .1 07 .1 12 r iv . o go w e in g ab oo n 67 .5 0 e . h ol ot ae ni a_ 10 sy nt yp e 18 96 .5 .5 .1 07 .1 12 r iv . o go w e in g ab oo n 66 .0 0 e . h ol ot ae ni a_ 11 sy nt yp e 18 98 .1 2. 28 .8 -1 1 r iv . o go w e in g ab oo n 67 .3 0 e . h ol ot ae ni a_ 12 sy nt yp e 18 98 .1 2. 28 .8 -1 1 r iv . o go w e in g ab oo n 56 .0 0 e . h ol ot ae ni a_ 13 sy nt yp e 18 98 .1 2. 28 .8 -1 1 r iv . o go w e in g ab oo n 64 .0 5 e . h ol ot ae ni a_ 14 sy nt yp e 18 98 .1 2. 28 .8 -1 1 r iv . o go w e in g ab oo n 75 .2 5 e . h ol ot ae ni a_ 15 sy nt yp e 19 00 .2 .1 7. 13 2 r iv . o go w e in g ab oo n 54 .4 0 e . e ut ae ni a_ 1 le ct ot yp e 18 64 .7 .1 3. 52 -5 6 r iv . h ui la (m os sa m ed es ) i n a ng ol a 70 .7 0 e . e ut ae ni a_ 2 pa ra le ct ot yp e 18 64 .7 .1 3. 53 -5 6 r iv . h ui la (m os sa m ed es ) i n a ng ol a 56 .7 5 e . e ut ae ni a_ 3 pa ra le ct ot yp e 18 64 .7 .1 3. 53 -5 6 r iv . h ui la (m os sa m ed es ) i n a ng ol a 66 .3 5 e . e ut ae ni a_ 4 pa ra le ct ot yp e 18 64 .7 .1 3. 53 -5 6 r iv . h ui la (m os sa m ed es ) i n a ng ol a 67 .7 0 e . e ut ae ni a_ 5 pa ra le ct ot yp e 18 64 .7 .1 3. 53 -5 6 r iv . h ui la (m os sa m ed es ) i n a ng ol a 48 .2 5 e . e ut ae ni a_ 6 pa ra le ct ot yp e 18 64 .7 .1 3. 53 -5 6 r iv . h ui la (m os sa m ed es ) i n a ng ol a 54 .8 0 e . k er st en ii_ 1 le ct ot yp e z m b 1 6 84 8 c oa st o pp os ite to z an zi ba r ( i.e . p ro ba bl y b ag am ay o) 64 .3 1 e . b ra zz ai _1 ho lo ty pe m n h n 2 8 6 40 4 r iv . s an gh a in m ob ak a 78 .2 e . t sh op oe ns is _1 ho lo ty pe 88 -2 5p15 87 k is an ga ni , r iv . t sh op o, u ps tr ea m o f t he h yd ro el ec tr ic d am , z aï re 64 .3 5 e . t sh op oe ns is _2 pa ra ty pe 87 -4 2p10 98 k is an ga ni , r iv . t sh op o, d ow ns tr ea m o f t he h yd ro el ec tr ic d am , z aï re 74 .7 0 e . t sh op oe ns is _3 pa ra ty pe 88 -2 5p15 90 k is an ga ni , r iv . t sh op o, d ow ns tr ea m o f t he h yd ro el ec tr ic d am , z aï re 52 .5 5 e . t sh op oe ns is _4 pa ra ty pe 88 -2 5p15 91 -1 59 2 k is an ga ni , r iv . t sh op o, d ow ns tr ea m o f t he h yd ro el ec tr ic d am , z aï re 68 .9 5 e . t sh op oe ns is _5 pa ra ty pe 88 -2 5p15 94 -1 59 5 k is an ga ni , r iv . t sh op o, d ow ns tr ea m o f t he h yd ro el ec tr ic d am , z aï re 53 .2 0 e . t sh op oe ns is _6 pa ra ty pe 88 -2 5p16 05 k is an ga ni , r iv . t sh op o, d ow ns tr ea m o f t he h yd ro el ec tr ic d am , z aï re 48 .7 5 european journal of taxonomy 310: 1–32 (2017) 28 e . p el le gr in i_ 1 le ct ot yp e m n h n 3 5 80 l ak e k iv u 52 .9 0 e . p el le gr in i_ 2 pa ra le ct ot yp e m n h n 3 5 81 l ak e k iv u 37 .9 5 e . p el le gr in i_ 3 pa ra le ct ot yp e m n h n 3 5 82 l ak e k iv u 36 .1 5 e . p el le gr in i_ 4 pa ra le ct ot yp e m n h n 1 99 6 26 5 l ak e k iv u 43 .2 5 e . a tr om ac ul at us _1 pa ra ty pe a m n h 3 5 84 5 r iv . y ak ul uk u, l ow er c on go 41 .1 4 e . a tr om ac ul at us _2 pa ra ty pe a m n h 6 21 4 r iv . y ak ul uk u, l ow er c on go 40 .6 5 1 m us eu m fü r n at ur ku nd e, b er lin , g er m an y. 2 m us éu m n at io na l d ’h is to ir e na tu re lle , p ar is , f ra nc e. 3 a m er ic an m us eu m o f n at ur al h is to ry , n ew y or k, u sa . van ginneken m. et al., unexpected cryptic diversity in enteromius 29 appendix 2. pc loadings of the pca in fig. 3. most important loadings indicated in bold. pc1 pc2 pc3 logsl 0.229 0.158 -0.033 loghl 0.236 0.018 -0.037 loged 0.271 -0.626 -0.661 logsnl 0.279 -0.060 0.230 logpropd 0.244 -0.062 -0.003 logiow 0.232 0.163 -0.091 logprdd 0.229 0.210 -0.148 logpodd 0.238 0.289 0.068 logdofbl 0.242 -0.310 0.395 logdofl 0.216 -0.310 0.528 logprpecd 0.224 0.030 -0.005 logprpeld 0.233 0.074 -0.011 logprad 0.236 0.061 -0.096 logpoad 0.223 0.460 -0.160 logbd 0.275 0.060 0.005 logmxcpd 0.260 -0.057 0.045 logmncpd 0.247 0.024 0.043 appendix 3. pc loadings of the pca in fig. 4. most important loadings indicated in bold. pc1 pc2 pc3 ll sc 0.242 0.538 0.195 d-l sc 0.440 -0.005 -0.196 pd sc 0.311 0.376 -0.047 cp sc 0.288 0.327 -0.043 lp sc 0.371 -0.339 0.376 l-b sc 0.441 -0.213 0.397 dfr -0.258 -0.285 0.213 afr -0.056 0.090 -0.513 pecfr -0.248 0.312 0.224 pelfr -0.325 0.341 0.513 european journal of taxonomy 310: 1–32 (2017) 30 appendix 4. pc loadings of the pca in fig. 5. most important loadings indicated in bold. pc1 pc2 pc3 ll sc 0.940 -0.040 -0.023 d-l sc 0.139 -0.097 -0.084 pd sc 0.136 -0.226 0.855 cp sc 0.192 -0.150 -0.410 lp sc 0.044 -0.033 -0.030 l-b sc 0.140 -0.174 -0.117 dfr <0.001 <0.001 <-0.001 afr 0.029 0.238 0.279 pecfr 0.142 0.910 0.039 pelfr 0.000 0.000 0.000 appendix 5. pc loadings of the pca in fig. 6. most important loadings indicated in bold. pc1 pc2 pc3 llsc 0.480 0.251 -0.120 d-lsc 0.395 0.189 0.404 pdsc 0.416 0.226 -0.519 cpsc 0.512 0.003 -0.020 l-bsc 0.317 -0.283 0.604 dfr 0.221 -0.584 0.005 afr 0.071 0.274 -0.040 pecfr -0.154 0.595 0.430 van ginneken m. et al., unexpected cryptic diversity in enteromius 31 appendix 6. pc loadings of the pca in fig. 7. most important loadings indicated in bold. pc1 pc2 pc3 ll sc 0.740 -0.164 0.202 d-l sc <-0.001 <-0.001 <-0.001 pd sc -0.011 -0.097 0.402 cp sc -0.105 0.552 0.767 lp sc 0.099 -0.403 0.208 l-b sc 0.132 -0.501 0.284 dfr 0.031 -0.054 -0.034 afr -0.071 0.050 0.007 pecfr 0.457 0.455 -0.290 pelfr 0.447 0.182 0.026 appendix 7. pc loadings of the pca in fig. 8. most important loadings indicated in bold. pc1 pc2 pc3 ll sc 0.971 -0.119 -0.142 d-l sc 0.144 0.223 -0.096 pd sc 0.052 0.219 -0.187 cp sc 0.028 0.902 -0.222 lp sc <-0.001 <-0.001 <0.001 l-b sc -0.015 0.053 -0.050 dfr <-0.001 <0.001 <-0.001 afr <-0.001 <-0.001 <0.001 pecfr 0.179 0.267 0.928 pelfr 0.006 0.001 0.152 european journal of taxonomy 310: 1–32 (2017) 32 appendix 8. pc loadings of the pca in fig. 9. most important loadings indicated in bold. pc1 pc2 pc3 ll sc 0.971 -0.073 -0.141 d-l sc 0.175 0.047 0.144 pd sc 0.051 -0.322 -0.092 cp sc 0.118 -0.085 0.953 lp sc <0.001 <0.001 <-0.001 l-b sc <-0.001 <-0.001 <0.001 dfr <-0.001 <-0.001 <0.001 afr 0.000 0.000 0.000 pecfr 0.092 0.904 0.090 pelfr 0.024 0.254 -0.185 improving nomenclatural consistency: a decade of experience in the world register of marine species tammy horton 1,*, serge gofas 2, andreas kroh 3, gary c.b. poore 4, geoffrey read 5, gary rosenberg 6, sabine stöhr 7, nicolas bailly 8, nicole boury-esnault 9, simone n. brandão 10, mark j. costello 11, wim decock 12, stefanie dekeyzer 13, francisco hernandez 14, jan mees 15, gustav paulay 16, leen vandepitte 17, bart vanhoorne 18 & sofie vranken 19 1 national oceanography centre, european way, southampton so14 3zh, uk. 2 departamento de biología animal, university of málaga, faculty of sciences, campus de teatinos, 29071 málaga, spain. 3 natural history museum vienna, department of geology and palaeontology, burgring 7, 1010 vienna, austria. 4 museums victoria, gpo box 666, melbourne, vic. 3001, australia. 5 national institute of water & atmosphere (niwa), 301 evans bay parade, greta point, wellington, new zealand. 6 academy of natural sciences, drexel university, 1900 benjamin franklin parkway, philadelphia, pa 19103-1195, usa. 7 swedish museum of natural history, dept. of zoology, box 50007, se-10405 stockholm, sweden. 8 fishbase information and research group (fin), c/o irri, khush hall, los baños, laguna 4031, philippines. 8 hellenic centre for marine research, gournes pediados, p.c. 71500, gouves, heraklion, crete, greece. 9 institut méditerranéen de biodiversité et d’écologie marine et continentale, umr 7263 cnrs, ird, aix-marseille university, station marine d’endoume, rue batterie des lions, marseille 13007, france. 10 programa de pós-graduação em sistemática e evolução, laboratório de geologia e geofísica marinha e monitoramento ambiental, universidade federal do rio grande do norte , campus universitário lagoa nova, cep 59072-970 – natal, rn – brazil. 11 institute of marine science, university of auckland, p. bag 92019, auckland 1142, new zealand. 12,13,14,15,17,18,19 flanders marine institute (vliz) – innovocean site – wandelaarkaai 7, 8400, oostende, belgium. 16 florida museum of natural history, gainesville, florida, 32611-2710, usa. * corresponding author: tammy.horton@noc.ac.uk 2 email: sgofas@uma.es 3 email: andreas.kroh@nhm-wien.ac.at 4 email: gpoore@museum.vic.gov.au 5 email: geoffrey.read@niwa.co.nz 6 email: rosenberg.ansp@drexel.edu 7 email: sabine.stohr@nrm.se european journal of taxonomy 389: 1–24 issn 2118-9773 https://doi.org/10.5852/ejt.2017.389 www.europeanjournaloftaxonomy.eu 2017 · horton t. et al. this work is licensed under a creative commons attribution 3.0 license. o p i n i o n p a p e r urn:lsid:zoobank.org:pub:6b09ffac-ea15-4e4f-8042-94c85bc85118 1 mailto:tammy.horton%40noc.ac.uk?subject= mailto:sgofas%40uma.es?subject= mailto:andreas.kroh%40nhm-wien.ac.at?subject= mailto:gpoore%40museum.vic.gov.au?subject= mailto:geoffrey.read%40niwa.co.nz?subject= mailto:rosenberg.ansp%40drexel.edu?subject= mailto:sabine.stohr%40nrm.se?subject= https://doi.org/10.5852/ejt.2017.389 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:6b09ffac-ea15-4e4f-8042-94c85bc85118 8 email: nbailly@hcmr.gr 9 email: nicole.boury-esnault@orange.fr 10 email: brandao.sn.100@gmail.com 11 email: m.costello@auckland.ac.nz 12 email: wim.decock@vliz.be 13 email: stefanie.dekeyzer@vliz.be 14 email: francisco.hernandez@vliz.be 15 email: jan.mees@vliz.be 16 email: paulay@flmnh.ufl.edu 17 email: leen.vandepitte@vliz.be 18 email: bart.vanhoorne@vliz.be 19 email: sofie.vranken@vliz.be 1 urn:lsid:zoobank.org:author:42a8bb0a-e193-4258-8857-266108dcff51 2 urn:lsid:zoobank.org:author:c7a4b3d3-ae51-4376-ba89-249cd1f1632d 3 urn:lsid:zoobank.org:author:386bd5af-0ca9-4c2e-a421-c1d3576e55a4 4 urn:lsid:zoobank.org:author:893860fe-e71b-4f69-a46a-8b89834579fc 5 urn:lsid:zoobank.org:author:d18accd9-291d-4b1a-b960-5a2f03695bdc 6 urn:lsid:zoobank.org:author:fea0bfc5-d865-4e3c-8edf-0ecce1d24411 7 urn:lsid:zoobank.org:author:412800eb-aace-4313-9810-61f89b740405 8 urn:lsid:zoobank.org:author:1da532e6-fa75-4dc1-9e29-6eecd4dfda87 9 urn:lsid:zoobank.org:author:8dac0752-82c3-4bb8-8ca5-9da491302533 10 urn:lsid:zoobank.org:author:27041e84-cc1d-456a-baee-928dfc2021c7 11 urn:lsid:zoobank.org:author:9f38cee2-3009-4368-9b4b-963c00e900f0 12 urn:lsid:zoobank.org:author:9a3bdba9-343b-4b5d-b978-edb46ad4353c 13 urn:lsid:zoobank.org:author:401d1c89-c0f8-408f-8035-0ad2e2c25dcb 14 urn:lsid:zoobank.org:author:1d813f3e-2cda-482b-a9be-d8eadfdb09d8 15 urn:lsid:zoobank.org:author:891eefda-63c2-4098-a6d9-26c5c99fc825 16 urn:lsid:zoobank.org:author:a2f155e4-7958-4e63-b36a-cab23f190a07 17 urn:lsid:zoobank.org:author:0c024f0f-c9e6-4398-94d5-b462b43e554e 18 urn:lsid:zoobank.org:author:3708ef86-f640-4ddf-87a3-2d1a1c118890 19 urn:lsid:zoobank.org:author:84c998a6-886d-4421-8645-00d303af3c24 abstract. the world register of marine species (worms) has been established for a decade. the early history of the database involved compilation of existing global and regional species registers. this aggregation, combined with changes to data types and the changing needs of worms users, has resulted in an evolution of data-entry consistency over time. with the task of aggregating the accepted species names for all marine species approaching completion, our focus has shifted to improving the consistency and quality of data held while keeping pace with the addition of > 2000 new marine species described annually. this paper defines priorities and longer-term aims that promote standardisation within and  interoperability among biodiversity databases, provides editors with further information on how to input nomenclatural data in a standardised way and clarifies for users of worms how and why names are  represented as they are. we 1) explain the categories of names included; 2) list standard reasons used to explain why a name is considered ‘unaccepted’ or ‘uncertain’; 3) present and explain the more difficult  situations encountered; 4) describe categories of sources and notes linked to a taxon; and 5) recommend how type material, type locality and environmental information should be entered. keywords. world register of marine species, worms, taxonomic databases, taxonomy, nomenclature. european journal of taxonomy 389: 1–24 (2017) 2 mailto:nbailly%40hcmr.gr?subject= mailto:nicole.boury-esnault%40orange.fr?subject= mailto:brandao.sn.100%40gmail.com?subject= mailto:m.costello%40auckland.ac.nz?subject= mailto:wim.decock%40vliz.be?subject= mailto:stefanie.dekeyzer%40vliz.be?subject= mailto:francisco.hernandez%40vliz.be?subject= mailto:jan.mees%40vliz.be?subject= mailto:paulay%40flmnh.ufl.edu?subject= mailto:leen.vandepitte%40vliz.be?subject= mailto:bart.vanhoorne%40vliz.be?subject= mailto:sofie.vranken%40vliz.be?subject= http://zoobank.org/urn:lsid:zoobank.org:author:42a8bb0a-e193-4258-8857-266108dcff51 http://zoobank.org/urn:lsid:zoobank.org:author:c7a4b3d3-ae51-4376-ba89-249cd1f1632d http://zoobank.org/urn:lsid:zoobank.org:author:386bd5af-0ca9-4c2e-a421-c1d3576e55a4 http://zoobank.org/urn:lsid:zoobank.org:author:893860fe-e71b-4f69-a46a-8b89834579fc http://zoobank.org/urn:lsid:zoobank.org:author:d18accd9-291d-4b1a-b960-5a2f03695bdc http://zoobank.org/urn:lsid:zoobank.org:author:fea0bfc5-d865-4e3c-8edf-0ecce1d24411 http://zoobank.org/urn:lsid:zoobank.org:author:412800eb-aace-4313-9810-61f89b740405 http://zoobank.org/urn:lsid:zoobank.org:author:1da532e6-fa75-4dc1-9e29-6eecd4dfda87 http://zoobank.org/urn:lsid:zoobank.org:author:8dac0752-82c3-4bb8-8ca5-9da491302533 http://zoobank.org/urn:lsid:zoobank.org:author:27041e84-cc1d-456a-baee-928dfc2021c7 http://zoobank.org/urn:lsid:zoobank.org:author:9f38cee2-3009-4368-9b4b-963c00e900f0 http://zoobank.org/urn:lsid:zoobank.org:author:9a3bdba9-343b-4b5d-b978-edb46ad4353c http://zoobank.org/urn:lsid:zoobank.org:author:401d1c89-c0f8-408f-8035-0ad2e2c25dcb http://zoobank.org/urn:lsid:zoobank.org:author:1d813f3e-2cda-482b-a9be-d8eadfdb09d8 http://zoobank.org/urn:lsid:zoobank.org:author:891eefda-63c2-4098-a6d9-26c5c99fc825 http://zoobank.org/urn:lsid:zoobank.org:author:a2f155e4-7958-4e63-b36a-cab23f190a07 http://zoobank.org/urn:lsid:zoobank.org:author:0c024f0f-c9e6-4398-94d5-b462b43e554e http://zoobank.org/urn:lsid:zoobank.org:author:3708ef86-f640-4ddf-87a3-2d1a1c118890 http://zoobank.org/urn:lsid:zoobank.org:author:84c998a6-886d-4421-8645-00d303af3c24 horton t., gofas s., kroh a., poore g.c.b., read g., rosenberg g., stöhr s., bailly n., boury-esnault n., brandão s.n., costello m.j., decock w., dekeyzer n., hernandez f., mees j., paulay g., vandepitte l., vanhoorne b. & vranken s. 2017. improving nomenclatural consistency: a decade of experience in the world register of marine species. european journal of taxonomy 389: 1–24. https://doi.org/10.5852/ejt.2017.389 introduction the world register of marine species (worms) is a biodiversity information system designed as a global open-access inventory of the names of marine taxa. worms was established ten years ago, in 2007, and has been growing in both content and popularity ever since (costello et al. 2013). worms currently contains 613 499 taxon names (from infraspecies to kingdom), 479 092 of which are species names and 243 081 are accepted marine species names (data accessed 18 dec. 2017). these names are entered, edited and revised as necessary by 278 current voluntary taxonomic editors (worms editorial board, see horton et al. 2017). the database is hosted on a ms sql database platform known as aphia (vandepitte et al. 2015). a regularly updated technical aphia manual is openly available to editors and users on the worms website (http://www.marinespecies.org/aphia.php?p=manual). the worms database was inaugurated in 2008, but was actually established as a project in late 2007 (appeltans et al. 2008, 2011; vandepitte et al. 2015). at the beginning, in order to facilitate the creation of a useful resource, and to ensure that taxonomist experts were not asked to repeat the work of collating information, a number of already existing species registers were incorporated into worms (costello et al. 2013). these included six all-taxon marine species lists at the regional level (e.g., the european register of marine species, erms (costello et al. 2000); register of antarctic marine species, rams (de broyer et al. 2017)), and 14 global species databases covering entire taxonomic groups at a global level (e.g., world list of marine, freshwater and terrestrial crustacea isopoda (boyko et al. 2017), world list of ophiuroidea (stöhr et al. 2017)). the aphia database also holds numerous non-marine taxa as part of global species databases, for those taxa also found in freshwater and/or terrestrial environments, and some orphaned non-marine taxon datasets, although these are not visible through the worms portal by default (costello et al. 2013; vandepitte et al. 2015). at the time of the inauguration in 2008, a press release highlighted that there were already 122 500 accepted marine species names in worms, and the 137 taxonomic experts involved at the time estimated that there were about 230 000 marine species known to science (census / worms news release 2008). by 2012, after five years of progress, worms  contained 460 000 taxon names (from infraspecies to kingdom), of which 368 000 were species names, of which 215 000 were accepted (costello et al. 2013). the original estimate of 230 000 accepted marine species was reached on 15 july 2015. with the task of aggregating the accepted species names for all marine species approaching completion, a list of new priorities was drawn up by the steering committee (a group of 12 editors elected for three-year terms to govern current and future development of worms, http://www.marinespecies.org/about.php#governance). these priorities focus on improving the consistency and quality of the data already held for each species whilst also keeping the database up-todate with the addition of ca 2000 new marine species described annually. the aggregation of global and regional datasets into worms, and the revisionary efforts of over 300 voluntary expert taxonomists over the last decade, have resulted in slightly diverging current editorial practices between taxa. this can influence management of the database and data use in the future.  while new taxonomic editors are provided with access to a technical training manual on the process of editing worms, there has not been, until now, any guidance on the best practice for entering and editing taxonomic names in worms. in order to provide a standard set of procedures for the most commonly encountered situations in worms, the steering committee decided that a guidance document should be written for use by the editors, illustrated by examples of best practice where possible. it was anticipated that this guidance document should promote consistency between the application of the international horton t. et al., best practice guidelines for worms taxonomic editors 3 https://doi.org/10.5852/ejt.2017.389 http://www.marinespecies.org/aphia.php?p=manual http://www.marinespecies.org/about.php#governance code of zoological nomenclature (iczn) and the international code of nomenclature for algae, fungi, and plants (icn, mcneill et al. 2012), and in the addition and editing of names in worms. by publishing a version of this guidance, we aim to make worms’ methodology available to other taxonomic name databases and worms users, to further promote consistency, and to thus facilitate the use of worms data in downstream analyses. users of taxonomic information are faced with numerous challenges in navigating taxonomic databases, including worms. these challenges include determining how taxonomic issues such as misspellings of names, misapplications of taxon concepts, synonymisations and name recombinations are represented in a taxonomic name database. these concepts were discussed in detail by costello et al. (2013). we make use of nomenclatural terms already covered extensively by hawksworth (2010) and refer to information and discussions in the comprehensive guidelines prepared by welter-schultes (2012) for the gbif database. although we refer specifically to the use of these terms and methods in relation to editing of  the worms database, it is anticipated that the examples and recommendations for consistency presented here will be applied by other database managers, users and custodians of collections using worms. rationale and application of the world register of marine species the aim of worms is to provide an authoritative and comprehensive list of names of marine organisms arranged in the most up-to-date and stable hierarchy as determined by the appropriate editor. the worms hierarchy incorporates, without necessarily being driven by, the very latest phylogeny such as in the tree of life (www.tolweb.org), but attempts to reflect a stable and practical classification. ‘match  taxa’, an online service on the worms website furthermore allows to upload taxon lists and retrieve the most-up-to-date names and classifications for these taxa. the primary goals of worms are: (1) to promote consistency and stability of names this upholds the goal stated in the preamble of the iczn “to promote stability and universality in the scientific names of animals and to ensure that the name of each taxon is unique and distinct”. consistency  and stability are of particular concern in the case of species important to society, be they of commercial or medical use, harmful to humans or endangered species listed in legal texts (garnett & christidis 2017). since the launch and development of worms, the accuracy of names used in non-taxonomic works has significantly improved. for example, prior to 2007 the manila clam ruditapes philippinarum (adams & reeve, 1845), a commercially important species farmed and introduced in many parts of the world, was recorded as ruditapes philippinarum in the title of 1270 publications listed in google scholar, and in 87 further publications (6.4%) with the invalid synonym ruditapes semidecussatus (reeve, 1864). in contrast, for the period 2007–2017, the species name philippinarum was used in 1040 titles and semidecussatus was not used in any article title. (2) to serve as a guide to interpret the taxonomic literature the original primary goal of worms was to collate a list of accepted species names, but also to include important synonyms. including information on unaccepted names (synonyms, misspellings, old combinations etc.) and linking these to the accepted name provides the user with an interpretation of the taxonomic history of a species and some understanding of the reason(s) for non-acceptance of certain names. resolving taxonomic issues requires informed individuals who understand the problems and how they have arisen, and who know the rules of the nomenclatural codes (costello et al. 2013). the addition of this expert-level interpretation of the taxonomic literature is the main difference between the worms database and other nomenclators that aim to serve a list of all names regardless of their taxonomic status. european journal of taxonomy 389: 1–24 (2017) 4 www.tolweb.org an additional priority of worms is to support each name in the database with a published, citable source containing the name that is linked, whenever possible, to the publisher or journal website, portals of digitisation efforts and/or an accessible pdf. providing these direct links to the taxonomic literature is another progressive feature of worms and is much appreciated by users. species have been described in thousands of books and journals and gathering this literature is time-consuming and difficult. again, expert taxonomists with knowledge of the literature for their taxonomic group are in  the best position to undertake this task. parallel with the development of worms, the biodiversity heritage library (bhl, http://www.biodiversitylibrary.org/) and many other repositories were set up, providing free access to the full texts of most publications prior to 1921 and to some later ones. the task for taxonomic editors is to determine which work is the original description, and where appropriate, which is a source that synonymises the species, or is relevant for another reason. important taxonomic references can therefore be linked directly to the taxon page by editors to aid interpretation of the literature by users. (3) to provide access to additional information regarding the taxon the minimum requirements for an entry in worms is the full scientific name (i.e., combination of  genus and specific epithet, author, year) placed in an accepted taxonomic hierarchy, annotated with  an  environment  flag  (e.g.,  marine/brackish/terrestrial/freshwater,  see  section  on  environment  later)  and supported by a reference. further information is both possible and recommended to be included, and the addition of the following has been agreed by the worms steering committee according to a priority list: original name, original description, type locality, and type species of genera. some species pages also contain considerable additional information (e.g., type materials, distributions, feeding types, habitats) and there are plans for more. a dataset integrated with worms known as marine species traits (http://www.marinespecies.org/traits/about.php) has now been established and certain traits were prioritized for inclusion, namely environment, geography, depth, substratum, mobility, skeleton, diet, body size and reproduction (costello et al. 2015). (4) to provide an estimate of the number of marine species the answer to the question “how many species are there on this planet?” including those species that remain to be discovered, remains disputed (bouchet et al. 2016). the worms database has been put to repeated use to make estimates of marine (and global) species numbers. these estimates commonly rely on the extrapolation of limited datasets under particular assumptions. mora et al. (2011) based estimations of the number of global species on the catalogue of life (col, http://www.catalogueoflife.org/) and the number in the ocean on the world register of marine species (worms), and concluded that the number of marine species was ca 2.2 million (±0.18 million se), of which 91 % are yet to be described. appeltans et al. (2012) analysed data from worms, used statistical models, reviewed the number of undescribed species in 100 field studies, and compiled the opinions of 122 authors to estimate “the  magnitude of global marine species diversity”. they concluded that fewer than 222 000–230 000 marine  species have already been described, predicted between one-third and two-thirds may be undescribed, and reported that fewer than one million marine species exist. poore et al. (2014) studying bathyal fauna from australia showed that the combination of high species richness, higher endemism than presently acknowledged, low proportions of described species, and the vast extent of unexplored bathyal and abyssal environments will lead to further accumulation of new species as more deep-sea regions are explored. they concluded that the estimates made by appeltans et al. were too low and that all estimates of global species, however calculated, are based on limited data. a recent paper by (costello & chaudhary 2017) concludes that there are likely 0.3 million species in the ocean and that two-thirds of species are now described. it is clear that the wide variation and disagreement in these estimates indicates that the actual number of marine species remains to be determined. however, the provision of horton t. et al., best practice guidelines for worms taxonomic editors 5 http://www.biodiversitylibrary.org/ http://www.marinespecies.org/traits/about.php http://www.catalogueoflife.org/ a comprehensive global list of accepted species by worms provides a basis for some species estimate methods, particularly those based on the ratio of known to unknown species in samples. once these goals are achieved, users can expect worms to help: provide a currently accepted name from a list of possible alternative names in use check the accuracy of names used in a taxonomic listing (the online ‘match taxa’ tool automatically matches a taxon list from a spreadsheet with worms (vanhoorne et al. 2008; vandepitte et al. 2015) cross-check names used elsewhere (e.g., other databases) and draw attention to mismatches. this is a two-way interaction and if names are incorrectly entered or missing in worms they will be revised or added users of worms vary from individuals to organisations engaged in various activities, which require the correct and consistent names for marine species. the worms webservices are now being used by 65 organisations from 20 countries, and since permission is not required, there are likely additional users we are unaware of. copies of the database have been licensed to 157 organisations and individuals in 33 countries and demand is growing steadily. since 2007, all metrics on website use show a steady increase (see http://www.marinespecies.org/users.php for up-to-date figures). we have recorded 1933 publications  making an explicit statement that the taxonomy is based on worms or any of its subregisters (www. marinespecies.org/wormsliterature.php), and the database has been cited in the references of over 300 peer-reviewed publications since 2007. the above goals and the many and varied uses of worms all require editorial consistency according to standardised guidelines, the provision of which is the subject of this paper. entering names in worms worms is a recording tool for published data and information. therefore, all names entered should be based on at least one published reference source in which that particular name string (species rank and below) or rank (supraspecific names) has been used. a source should be linked to the name at the time of  entry into the database and thus provides a reference to the published taxonomic literature (see section on ‘source categories’). entries in worms do not reflect unpublished taxonomic opinions nor propose  new combinations for species names (for exceptions see section on ‘new combination reference’). most worms taxa are governed by two nomenclatural codes; the international code of zoological nomenclature (iczn) and the international code of nomenclature for algae, fungi, and plants (icn). since 2012, worms has also incorporated marine viruses following a collaboration with the international committee on taxonomy of viruses (ictv) (king et al. 2012) and holds marine bacteria according to the international code of nomenclature of prokaryotes (prokaryotic code (2008 revision), parker et al. 2015). to avoid misunderstandings between their different terminologies, worms uses a neutral terminology that encompasses that used in these codes. differences are pointed out below, as necessary. the definitions provided were derived after careful consideration of nomenclatural rules. they may  serve as guidance to non-taxonomists and students, beyond their application in worms. to achieve most rapidly a list of all marine species and taxa, worms editors were not requested to enter original names from the outset. nomenclators aim at recording all original names for a taxonomic group with their nomenclatural status, including the type information, but not their taxonomic status (e.g., accepted or unaccepted). managing scientific names in a database is facilitated if original names are  also recorded because all subsequent forms of the name can be linked to one objective name that will, in principle, never change (pyle & michel 2008). therefore, in 2015 an original name field was added to  worms, to which all subsequent names can be linked. it is now a priority for editors to enter original names in worms and to link all combinations to the original name. this task is eased by existing nomenclators european journal of taxonomy 389: 1–24 (2017) 6 http://www.marinespecies.org/users.php www.marinespecies.org/wormsliterature.php www.marinespecies.org/wormsliterature.php in different domains. there are multiple digital sources of names in existence (see rees et al. 2017 for an overview and comparison of major nomenclators). in 2005, the international commission of zoological nomenclature developed zoobank as the nomenclator for groups covered by the iczn (polaszek et al. 2005; pyle & michel 2008). zoobank has now been defined in the iczn as the repository of all original  names published in zoology (iczn 2012). additionally, since 1 january 2012, registration in zoobank of published works containing new taxonomic names is mandatory for electronic publications and strongly recommended for paper ones, including from the legacy literature (krell & pape 2015). in the future, it is anticipated that there will be progress towards the exchange of data on original names between zoobank and worms (and other databases) to avoid duplication of effort. name status terminology in the published literature, it is not always clear whether a name is currently valid or not, either for nomenclatural or taxonomic reasons. in the interest of clarity and consistency, when a name is entered in worms it must be given a status from a dropdown menu. the options for these in worms are: 1. accepted name 2. unaccepted name 3. temporary name 4. uncertain name 5. alternate representation 1. accepted name the primary goal of worms is to hold the accepted name of each marine species, because most nonspecialist users need to find the correct, current name of their taxon of interest. accepted names are  denoted by the status ‘accepted’ on the taxon page. this is the name considered by the editor to be the valid (zoology) or correct (plants, algae and fungi) name of the taxon. a newly published taxon is entered to worms as ‘accepted’ (unless failing to fulfil criteria of availability (zoology) or not validly  published (plants, algae and fungi)). the status of an original combination will only be changed to ‘unaccepted’ following publication of the name as a new combination or as a junior synonym. one or several sources should be linked to the name to support its inclusion in the database. the accepted name is selected by the editor after consideration of the most up-to-date revision available. if contemporary works give different generic allocations or different rankings of a taxon, the editor will need to make a choice, which may not necessarily be the latest published status, but the one which he/ she considers to be the most soundly supported. this is an important point for worms editors and a place where there have been differences of opinion in how best to represent the current accepted name. how should an editor decide on the accepted name? how can we provide consistency in this decision-making for the users of worms? editors may decide to represent the most up-to-date revision in worms (which may not be the most authoritative), or they may choose to represent one that is perhaps a few years older, but which is considered by that editor to have stronger support. for example, a thorough revision supported by sequence data could be considered to have greater merit than a more recent revision that consists of new combinations provided in a list without discussion. an editor may therefore decide to represent the older revision as accepted. in such cases there should always be a ‘taxonomic remark’ note explaining the details of the case, and the opposite view should be included and be either flagged as ‘unaccepted’ or ‘alternate representation’ according to the editor’s decision. names should be linked to their original combination (zoology) or basionym (plants, algae and fungi), i.e., the name (genus, species and subspecies) used in the original publication (and correctly spelled under iczn or icn rules) and which may or may not be the same as the accepted name. in worms horton t. et al., best practice guidelines for worms taxonomic editors 7 this is denoted the ‘original name’ in order to encompass and avoid conflict with the terms ‘original  combination’ and ‘basionym’. both accepted and original names should be added to worms (as well as any superseded combinations that the editor sees fit). when a new taxon is entered and the authority is added without brackets (i.e., in  its original combination), the ‘original name’ field does not need to be filled, as the taxon name is given  this status automatically. to indicate that a combination previously added to the database is the original name, the editor can click on ‘mark as original name’, which is an option located after the taxon name string on each taxon page when an editor is logged in to worms. linking of non-original combinations with the original name is achieved by filling the ‘original name’ field on the taxon edit page (click ‘edit  taxon’). the original name can be selected from the list generated when the editor begins typing in this field. if the original name is not yet in worms then it will need to be added before this step.   the type species of a genus should always be entered in the form of its original combination, even if it was cited in another combination at the time of designation (recommendation 67b (iczn 1999); article 10, ex. 3 (icn)). 2. unaccepted name a name that has been in use at any time but is not currently accepted. this includes names which are invalid (e.g., junior synonym, preoccupied, rejected), specific names in generic combinations which are  no longer accepted, specific names written with the wrong grammatical agreement and misspellings. in  worms an unaccepted name must point to a currently accepted name. there are some special cases, including unavailable names, where a name does not point to a currently accepted name, e.g., some cases of nomina nuda and names that are published online only without registration in zoobank (e.g., goniopygus annularis vaziri & arab, 2012: http://marinespecies.org/aphia.php?p=taxdetails&id=738293). in these cases, the editor should contact the data management team (dmt, info@marinespecies.org) to have the name entered without linking to an accepted name. unaccepted names should be linked to a published source in which the status of the name is justified.  editors should not make subjective changes on worms that have not been published. exceptions can be made for homonyms, nomina nuda and nomina oblita, misspellings, incorrect original spellings and grammatical agreement of specific epithets. in such cases editors should use the ‘notes’ field to identify  the change and explain their reasoning. for unaccepted names, editors must include a statement in the ‘unacceptreason’ field. this statement,  which explains the reason for non-acceptance of the taxon, will be displayed in brackets after the status. the word ‘invalid’ should not be used in this field because this term has its own meaning in  bionomenclature, and is implicit from the ‘unaccepted’ status. a dropdown list of the most commonly used reasons is available to encourage consistency of usage for these terms. these are listed and explained below. a reason for non-acceptance should be chosen from the available formats, if applicable, although a free text field is also provided to allow full explanation  of complex cases. a. superseded combination this is to be used when a species is transferred to another genus, and is also known as a new combination. the term synonym should not be used for recombinations, because it has its own specific meaning (see  below). european journal of taxonomy 389: 1–24 (2017) 8 http://marinespecies.org/aphia.php?p=taxdetails&id=738293 info@marinespecies.org further detail may be provided by using the following terms in the free text field if required. superseded original combination superseded recombination see, e.g., http://www.marinespecies.org/aphia.php?p=taxdetails&id=209687 b. junior homonym to be used to indicate the invalid status of the junior, or later established name, or in the case of simultaneous establishment, the name not given precedence by a first reviser or by an iczn or icn  ruling. c. junior subjective synonym this is to be applied as the reason for the unaccepted status for the junior, or later established, subjective synonym (a published opinion that two names apply for the same taxon). this is known as an allotypic synonym in the icn. d. junior objective synonym this can be applied to the later established name in cases where two or more scientific names have the same type material (this is known as a homotypic synonym in the icn). for example, when two different authors name a new species based on the same specimen or in more deliberate cases, where neotypes are selected to ensure that two different nominal taxa are synonymized by anchoring to the same primary type. e. nomen nudum a latin term (meaning ‘naked name’) applied to those published names that do not comply with the name requirements of the codes, e.g. lack of a description or diagnosis (see iczn glossary). nomina nuda are not available names, and therefore an identical name may be made available later for the same or a different concept. this can be used when the taxon is indicated as a nomen nudum in a revisionary work (which is then linked as the ‘status source’), but it may also be used when an editor discovers a name with such status. if this status is not yet published, then the editor should add a clear ‘status note’ to explain the problem. f. nomen oblitum a latin term (meaning ‘forgotten name’) applied after 1 january 2000 to a name, unused since 1899, which as a result of an action taken under iczn article 23.9.2 does not take precedence over a younger synonym or homonym in prevailing usage; the younger name which takes precedence over the nomen oblitum may be called a nomen protectum (meaning ‘protected name’). the term nomen oblitum was also applied to a disused senior synonym rejected between 6 november 1961 and 1 january 1973 under iczn article 23b of the code editions then in force (see iczn article 23.12.2). nomina oblita remain available names; see iczn articles 23.9 and 23.12 for conditions controlling their use as valid names. see example here: http://www.marinespecies.org/aphia.php?p=taxdetails&id=541681 spelling variations there is a thorough discussion of “how is the name correctly spelled?” in welter-schultes (2012), and  editors are strongly encouraged to read section 8 of those guidelines. some sections are modified and  included here to indicate how those guidelines should be applied to editing taxa in worms. worms editors should always verify the correct spelling of any names used. as noted in welterschultes (2012) “in many cases spellings are not consistent with the original spellings, but appear in a slightly modified form. often these modified spellings are correct (justified emendations), but in other  horton t. et al., best practice guidelines for worms taxonomic editors 9 http://www.marinespecies.org/aphia.php?p=taxdetails&id=209687 http://www.marinespecies.org/aphia.php?p=taxdetails&id=541681 cases they turn out to be incorrect and not in accordance with the original description and the rules of nomenclature (unjustified emendations). in most cases it is convenient to use the correct original  spelling for a name, especially if names are not very commonly used”. in such cases of unjustified  emendations it is therefore recommended that the editor use the correct original spelling for the accepted name, particularly where the name has been used in very few publications, and is therefore not in prevailing use. g. incorrect original spelling corrections to an incorrect original spelling are classed by the iczn as a justified emendation and  the iczn rules and recommendations must be followed (see section 8 in welter-schultes 2012). if a correction to an accepted name has been made under the provisions of iczn article 32.5, the corrected form is added to the database as ‘accepted’. the original incorrect spelling is an unavailable name and should be entered in worms with status ‘unaccepted’ and the reason for non-acceptance as ‘unavailable name: incorrect original spelling’. it is left at the discretion of the editor whether or not to include original names with hyphens/diacritical marks or specific names with upper case initial  letters. h. incorrect subsequent spelling an ‘incorrect subsequent spelling’ is either a misspelling (also known as a lapsus calami or denoted ‘sic’ in the published literature), or an unjustified emendation (a deliberately different spelling with an  explicit statement of intention or having cited both the original and the changed spellings, see iczn article 33.2). i. misspelling misspellings may be entered in worms for erroneous names in the literature that need to be captured, e.g., when they are widespread (e.g,. arca noe for arca noae), or when they are needed in the database as parents for species names introduced in combination with a misspelled genus name (when the record must appear exactly as published). misspellings should be entered into worms without an authority and with the status ‘unaccepted’. the term ‘misspelling’ (or ‘misspelling of genus species author, date’) should be used as the reason for nonacceptance. the publication containing the misspelling should be linked to this unaccepted taxon as a source ‘basis of record’ and not as ‘original description’. misspelled names should link to the correctly spelled name, which may not necessarily be the accepted name. see, e.g., http://www.marinespecies.org/aphia.php?p=taxdetails&id=381235 j. unjustified emendation unjustified  emendations  are  nomenclatural  acts  under  iczn  article  33.2  and  must  strictly  be  a  ‘demonstrably intentional change’ of the original spelling. cases where both the original spelling and the changed spelling are cited, with the latter used as valid, are unjustified emendations, and therefore  become available names. unjustified emendations take as authority the author of the emendation. the  term ‘unjustified emendation of genus species author, date’ should be added as free text as the reason for non-acceptance, with the source linked as ‘original description’ (since unjustified emendations are  available names). when only the changed spelling is given, and an intention is not explicit, the name should be treated as a misspelling. k. incorrect grammatical agreement of specific epithet to be used in the case of mandatory changes in spelling consequent upon changes in rank or combination. see welter-schultes (2012) section 8.1.6 on gender agreement, which contains a comprehensive discussion on this topic. if a correction has been made to an accepted name under the provisions european journal of taxonomy 389: 1–24 (2017) 10 http://www.marinespecies.org/aphia.php?p=taxdetails&id=381235 of iczn article 34, which covers gender agreement or a change of suffix with change of rank, the  corrected form is to be listed as ‘accepted’ with an appropriate source linked. the original name with the wrong gender agreement should be added to the database with status ‘unaccepted’ and reason for nonacceptance ‘incorrect grammatical agreement of specific epithet’. editors are encouraged to introduce  correct spellings as new, accepted taxa when such errors are spotted (in accepted names only, superseded combinations should be written as encountered and should not be corrected). a ‘nomenclature note’ should be added to explain that this is the first time the correct spelling is used. see example here:  http://www.marinespecies.org/aphia.php?p=taxdetails&id=103195 l. misapplication there are many examples in taxonomic nomenclature when an accepted name has been misapplied to an incorrectly identified specimen. such misapplications of the name (also known as misidentifications)  are, in worms, referred to as misapplications. misapplications are not nomenclatural acts and therefore do not need to be systematically entered in synonymy lists. they need only be entered in the database when one or more of the following conditions is met: -  the misapplication is given in an influential  textbook or identification guide and has initiated an  incorrect usage in many subsequent works. the misapplication leads to species being recorded outside their known geographical area. the misapplication has been unduly treated as a separately available name (i.e., with separate authorship and as an apparent homonym of the original name) by later authors. -  the misidentified specimen has been explicitly cited as a basis for the description of a subsequently  available name. -  the misapplication is involved in the typification of a genus so as to become relevant under iczn  article 69.2.3 or 70.3 (but note that a misapplication explicitly applied as type of a new genus is a nomenclatural act under iczn article 67.13). misapplications need a separate entry, distinct from the correctly identified ‘genus species author, date’, because they link to a different accepted name. the format for such cases should be: genus species author, date sensu author of misapplication, date the reason for non-acceptance field should contain ‘misapplication’. for an example of this and how to  represent it in worms see: http://www.marinespecies.org/amphipoda/aphia.php?p=taxdetails&id=510827 m. unavailable names in addition to those already mentioned, there are numerous complex reasons for a name to be unavailable, therefore a free text field is provided for use in these cases. some examples of such usages are given below: published as ‘variety’ or ‘form’ after 1960 (iczn 15.2). http://www.marinespecies.org/aphia.php?p=taxdetails&id=758580 placed on the official index by iczn opinion # http://www.marinespecies.org/aphia.php?p=taxdetails&id=266910 editors can use the search function to see how other similar entries have been made in worms by typing, e.g., ‘iczn opinion’ in the ‘unacceptreason contains’ field on the aphia taxon search page. horton t. et al., best practice guidelines for worms taxonomic editors 11 http://www.marinespecies.org/aphia.php?p=taxdetails&id=103195 http://www.marinespecies.org/amphipoda/aphia.php?p=taxdetails&id=510827 http://www.marinespecies.org/aphia.php?p=taxdetails&id=758580 http://www.marinespecies.org/aphia.php?p=taxdetails&id=266910 n. interim unpublished the definition of this is “an as yet unavailable name (until in a print issue), which has been published  online only, in a work that does not show evidence of zoobank registration (iczn article 8.5)”. this  status is only needed for taxa published according to the iczn and is not applicable to icn governed taxa. a major difference in the implementation of electronic-only publication in the nomenclatural codes is that in zoology (iczn), this must be accompanied by registration in zoobank, while for algae, fungi and plants (icn), it is considered just another publication type and is not necessarily linked to registration (fungal names, however, must be registered to be validly published) (nicolson et al. 2017). this status is needed in worms because some print journals publishing taxonomy make articles containing new taxa available online well before appearance of the print copy. these have delays of up to a year before being assigned to a journal issue and being printed, thus becoming conventionally iczn print published, and making the names available under the provisions of the iczn. names published in these advanced versions of articles that were not registered in zoobank are unavailable according to iczn regulations, until a version (online or print) that satisfies the requirements of the code is published. such names  should be entered to worms with the reason for non-acceptance as ‘interim unpublished’, and they will not point to an accepted name. these names can be monitored and when they are validly published, they can be changed to ‘accepted’ and be linked to the iczn-compliant version of the respective work. when editors add the publication containing the original description, it should be added with the date of the print edition, but the date of publication of the online edition should also be included as a note, so users can see that there is no oversight from the part of the editor, e.g., neopycnodonte zibrowii gofas, salas & taviani in wisshak, lópez correa, gofas, salas, taviani, jakobsen & freiwald, 2009. the source states: deep sea research i 56 (3): 374–404 [published online 29 october 2008; printed edition of the journal dated march 2009]. see an example here: http://www.marinespecies.org/aphia.php?p=sourcedetails&id=126164 3. temporary name this name status can be used to create ad-hoc higher rank taxa of convenience to accommodate child taxa for which the classification is not yet finalised. this is usually applied to family and generic names.  if, for example, there is a taxon (any rank) that does not fit in the current classification, the editor can  create a temporary name ‘x incertae sedis’, where ‘x’ is the name of the parent taxon. there are many examples of this usage within worms, see, e.g., lysianassoidea incertae sedis. incertae sedis means ‘of an uncertain seat’; and should be applied only to taxa of uncertain systematic position. when editors create these names, the status ‘temporary name’ should be applied and the placement of taxa within this category should, whenever possible, be backed up with a ‘status source’. in some cases, placeholder names may be used to indicate that we know which clade a taxon belongs to in the phylogeny and we also know that it does not belong to the crown group. these can be represented using [unassigned] in front of the taxon name, see an example here: http://www.marinespecies.org/aphia.php?p=taxdetails&id=382219 temporary names can also be used as a way to record unnamed cryptic species in worms. in many cases there is a considerable time lag between discovery of cryptic species (e.g., during phylogeographic or barcoding studies) and their formal taxonomic description. informal names, which are attributed to individual cryptic clades, are sometimes used again in subsequent papers. these names thus become unofficial names for the taxa involved. when this occurs over an extended interval, it may be desirable  to record these names in worms. an excellent example for such a case is the tropical sea urchin genus echinometra, which contains six accepted, formally described extant species, and several yet unnamed cryptic species. these were discovered by cross-fertilization experiments in the 1980s european journal of taxonomy 389: 1–24 (2017) 12 http://www.marinespecies.org/aphia.php?p=sourcedetails&id=126164 http://www.marinespecies.org/aphia.php?p=taxdetails&id=731859 http://www.marinespecies.org/aphia.php?p=taxdetails&id=382219 (uehara & shingaki 1985), and confirmed by genetic studies (matsuoka & hatanaka 1991; mccartney  et al. 2000). formal descriptions, however, are still missing, in part because the relationship between the four cryptic species and available historical taxon names is not well understood. the informal names echinometra sp. a, and echinometra sp. c, established in one of the first papers dealing with the  subject, are widely used (and consistently applied) by the scientific community and, therefore, have been  captured in worms (http://www.marinespecies.org/echinoidea/aphia.php?p=taxdetails&id=994686; http://www.marinespecies.org/echinoidea/aphia.php?p=taxdetails&id=994687). 4. uncertain name this status is used to indicate taxonomic or nomenclatural uncertainty for cases that cannot be classed as either ‘accepted’ or ‘unaccepted’. in worms, an unaccepted name must point to a currently accepted name. uncertain names, however, commonly do not point to an accepted name and the editor should contact the data management team to have this name entered without linking to an accepted name. for uncertain names, editors must include a statement of the reason for non-acceptance. there are three options available for uncertain names to encourage consistency of usage for these terms. these are listed and explained below. a. nomen dubium a latin term (meaning ‘doubtful name’) which is used to refer to a name of unknown or doubtful application. this applies to a name which is available (animals) or validly published (plants, algae and fungi) but of uncertain taxonomic significance despite the attention given, usually, for example, because  types have been lost (or are poorly preserved), or if the description does not allow us to know with certainty which species was really meant. nomen dubium should only be used when given this status in a revisionary work, not when an editor deems it so, and it should be supported by the source flagged as  ‘status source’. see here: http://www.marinespecies.org/aphia.php?p=taxdetails&id=851739 if an editor has doubts about a name in recent use that has not been stated in the literature to be a nomen dubium, a note could be added to the name but it must remain ‘accepted’. see here: http://www.marinespecies.org/aphia.php?p=taxdetails&id=391348; http://www.marinespecies.org/aphia.php?p=taxdetails&id=737113 b. taxon inquirendum a taxon may be of uncertain taxonomic significance because it has not been revised nor used in the  recent literature; contrary to the nomen dubium which resists revision because the supporting data are deficient, the taxa treated as inquirenda may still have extant type specimens or be adequately described, but just not have been reassessed; their generic assignment may be outdated, and they may be overlooked synonyms of valid names. this uncertain status should have a ‘status note’ added as explanation, and should preferably only be used when given this status in a revisionary work, not when an editor deems it so. the information should be supported by a source flagged as ‘status source’. see here:  http://www.marinespecies.org/aphia.php?p=taxdetails&id=711728 c. unassessed this is used to refer to names coming from nomenclators (lists of names) or from, e.g., museum collection databases, where the status might be known in the published literature, but for which the status has not yet been researched, assessed and documented in worms by a taxonomic editor. when such lists are integrated into worms, the names should be labelled as ‘unassessed’. for these names, the priority will be to assess their current status and to document this with an appropriate source flagged as ‘status  source’. horton t. et al., best practice guidelines for worms taxonomic editors 13 http://www.marinespecies.org/echinoidea/aphia.php?p=taxdetails&id=994686 http://www.marinespecies.org/echinoidea/aphia.php?p=taxdetails&id=994687 http://www.marinespecies.org/aphia.php?p=taxdetails&id=851739 http://www.marinespecies.org/aphia.php?p=taxdetails&id=391348 http://www.marinespecies.org/aphia.php?p=taxdetails&id=737113 http://www.marinespecies.org/aphia.php?p=taxdetails&id=711728 5. alternate representation this is any combination that appears in the literature, is distinct from the ‘accepted name’, but is not currently wrong or unaccepted. alternate representation can be used to deal with contradictory taxonomic treatments. one of the names must be ‘accepted’ but the alternative treatment may be stated as ‘alternate representation’ instead of ‘unaccepted’. another use for this status is to deal with subgenera, subfamilies, etc. names of species with an interpolated subgenus name – genus (subgenus) species – may be given the status of ‘alternate representation’ to the ‘accepted’ plain binomen genus species. see, e.g., http://marinespecies.org/aphia.php?p=taxdetails&id=138995 this can be extended to the situation where the subgenus is raised to genus status, but where there is, as yet, no consensus in the literature (then subgenus species is also ‘alternate representation’ to genus species). when a consensus emerges or when a definitive revision is published, one of the alternate  representations should become ‘unaccepted’ and the other one ‘accepted’. problematic entries linking an unaccepted name to more than one taxon it often happens that a higher taxon becomes unaccepted and its ‘children’ are spread over several other taxa. in a taxonomic publication, this is represented by ‘in part’ or ‘pro parte’. it is not possible to represent this in worms, since this makes the database structure highly complex. in such cases, editors can contact the database management team to arrange to have no accepted name, and should add a clear taxonomic remark note. see an example here: http://www.marinespecies.org/aphia.php?p=taxdetails&id=106675 authorities the international plant names index (ipni) has a standardised list of authors and their abbreviations (with strong recommendations to use these). authors of plant, algal and fungal taxa should be cited following the standard form indicated in http://www.ipni.org/ipni/authorsearchpage.do. the iczn code does not provide an official guide for the name of the author. according to welterschultes (2012), researchers should use the authority as written in the publication, we advise editors to follow this recommendation. in accordance with iczn art. 50.1, where the authority of a taxon differs from the author(s) listed on a publication, editors should use the following format: ‘author 1 in author 2, date’. authors of zoological names are usually cited without their initials. in some cases, when more than one homonymous author is active within the same taxon on overlapping dates, they can be cited using the format ‘e.a. smith’. if these authors also have the same initials, use as many letters of the first name as  needed to discriminate them (e.g., ev. marcus/er. marcus, ant. bivona/and. bivona). in case of doubt, editors can check the current usage in worms with an authority search. nomen protectum a latin term (meaning ‘protected name’) applied to a name that has been given precedence over its unused senior synonym or senior homonym relegated to the status of nomen oblitum (see iczn article 23.9.2). if this status is given to a name in the literature, then a ‘status note’ should be written detailing the information regarding the case and backed up by the ‘status source’. a nomen protectum is protected against an older synonym or homonym but can become unaccepted european journal of taxonomy 389: 1–24 (2017) 14 http://marinespecies.org/aphia.php?p=taxdetails&id=138995 http://www.marinespecies.org/aphia.php?p=taxdetails&id=106675 http://www.ipni.org/ipni/authorsearchpage.do (e.g., by further synonymy with another name than the corresponding nomen oblitum). see here: http://www.marinespecies.org/aphia.php?p=taxdetails&id=540396 source categories when a source (reference/publication) is linked to a taxon, a relevant source category must be chosen. the addition of certain sources such as ‘original description’, ‘new combination reference’ or ‘status source’, are necessary, but further choice of sources under a species entry is at the discretion of the editor. some common species trigger a large literature and the editor should select the most relevant sources. sources providing only the species name in a regional checklist or database should not be linked to the taxon name, but as a source for the corresponding ‘distribution’ entry. the following source categories (in alphabetical order) are available: a. additional source this can be used for any publication that contains additional information about the present taxon, which cannot be categorised as another source category. b. basis of record this is the default category for the source which has triggered the entry in worms, either as an accepted name or an unaccepted name. unavailable (unaccepted) names (e.g., nomina nuda; misspellings, misapplications) to which ‘original description’ cannot apply will also use this source category. c. biology source a publication that gives information about the biology or natural history of the present taxon, e.g., morphology, feeding, growth, reproduction, etc. d. classification source this should be used for publications that contain information about the classification of the present  taxon. it can relate to the phylogenetic classification of a taxon and encompasses both molecular and  morphological approaches. e. context source a publication that is used to link a taxon to a geographic context (e.g., the european register of marine species), or to a thematic context (sub-collection) of aphia, e.g., to a thematic species database (tsd) such as the world register of deep-sea species (wordss). f. ecology source a publication that gives information about the relations and interactions between the present taxon and its environment, including other organisms. g. identification resource this can be applied to papers or websites which include keys to species in a genus, genera in a family or a more general identification guide. the usage will vary across taxa. h. new combination reference this should be used to indicate the first source to propose the new combination; for animals this is  often difficult to trace and not explicit, except in recent publications; for botanical names the author of  this source is part of the authority cited with the valid name. the new combination reference should be linked to the unaccepted name and will then also be visible on the accepted name page. in cases where the new combination is implied by a published work, but the taxon is not specifically mentioned (e.g.,  if a subgenus is raised to genus status or sinking a species-rich genus into another), the included species horton t. et al., best practice guidelines for worms taxonomic editors 15 http://www.marinespecies.org/aphia.php?p=taxdetails&id=540396 should be combined accordingly in their ‘accepted’ name. in these cases, a ‘taxonomic remark’ such as: “new combination herein, to be consistent with synonymisation of [genus 1] with [genus 2] by [author, year]”, should be added by the editor. i. original description this is the publication in which the nominal taxon was originally described. original description sources should be added to the ‘original name’ (not to recombinations) and tagged as ‘original description’ using the source drop-down menu. page references are not needed where a paper is entirely dedicated to the description of the referred taxon, but they are useful additions in larger works. ideally, the page field  should contain the interval of pages dealing with the taxon, as well as the reference to plates if these are separate from the text, and may include references to text-figures as well. j. original description (unavailable nomenclaturally) this source category is intended for cases (common in the 17th and early 18th century) where names were made available in a binominal work by a mere reference to an earlier source (e.g., pre-linnean, or rejected) where the species is figured and described (see iczn art. 12.2.1). in those cases, this earlier  source should be linked as: ‘original description (unavailable nomenclaturally)’. this can also be used more generally for sources where new taxa were proposed but failed to fulfil the conditions for  availability (e.g., varieties for animals after 1960; species described without a type after 1999), where the conditions for availability were fulfilled later. k. redescription this is a publication in which the present taxon is redescribed. l. source of synonymy this should be used for those publications that contain information on the synonymy of a taxon (i.e., where one taxon is declared a junior subjective synonym of another). it should not be used to indicate those papers from which a ‘full synonymy’ listing has been taken. it should not be confused with ‘new combination reference’. m. status source this is used for a publication in which the current status of the taxon is confirmed as valid, or pulled  out of synonymy, or stated as ‘nomen dubium’ or ‘taxon inquirendum’, after a revision. at supraspecific  level it may refer to a source where the taxon was assigned a particular rank (e.g., subfamily to family). iczn opinions are treated as regular papers and cited as ‘status source’ for the taxa they rule on. for downgrading to a junior synonym ‘source of synonymy’ is used instead. for transfers to another genus ‘new combination reference’ is used. n. subsequent type designation this is used for a publication in which the type species or type specimen of the nominal taxon was designated. this source is needed at the genus level when the type species field indicates ‘subsequent  designation’. at the species level it would record any lectotype or neotype designation. o. toxicology source this source category is used in the ioc-unesco taxonomic reference list of harmful micro algae (http://www.marinespecies.org/hab/) and indicates a publication that contains information on the toxic nature of the present taxon. european journal of taxonomy 389: 1–24 (2017) 16 http://www.marinespecies.org/hab/ note categories where further information needs to be provided for clarification, a variety of notes are available to the  editor. if the information is related to a particular linked source, then it is recommended that the note be attached to the source itself. when adding a note, the editor can also link a source to back up the note, which is then visible when clicking on the details tab. the following note categories (in alphabetical order) remain for narrative text that should be visibly associated with the taxon: a. altitude this note is for recording the altitude of a taxon. it is only used in certain global species databases which include non-marine taxa, e.g., molluscabase, or amphidromous and diadromous species that enter rivers, or landlocked populations in lakes, e.g., fishbase. b. depth this note is for recording a depth record or depth range of a taxon. c. description this note allows editors to input diagnoses for taxa. d. distribution this note is for any additional information on the distribution of a taxon that is not placed in the distribution module. e. etymology this note allows editors to input information regarding the etymology of a name. f. fossil range this note is for additional narrative information concerning the fossil range of a taxon; referring to presence in specific time intervals. g. lsid this is a place to provide a link to another database where life-science identifiers (lsids) are used. h. miscellaneous this note is for additional narrative information that does not fit in any of the other categories. i. nomenclature this note is for additional narrative information about the nomenclature of a taxon. it can include information on, e.g., original spellings and misspellings, homonymy, grammatical gender, authority and publication date, etc. j. stratigraphy this note is for additional narrative information concerning the stratigraphy (presence of fossil taxa in particular rock units). k. status this note category should be used to provide additional information regarding the status of a nominal taxon, for example, where it has been indicated in the literature as, e.g., nomen protectum, taxon inquirendum, nomen dubium, or has been suppressed or conserved by an iczn opinion, etc. horton t. et al., best practice guidelines for worms taxonomic editors 17 l. type material this note can be used to cover all narrative information regarding the type material of a nominal taxon, including notes on holotype, syntype, paratype, neotype and lectotype specimens, type designation, and museum deposition information. m. type species this note is for additional narrative information about the designation of type species that is not possible to indicate in the type species field. n. type locality this note is for additional narrative information about the type locality of a taxon, e.g. an incorrectly spelled or ambiguous type-locality descriptor could be input here. o. taxonomic remark this note is for any additional taxonomic remarks on a taxon. it can include information on synonymy, new combinations, classification and identification. entering type material and type locality data it is important to clarify here the difference between a type locality and a distribution. the type locality is a unique locality (more rarely localities) associated with the holotype, syntypes, neotype or lectotype of an available species-group name. a distribution is a property of a taxon and synthesises the type localities of all its synonyms and all published records. the type locality can be defined as the geographical and stratigraphical attributes of the name-bearing  type(s) stated when a new species is described. it is considered as specimen-related data. it is therefore strongly recommended that type material data (e.g., institution/collection, registration number, nature of type) and type locality data (e.g., latitude, longitude, depth, date) are entered using the specimen module. all data concerning name-bearing types should be entered in association with the original name in worms, not with a recombination. the information will then be automatically associated with all names linked to this original name. type localities should be entered as close to verbatim as possible from the original publication providing all available details, e.g., geographical place name, depth, latitude, longitude. the type locality entry must be based on its original literature source, which should be linked accordingly. type locality may change over time, being restricted by lectotype designation or superseded by neotype designation or iczn opinion. in these cases, further literature sources can be linked to these changes (in the appropriate field in the specimen module). see an example for a neotype entry here: http://www.marinespecies.org/aphia.php?p=specdetails&id=67634&tid=124332 the online input interface offers the option to enter coordinates for type localities in degrees-minutesseconds, which are then re-calculated to decimal degrees. when editors enter decimal degrees, they need to be aware of the use of the minus-sign and use it appropriately (east and north = ‘+’; west and south = ‘-‘). environment it is now mandatory when adding a new taxon to indicate the environment of the taxon. if the name has already been created, editors can add this information retrospectively. in worms, four flags are  available (marine, freshwater, brackish, terrestrial) and any combination of these can be checked. each flag has three options (yes, no and unknown). european journal of taxonomy 389: 1–24 (2017) 18 http://www.marinespecies.org/aphia.php?p=specdetails&id=67634&tid=124332 as a default, the database shows only marine and/or brackish water species based on the environment flags (where marine = ‘yes’ or ‘unknown’ and/or brackish = ‘yes’ or ‘unknown’). therefore, to ensure  that pure freshwater and terrestrial species are not shown on worms, it is important that ‘no’ is checked for both the marine and brackish environment boxes. the generally agreed definitions are: freshwater: species occur at a salinity range of 0–0.5 ppt brackish water: species occur at a salinity range of 0.5–30 ppt marine: species occur at salinities higher than 30 ppt further distinctions of salinities (e.g., oligohaline (0.5–5 ppt), mesohaline (5–18 ppt) and polyhaline (18–30 ppt)), can be added in a note on the taxon page, if needed. when entering more difficult-to-define environments such as for those species inhabiting strandline  habitats, which flags should be checked? some groups have chosen terrestrial only for such species.  in the world amphipoda database, this was agreed as follows: ‘beach hoppers’ (talitridae) recorded in the marine environment should be flagged ‘marine only’ as they depend on the marine environment.  ‘land  hoppers’  should  be  flagged  ‘terrestrial  only’ as they occur away from marine or freshwater sources and ‘freshwater hoppers’ should be flagged ‘freshwater only’ as they are recorded, e.g., on the  beaches of lakes and rivers and therefore depend on the freshwater environment. other taxonomic group editors may choose to combine environment flags, e.g., insects and birds. for seabirds, both marine and  terrestrial are checked, as all seabirds also depend on the land for reproduction. similarly, both ‘marine’ and ‘terrestrial’ flags are checked for certain insects that live on land but are truly dependent on the  marine environment for at least part of their life cycle. for freshwater species, similar combinations of ‘fresh’ and ‘terrestrial’ can be made. for anadromous/catadromous fish, both ‘marine’ and ‘fresh’ may  be checked. it is important that editors use these flags consistently within a taxon. higher classification a  higher  level  classification  of  all  living  organisms was published by ruggiero et al. (2015). this paper provides a management classification for all living organisms and will form the basis for the  catalogue of life higher classification. a comparison between this paper and the higher classification in  worms is being made. where inconsistencies appear, the dmt consults with the responsible editors to discuss if changes are needed. worms aims to reflect current knowledge and therefore will allow newer  publications to be applied to higher level taxa if the editor deems it necessary and scientific consensus  has been reached. if no editor is available for a group, the classification by ruggiero  et al. (2015) will be followed. ruggiero et al. (2015) cover the classification from superkingdom to order level.  below the rank of order (e.g., suborder, infraorder, parvorder, superfamily, family, subfamily, genus) the classification is the responsibility of the editor of the taxon. decisions about making changes at these levels may await scientific consensus if there is dispute or the  editor is uncertain of the validity of the proposed changes. in such cases it is recommended that the editor add a ‘taxonomic remark’ note indicating that the classification is currently under scrutiny and providing  one or more ‘classification source’ links to the relevant scientific literature for the user. an editor may  also contact the catalogue of life team for further discussion to update the management hierarchy. conclusion worms has now existed for a decade and has been edited by more than 300 taxonomic experts during this period. naturally, it could not be foreseen from the start how the data would be entered, which types of data would be needed and how best to ensure consistency. experience has shown that researchers horton t. et al., best practice guidelines for worms taxonomic editors 19 working with different groups of organisms follow different traditions for handling taxonomic data. differences exist not only between, e.g., botanists and zoologists, but also at the lower taxonomic level. where no guidance existed, editors have found their own way of doing things, which has resulted in the proliferation of a confusing variety of terms, each of them well motivated but essentially similar to others. the need for standardisation has become apparent in recent years and has now led to the restriction of data field inputs to a choice of pre-defined values (e.g., status of a name, reason for non-acceptance  or category of source or note). this streamlining of data terms has allowed for similar meanings to be grouped. unclear terms have been refined and clarified. at the same time, in working with such a diverse  team of editors and taxa, worms recognises that there will always be more complex and difficult cases  and therefore strives to allow editors the freedom needed to record and explain these. increasing usage and application of worms in other taxonomic databases, institutes, museums, herbaria, environmental agencies, fisheries industry and research, clearly demonstrates its usefulness. the revised priorities for  data entry and improved consistency will further strengthen its usage in the biodiversity informatics arena and places greater emphasis on the importance of expert knowledge as applied in worms. acknowledgements the authors would like to acknowledge the important contributions of past members of the steering committee and the past and present members of the worms editorial board in the development of the world register of marine species. the creation and continuous further development of the worms database  has  benefited  from  funding  from  numerous  projects,  most  recently  emodnet  biology  i  (mare/2008/03—lot 4 biology—contract no. si2.531562; 2009–2011), emodnet biology ii (mare/2012/10—lot 5 biology—contract no. si2.657705; 2012–2016) and lifewatch (2012–2016). the first author (th) is supported by the uk natural environment research council (nerc). references appeltans w., costello m.j., vanhoorne b., decock w., vandepitte l., hernandez f., mees j. & vanden berghe e. 2008. aphia for a world register of marine species (worms). in: mees j. & seys j. 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(eds). proceedings of the international marine data and information systems conference imdis-2008, athens, greece, 31 march–2 april 2008. ioc workshop report no. 219, paris. horton t. et al., best practice guidelines for worms taxonomic editors 23 https://doi.org/10.1046/j.1365-294x.2000.01022.x https://doi.org/10.1371/journal.pbio.1001127 https://doi.org/10.1186/s12862-017-0961-8 https://doi.org/10.1099/ijsem.0.000778 https://doi.org/10.1007/s12526-014-0255-y https://doi.org/10.17161/bi.v12i0.6522 https://doi.org/10.1371/journal.pone.0119248 http://www.marinespecies.org/ophiuroidea https://doi.org/10.3390/jmse3041448 welter-schultes f.w. 2012. guidelines for the capture and management of digital zoological names information. version 1.1 released on march 2013. global biodiversity information facility, copenhagen. available from http://www.gbif.org/document/80625 [accessed 5 dec. 2017]. manuscript received: 8 march 2017 manuscript accepted: 18 july 2017 published on: 28 december 2017 topic editor: koen martens desk editor: kristiaan hoedemakers printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, spain. european journal of taxonomy 389: 1–24 (2017) 24 http://www.gbif.org/document/80625 european journal of taxonomy 75: 1-123 issn 2118-9773 http://dx.doi.org/10.5852/ejt.2014.75 www.europeanjournaloftaxonomy.eu 2014 · david g. notton this work is licensed under a creative commons attribution 3.0 license. m o n o g r a p h urn:lsid:zoobank.org:pub:e02c736e-661d-407f-9322-24bfb95e233e 1 a catalogue of the types of diapriinae (hymenoptera, diapriidae) at the natural history museum, london david g. notton department of entomology, the natural history museum, cromwell road, london, sw7 5bd, united kingdom d.notton@nhm.ac.uk urn:lsid:zoobank.org:author:2e452ef2-703c-47c0-8432-6ab9c05ac06a abstract. the types of nominal species of diapriinae in the collection of the natural history museum, london, are catalogued. lectotypes are designated for the following taxa: diapria peraffinis ashmead, 1896; d. smithii ashmead, 1896; galesus bipunctatus ashmead,1894; g. (g.) foersteri var. nigricornis kieffer, 1911; g. sexpunctatus ashmead, 1893; g. walkeri kieffer, 1907; idiotypa nigriceps kieffer, 1909; i. nigriceps kieffer, 1911; i. pallida ashmead, 1893; i. pallida ashmead in riley, ashmead & howard, 1894; paramesius angustipennis kieffer, 1911; p. cameroni kieffer, 1911; phaenopria cameroni kieffer, 1911; p. halterata kieffer, 1911; p. magniclavata ashmead, 1896; tropidopsis clavata ashmead, 1893; t. clavata ashmead in riley, ashmead & howard, 1894. new combinations are proposed: aneuropria bifurcata comb. nov. for mantara bifurcata dodd, 1920; basalys quadridens comb. nov. for microgalesus quadridens kieffer, 1912; coptera cratocerus comb. nov. for galesus cratocerus cameron, 1912; coptera sexpunctata comb. nov. for galesus sexpunctatus ashmead, 1893; doliopria magniclavata comb. nov. for phaenopria magniclavata ashmead, 1896; spilomicrus aterrimus comb. nov. for hoplopria aterrima dodd, 1920; spilomicrus campbellanus comb. nov. for antarctopria campbellana yoshimoto, 1964; spilomicrus coelopae comb. nov. for antarctopria coelopae early, 1978; spilomicrus diomedeae comb. nov. for antarctopria diomedeae early, 1978; spilomicrus helosciomyzae comb. nov. for malvina helosciomyzae early & horning, 1978; spilomicrus insulae comb. nov. for malvina insulae early, 1980; spilomicrus latigaster comb. nov. for antarctopria latigaster brues in tillyard, 1920; spilomicrus punctatus comb. nov. for malvina punctata cameron, 1889; spilomicrus rekohua comb. nov. for antarctopria rekohua early, 1978; trichopria bouceki comb. nov. for oxypria bouceki masner, 1959; trichopria nigriceps comb. nov. for tropidopria nigriceps ashmead in riley, ashmead & howard, 1894; trichopria nigriceps comb. nov. for xyalopria nigriceps kieffer, 1907; trichopria spinosiceps comb. nov. for acidopria spinosiceps dodd, 1920; trichopria walkeri comb. nov. for diapria walkeri dalla torre, 1890. new replacement names are proposed: coptera mosselensis nom. nov. for c. nigricornis nixon, 1930 preocc.; coptera pijiguaorum nom. nov. for c. sexpunctata montilla & garcía, 2008 preocc.; spilomicrus kozlovi nom. nov. for s. punctatus kozlov, 1978 preocc.; trichopria fluminis nom. nov. for t. nigriceps (kieffer, 1907) preocc.; t. thermarum nom. nov. for t. nigriceps (kieffer, 1913) preocc. new specific synonyms are proposed: basalys cursitans (kieffer, 1911) = b. pedisequa (kieffer, 1911) syn. nov. (the former removed from synonymy with b. parvus thomson, 1858); b. iphicla nixon, 1980 = b. macroptera (kieffer, 1911) syn. nov.; coptera bipunctata (ashmead in riley, ashmead & howard, 1894) = c. sexpunctata (ashmead, 1893) syn. nov.; idiotypa nigriceps kieffer, 1911 = i. nigriceps kieffer, 1909 syn. nov.; i. pallida ashmead in riley, ashmead & howard, 1894 = i. pallida ashmead, 1893 syn. nov.; psilus nigricornis (kieffer, http://dx.doi.org/10.5852/ejt.2014.75 http://www.europeanjournaloftaxonomy.eu https://creativecommons.org/licenses/by/3.0/ http://zoobank.org/urn:lsid:zoobank.org:pub:e02c736e-661d-407f-9322-24bfb95e233e mailto:d.notton%40nhm.ac.uk?subject= http://zoobank.org/urn:lsid:zoobank.org:author:2e452ef2-703c-47c0-8432-6ab9c05ac06a european journal of taxonomy 75: 1-123 (2014) 2 1911) = p. fuscipennis (curtis, 1831) syn. nov.; p. walkeri (kieffer, 1907) = p. fuscipennis (curtis, 1831) syn. nov.; t. bouceki (masner, 1959) = t. conotoma (kieffer, 1911) syn. nov.; trichopria halterata (kieffer, 1911) = t. halterata (kieffer, 1909) syn. nov. new generic synonyms are proposed: antarctopria brues in tillyard, 1920 = spilomicrus westwood, 1832 syn. nov.; malvina cameron, 1889 = spilomicrus westwood, 1832 syn. nov.; mantara dodd, 1920 = aneuropria kieffer, 1905 syn. nov.; microgalesus kieffer, 1912 = basalys westwood, 1833 syn. nov.; xyalopria kieffer, 1907 = trichopria ashmead, 1893 syn. nov. (xyalopria is removed from synonymy with megaplastopria ashmead, 1903). a brief account of some aspects of the history of these types is given. keywords. hymenoptera, diapriidae, diapriinae, types, the natural history museum (london). notton d.g. 2014. a catalogue of the types of diapriinae (hymenoptera, diapriidae) at the natural history museum, london. european journal of taxonomy 75: 1-123. http://dx.doi.org/10.5852/ejt.2014.75 introduction diapriinae haliday, 1833 is a cosmopolitan subfamily of diapriid wasps including about 1000 described species (johnson 1992). most are pupal or puparial endoparasitoids of diptera or more rarely coleoptera or formicidae. diapriinae are often a major component of the microhymenopteran fauna attacking diptera in a range of habitats, but despite this they remain poorly known. the location and recognition of type specimens and their placement in currently recognised taxa is a necessary preliminary step to revisionary studies which, in order that they have lasting value, must be based on knowledge of species described already. this problem is particularly severe for neglected families such as the diapriidae where there is a large and fragmented literature, with the older descriptions not always mentioning type depositories. there are a number of publications which refer to types in the natural history museum, london (bmnh), most usefully masner (1965), which mentions types of 135 nominal species of diapriinae, and johnson’s (1992) world catalogue, which cites publications mentioning types. a number of types have been added to the bmnh collection since 1965 thanks to the generosity of lubomír masner and other colleagues, and some types have been discovered which were previously overlooked. the current study aims to catalogue all types of diapriinae in london in the context of the history of those types, modern assessments of type status and current ideas on classification of diapriinae, in order to make them accessible to future workers as a basis for future revisions of world diapriinae. the types of more than 250 nominal species are treated, a significant proportion of the world fauna when compared to the circa 1200 valid species listed in the last world catalogue (johnson 1992). the concept of diapriinae followed here is that of masner & garcía (2002) and notton (2004), including all the genera traditionally placed in psilini and some genera for which tribal placement is problematic. a brief history of some of the people associated with these types is given here. william harris ashmead (1855-1908) william ashmead, was assistant curator, division of insects at the united states national museum, (smithsonian institution, washington d.c.) and one of the foremost american workers of his time in the systematics of hymenoptera (anon. 1908; 1909; bethune 1908, 1909; crawford 1909; horn 1909; howard 1908; howard, crawford & banks 1909; semenov-tian-shanskii 1909; smith 1910). types of ashmead’s species of diapriinae found in the bmnh are from material collected by herbert huntingdon smith (1852-1919) in the west indies as part of a project coordinated by the west india exploration committee. the bmnh entomology registers include donations of numerous insects of all kinds from the committee and its agents; the record for lot 1899-331 shows the committee presented 2205 hymenoptera from st. vincent and grenada, including 610 species and 347 types. new taxa of hymenoptera were described in a series of papers, mainly by ashmead (ashmead 1893, 1896, 1900; howard 1897; riley, ashmead & howard 1894). it is likely that the intention of the committee was to notton d.g., diapriinae types at the natural history museum, london 3 deposit the majority of the specimens at the bmnh since the committee was a british endeavour set up by the royal society and the british association, and st vincent and grenada were longstanding british colonies at the time. published reports on the parasitica do not mention type repositories, except for ashmead (1893), which generally has series split between london and washington. on the basis of what has been found in the bmnh collection, unique specimens and most important individuals are generally in the bmnh, and any remaining specimens appear to have been split between london and washington in recognition of ashmead’s effort in working up the material, although only a few were recognised by masner & muesebeck (1968). ashmead did not designate holotypes, so type series are syntypic. where there are series, ashmead’s type labels cannot be taken as evidence that the specimens are holotypes, as he did not consistently label types or add determination labels. consequently, all available undetermined h.h. smith material has been assessed for type status, and it has become apparent that much previously unlabelled material in the bmnh is syntypic. unfortunately, much h.h. smith material was mounted on short bendy pins and a number of specimens have been damaged in the past as a result. many of these were repinned during the current project so they could be examined safely. lectotype designations have been based on bmnh specimens by previous authors, and it seems pragmatic to continue this, insofar as bmnh specimens are taxonomically suitable. peter cameron (1847-1912) peter cameron was an energetic amateur student of hymenoptera, describing many species despite meagre means and poor health, and donating much of his type material to the bmnh (anon. 1913; gmw 1913; morice 1913; morley 1913). cameron is an important figure not just because of the species of diapriinae he described himself, but also because he loaned material to kieffer (q.v. below) to describe, mainly from scotland and england. in his own descriptions, cameron did not always designate holotypes or give a clear indication of the numbers of specimens on which he based his descriptions (notton, buffington & van noort 2009; quinlan 1974); in these cases the specimen(s) have been treated as syntypes (iczn 1999: recommendation 73f) even when there is only one in the bmnh, since cameron is known to have dispersed syntype series. abbé jean-jacques kieffer (1856-1925) jean-jacques kieffer was a catholic priest and teacher at a secondary school in bitche in lorraine, northern france. in his spare time he published hundreds of taxonomic papers on parasitic hymenoptera and gall midges (cecidomyiidae) (nominé 1925, 1926; kelner-pillault 1958; gagné 1994; notton 2004; vlug 1995). as kieffer described many of the species covered in the catalogue below, it is worth mentioning the history of his types. kieffer based his descriptions on specimens in his own collection, but also on material borrowed from other collectors and museums. much of kieffer’s personal collection, at least the diapriidae, has survived at the muséum national d’histoire naturelle, paris, france, despite the doubt cast on the survival of kieffer material belonging to some other taxa (gagné 1994 and references therein; vlug 1995). in brief, p.l.g. benoit, head of the invertebrates section at the musée du congo belge at tervuren, found kieffer’s collection at the collège de bitche where kieffer taught. thanks to the recteur, r.p.p.j. schmitt, the collection was transferred to the entomology laboratory of the muséum in 1957 (kelner-pillault 1958). otherwise, types which kieffer described from material received from collectors or museums were usually returned to them and so their current location depends on the fate of their collection of origin. thus, many types can be traced where kieffer states the collector in original descriptions. for example, some of those based on du buysson and de gaulle material are now in paris, whereas those based on cameron specimens are in london (notton 1995, 2004 and this study). kieffer also exchanged specimens of myrmecophile hymenoptera with e. wasmann; hence, some syntype series of wasmann and kieffer myrmecophile diapriids are split between paris and maastricht (dessart 1975; notton 2004). european journal of taxonomy 75: 1-123 (2014) 4 many of kieffer’s diapriid types in bmnh were described from scottish and english specimens from the collection of peter cameron, q.v. above. cameron posted the specimens to kieffer, who numbered and described or identified them and sent them back. rather than adding determination labels, kieffer wrote lists of specimen numbers and identifications on postcards and sent them back to cameron separately. the specimens and postcards were given to the bmnh by cameron, with the addition of a list written by him of the species represented, and the specimens were registered as lots 1910-55 and 1910-302. the specimens can be recognised today by kieffer’s number labels and one of the two register numbers, and they may also have labels in cameron’s handwriting or identification labels added later by bmnh curators based on kieffer’s lists. the postcards sent by kieffer, together with secondary lists based on them prepared by cameron and bmnh curators, are now held by the bmnh entomology library (entomology accession registers and insect room lists: 29, 32, 82, 83, 84). all specimens which could be recognised from these lists were reassessed for their type status, giving most weight to kieffer’s original number labels and postcard lists. cameron had very poor handwriting, did not always label his specimens, and never labelled them with the country which created problems, both for him and for kieffer, when it came to reporting localities correctly in publications. the documentary evidence of individually numbered specimens and kieffer’s numbered lists make it certain that kieffer and cameron were working from the same material, and yet there are many differences between the localities reported by cameron (1910), the type localities given by kieffer for species described from cameron’s material and the actual localities written on the specimens concerned, where it has been possible to decipher them. it appears that kieffer assumed that most of cameron’s specimens came from scotland when in fact many really came from england. consequently, a number of type localities have been revised in the catalogue below where it is evident that the data on the specimens does not match the published type locality. even without the problems of cameron’s labels, it appears kieffer had a poor grasp of geography because the postcards he wrote to cameron at whitle, new mills, near stockport in england are addressed to scotland. a further complication for kieffer’s names is that cameron (1910) published many as nomina nuda before kieffer made them available. similarly, it appears that kieffer assumed that walker material of galesus walkeri was from england when it was actually from norway. another major group of kieffer types is based on material collected by the percy sladen trust expedition to the seychelles (kieffer 1912a; huggert 1979) and is split between the bmnh and the zoology museum, university of cambridge, cambridge, uk. the existence of the cambridge specimens only became apparent at a late stage in the preparation of this work, and unfortunately there was not time to include them in the current work. on the basis of dating evidence presented by evenhuis (1994), kieffer’s report on material from the sladen expedition in the transactions of the linnean society (kieffer 1912a) was published before his genera insectorum v. 124 (kieffer 1912b). this conclusion is supported by the fact that kieffer quoted page references from the transactions in genera insectorum. lastly, another problem with kieffer’s names is that he sometimes described the same species twice by mistake. in these cases, to avoid future nomenclatural problems lectotype designations are used here to ensure that one specimen is the primary type for both nominal species, thus creating objective synonymy. material and methods: notes on the recognition of types and the arrangement and format of the catalogue the status of nominal species represented by type material was considered and details of type material given. within the catalogue, original combinations are arranged alphabetically by author, then in order of publication date and page number. full label data are quoted for primary types, except where illegible ([illeg.]), slashes are used to indicate the end of a line (/), where a slash occurs in the label data this has been replaced by a dash (-), a semicolon is used between labels (;) and a full stop between notton d.g., diapriinae types at the natural history museum, london 5 mounts (.). specimen condition is noted where this may help in the recognition of primary types, as well as comparative notes on original descriptions. each specimen was assessed for type status. for difficult cases, the criteria of notton (2004) have been followed when recognising syntypes. earlier authors did not always designate holotypes or give a clear indication of the numbers of specimens on which they based descriptions; in these cases the specimen(s) have been treated as syntypes (iczn 1999: recommendation 73f) even when there is only one in the bmnh, since types series have often been dispersed. inevitably, some specimens were found to bear labels with incorrect type status. a degree of latitude was necessary when matching specimens with the descriptions of kieffer and ashmead, which are sometimes inaccurate. the species described by borgmeier (1939) from type series (i.e. two or more specimens) are all considered to be based on syntypes, even where these were labelled “paratype” or “cotype”, because he did not designate a holotype in the original publication, instead referring to all type specimens equally as “typos” (types). of course, the mention of the word “type” or equivalent expression on a label is not necessarily evidence that the specimen is fixed as a holotype (article 72.4.7). similarly, the species described by nixon (1930) from type series are all considered to be based on syntypes even where these were labelled by nixon as “type” or “paratype”, because he did not designate a holotype in the original publication. in cases where type series included both sexes, nixon labelled one specimen of each sex as “type”, so in this particular work he clearly did not use the word type consistently to mean holotype. the identity of each nominal species is given with species placed in currently recognised genera, using the generic keys of nixon (1980), masner & garcía (2002) and various other sources cited in notton (2004). antennal segments are referred to as follows: antennal segment 1 (scape) = a1; segment 2 (pedicel) = a2; segment 3 (flagellar segment 1) = a3; and so on. article numbers cited refer to the international code of zoological nomenclature (iczn 1999). lastly, a nomenclatural summary is given. nominal taxa mentioned in the catalogue are listed. more complete synonymies will be found in johnson (1992) and other published sources. the tribe psilini is available from fallén (1812) and although it predates diapriidae haliday, 1833, diapriidae retains priority (article 35.5) which, in the interest of stability, gives precedence to names in use at a higher rank before 1999. muesebeck & walkley (1956) and johnson (1992) have been consulted to confirm the type species of genera involved in new generic synonymy. diapriid genus names are often based on the ending –pria, of which the first was diapria latreille, 1796. the derivation of this name is from the greek διαπριω, to saw in two, a reference to the incision at the base of the large tergite. this name is based on a greek word latinized with a change of ending to the feminine latin gender ending –a, and so takes the feminine gender (iczn 1999: art. 30.1.3). later names ending in –pria are also based on πριων, to saw, by analogy with diapria and similarly have feminine gender. basalys westwood, 1833 is derived from greek βασις (foot) + αλυς (listlessness); an appropriate name for a wasp which will not sit still. since αλυς is masculine this genus has masculine gender (m.a.alonso-zarazaga pers. comm.; iczn 1999: article 30.1.2). westwood was somewhat of a classical linguist and commonly used greek words when composing the names of genera, so while the species of this genus usually have a distinct basal vein, linguistically basalys is not connected with the latin word basalis (basal) except possibly that westwood may have intended it as a homophonic pun. the latin epithets pedisequus (variant spelling pedissequus) a male foot servant or lackey and pedisequa (or pedissequa) a waiting-lady are nouns, and so do not decline. all names considered here which were originally described as varieties are deemed to be available at subspecific rank from their date of publication (article 45.6.4). all taxonomic acts are attributable to the author except for the new name coptera pijiguaorum nom. nov. which is attributable to notton, montilla and garcía. european journal of taxonomy 75: 1-123 (2014) 6 results type catalogue tropidopsis clavata ashmead, 1893: 402, pl. xvii, figs 2, 2a. valid name basalys clavatus (ashmead, 1893). summary of types lectotype ♀, bmnh number 9.916. lectotype here designated. primary type data st. vincent/ w.i./ h.h. smith/ 230; tropidopsis/ clavatus/ ♀ ashm.; w.indies/ 99-331. type locality west indies, st vincent. remarks this species was described from an unspecified number of females and males. although ashmead states that the types are in the smithsonian institution, washington d.c., only one male syntype is there (masner & muesebeck 1968) and a female syntype is in the bmnh. another specimen in the bmnh was labelled by ashmead as “var.” and is much darker than the description and so is not considered part of the type series. since the male syntype is a trichopria ashmead, 1893 and the female syntype is a basalys, the female is designated here as lectotype in order to stabilise the current generic placement established by masner and muesebeck. the lectotype is mounted on a card point, with the right antenna lost and left antenna lost beyond a8. idiotypa pallida ashmead, 1893: 403, 463, pl. 17, fig. 3. valid name idiotypa pallida ashmead, 1893. summary of types lectotype ♀, bmnh number 9.1004. lectotype here designated. paralectotype ♂. primary type data st. vincent/ w.i./ h.h. smith/ 158; idiotypa/ pallida/ ♀ ashm.; w.indies/ 99-331. type locality west indies, st vincent. remarks this species is named at the end of the generic description of idiotypa with a reference to the figure of the female; the figure serves as an indication to make this species name available (article 12.2.7). the explanation of the plates (ashmead 1893: 463) shows that ashmead saw both sexes and it is reasonable to suppose that the syntype series is the same as for idiotypa pallida ashmead in riley, ashmead & howard, 1894. the same specimen is designated as the lectotype for both i. pallida ashmead, 1893 and notton d.g., diapriinae types at the natural history museum, london 7 i. pallida ashmead in riley, ashmead & howard, 1894 q.v. to ensure objective synonymy. the lectotype is mounted on a card point, is missing part of the right hind leg, and has the wings stuck together. galesus sexpunctatus ashmead, 1893: 408, 463, pl. 17, fig. 6. valid name coptera sexpunctata (ashmead, 1893) comb. nov. summary of types lectotype ♀, bmnh number 9.1003, here designated. paralectotype ♂. primary type data st. vincent/ w.i./ h.h. smith; galesus/ bipunctatus/ ♀ ashm.; w.indies/ 99-331. type locality west indies, st vincent. remarks this species is named at the end of the generic description of galesus haliday in curtis 1829 with a reference to the figure of the female and male; the figure serves as an indication to make this species name available (article 12.2.7). although ashmead (1893) says this species was from st. vincent, in his study of st. vincent diapriids the following year, the only species of galesus mentioned is g. bipunctatus ashmead in riley, ashmead & howard, 1894. it is reasonable to assume that this is one and the same species redescribed under another name because: a) both species were described from st. vincent from type series including both sexes; b) both species are referred to as having six punctures; g. sexpunctatus in the specific epithet and g. bipunctatus in its original description which includes the phrase “vertex with six small punctures”; c) an examination of ashmead’s labels shows how the change of names could have happened as 6-punctatus and bipunctatus look quite similar in ashmead’s handwriting. minor differences between the figures of g. sexpunctatus in ashmead (1893) and the description of g. bipunctatus ashmead in riley, ashmead & howard (1894) may be discounted because the figures in ashmead (1893) are often inaccurate in details. it is reasonable therefore to regard the syntype series of g. sexpunctatus as the same as that of g. bipunctatus. the original description of g. bipunctatus was based on two females and two males, of which a pair was found in the bmnh. the same specimen is designated lectotype for both g. sexpunctatus ashmead, 1893 and g. bipunctatus ashmead in riley, ashmead & howard, 1894 q.v. to ensure objective synonymy. the lectotype is glued to a micropin and is entire. g. sexpunctatus belongs in the genus coptera. coptera sexpunctata montilla & garcía, 2008 is now a secondary junior homonym and the new replacement name coptera pijiguaorum notton, montilla & garcía nom. nov. is proposed. pijiguaorum is the genitive plural of a latinized form of the name of the pijiguaos people of venezuala. idiotypa pallida ashmead in riley, ashmead & howard, 1894: 243. valid name idiotypa pallida ashmead, 1893 syn. nov. summary of types lectotype ♀, bmnh number 9.1004, here designated. paralectotype ♂. european journal of taxonomy 75: 1-123 (2014) 8 primary type data st. vincent/ w.i./ h.h. smith/ 158; idiotypa/ pallida/ ♀ ashm.; w.indies/ 99-331. type locality west indies, st vincent. remarks although already described under the same name in 1893, ashmead redescribed this species as new in 1894, from two females and one male, of which a pair was found. it is reasonable to assume that the syntype series is the same for both i. pallida ashmead, 1893 and i. pallida ashmead in riley, ashmead & howard, 1894, and the same specimen is designated lectotype for both names to ensure objective synonymy. the lectotype is mounted on a card point, and is missing part of the right hind leg, and has the wings stuck together. hemilexis latipennis ashmead in riley, ashmead & howard, 1894: 244. valid name entomacis latipennis (ashmead in riley, ashmead & howard, 1894). summary of types holotype ♂, bmnh number 9.687, by monotypy. primary type data 1500 feet; st. vincent/ w.i./ h.h. smith; type/ h.t.; hemilexis/ latipennis/ ♂ type ashm./ unique; type lost and/ and absolutely no fragments in/ the drawer/ n.d.m.f. 23-iii-1977. type locality west indies, st vincent. remarks only the mount and labels of the holotype remain. it has been missing since before 1965 (masner 1965). on the basis of ashmead’s description there is no doubt that this species belongs to the genus entomacis. hemilexodes filiformis ashmead in riley, ashmead & howard, 1894: 244. valid name entomacis filiformis (ashmead, 1984). summary of types holotype ♂, bmnh number 9.917, by monotypy. primary type data leeward side/ st. vincent, w.i./ h.h. smith/ 201; w.indies/ 99-331; entomacis m/ filiformis ash. type locality west indies, st vincent. notton d.g., diapriinae types at the natural history museum, london 9 remarks the holotype is mounted on a card point with the left antenna missing beyond scape. tropidopsis clavata ashmead in riley, ashmead & howard, 1894: 245. valid name basalys clavatus (ashmead, 1893). summary of types lectotype ♀, bmnh number 9.916, here designated. primary type data st. vincent/ w.i./ h.h. smith/ 230; tropidopsis/ clavatus/ ♀ ashm.; w.indies/ 99-331. type locality west indies, st vincent. remarks this species was redescribed in error from the same type series as tropidopsis clavata ashmead, 1893 q.v. the lectotype of t. clavata ashmead, 1893 q.v. is also designated as lectotype of t. clavata ashmead in riley, ashmead & howard, 1894 to create objective synonymy. hence, the condition of the lectotype and generic placement of t. clavata ashmead, 1984 are the same as for t. clavata ashmead, 1893. paramesius thoracicus ashmead in riley, ashmead & howard, 1894: 245. valid name paramesius thoracicus ashmead in riley, ashmead & howard, 1894. summary of types syntypes 3 ♀♀, 2 ♂♂. primary type data leeward side/ st. vincent, w.i./ h.h. smith/ 201; paramesius/ thoracicus/ ♀ ashm.; w.indies/ 99331 (syntype ♀). st. vincent/ w.i./ h.h. smith/ 198; w.indies/ 99-331 (syntype ♀). leeward side/ st. vincent, w.i./ h.h. smith/ 201; w.indies/ 99-331 (syntype ♀). 1500 feet; st. vincent,/ w.i./ h.h. smith; paramesius/ thoracicus/ ♂ ashm.; w.indies/ 99-331 (syntype ♂). 1000 feet; st. vincent,/ w.i./ h.h. smith; w.indies/ 99-331 (syntype ♂). type locality west indies, st vincent. remarks this species was described from four females and four males, of which three females and two males were found. another female specimen labelled by ashmead as this species is from grenada, so is not syntypic. the three female syntypes are mounted on card points, the one with ashmead’s determination label is entire, the one numbered 198 is missing its head, the third is missing its head and right fore leg. the two male syntypes are glued to the points of micro-pins, both are entire. european journal of taxonomy 75: 1-123 (2014) 10 spilomicrus aneurus ashmead in riley, ashmead & howard, 1894: 246. valid name spilomicrus aneurus ashmead in riley, ashmead & howard, 1894. summary of types syntypes ♀, 3 ♂♂. primary type data st. vincent/ w.i./ h.h. smith; spilomicrus/ aneurus/ ♀ ashm.; w.indies/ 99-331 (syntype ♀). st. vincent/ w.i./ h.h. smith/ 237; spilomicrus/ aneurus/ ♂ ashm.; w.indies/ 99-331 (syntype ♂). sea level; st. vincent/ w.i./ h.h. smith; w.indies/ 99-331 (syntype ♂). 1000 feet; st. vincent/ w.i./ h.h. smith; w.indies/ 99-331 (syntype ♂). type locality west indies, st vincent. remarks this species was described from a female and five males, of which one female and three males were found. the female syntype is glued to a micropin and is entire; the male with ashmead’s determination label is mounted on a card point and is entire; the male labelled “sea level” is glued to a micropin and is missing the ends of both antennae; the male labelled “1000 feet” is micropinned and is entire. spilomicrus vulgaris ashmead in riley, ashmead & howard, 1894: 246, 247. valid name spilomicrus vulgaris ashmead in riley, ashmead & howard, 1894. summary of types syntypes 11 ♀♀, 16 ♂♂. primary type data mountain forest. 3000ft/ in rotting leaves, march; st. vincent/ w.i./ h.h. smith; spilomicrus/ vulgaris/ ♀ ashm.; w.indies/ 99-331 (syntype ♀). 2000 feet; st. vincent/ w.i./ h.h. smith; spilomicrus/ vulgaris/ ♀ ashm.; w.indies/ 99-331 (syntype ♀). 2000 ft; st. vincent/ w.i./ h.h. smith; w.indies/ 99-331 (syntypes 4♀♀). st. vincent/ w.i./ h.h. smith; w.indies/ 99-331 (syntypes 2♀♀, 5♂♂). st. vincent/ w.i./ h.h. smith/ 158; w.indies/ 99-331 (syntype ♀). st. vincent/ w.i./ h.h. smith/ 227; w.indies/ 99-331 (syntype ♀). st. vincent/ w.i./ h.h. smith/ 244; w.indies/ 99-331 (syntype ♀). leeward side/ st. vincent,/ w.i./ h.h. smith/ 45; spilomicrus/ vulgaris/ ♂ ashm.; w.indies/ 99-331 (syntype ♂). 1500 feet; st. vincent/ w.i./ h.h. smith; w.indies/ 99-331 (syntypes 6♂♂). 500 feet; st. vincent/ w.i./ h.h. smith; w.indies/ 99-331 (syntype ♂). sea level; st. vincent/ w.i./ h.h. smith; w.indies/ 99331 (syntype ♂). leeward side/ st. vincent, w.i./ h.h. smith/ 240; w.indies/ 99-331 (syntype ♂). st. vincent, w.i./ h.h. smith/ 241; w.indies/ 99-331 (syntype ♂). type locality west indies, st vincent. notton d.g., diapriinae types at the natural history museum, london 11 remarks this species was described from 14 females and 22 males, most of which were found. one specimen labelled by ashmead as this species is excluded from the type series because it is from grenada. most of the syntypes are entire. galesus bipunctatus ashmead in riley, ashmead & howard, 1894: 248. valid name coptera sexpunctata (ashmead, 1893) syn. nov. summary of types lectotype ♀, bmnh number 9.1003, here designated. paralectotype ♂. primary type data st. vincent/ w.i./ h.h. smith; galesus/ bipunctatus/ ♀ ashm.; w.indies/ 99-331. type locality west indies, st vincent. remarks the original description was based on two females and two males, of which a pair was found. as explained above it is reasonable to suppose that the syntype series of g. bipunctatus and g. sexpunctatus are exactly the same specimens. the same specimen is designated lectotype for both g. sexpunctatus ashmead, 1893 q.v. and g. bipunctatus ashmead in riley, ashmead & howard, 1894 to ensure objective synonymy. the lectotype is glued to a micropin and is entire. this species is a coptera, and coptera sexpunctata is a new combination. loxotropa thoracica ashmead in riley, ashmead & howard, 1894: 249. valid name basalys thoracicus (ashmead in riley, ashmead & howard, 1894). summary of types holotype ♀, bmnh number 9.740, by monotypy. primary type data leeward side/ st vincent, w.i./ h.h. smith/ 242; type/ h.t.; loxotropa/ thoracica/ ♀ type ashm./ unique. type locality west indies, st vincent. remarks the holotype is mounted on a card point, the head and tips of the fore wings are lost. tropidopria triangularis ashmead in riley, ashmead & howard, 1894: 249. valid name acanthopria triangularis (ashmead in riley, ashmead & howard, 1894). european journal of taxonomy 75: 1-123 (2014) 12 summary of types lectotype ♂, bmnh number 9.937, selected by masner (1965). primary type data st. vincent/ w.i./ h.h. smith; tropidopria/ triangularis/ ♂ ashm.; w.indies/ 99-331. type locality west indies, st vincent. remarks the lectotype is glued to a micropin, and is a little faded, with the left antenna missing beyond the scape. this species belongs to acanthopria ashmead, 1896 where it was placed by masner (1965). tropidopria nigriceps ashmead in riley, ashmead & howard, 1894: 249, 250. valid name trichopria nigriceps (ashmead in riley, ashmead & howard, 1894) comb. nov. summary of types syntypes 3 ♂♂. primary type data 1500 feet; st. vincent/ w.i./ h.h. smith; tropidopria/ nigriceps/ ♂ ashm.; w.indies/ 99-331 (syntype ♂). 1000 feet; st vincent/ w.i./ h.h. smith; w.indies/ 99-331 (syntype ♂). st vincent/ w.i./ h.h. smith/ 158; w.indies/ 99-331 (syntype ♂). type locality west indies, st vincent. remarks this species was described from a female and four males. the three male syntypes agree well with the description except that the scutellum is weakly to obscurely carinate; however, there is the superficial appearance of a strong carina because the cuticle is transparent so that a dark gap between the internal scutellar muscle blocks is visible in the place where a carina might be, so probably ashmead misinterpreted this character and this should not exclude these males from the type series. one additional female standing over this name is not considered syntypic as it disagrees with the description in several points, most notably the antennal club which is sub-four-segmented and has the apical four segments darkened. the syntype labelled “1500 feet” is glued on a micropin and the left and right fore legs and left mid leg are missing. the syntype labelled “1000 feet” is glued on a micropin and both antennae are missing beyond a4. the syntype numbered “158” is mounted on a card point, with the right antenna missing beyond a3 and the right hind leg missing. the new combination trichopria nigriceps (ashmead in riley, ashmead & howard, 1894) is a senior secondary homonym of trichopria nigriceps (kieffer, 1913), so kieffer’s species is given the new replacement name trichopria thermarum nom. nov. thermarum is the genitive plural of therma, latin for a thermal bath, or hot spring, and refers to the type locality of los baños. notton d.g., diapriinae types at the natural history museum, london 13 tropidopria pallida ashmead in riley, ashmead & howard, 1894: 249, 250. valid name trichopria pallida (ashmead in riley, ashmead & howard, 1894). summary of types syntypes 10 ♀♀, 3 ♂♂. primary type data st. vincent/ w.i./ h.h. smith; w.indies/ 99-331 (syntypes 5♀♀, ♂). st. vincent/ w.i./ h.h. smith/ 158; tropidopria/ pallida/ ♀ ashm.; w.indies/ 99-331 (syntype ♀). st. vincent/ w.i./ h.h. smith/ 158; w.indies/ 99-331 (syntypes 2♀♀). st. vincent/ w.i./ h.h. smith/ 201; w.indies/ 99-331 (syntype ♀). st. vincent/ w.i./ h.h. smith/ 6; w.indies/ 99-331 (syntype ♀). may; st. vincent/ w.i./ h.h. smith; tropidopria/ pallida/ ♂ ashm.; w.indies/ 99-331 (syntype ♂). st. vincent/ w.i./ h.h. smith/ 73; w.indies/ 99-331 (syntype ♂). type locality west indies, st vincent. remarks this species was described from twelve females and six males, of which ten females and three males were found. the three females labelled “158” are mounted on card points, two are entire and one is missing the antennal tips; female syntype labelled “201” is mounted on a card point, and is missing the antennal tips; five remaining female syntypes are glued to micropins, four are entire and one is missing its head; male syntype labelled “may” is glued to a micropin and is entire; male syntype labelled “73” is mounted on a card point and is entire; third male is glued to a micropin and is missing its front legs. diapria mellea ashmead in riley, ashmead & howard, 1894: 251. valid name trichopria mellea (ashmead in riley, ashmead & howard, 1894). summary of types syntypes 2 ♀♀, ♂. primary type data st. vincent/ w.i./ h.h. smith/ 158; diapria/ mellea/ ♀ ashm.; w.indies/ 99-331 (syntype ♀). st. vincent/ w.i./ h.h. smith/ 158; w.indies/ 99-331 (syntype ♀). windward side/ st. vincent, w.i./ h.h. smith; diapria/ mellea/ ♂ ashm.; w.indies/ 99-331 (syntype ♂). type locality west indies, st vincent. remarks this species was described from three females and two males, of which two females and one male were found. the female syntype with ashmead’s determination label is mounted on a card point and is entire. the other female syntype is mounted on a card point and is missing most of the antennae. the male european journal of taxonomy 75: 1-123 (2014) 14 syntype was glued to a micropin, and has evidently fallen off and been reglued on the same micropin at some point, so that the tip of the left antenna and tips of some of the legs are glued separately. trichopria insularis ashmead in riley, ashmead & howard, 1894: 251, 252. valid name trichopria insularis ashmead in riley, ashmead & howard, 1894. summary of types syntype ♀. primary type data st. vincent/ w.i./ h.h. smith; trichopria/ insularis/ ♀ ashm.; w.indies/ 99-331. type locality west indies, st vincent. remarks this species was described from one specimen of each sex. one female syntype was found; it is glued to the end of a micropin, and is entire. trichopria pleuralis ashmead in riley, ashmead & howard, 1894: 251, 252. valid name trichopria pleuralis ashmead in riley, ashmead & howard, 1894. summary of types syntype ♂. primary type data leeward side/ st. vincent, w.i./ h.h. smith/ 239; trichopria/ pleuralis/ ♂ ashm.; w.indies/ 99-331. type locality west indies, st vincent. remarks this species was described from one specimen of each sex, of which the male was found. the male syntype is mounted on a card point and has much of the right flagellum missing; it has a fine but distinct basal vein and triangular marginal vein and belongs to the genus basalys. it appears that the syntype series includes two species because ashmead says the female syntype has a four-segmented antennal club, so it is unlikely to be a basalys and more likely a trichopria. it will therefore be necessary at some point to designate a lectotype to determine the application of the name trichopria pleuralis; however, no designation is made here since designating the male would create homonymy elsewhere and change the established generic combination, so future revisers are recommended to locate and consider the female syntype if possible before resolving this problem. the established generic combination is retained for now. notton d.g., diapriinae types at the natural history museum, london 15 trichopria atriceps ashmead in riley, ashmead & howard, 1894: 251, 253. valid name trichopria atriceps ashmead in riley, ashmead & howard, 1894. summary of types syntypes ♀, ♂. primary type data leeward side/ st. vincent, w.i./ h.h. smith/ 72; trichopria/ atriceps/ ♀ ashm.; w.indies/ 99-331 (syntype ♀). st. vincent/ w.i./ h.h. smith/ 158; trichopria/ atriceps/ ♂ ashm.; w.indies/ 99-331 (syntype ♂). type locality west indies, st vincent. remarks this species was described from two females and one male. a pair of syntypes was found; the female syntype is mounted on a card and is entire; the male is mounted on a card point and is entire. phaenopria subclavata ashmead in riley, ashmead & howard, 1894: 253, 254. valid name trichopria subclavata (ashmead in riley, ashmead & howard, 1894). summary of types syntypes ♀, ♂. primary type data st. vincent/ w.i./ h.h. smith/ 238; phaenopria/ subclavata/ ♀ ashm.; w.indies/ 99-331 (syntype ♀). leeward side/ st. vincent, w.i./ h.h. smith/ 239; phaenopria/ subclavata/ ♂ ashm.; w.indies/ 99-331 (syntype ♂). type locality west indies, st vincent. remarks this species was described from two females and two males, of which a pair was found. thirteen other females standing over this name, collected by h.h. smith, are not syntypic as they are from grenada. both syntypes are mounted on card points: the female is missing the left flagellum and right fore leg; the male is entire. phaenopria simillima ashmead in riley, ashmead & howard, 1894: 253, 254. valid name trichopria simillima (ashmead in riley, ashmead & howard, 1894). summary of types syntype ♀. european journal of taxonomy 75: 1-123 (2014) 16 primary type data st. v., west/ indies, smith; phaenopria/ simillima/ ♀ ashm.; w.indies/ 99-331. type locality west indies, st vincent. remarks this species was described from one female and one male. one female syntype was found; this is glued to the end of a micropin and is entire. loxotropa pleuralis ashmead, 1896: 803. valid name basalys pleuralis (ashmead, 1896). summary of types holotype ♀, bmnh number 9.742, by monotypy. primary type data balthazar/ windward side/ grenada, w.i./ h.h. smith; type/ h.t.; loxotropa/ pleuralis/ ♀ type ashm. type locality west indies, grenada, balthazar. remarks this species was described from a single female. one specimen was found which has quite a small scutellar pit; however, it agrees well with the description in all other respects and is certainly the holotype. the holotype is mounted on a card and is entire. loxotropa grenadensis ashmead, 1896: 803. valid name basalys grenadensis (ashmead, 1896). summary of types holotype ♀, bmnh number 9.741, by monotypy. primary type data balthazar/ windward side/ grenada, w.i./ h.h. smith/ 20; type/ h.t.; loxotropa/ grenadensis/ ♀ type ashm. type locality west indies, grenada, balthazar. remarks this species was described from one female. the holotype is mounted on a card point and is entire. the type is a basalys with 12-segmented antenna, an abrupt three-segmented club, and fore wing with a basal vein. notton d.g., diapriinae types at the natural history museum, london 17 acanthopria crassicornis ashmead, 1896: 804. valid name acanthopria crassicornis ashmead, 1896. summary of types syntype ♀, bmnh number 9.691. syntypes 2 ♂♂. primary type data type/ h.t.; st. john’s riv./ leeward side/ grenada, w.i./ h.h. smith/ 19; acanthopria/ crassicornis/ ♀ type ashm. (syntype ♀). mount gay est./ leeward side/ grenada, w.i./ h.h. smith/ 26; type; acanthopria/ crassicornis/ ♂ type ashm.; w.indies/ 99-331 (syntype ♂). mount gay est./ leeward side/ grenada, w.i./ h.h. smith/ 25; w.indies/ 99-331 (syntype ♂). type locality west indies, grenada, balthazar and mount gay estate. remarks this species was described from one female and four males, of which one female and two males were found. the female differs from the description in that it does not have two faint grooves posteriorly on the mesonotum, but the mesoscutal setae are stuck down in a way which suggests this, so this most likely an error of observation on ashmead’s part and this specimen can still be considered syntypic. also this specimen is labelled st john’s river but still is considered syntypic since this river is near the mount gay estate. all three syntypes are mounted on card points, the female is entire, the male numbered “26” has the right hind leg missing, the male numbered “25” is entire. diapria smithii ashmead, 1896: 804, 805. valid name trichopria smithii (ashmead, 1896). summary of types lectotype ♀, bmnh number 9.710, here designated. paralectotype ♂. primary type data grenada, w.i./ h.h. smith/ 51; type/ h.t.; diapria/ smithii/ ♀ type ashm. type locality west indies, grenada. remarks this species was described from a female and a male specimen, both of which were found. the female is a trichopria and the male is an acanthopria. the female is designated as lectotype to maintain the current combination (kieffer 1916; masner 1965). the original description included the multiple original spellings smithii and smithi; smithii is to be considered the correct original spelling following the first reviser action of dalla torre (1898). consequently the spelling smithi is unavailable. european journal of taxonomy 75: 1-123 (2014) 18 diapria grenadensis ashmead, 1896: 804, 805. valid name trichopria grenadensis (ashmead, 1896). summary of types lectotype ♀, bmnh number 9.727, designated by masner (1965). paralectotypes 5 ♀♀, ♂. primary type data balthazar/ windward side/ grenada, w.i./ h.h. smith/ 20; lecto/ type/ c.m.; w.indies/ 99-331; diapria/ grenadensis/ ♀ type ashm.; selected as/ lectotype by/ l. masner xi.61. type locality west indies, grenada, balthazar. remarks this species was described from nine females and two males, of which six female and one male syntypes were found. masner (1965) designated a lectotype under the unnecessary replacement name trichopria (planopria) grenadicola kieffer, 1916. the lectotype is mounted on a card point and is entire; it is a trichopria. diapria peraffinis ashmead, 1896: 805, 806. valid name trichopria peraffinis (ashmead, 1896). summary of types lectotype ♂, bmnh number 9.709a, here designated. paralectotype ♀, bmnh number 9.709b. primary type data balthazar/ windward side/ grenada, w.i./ h.h. smith/ 40; type; diapria/ peraffinis/ ♂ type ashm.; w.indies/ 99-331. type locality west indies, grenada, balthazar. remarks this species was described from a female and two males, of which one female and one male were found. the female syntype is a doliopria kieffer, 1910 and the male syntype is a trichopria. the female syntype has only eleven antennal segments, not twelve as ashmead described it, however it is very small and ashmead most probably miscounted, otherwise it agrees with the description and bears ashmead’s type label, so is considered to be syntypic here. the male is designated as lectotype in order to maintain the current generic combination. the lectotype is mounted on a card point and is missing the apex of the right hind tarsus. notton d.g., diapriinae types at the natural history museum, london 19 diapria melanopleura ashmead, 1896: 805, 806. valid name trichopria melanopleura (ashmead, 1896) stat. rev. summary of types holotype ♂, bmnh number 9.726, by monotypy. primary type data balthazar/ windward side/ grenada, w.i./ h.h. smith; type/ h.t.; diapria/ melanopleura/ ♂ type ashm. type locality west indies, grenada, balthazar. remarks the holotype is mounted on a card and has the tip of right hind tarsus missing. this species was included in xyalopria kieffer, 1907 by masner (1965), then megaplastopria ashmead, 1903 by arias-penna (2003); however, this species does not belong to megaplastopria. while it has a slender propodeal spine, it lacks key characters of xyalopria such as the occipital spine and carina, grooved scape and flared corners of the metasomal tergites, and is returned here to trichopria. it belongs to the group of species in which the males have whorled hairs on the antennal flagellum. diapria unicolor ashmead, 1896: 805, 806. valid name trichopria unicolor (ashmead, 1896). summary of types holotype ♀, bmnh number 9.708, by monotypy. primary type data balthazar/ windward side/ grenada, w.i./ h.h. smith/ 38; type/ h.t.; diapria/ unicolor/ ♀ type ashm. type locality west indies, grenada, balthazar. remarks the holotype is mounted on a card point and is entire. ceratopria grenadensis ashmead, 1896: 807. valid name basalys grenadae (kieffer, 1912). summary of types syntype ♀, bmnh number 9.745. european journal of taxonomy 75: 1-123 (2014) 20 primary type data mount gay est./ leeward side/ grenada, w.i./ h.h. smith/ 25; type/ h.t.; ceratopria/ grenadensis/ ♀ type ashm.; w.indies/ 99-331. type locality west indies, grenada, balthazar and mount gay estate. remarks described from two females of which one syntype was found. the syntype is mounted on a card point and is entire. it is a basalys with the antenna 12-segmented, an abrupt three-segmented club, and fore wing with basal vein. ceratopria flavipes ashmead, 1896: 807. valid name basalys flavidipes (kieffer, 1912). summary of types holotype ♀, bmnh number 9.744, by monotypy. primary type data balthazar/ windward side/ grenada, w.i./ h.h. smith/ 40; type/ h.t.; ceratopria/ flavipes/ ♀ type ashm. type locality west indies, grenada, balthazar. remarks this species was described from a single female. the holotype is mounted on a card point and is entire. trichopria grenadensis ashmead, 1896: 807, 808. valid name trichopria ashmeadi kieffer, 1912. summary of types syntype ♀, bmnh number 9.712. syntype ♂. primary type data grand etang/ windward side/ 1900ft/ grenada, w.i./ h.h. smith/ 13; type/ h.t.; trichopria/ grenadensis/ ♀ type ashm. (syntype ♀). balthazar/ windward side/ grenada, w.i./ h.h. smith/ 37; type; trichopria/ grenadensis/ ♂ type ashm.; w.indies/ 99-331 (syntype ♂). type locality west indies, grenada, balthazar and grand étang. notton d.g., diapriinae types at the natural history museum, london 21 remarks this species was described from one female and two males, of which one female and one male syntype was found. both syntypes are mounted on card points, are entire, and both belong to trichopria. trichopria bifoveata ashmead, 1896: 808. valid name trichopria bifoveata ashmead, 1896. summary of types syntype ♀, bmnh number 9.711. primary type data balthazar/ windward side/ grenada, w.i./ h.h. smith/ 20; type/ h.t.; trichopria/ bifoveata/ ♀ type ashm. type locality west indies, grenada, balthazar. remarks this species was described from two females, of which one was found. the syntype is mounted on a card point, is entire, and has the wings matted and dirty. trichopria affinis ashmead, 1896: 808. valid name trichopria neotropica masner, 1965. summary of types holotype ♀, bmnh number 9.707, by monotypy. primary type data balthazar/ windward side/ grenada, w.i./ h.h. smith/ 20; type/ h.t.; trichopria/ affinis/ f type ashm. type locality west indies, grenada, balthazar. remarks this species was described from a single specimen. it is mounted on a card point and is entire. phaenopria nigricornis ashmead, 1896: 810. valid name trichopria obscura masner, 1965. summary of types holotype ♂, bmnh number 9.722, by monotypy. european journal of taxonomy 75: 1-123 (2014) 22 primary type data st. john’s riv./ leeward side/ grenada, w.i./ h.h. smith/ 16; type/ h.t.; phaenopria/ nigricornis/ ♂ type ashm. type locality west indies, grenada, st john’s river. remarks this species was described from one male. the specimen agrees well with the description in all respects, except that a4 is angulate; probably ashmead missed this character. it is mounted on a card point and is entire. phaenopria angulifera ashmead, 1896: 810 lines 13, 29. valid name trichopria angulifera (ashmead, 1896). summary of types holotype ♂, bmnh number 9.718, by monotypy. primary type data balthazar/ windward side/ grenada, w.i./ h.h. smith; type/ h.t.; phaenopria/ angulifera/ m type ashm. type locality west indies, grenada, balthazar. remarks described from one male specimen; it is mounted on a card, is entire, and the wings are matted. phaenopria grenadensis ashmead, 1896: 809 (as p. grenadensis), 810 line 44 (as p. angulifera [sic]). valid name trichopria confusa masner, 1965. summary of types syntype ♀, bmnh number 9.713. syntype ♀. primary type data balthazar/ windward side/ grenada, w.i./ h.h. smith; type/ h.t.; phaenopria/ grenadensis/ ♀ type ashm. (syntype ♀). balthazar/ windward side/ grenada, w.i./ h.h. smith/ 20.; w.indies/ 99-331 (syntype ♀). type locality west indies, grenada, balthazar and mount gay estate. notton d.g., diapriinae types at the natural history museum, london 23 remarks this species was described from six females, of which two syntypes were found. the specimen numbered 9.713 is mounted on a card and is missing part of the right fore leg; the other is mounted on a card point and is missing the tip of the right hind leg. both belong to trichopria. in his treatment of this species ashmead used the specific name grenadensis in the key (p. 809) and angulifera heading the description (p. 810 line 44 species number 3) as is evident from the correspondence between the characters used in the key and the description, hence p. angulifera ashmead, 1896: 810 line 44 and p. grenadensis are multiple original spellings. ashmead (1900) is the first reviser under art. 24.2.4, and he chose p. grenadensis as the correct original spelling. p. angulifera ashmead, 1896: 810 line 44 is thus an incorrect original spelling, has no separate availability and cannot enter into homonymy with p. angulifera ashmead, 1896: 810 lines 12, 29 (species number 2) q.v. above. phaenopria nigriclavata ashmead, 1896: 809, 811. valid name trichopria nigriclavata (ashmead, 1896). summary of types holotype ♀, bmnh number 9.721, by monotypy. primary type data chantilly est./ windward side/ grenada, w.i./ h.h. smith/ 14; type/ h.t.; phaenopria/ nigriclavata/ ♀ type ashm. type locality west indies, grenada, chantilly estate. remarks this species was described from one female. the holotype is mounted on a card point and is entire. phaenopria balthazari ashmead, 1896: 809, 811. valid name trichopria balthazari (ashmead, 1896). summary of types syntype ♀, bmnh number 9.719. primary type data balthazar/ windward side/ grenada, w.i./ h.h. smith/ 20; type/ h.t.; phaenopria/ balthazari/ ♀ type ashm. type locality west indies, grenada, balthazar. remarks this species was described from two females, of which one was found. the syntype is mounted on a card point and is entire. european journal of taxonomy 75: 1-123 (2014) 24 phaenopria magniclavata ashmead, 1896: 809, 812. valid name doliopria magniclavata (ashmead, 1896) comb. nov. summary of types lectotype ♀, bmnh number 9.720, here designated. paralectotype ♀. primary type data grand etang/ windward side 1900ft./ grenada, w.i./ h.h. smith/ 13; type/ h.t.; phaenopria/ magniclavata/ ♀ type ashm. type locality west indies, grenada, grand étang. remarks this species was described from three females, of which two were found. the two syntypes differ from the description in having an eleven-segmented antenna and belong to doliopria; however, this discrepancy is accounted for by the fact that the antennae are covered with glue which obscures the number of segments and there is no reason not to consider them syntypes. a lectotype is designated in order to stabilise the new combination. the lectotype is mounted on a card point and is entire. zacranium oahuense ashmead, 1901: 295. valid name zacranium oahuense ashmead, 1901. summary of types holotype ♀, bmnh number 9.680, by monotypy. primary type data 29; type/ h.t.; waianae mts.,/ oahu, ft/ perkins. 4.1892; sandwich is./ 1912-215; zacranium/ oahuense/ ♀ type ashm. type locality hawaii, oahu, waianae mountains. remarks this species was described from one female. the holotype agrees well with the description and has ashmead’s type label. although ashmead says it was collected in june, the label says april, an error on ashmead’s part. the holotype is mounted on a card and is entire. zacranium ashmead, 1901 has a slender fore tibial spine and is most probably allied to the diapria genus group. platymischoides molokaiensis ashmead, 1901: 296. valid name platymischoides molokaiensis ashmead, 1901. notton d.g., diapriinae types at the natural history museum, london 25 summary of types syntype ♀, bmnh number 9.679. syntype ♀. primary type data 163; type/ h.t.; molokai mts.,/ ?4000 ft+/ perkins ix1893.; sandwich is./ 1912-215; platymischoides/ molokaiensis/ ♀ type ashm. (syntype ♀). hawaii: molokai/ mountains/ ?4000 ft+: r.c.l./ perkins: ix.1893; sandwich isles/ bmnh(e)/ 1912-215; ♀ platymischoides/ molokaiensis ashm./ type det. ashmead (syntype ♀). type locality hawaii, molokai. remarks described from three females, of which two were found, agreeing with the description except they were collected in september, not october. this is probably an error on ashmead’s part as this part of the label is not as clear as it could be. the two syntypes were originally mounted together on the same card, but one has been remounted on a card point; the specimen still on the original card has a red ink circle around it and has the left apical flagellomere separated but present; the other specimen is entire. platymischoides ashmead, 1901 and platymischodes ashmead, 1901 are multiple original spellings. in choosing platymischoides, ashmead (1903) appears to be the first reviser and platymischodes is consequently unavailable (article 24.2.4). phaenopria hawaiiensis ashmead, 1901: 296. valid name trichopria hawaiiensis (ashmead, 1901). summary of types syntype ♀, bmnh number 9.723. syntype ♂. primary type data type/ h.t.; molokai mts.,/ 3000ft./ perkins 19 & 20ix1893.; sandwich is./ 1912-215; phaenopria/ hawaiiensis/ ♀ type (syntype ♀). type/ molokai mts.,/ 8000ft./ perkins 19 & 29ix1893; sandwich is./ 1912-215; phaenopria/ hawaiiensis/ ♂ type ashm. (syntype ♂). type locality hawaii, molokai. remarks this species was described from one female and one male. although ashmead says july, both specimens were collected in september. this is probably an error on ashmead’s part since the dates on the labels are almost illegible. similarly, although ashmead says 3500 ft, one specimen is labelled 3000 ft (or possibly 5000 ft) and the other 8000 ft; again the altitudes are written on the labels in small, barely legible handwriting, and in one case the number is transfixed by the pin, so probably this was an error in transcription on ashmead’s part. the syntypes are both carded: the female is entire; the male is missing its left antenna beyond a5 and its right mid and hind legs beyond the coxae. european journal of taxonomy 75: 1-123 (2014) 26 mimopria comes borgmeier, 1939: 532, figs 1, 13. valid name mimopria comes borgmeier, 1939. summary of types syntype ♀. primary type data campinas, goiaz/ schwarzmaier/ 24.t.30/ e. crassicorne; cotypus; mimopria/ comes borgm./ paratype/ det. borgmeier; brit. mus./ 1950-553. type locality brazil, goiás, campinas and inhumas; são paulo, barrettos; pernambuco, tapera. remarks this species was described from “numerous” females. other syntypes were apparently in borgmeier’s collection. one female was found agreeing with borgmeier’s description, and although this specimen is labelled as a “paratype” and “cotype”, since neither a unique holotype or lectotype has yet been fixed, the type series is still syntypic (article 73.2). the syntype is mounted on a card point and has the head missing. mimopria barbata borgmeier, 1939: 534, figs 14, 19. valid name mimopria barbata borgmeier, 1939. summary of types syntype ♀. primary type data eciton/ crassicorne; campinas/ goiás 9.iii.36/ schwarzmaier; cotypus; mimopria/ barbata borg./ paratype/ det. borgmeier; brit. mus./ 1950-553. type locality brazil, goiás, campinas. remarks this species was described from ten females of which one was found agreeing with borgmeier’s description, and although this specimen is labelled as a “paratype” and “cotype”, since neither a unique holotype or lectotype has yet been fixed, all of the type series is still syntypic (article 73.2). the syntype is mounted on a card point and entire. mimopria splendens borgmeier, 1939: 534, figs 1, 15. valid name mimopriella splendens (borgmeier, 1939). notton d.g., diapriinae types at the natural history museum, london 27 summary of types syntype ♀. primary type data campinas, goyaz/ e. göldii 756/ 15.4.36; cotypus; mimopria/ splendens b./ paratype/ det. borgmeier; brit. mus. 1950-553. type locality brazil, goiás, campinas and trindade. remarks this species was described from seven females of which one was found agreeing with borgmeier’s description, and although this specimen is labelled as a “paratype” and “cotype”, since neither a unique holotype or lectotype has yet been fixed, all of the type series is still syntypic (article 73.2). the syntype is micropinned and entire. this species was transferred to mimopriella masner & garcía, 2002 by masner & garcía (2002). philolestes pronotalis borgmeier, 1939: 536, figs 5, 11, 18. valid name notoxoides pronotalis (borgmeier, 1939). summary of types syntype ♀. primary type data eciton/ dulcius; campinas/ goiás 5.xi.33/ schwarzmaier; cotypus; philolestes/ pronotalis borgm./ paratype/ det. borgmeier; brit. mus./ 1950-553. type locality brazil, goiás, campinas. remarks this species was described from numerous dealate females, one alate female and four males of which one female was found agreeing with borgmeier’s description. although this specimen is labelled as a “paratype” and “cotype”, since neither a unique holotype or lectotype has yet been fixed, all of the type series is still syntypic (article 73.2). the syntype is micropinned, is missing its head, and is mounted together with an ant. philolestes kieffer, 1922 was synonymized with notoxoides ashmead, 1903 by masner (1977). notoxopria pedissequa borgmeier, 1939: 538, figs 2, 12. valid name notoxoides pedissequa (borgmeier, 1939). summary of types syntype ♀. european journal of taxonomy 75: 1-123 (2014) 28 primary type data campinas, goyas/ schwarzmaier/ 17.xii.28/ e. pseudops; cotypus; notoxopria/ pedissequa/ borgm. paratype/ det. borgmeier; brit. mus. 1950-553. type locality brazil, goiás, campinas. remarks this species was described from numerous dealate females and one alate female, of which one was found agreeing with borgmeier’s description. although this specimen is labelled as a “paratype” and “cotype”, since neither a unique holotype or lectotype has yet been fixed, all of the type series is still syntypic (article 73.2). the syntype is micro-pinned and entire and it is mounted together with an ant. notoxopria kieffer, 1910 was synonymized with notoxoides by masner (1977). the epithet “pedissequa” is a noun in apposition and is indeclinable. asolenopsia schwarzmaieri borgmeier, 1939: 542, fig. 10. valid name asolenopsia schwarzmaieri borgmeier, 1939. summary of types syntype ♀. primary type data campinas/ goyaz/ e. pseudops 686/ 4.2.36; cotypus; campinas/ goiás/ schwarzmaier; asolenopsia/ schwarzmaieri/ borgm. paratype/ det. borgmeier; brit. mus./ 1950-553. type locality brazil, goiás, campinas. remarks this species was described from numerous females, of which one was found agreeing with borgmeier’s description. although this specimen is labelled as a “paratype” and “cotype”, since neither a unique holotype or lectotype has yet been fixed, all of the type series is still syntypic (article 73.2). the syntype is mounted on a card and is entire. neivapria penicillata borgmeier, 1939: 544, figs 3, 4, 6, 7. valid name neivapria penicillata borgmeier, 1939. summary of types syntype ♀. primary type data campinas, goyas/ schwarzmaier/ t22 mimense/ 11.3.36; cotypus; neivapria/ penicillata/ borgm. paratype/ det. borgmeier. notton d.g., diapriinae types at the natural history museum, london 29 type locality brazil, goiás, campinas. remarks this species was described from numerous females of which one was found agreeing with borgmeier’s description. although this specimen is labelled as a “paratype” and “cotype”, since neither a unique holotype or lectotype has yet been fixed, all of the type series is still syntypic (article 73.2). the syntype is mounted on a card point and the head is lost. loxotropa fuliginosi box, 1921: 16. valid name basalys pedisequa (kieffer, 1911). summary of types syntype ♀, bmnh number 9.762. primary type data loxotropa/ fuliginosi/ woking/ 30.v.20; assoc. with/ lasius fuliginosus; ex coll./ donisthorpe/ b.m.1934-4; loxotropa/ fuliginosi box; holo-/ type. type locality england, woking. remarks this species was described from an unspecified number of females from donisthorpe’s collection. the sole surviving syntype is mounted on a card and is entire. nixon’s (1980) interpretation of basalys pedisequa (kieffer, 1911) is followed here. aulatopria tucumana brèthes, 1927a: 164. valid name bruchopria tucumana (brèthes, 1927). summary of types syntypes ♀, ♂. primary type data reared; rosenfeld y/ barber, colls.; est. exp./ a.c. no. 115; tucumán/ i-22-1913; aulatopria/ tucumana brèthes; pres. by imp. bur. ent./ brit. mus./ 1928-347 (syntypes ♀, ♂). type locality argentina, tucumán. remarks the description for this species was published twice, in the revista industrial y agrícola de tucumán (brèthes 1927a) and translated into english in the bulletin of entomological research (brèthes 1927b). it is most probable that the revista came first, as this work was published every two months at that european journal of taxonomy 75: 1-123 (2014) 30 time; volume 17(7-8) contains weather reports up to 31 january 1927 and was probably published shortly afterwards, whereas the english translation in the bulletin has the cover date december 1927. brèthes mentions that the specimens were reared during studies of the sugar-cane moth borer diatraea saccharalis fabricius, 1794 by rosenfeld and barber and published in 1914 also in revista industrial y agrícola de tucumán. rosenfeld and barber’s paper could not be traced during this study, but is mentioned here since it may throw further light on the composition of the type series. two syntypes were found which had been reared by rosenfeld and barber in 1913, labelled by brèthes, and received at the imperial bureau of entomology in london, shortly after the date of the description. both syntypes are mounted on card points and are somewhat dirty: the female is missing the left fore wing and has the right fore wing chipped; the male is entire. this species is now placed in bruchopria kieffer, 1921 following masner & garcía (2002). paramesius fasciatipennis cameron, 1888: 437, pl. 18, fig. 18. valid name spilomicrus fasciatipennis (cameron, 1888). summary of types syntype ♀, bmnh number 9.639. syntypes ♀, ♂. primary type data bugaba,/ panama/ champion; type/ h.t.; b.c.a. hymen. i./ paramesius/ fasciatipennis/ cam.; paramesius/ fasciatipennis/ cam. type (syntype ♀). fasciati-/ pennis; v.de chiriqui/ 25-4000ft/ champion; p.cameron coll./ 1914-110 (syntype ♀). maculi-/ fasciati-/ pennis; v.de chiriqui/ 25-4000ft/ champion; p.cameron coll./ 1914-110 (syntype ♂). type locality panama, bugaba, volcán de chiriquí. remarks this species was described from at least one specimen of each sex. two female and one male syntypes were found. cameron appears to have later changed his mind about the identification of the male, crossing out the name fasciatipennis and replacing it with maculipennis; however, this specimen is clearly the male referred to as it agrees so well with the description, and not with that of maculipennis. another female labelled “fasciatipennis” differs significantly from the description in having shallow notauli, a shorter petiole, different antennal proportions, the altitude is wrong, and it is not considered syntypic. all three syntypes are mounted on cards and are entire. paramesius maculipennis cameron, 1888: 438, pl. 18, figs 11-12. valid name spilomicrus maculipennis (cameron, 1888). summary of types syntype ♀, bmnh number 9.640b. primary type data fasciati/ maculi-/ pennis; bugaba/ panama/ champion; p.cameron coll./ 1914-110 (syntype ♀). notton d.g., diapriinae types at the natural history museum, london 31 type locality panama, bugaba. remarks this species was described from at least one specimen of each sex. one female was found which agreed exactly with the description and had been labelled by cameron as “maculipennis”. the male and female, mounted together, from vera paz formerly regarded as types (masner 1965) are not considered syntypic because they are not from the type locality. a fourth specimen labelled “maculipennis” did not agree with the description and is considered here to be a syntype of p. fasciatipennis q.v. which cameron later incorrectly reidentified. the syntype is mounted on a card and is entire. paramesius canaliculatus cameron, 1888: 439, pl. 18, fig. 24. valid name spilomicrus canaliculatus (cameron, 1888). summary of types syntype ♀, bmnh number 9.638. primary type data cordova; mexico/ salle coll.; type/ h.t.; b.c.a. hymen. i/ paramesius/ canaliculatus/ cam. type. type locality mexico. remarks this species was described from an unspecified number of females, of which one syntype was found. the syntype is mounted on a card and is entire. paramesius chiriquensis cameron, 1888: 439, pl. 18, fig. 15. valid name pentapria chiriquensis (cameron, 1888). summary of types syntype ♀, bmnh number 9.641. primary type data v.de chiriqui/ 8000ft/ champion; b.c.a. hymen. i./ paramesius/ chiriquensis/ cam.; type/ h.t.; paramesius/ chiriquensis/ p.439 cam. type locality panama, volcán de chiriquí. remarks this species was described from an unspecified number of females of which one was found. the syntype agrees with the description except that cameron miscounted the number of keels on the petiole there are seven. the syntype is mounted on a card and is entire. european journal of taxonomy 75: 1-123 (2014) 32 spilomicrus tinctipennis cameron, 1888: 440, pl. 18, fig. 17. valid name idiotypa tinctipennis (cameron, 1888). summary of types holotype ♀, bmnh number 9.747, by monotypy. primary type data bugaba/ panama/ champion; b.c.a. hymen. i./ spilomicrus/ tinctipennis/ cam.; type/ h.t.; spilomicrus/ tinctipennis/ cam./ b.c.a. ii 440. type locality panama, bugaba. remarks the description of this species was based on an unspecified number of females. one specimen was found which agreed with the description, except that it did not have a blunt, projecting, basal tooth on the hind femur; however, a piece of dirt was found on the base of the right hind femur which gave the appearance of a tooth (the base of the left femur was obscured). it is therefore considered that cameron’s description is incorrect on this point, and should not prevent the specimen from being a type. it is most unlikely there would be another specimen with a piece of dirt on the base of the hind femur giving the appearance of a tooth, so it is considered that this was the only specimen cameron saw and is therefore a holotype by monotypy. the holotype is mounted on a card and is dirty with the left wings crumpled. the piece of dirt which gave the appearance of a tooth on the femur has been removed, but the specimen is now clearly labelled so it can still be recognised. malvina punctata cameron, 1889: 13. valid name spilomicrus punctatus (cameron, 1889) comb. nov. summary of types syntype ♀, bmnh number 9.681. syntypes 3 ♀♀. primary type data type; greymouth; cameron coll./ 98-124; malvina/ punctata/ cam. type/ new zealand (syntype ♀). n.zeal. (syntype ♀). cameron./ 99-30.; n.zeal. (syntypes 2♀♀). type locality new zealand, greymouth. remarks the description was made from an unspecified number of females, certainly more than one since cameron says “coxae usually black”, and since the specimen labelled by cameron as type has the petiole obscured by its wings he must have seen another to describe the petiole. besides the specimen labelled by cameron as a type, three other specimens were found in the collection, two with register labels saying they were from cameron’s collection, and the third with a similar mounting style and locality notton d.g., diapriinae types at the natural history museum, london 33 apparently written in cameron’s hand. all four are considered syntypic. in a later paper cameron (1898) noted there were four specimens in this collection. the syntype with register number 98-124 is mounted on a card and is missing its left hind leg; the syntype without a register number is mounted on a card point and is missing the right antenna beyond a2; of the two syntypes with register number 99-30, one is mounted on a card point and is missing parts of all the tarsi and the other is mounted on a card and is missing the right hind tarsus. malvina punctata appears to be a part of a small group of related species with pronounced punctation and sometimes striation on the head and mesosoma; these appear to be little more than a derived species group of spilomicrus westwood, 1832. cameron (1889) noted that this species has a metanotal spine, which is a rare character in diapriinae; however, it is not the metanotum, but the anterior propodeum which is spinose, a feature found in some other spilomicrus. this species is transferred to spilomicrus here. since m. punctata is the type species of malvina cameron, 1889, this genus becomes a synonym of spilomicrus syn. nov. the pronotal groove, which is supposed to be a feature of malvina according to early (1980), is virtually absent in the females of the type species and is found in some other species of spilomicrus which had not been included in malvina. the other species currently placed in malvina can be accounted for as follows: m. quadriceps (smith, 1878) is returned to spilomicrus stat. rev., q.v. below; m. helosciomyzae early & horning, 1978 and m. insulae early, 1980 are both transferred to spilomicrus (new combinations). spilomicrus punctatus (cameron, 1889) comb. nov. is a senior secondary homonym of spilomicrus punctatus kozlov, 1978; the latter is given the replacement name spilomicrus kozlovi nom. nov. loxotropa tricarinata cameron, 1912: 69. valid name trichopria tricarinata (cameron, 1912). summary of types holotype ♀, bmnh number 9.751, by monotypy. primary type data type; z13; kuching/ j.h./ 100; p.cameron coll./ 1914-110; loxotropa/ 3-carinata/ cam. type/ borneo. type locality malaysia, borneo, sarawak, kuching. remarks the description of this species was made from at least one female. the type specimen agrees broadly with the description; there are a number of errors in the description, in particular cameron’s erroneous observation of a non-existent basal cell in the fore wing, which may be explained by the fact that the venter, metapleura, propodeum, petiole and wings are all covered with glue and the wings are folded together, obscuring part of the body. since the errors of observation relate to the particular condition of this specimen it is considered that cameron based his description only on this specimen; hence it is a holotype by monotypy. the holotype is mounted on a card and is entire. galesus cratocerus cameron, 1912: 69. valid name coptera cratocerus (cameron, 1912) comb. nov. european journal of taxonomy 75: 1-123 (2014) 34 summary of types syntype ♀, bmnh number 9.672. primary type data type; kuching/ j.h.; p.cameron coll./ 1914-110; galesus/ cratocerus/ cam. borneo. type locality malaysia, borneo, sarawak, kuching. remarks this species was described from an unspecified number of females. one syntype was found, although there may have been others as cameron made detailed observations on the length of the hairs on the antennae, propodeum and petiole, which in the surviving syntype are all covered with glue. the syntype is mounted on a card, much matted with glue, right antenna missing beyond a5, and left antenna obscured by glue but probably missing beyond the scape. hoplopria curvispina cameron, 1913: 136. valid name spilomicrus curvispina (cameron, 1913). summary of types syntype ♂, bmnh number 9.918. primary type data type; 359; p.cameron coll./ 1914-110; hoplopria/ curvispina/ cam. type/ br. guiana. type locality guyana (as british guiana). remarks this species was described from an unspecified number of males. one male was found. reference to the “male” on p. 135 could equally mean the male sex, so is not clear evidence that there was only one specimen, and the type is treated here as a sole surviving syntype. the syntype is mounted on a card, with the head reglued upside-down, the left hind wing has become detached and is now stuck to the scapes, and the left antenna beyond a6 and parts of the left fore and mid tarsi are missing. the name curvispina does not change its ending in the new combination since it can be considered based on the latin noun spina, meaning a thorn (article 31.2.2). hoplopria picicornis cameron, 1913: 136. valid name spilomicrus picicornis (cameron, 1913). summary of types syntypes 2 ♀♀, bmnh number 9.647. notton d.g., diapriinae types at the natural history museum, london 35 primary type data type; 358; p.cameron coll./ 1914-110; hoplopria/ picicornis/ cam. type/ br. guiana (syntypes 2 ♀♀). type locality guyana (as british guiana). remarks the description is based on an unspecified number of females; although cameron refers to “the female” on p. 135, this could refer to the female sex and is not clear evidence for a unique type. two female syntypes were found, mounted on the same card, the left specimen is entire, and the right specimen is missing its head and the tip of the right hind tarsus. hoplopria aterrima dodd, 1920: 373. valid name spilomicrus aterrimus (dodd, 1920) comb. nov. summary of types holotype ♀, bmnh number 9.635, by original designation. paratype ♂. lectotype designation by masner (1965) invalid. primary type data lecto/ type/ l.m.; kuching/ j.h.; p.cameron coll./ 1914-110; brachyaulax/ striaticollis/ cam. type/ borneo; hoplopria/ aterrima dodd/ ♀; selected as lectotype by/ l. masner 25.xi.01. type locality malaysia, borneo, sarawak, kuching. remarks this species was described from one specimen of each sex, of which dodd said one was type and the other cotype (effectively holotype and paratype). there is, however, no clear evidence to say which of the two specimens is the type either from dodd’s description or labels. masner (1965) tried to resolve the situation by designating a lectotype; however, this action is invalid since only syntypes can be designated as lectotypes and dodd had already designated a holotype. it is prudent to recognise the female specimen as dodd’s holotype because a) dodd described the female first and in more detail and so it is most likely he considered the female specimen the type, and b) it is in the interests of stability to follow masner in recognising the female as the name bearing type. the holotype is mounted on a card and is entire. following the synonymy of hoplopria ashmead, 1893 with spilomicrus presented by masner & garcía (2002), the position of this species in spilomicrus is confirmed. hoplopria obsoleta dodd, 1920: 375. valid name spilomicrus obsoletus (dodd, 1920). summary of types holotype ♀, bmnh number 9.636, by monotypy. european journal of taxonomy 75: 1-123 (2014) 36 primary type data omilteme/ guerrero/ 8000ft./ aug. h.h. smith; godman-salvin/ coll. 1904.-1.; type/ h.t.; hoplopria/ obsoleta/ dodd ♀. type locality mexico, guerrero, omilteme. remarks this species was described from one female. the holotype is mounted on a card with the right fore and hind wings missing. hoplopria affinis dodd, 1920: 375. valid name spilomicrus affinis (dodd, 1920). summary of types holotype ♀, bmnh number 9.637, by original designation. paratype ♀. primary type data teapa/ tabasco/ march. h.h.s.; godman-salvin/ coll. 1904.-1.; type/ h.t.; hoplopria/ affinis/ dodd ♀. type locality mexico, tabasco, teapa. remarks this species was described from two females, which dodd designated type and cotype, of which both were found. the holotype is mounted on a card and has part of the left hind tarsus missing. hemilexis paucisetis dodd, 1920: 377. valid name spilomicrus paucisetis (dodd, 1920). summary of types holotype ♀, bmnh number 9.724, by original designation. primary type data mt. wellington,/ s.tasmania./ 12-21 mch. 1913.; 1,300 to 2,300 ft./ r.e.turner./ 1913-212.; hemilexis/ paucisetis/ dodd ♀. type locality australia, tasmania, mount wellington. remarks the holotype is mounted on a card point, with the head, left fore wing and part of right hind tarsus missing. notton d.g., diapriinae types at the natural history museum, london 37 bakeria rugosa dodd, 1920: 377. valid name doddius rugosus (dodd, 1920). summary of types holotype ♀, bmnh number 9.676, by original designation. primary type data omilteme/ guerrero/ 8000 ft./ aug., h.h. smith.; godman-salvin/ coll. 1904.-1.; type; bakeria/ rugosa/ dodd, ♀. type locality mexico, guerrero, omilteme. remarks the holotype is mounted on a card, all wings missing except left hind wing, right antenna beyond scape mounted separately, and left fore leg and both mid legs damaged. masner & garcía (2002) described the genus doddius with bakeria rugosa as the type species. arias-penna (2003) placed this species simultaneously in doddius and pentapria kieffer, 1905; the latter is an error. paramesius longior dodd, 1920: 378. valid name paramesius longior dodd, 1920. summary of types holotype ♀, bmnh number 9.663, by original designation. paratype ♂. primary type data type; hewitt/ kuching; j2; p.cameron coll./ 1914-110; brachyaulax/ rufipes cam. ♀/ type borneo; paramesius ♀/ longior dodd. type locality malaysia, borneo, sarawak, kuching. remarks this species was described from a specimen of each sex which had been labelled with the manuscript name “brachyaulax rufipes” by cameron. one female and four males were found labelled “b. rufipes” in cameron’s hand. of these two of the males were mounted together; these could not have been seen by dodd as he only mentioned one male. a third male is labelled as a type but is not, as the antennae are the wrong colour. the female and remaining male are the types. the female is labelled “type” and the male “cotype”, following dodd’s published, designation so the female is the holotype by original designation. the holotype is mounted on a card, and has its wings torn and partly missing. european journal of taxonomy 75: 1-123 (2014) 38 mantara bifurcata dodd, 1920: 380. valid name aneuropria bifurcata (dodd, 1920) comb. nov. summary of types holotype ♀, bmnh number 9.677, by original designation. paratypes ♀, ♂. primary type data 1081.; type; madeira./ wollaston./ 55.7; mantara/ bifurcata/ dodd ♀; holotype ♀/ mantara/ bifurcata dodd/ l.huggert-81. type locality madeira. remarks this species was described from two females and a male, referred to in the original description as a “type” and “cotypes”. three specimens matching the description were found, one of the females has dodd’s identification label, and this is considered to be the holotype by original designation. the holotype is mounted on a card, with the left antenna mounted separately. mantara bifurcata is actually a species of aneuropria kieffer, 1905. since mantara bifurcata is the type species of mantara, this means mantara dodd, 1920 is a junior synonym of aneuropria kieffer, 1905 syn. nov. aneuropria bifurcata comb. nov. is clearly a different species (in head shape, antennal proportions and the lack of notauli) from the widespread european a. foersteri kieffer, 1910, which is also found in madeira. acidopria spinosiceps dodd, 1920: 380. valid name trichopria spinosiceps (dodd, 1920) comb. nov. summary of types holotype ♀, bmnh number 9.692, by original designation. paratype ♀. primary type data type; kuching/ j.h.; p.cameron coll./ 1914-110; brachyaulax/ erythrocerus/ cam. type borneo; acidopria/ spinosiceps/ dodd ♀ type. type locality malaysia, borneo, sarawak, kuching. remarks this species was described from two females referred to as “type” and “cotype”. two specimens were found corresponding to the description, one of which had dodd’s type label and is regarded as the holotype by original designation. the holotype is mounted on a card and has parts of five legs missing. this species was placed by dodd in acidopria kieffer, 1913 (now a junior synonym of basalys) because of the spines on the head; however, it is a species of trichopria. it differs from those species of basalys with spinose heads in having the spines behind the ocellar triangle rather than in front of it. a non-type notton d.g., diapriinae types at the natural history museum, london 39 male was found which agrees closely with the females and it shows that this species belongs to the group of trichopria with whorled hairs on the male antenna. galesus muscidorum dodd, 1920: 381. valid name coptera muscidorum (dodd, 1920). summary of types holotype ♀, bmnh number 9.654, by original designation. primary type data type/ h.t.; reared/ from pupa found/ in palpalis/ breeding ground/ similar holes can/ often be found in/ palpalis pupae/ g.d.h.carpenter/ uganda/ aug 10.1910.; galesus/ muscidorum/ dodd ♀. type locality uganda. remarks the holotype is micropinned, with the left antenna missing beyond a8 and mounted with a fly puparium, presumably the host. the generic placement established by masner (1965) is confirmed here. aneurhynchus indicus dodd, 1920: 382. valid name aneurhynchus indicus dodd, 1920. summary of types holotype ♀, bmnh number 9.634, by original designation. primary type data type; kangra valley/ 4,500 ft./ apr. 1899/ dudgeon.; punjab./ g.c.dudgeon./ 1903-37.; aneurhynchus/ indicus dodd ♀. type locality india, punjab province, kangra valley. remarks this species was described from a single female. the holotype is mounted on a card and is entire. antarctopria coelopae early, 1978: 209, 210, figs 1-9. valid name spilomicrus coelopae (early, 1978) comb. nov. summary of types paratypes 12 ♀♀, 17 ♂♂. european journal of taxonomy 75: 1-123 (2014) 40 remarks an additional male from the otago peninsula labelled as a paratype is not a type; specimens from this locality are listed under “other material” and so are not types. all current antarctopria species, a. coelopae, a. campbellana yoshimoto, 1964, a. diomedeae early, 1978, a. latigaster brues in tillyard, 1920 and a. rekohua early, 1978 appear to belong to a derived species group of spilomicrus showing pronounced wing reduction and the associated suite of characters that go with it. extra setae on the gastral tergites are found on wing-reduced representatives of a number of other diapriid genera and is not considered here a character of generic value. consequently, these five antarctopria species are transferred to spilomicrus (new combinations), and since antarctopria latigaster is the type species of antarctopria, this genus is now a synonym of spilomicrus syn. nov. antarctopria diomedeae early, 1978: 210, 223, figs 28-32. valid name spilomicrus diomedeae (early, 1978) comb. nov. summary of types paratypes 2 ♂♂. remarks for notes on the new generic combination, see a. coelopae above. spilomicrus pilgrimi early, 1978: 210, 224, figs 33-38. valid name spilomicrus pilgrimi early, 1978. summary of types paratype ♀. hemilexomyia spinosa early, 1980: 154, 160, figs 12-20, 23-24. valid name hemilexomyia spinosa early, 1980. summary of types paratypes 3 ♀♀, ♂. entomacis subaptera early, 1980: 154, 163, figs 21-22, 25. valid name entomacis subaptera early, 1980. summary of types paratypes ♀, ♂. notton d.g., diapriinae types at the natural history museum, london 41 spilomicrus barnesi early & horning, 1978: 231, figs 1-6, 12. valid name spilomicrus barnesi early & horning, 1978. summary of types paratypes 6 ♀♀, 4 ♂♂. rostropria simplex early & naumann, 1990: 529, 530, figs 1-18. valid name rostropria simplex early & naumann, 1990. summary of types paratypes ♀, ♂. rostropria spiniventris early & naumann, 1990: 529, figs 19-38. valid name rostropria spiniventris early & naumann, 1990. summary of types paratypes ♀, ♂. trichopria tachinidarum ferrière, 1933b: 104, fig. 11a-b. valid name trichopria tachinidarum ferrière, 1933. summary of types syntype ♀, bmnh number 9.714. syntypes 3 ♀♀, 2 ♂♂. primary type data type; java/ buitenzorg/ xi.1930/ r.w.paine/ par. ‘u’; hyperparasite/ from puparium/ of tachinid ‘b’/ of tirathaba; pres. by/ imp. inst. ent./ b.m.1933-375; trichopria/ tachinidarum ♀/ ch. ferriere type. (syntype ♀). java/ buitenzorg/ xi.1930/ r.w.paine/ par. ‘u’; buitenzorg: xi-30. r.w.p./ diapriid, trichopria sp.?; hyperparasite from puparium of/ tachinid ‘b’ of tirathaba.; pres. by/ imp. inst. ent./ b.m.1933-375; trichopria/ tachinidarum ♀/ ch. ferriere cotype (syntype ♀). java/ buitenzorg/ xi.1930/ r.w.paine/ par. ‘u’; hyperparasite from puparium of/ tachinid ‘b’ of tirathaba.; pres. by/ imp. inst. ent./ b.m.1933-375; trichopria/ tachinidarum ♀/ ch. ferriere cotype (syntypes 2 ♀♀). java/ buitenzorg/ xi.1930/ r.w.paine/ par. ‘u’; hyperparasite from puparium of/ tachinid ‘b’ of tirathaba.; pres. by/ imp. inst. ent./ b.m.1933-375; trichopria/ tachinidarum ♂/ ch. ferriere cotype (syntypes 2 ♂♂). type locality indonesia, west java, kota bogor (as java, buitenzorg). european journal of taxonomy 75: 1-123 (2014) 42 remarks since ferrière’s paper was published in two parts, the name of this species was introduced first as a nomen nudum (ferrière 1933a: 86) and made available shortly after (ferrière 1933b: 104). this species was described from six females and three males, of which four females and two males were found. although ferrière labelled one specimen “type” and five “cotype”, there is no type designation and all the types are syntypic. nor does the mention of a type and paratype by huggert (1982) count as an original holotype designation since it is not in the original description. all the syntypes are mounted on card points: the first female is entire; second female entire; third female missing head and gaster; fourth female missing left hind wing and left fore wing and antenna mounted on a microslide attached to the same pin; first male missing left antenna beyond a5; second male missing right antenna, gaster and hind legs, the antenna and dissected gaster are mounted on a separate slide. trichopria capensis var. robustior ferrière, 1935: 329, 338. valid name trichopria capensis robustior ferrière, 1935. summary of types syntypes 17 ♀♀, 8 ♂♂. primary type data zululand/ empangen/ 16.v.1923/ r.h.harris; sent by/ div. of ent./ pretoria; primary parasite/ of glossina pallidipes/ puparia; ser. no./ 1186; trichopria/ capensis kieffer/ var. robustior, silv./ f.silvestri det. (syntype ♀). zululand:/ empangen: 16.v.1923/ primary parasite of/ glossina pallidipes/ puparium: r.h.harris; ser. no. div. of entom./ pretoria/ south africa/ 1186; pres. by/ imp. bur. ent./ brit. mus./ 1929-43 (syntype ♀). zululand:/ empangen: 16.v.1923/ primary parasite of/ glossina pallidipes/ puparium: r.h.harris; sth. africa/ pretoria/ div. of entom.; pres. by/ imp. bur. ent./ brit. mus./ 192943; trichopria/ capensis kff./ v. robustior s./ silvestri det. (syntype ♀). zululand:/ empangen: 16.v.1923/ primary parasite of/ glossina pallidipes/ puparium: r.h.harris; sent by division of/ entomology, pretoria,/ s.africa, ser. no. 1186; pres. by/ imp. bur. ent./ brit. mus./ 1929-43 (syntypes 5 ♀♀, 5 ♂♂). zululand:/ empangen: 16.v.1923/ primary parasite of/ glossina pallidipes/ puparium: r.h.harris; sent by division of/ entomology, pretoria,/ s.africa, ser. no. 1186:/ pres. by/ imp. bur. ent.:/ brit. mus./ 1929-43 (syntypes 9 ♀♀, 3 ♂♂). type locality south africa, kwazulu-natal province, empangeni (as natal, zululand, empangen). remarks this species was described from seventeen females and eight males. all specimens were located, together with a fly puparium, presumably the host. no holotype was designated so all are syntypes. all the syntypes are mounted on card points, and most are entire. this variety was attributed to silvestri by ferrière, but silvestri never published the name and does not appear to have been responsible for ferrière’s description, so it is attributable to ferrière (1935). abothropria lloydi ferrière, 1935: 329, 338, fig. 3a-c. valid name lepidopria lloydi (ferrière, 1935). notton d.g., diapriinae types at the natural history museum, london 43 summary of types syntype ♀, bmnh number 9.678. syntypes 10 ♀♀, 9 ♂♂. primary type data type; tanganyika/ bugambwa/ 7.vi.1933/ j.e.m.lloyd; ex glossina/ palpalis pup.; pres by/ imp. inst. ent./ b.m.1935-231; abothropria/ lloydi ♀/ ch. ferrière type (syntype ♀). tanganyika/ bugambwa/ 7.vi.1933/ j.e.m.lloyd; ex glossina/ palpalis pup.; pres by/ imp. inst. ent./ b.m.1935-231; abothropria/ lloydi ♀/ ch. ferrière cotype (syntypes 7 ♀♀). tanganyika/ bugambwa/ 7.vi.1933/ j.e.m.lloyd; ex glossina/ palpalis pup.; pres by/ imp. inst. ent./ b.m.1935-231; abothropria/ lloydi ♂/ ch. ferrière cotype (syntypes 3 ♂♂). tanganyika t./ bugambwa, 7.vi.1933/ j.e.m.lloyd/ ex glossina/ palpalis, pupa; diapriidae:/ abothropria/ lloydi cotype/ ch. ferrière ♀ (syntype ♀). tanganyika t./ bugambwa, 7.vi.1933/ j.e.m.lloyd/ ex glossina/ palpalis, pupa; diapriidae:/ abothropria/ lloydi cotype/ ch. ferrière ♂ (syntype ♂). tanganyika t./ bugambwa, 7.vi.1933/ j.e.m.lloyd/ ex glossina/ palpalis, pupa; diapriidae:/ abothropria/ lloydi cotypes/ ch. ferrière 5♂ 2♀ (syntypes 2 ♀♀, 5 ♂♂). type locality tanzania, mara region, buganbwe island (as tanganyika, bugambwa). remarks this species is said to have been described from 11 ♀♀ and 11 ♂♂, of which the following were found: 8 ♀♀, 3 ♂♂ pinned; 3 ♀♀, 6 ♂♂ on slides; and two glossina wiedemann, 1830 puparia, presumably the hosts. these are all syntypes as no holotype was designated. four other specimens of this species could not be assessed as they were on loan. the pinned syntypes, 8 ♀♀ and 3 ♂♂, are mounted on card points: the first female is entire; the second female has the left antenna missing and the head dented; the third female is entire; the fourth female has the right antenna mounted separately and is not fully developed – it appears to have been dissected from the host; the fifth female is entire; the sixth female is entire and has the head flattened; the seventh female has no wings; the eighth female has the left wings crumpled; the first male is not fully developed and has a teratological right antenna; the second male is entire; the third male has the right antenna missing beyond the scape and the left wing crumpled. the other 3 ♀♀ and 6 ♂♂ syntypes are slide mounted and mostly entire. this species was transferred from abothropria kieffer, 1913 to lepidopria kieffer, 1911 by notton (2004), but it may well deserve a genus of its own. spilomicrus kiefferi fouts, 1925: 150. valid name spilomicrus atriclavus ashmead, 1893. summary of types paratype ♀. trichopria (trichopria) tabanivora fouts in cameron, 1926: 39. valid name trichopria tabanivora fouts in cameron, 1926. summary of types paratypes ♀, ♂. european journal of taxonomy 75: 1-123 (2014) 44 loxotropa nigrescens fouts, 1927: 165, 169, fig. 1. valid name basalys nigrescens (fouts, 1927). summary of types paratypes 2 ♀♀. remarks two paratypes have come to the bmnh via the whittaker collection. paramesius laetus fouts, 1927: 170. valid name paramesius laetus fouts, 1927. summary of types paratype ♀. remarks the paratype has come to the bmnh via the whittaker collection. labolips innupta haliday, 1857: 173, pl. 10, figs 1-20. valid name labolips innupta haliday, 1857. summary of types syntype ♀, bmnh number 9.905. primary type data 59-4; labolips/ extincta ♀; britain/ purchased from/ t.v.wollaston/ named by haliday/ bm 1859.4; syntype ♀/ labolips/ innupta haliday/ det. d.notton, 2003. type locality ireland, county down, between belfast and holywood, knocknagoney, clifden house. remarks haliday evidently collected a long series of this species, much of which he dispersed before making a formal description in an effort to discover whether it was previously known or described (haliday 1857). hence, as well as the twelve syntypes in dublin, syntypic material of labolips innupta is also to be found in the collections of the natural history museum, london (one female, via t.v. wollaston) and in the dale collection at the hope department of entomology, oxford university museum of natural history (two females). the london and oxford syntypes are all labelled “labolips extincta”, a manuscript name haliday used for this species before settling on the published name. there may also be syntypes in the förster collection at the naturhistorisches museum wien, wien, austria, as the description of labolips förster, 1856 was made by förster from two females sent to him by haliday (förster 1856: 125; notton notton d.g., diapriinae types at the natural history museum, london 45 & o’connor 2004). the london syntype is mounted on a card and is set with the wings spread in a similar fashion to the other syntypes. trichopria stomoxydis huggert, 1977: 205, figs 1-7. valid name trichopria stomoxydis huggert, 1977. summary of types holotype ♀, bmnh number 9.801, by original designation. paratypes 20 ♀♀, 8 ♂♂. primary type data holo-/ type; uganda/ c.i.b.c./ 25.vii.1975/ c.i.e.a8124; ex pupa of/ stomoxys nigra/ from cut/ elephant grass; ♀ trichopria sp./ nr. inconspicua?/ det. z.bouček, 1975; holotypus/ trichopria ♀/ stomoxydis/ n. sp./ l.huggert-76. type locality uganda. remarks the holotype is mounted on a card point, the left hind wing is missing and the left fore wing and left antenna are mounted on a microslide on the same pin. trichopria pseudosaxatilis huggert, 1977: 214, figs 32-37. valid name trichopria pseudosaxatilis huggert, 1977. summary of types holotype ♂, bmnh number 9.802, by original designation. primary type data 5; silhouette, ‘08/ seychelles exp.; diapria/ saxatilis/ j.j.kieffer/ paratype; holotypus/ trichopria ♂/ pseudosaxatilis/ n. sp./ l.huggert-76. type locality seychelles, silhouette, high country near pot-à-eau. remarks this species was described from a single male. this holotype is also a paralectotype of diapria saxatilis kieffer, 1912 q.v. it is possible to be more precise about the type locality than huggert (1977) based on information given about this specimen by kieffer (1912a). the holotype is mounted on a card; the left wings and left antenna are mounted on a microslide on the same pin. trichopria nirva huggert, 1977: 215, figs 38-43. valid name trichopria nirva huggert, 1977. european journal of taxonomy 75: 1-123 (2014) 46 summary of types holotype ♀, bmnh number 9.803, by original designation. primary type data 145; mahe, ’08-9./ seychelles exp.; diapria/ saxatilis/ j.j.kieffer/ paratype; holotypus/ trichopria ♀/ nirva n. sp./ l.huggert-76. type locality seychelles, mahé. remarks this species was described from a single female. the holotype is also a paralectotype of diapria saxatilis q.v. the holotype is mounted on a card; the right antenna beyond the scape is missing and the left antenna and wings are mounted on a microslide on the same pin. trichopria euvulgaris huggert, 1979: 19, figs 22-27. valid name trichopria euvulgaris huggert, 1979. summary of types holotype ♂, bmnh number 9.806, by original designation. primary type data mahe, ’08-9./ seychelles exp./ 95; type; type/ h.t.; seychelles islands./ percy sladen trust/ expedition./ 1913-170.; rhopalopria/ vulgaris/ j.j.kieffer/ type; holotypus/ trichopria ♂/ euvulgaris/ n. sp./ l.huggert -77. type locality seychelles, mahé. remarks this species was described from a single male. the holotype is also a paralectotype of rhopalopria vulgaris kieffer, 1912 q.v. the holotype is mounted on a paper-covered cork block with the left hind wing missing and the left antenna and fore wing mounted on a microslide. trichopria ensia huggert, 1979: 23, figs 35-40. valid name trichopria ensia huggert, 1979. summary of types holotype ♀, bmnh number 9.807, by original designation. primary type data mahe, ’08-9/ seychelles exp/ 45; seychelles islands/ percy sladen trust/ expedition./ 1913170; paratype; diapria/ seychellensis/ j.j.kieffer/ paratype; holotypus ♀/ trichopria/ ensia n. sp./ l.huggert-77. notton d.g., diapriinae types at the natural history museum, london 47 type locality seychelles, mahé. remarks this species was described from a single female. the holotype is also a paralectotype of diapria seychellensis kieffer, 1912 q.v. the holotype is mounted on a paper-covered cork block, the left antenna is missing beyond scape, and the left wings and right antenna are mounted on a microslide on the same pin. trichopria percyi huggert, 1979: 24, figs 41-46. valid name trichopria percyi huggert, 1979. summary of types holotype ♂, bmnh number 9.808, by original designation. primary type data mahe, ’08-9/ seychelles exp/ 97; seychelles islands./ percy sladen trust/ expedition./ 1913170.; paratype; diapria/ seychellensis/ j.j.kieffer/ paratype; holotypus ♂/ trichopria/ percyi n. sp./ l.huggert-77. type locality seychelles, mahé. remarks trichopria percyi was described from a single male. the holotype is also a paralectotype of diapria seychellensis q.v. the holotype is mounted on a paper-covered cork block, the left mid leg is mounted separately, and the left antenna and wings are mounted on a microslide on the same pin. townesella marjoriae huggert & masner, 1983: 67, 70, 71, figs 14, 16, 18-20. valid name townesella marjoriae huggert & masner, 1983. summary of types paratype ♀. trichopria myrmicae huggert & masner, 1983: 68, 77, figs 39-49. valid name trichopria myrmicae huggert & masner, 1983. summary of types paratype ♀. european journal of taxonomy 75: 1-123 (2014) 48 neurogalesus carinatus kieffer, 1907: 298. valid name neurogalesus carinatus kieffer, 1907. summary of types holotype ♀, bmnh number 9.673, by monotypy. primary type data type; mackay/ 3.01; queensland./ r.e.turner./ 1907-94; neurogalesus/ carinatus ♀/ type. kieff./ named by/ j.j.kieffer. type locality australia, queensland, mackay. remarks this species was described from an unspecified number of females; however, kieffer describes the scutellum as “vorn beschädigt”, i.e., damaged in front, a rather unusual thing to say. the specimen recognised as the type has been pinned at exactly this point, destroying the basal scutellar pit. had kieffer seen any other specimens it is likely they would not have been damaged in precisely this place and he would have described the distinctive bifoveate scutellar pit, so it is considered that he saw only this specimen; hence it is a holotype by monotypy. the holotype is micropinned through the mesosoma and has the head missing. despite what kieffer says about its resemblance to a galesus, this species belongs to the spilomicrini. xyalopria nigriceps kieffer, 1907: 300. valid name trichopria fluminis nom. nov. summary of types syntype ♀, bmnh number 9.690. primary type data type; rio/ 59-21x; xyalopria/ nigriceps/ kieffer; named by j.j.kieffer. type locality brazil, rio de janeiro. remarks this species was described from an unspecified number of females, of which one was found. the syntype is mounted on a card, with part of the right mid tarsus missing and the left mid tarsus mounted separately. this species was included in megaplastopria by arias-penna (2003); however, it does not belong there. while it is large, elongate and has a long, slender propodeal spine, it lacks significant characters of megaplastopria such as the occipital spine and carina, grooved scape and flared corners of the metasomal tergites, and is here transferred to trichopria instead. since the new combination t. nigriceps (kieffer, 1907) is a junior secondary homonym of t. nigriceps (ashmead in riley, ashmead & howard, 1894), kieffer’s species is given the new replacement name trichopria fluminis nom. nov. the word fluminis, notton d.g., diapriinae types at the natural history museum, london 49 genitive of flumen (latin for river) refers to the type locality rio de janeiro. since x. nigriceps kieffer is the type species of xyalopria, xyalopria is removed from synonymy with megaplastopria and becomes a junior synonym of trichopria syn. nov. galesus punctulatus kieffer, 1907: 301. valid name coptera punctulata (kieffer, 1907). summary of types syntype ♂, bmnh number 9.653. primary type data type; pt/ natal/ 56-63; galesus/ punctulatus/ kieffer/ named by j.j.kieffer; bmnh(e)1856-63/ south africa:/ port natal:/ coll. plant:/ purch. stevens. type locality south africa, durban (as port natal). remarks this species was described from an unspecified number of males, of which one was found. the syntype is micropinned, the left fore wing is mounted separately and the metasoma, left hind wing and part of the right mid tarsus are missing. galesus walkeri kieffer, 1907: 301. valid name psilus fuscipennis (curtis, 1831) syn. nov. summary of types lectotype ♂, bmnh number 9.919, here designated. primary type data alten/ finnmark/ f.walker/ feb.37/ 201. type locality noted as “wohl england”, i.e., probably from england, but actually from norway, finnmark, alta. remarks this species was described from an unspecified number of males, said to be in the british museum by kieffer. one male specimen was found which agreed with the description, except that kieffer says the collector was j. walker, whereas the label says “f. walker” although the “f.” is written like a “j.” kieffer says the specimens were probably from england, suggesting that the label was difficult to read and that he assumed the specimen(s) was english because it was collected by walker. the label on the type is difficult to read but actually says “alten, finnmark”, no doubt a specimen collected by francis walker on his 1836-7 tour of scandinavia (christy & walker 1837). the close agreement with the description, kieffer’s mistake in reading the “j.” for “f.” because of the peculiar style of writing and the near illegibility of the label which caused him to guess the locality (erroneously) convince me that this european journal of taxonomy 75: 1-123 (2014) 50 is a type. this specimen belongs to psilus fuscipennis following the interpretation of nixon (1980) and the two species are synonymized here. a lectotype is designated to stabilise the synonymy. the lectotype is mounted on a card and is missing the left fore leg and the tips of three tarsi. galesus gracilipes kieffer, 1907: 302. valid name psilus cornutus panzer, 1801. summary of types syntype ♂, bmnh number 9.648. primary type data type; galesus/ gracilipes/ kieffer/ named by/ j.j.kieffer; milleri/ [illeg.]. type locality england. remarks this species was described from an unspecified number of males. the syntype is mounted on a card and has the left wings missing. loxotropa pedestris kieffer, 1907: 302. valid name basalys parvus thomson, 1858. summary of types holotype ♀, bmnh number 9.728, by monotypy. primary type data uxbridge; type; loxotropa/ pedestris/ kieffer/ named by/ j.j.kieffer. type locality england, uxbridge. remarks this species was described from an unspecified number of females, but kieffer (1911b: 926) says “type au musée de londres”, so evidently there was only one type, thus a holotype by monotypy. the holotype is mounted on a card and is entire. phaenopria halterata kieffer, 1909: 381. valid name trichopria halterata (kieffer, 1909). summary of types lectotype ♂, bmnh number 9.715, designated by notton (1995). notton d.g., diapriinae types at the natural history museum, london 51 primary type data galloway; type; 478; cameron coll./ 1910-55.; halterata k. type locality scotland, galloway. remarks this species was described from an unspecified number of males. the lectotype agrees well with the description, although it has a weak (and easily overlooked) scutellar pit. the lectotype is mounted on a card, left fore leg mounted separately, most tarsi broken. phaenopria cameroni kieffer, 1909: 382. valid name trichopria cameroni (kieffer, 1909). summary of types lectotype ♀, bmnh number 9.717, designated by notton (1995). primary type data m; type; suspecta nees; 231; cameron coll./ 1910-55.; cameroni k. type locality probably england, mickleham. remarks this species was described from an unspecified number of females and it is quite likely that the type series contained at least two specimens from clyde at newton and mickleham (cameron 1910). a single syntype was located which had been designated as the lectotype by notton (1995). since the lectotype has an “m” written on the mount, it is probably from mickleham in england. although kieffer gave the type locality as scotland, there is no reason to exclude the mickleham specimen from the type series because kieffer is known to have misinterpreted many of the localities on cameron’s specimens. the lectotype is mounted on a card and has become twisted on its mount, breaking the antennae, and a12 of the right antenna is missing. trichopria inermis kieffer, 1909: 386. valid name trichopria nigra (nees, 1834). summary of types lectotype ♂, bmnh number 9.705, designated by notton (1995). primary type data thornhill/ 11ix.; type; 238.; cameron coll./ 1910-55.; inermis k.; holotype; ♂ lectotype/ trichopria/ inermis/ kieffer, 1909/ des. d.notton, 1995. european journal of taxonomy 75: 1-123 (2014) 52 type locality scotland, thornhill. remarks this species was described from an unspecified number of males of which one specimen was found. the lectotype is mounted on a card and is entire. idiotypa nigriceps kieffer, 1909: 386. valid name idiotypa nigriceps kieffer, 1909. summary of types lectotype ♀, bmnh number 9.746, here designated. primary type data lochawe; type; 230.; cameron coll./ 1910-55.; nigriceps k. type locality scotland, lochawe village. remarks this species was described from an unspecified number of females. a single female syntype was found which is designated here as lectotype to ensure objective synonymy with idiotypa nigriceps kieffer, 1911 q.v. the lectotype is mounted on a card and is entire. paramesius inchoatus kieffer, 1911a: 756. valid name paramesius rufipes (fonscolombe, 1832). summary of types lectotype ♀, bmnh number 9.813, designated by nixon (1980). paralectotype ♀. primary type data q.bl.; 469.; paramesius/ inchoatus, k.; cameron coll./ 1910-302.; determined by/ dr. kieffer.; selected as/ type of/ paramesius/ inchoatus k/ by g.e.j.n. 1976. type locality scotland. remarks this species was described from an unspecified number of females. two specimens were located, of which one had been designated as a lectotype by nixon (1980). the lectotype is mounted on a card and is entire. notton d.g., diapriinae types at the natural history museum, london 53 paramesius subinermis kieffer, 1911a: 757. valid name paramesius rufipes (fonscolombe, 1832). summary of types lectotype ♂, bmnh number 9.812, designated by nixon (1980). paralectotype ♂. primary type data bishop; bishopton; 446.; paramesius/ subinermis, k.; cameron coll./ 1910-302.; determined by/ dr. kieffer; selected as/ type of/ paramesius/ subinermis/ k/ by g.e.j.n. 1976. type locality scotland, bishopton. remarks this species was described from an unspecified number of males. three specimens were found, of which one had been designated as lectotype by nixon (1980). the lectotype is mounted on a card and parts of the right mid and hind tarsi are disarticulated. paramesius angustipennis kieffer, 1911a: 762. valid name paramesius brachypterus (thomson, 1858). summary of types lectotype ♀, bmnh number 9.929, here designated. primary type data mugd/ 26-7; mugdock; 455.; paramesius/ angustipennis; cameron coll./ 1910-302.; determined by/ dr. kieffer. type locality scotland, mugdock. remarks this species was described from at least two females, of which one syntype was found. macek (2002) synonymized this species with p. brachypterus without seeing the type. the syntype belongs to p. brachypterus following the interpretation of this species by nixon (1980) and macek (2002) and macek’s synonymy is confirmed. a lectotype is designated to stabilise the application of the name and its synonymy. the lectotype is mounted on a card, with the left fore wing missing. the species epithet augustipennis is an incorrect original spelling, and is to be corrected (article 32.5.1), since there is clear evidence in the original publication of an inadvertent error, i.e., in the description kieffer says “ailes mince…”, i.e., wing narrow which when formed into a latin name from angustus (narrow or straight) + penna (a wing) would become angustipennis not augustipennis. european journal of taxonomy 75: 1-123 (2014) 54 paramesius nigricornis kieffer, 1911a: 764. valid name paramesius rufipes (fonscolombe, 1832). summary of types syntype ♀, bmnh number 9.661. primary type data galloway; type; 265.; cameron coll./ 1910-55.; nigricornis k. type locality scotland and norway, solum, overhaiden. remarks this species was described from an unspecified number of females, of which one was found. the syntype is mounted on a card and has the left hind wing disarticulated. paramesius cameroni kieffer, 1911a: 767. valid name paramesius brachypterus (thomson, 1858). summary of types lectotype ♂, bmnh number 9.927, here designated. paralectotypes 2 ♂♂. primary type data mugdock/ 9-8; mugdock; 445.; paramesius/ cameroni, k.; cameron coll./ 1910-302.; determined by/ dr kieffer. type locality scotland, mugdock. remarks this species was described from at least three males. three syntypes were located, of which one is designated a lectotype. macek (2002) synonymized this species with p. brachypterus without seeing the type. the lectotype belongs to p. brachypterus following the interpretation of this species by nixon (1980) and macek (2002), so macek’s synonymy is confirmed. the lectotype is designated to stabilise the application of the name and its synonymy. the lectotype is mounted on a card and is entire. paramesius dentatus kieffer, 1911a: 768. valid name paramesius rufipes (fonscolombe, 1832). summary of types syntype ♂, bmnh number 9.662. notton d.g., diapriinae types at the natural history museum, london 55 primary type data mug/ 26-7; type; 253.; cameron coll./ 1910-55.; dentatus k. type locality scotland. remarks described from an unspecified number of males. the syntype is mounted on a card and is entire. spilomicrus hemipterus var. pedissequus kieffer, 1911a: 774. valid name spilomicrus hemipterus marshall, 1868. summary of types syntype ♀, bmnh number 9.782. primary type data β.; type/ of var.; spilomicrus/ hemipterus,/ var.-/ pedissequus/ kief. ♀.; ?aneurhynchus sp./ in nest of lasius/ fuliginosus/ oxshott. β/ (15th-)12-4-01./ 27; holotype of/ spilomicrus/ pedisequus k./ n.d.m.fergusson, 1974. type locality england. remarks this species was described from an unspecified number of females. one specimen was found which agreed well with the description except for having the basal pit bifoveate. the syntype is mounted on a card, missing the tips of both antennae and the left mid leg; four tarsi are broken but parts of two of them are still present. it appears that pedissequus is the correct original spelling, as there is no evidence in the text of the original description that it is an inadvertent error (article 32.5.1). the epithet pedisequus is therefore an unjustified emendation attributable to kieffer (1916) and is a junior objective synonym. spilomicrus integer var. variicornis kieffer, 1911a: 777. valid name spilomicrus integer thomson, 1858. summary of types syntype ♀, bmnh number 9.644. primary type data [illeg.]/ glen. 20-5; type; 465.; cameron coll./ 1910-55.; integer k. v.; integer var./ variicornis, k. type locality scotland. european journal of taxonomy 75: 1-123 (2014) 56 remarks this species was described from an unspecified number of females, of which one was found. the syntype is mounted on a card and is entire. spilomicrus crassiclavis kieffer, 1911a: 788. valid name spilomicrus crassiclavis kieffer, 1911. summary of types lectotype ♀, bmnh number 9.645a, designated by notton (1999). primary type data mugdock/ 11-7; cameron coll./ 1910-302.; mugdock; determined by/ dr kieffer.; spilomicrus/ crassiclavis, k.; spilomicrus/ formosus janss./ g.nixon det. 1972; ♀ lectotype/ spilomicrus/ crassiclavis/ kieffer, 1911: 788/ des. d.notton, 1999. type locality scotland, mugdock. remarks this species was described from an unspecified number of females and males. the lectotype is mounted on a card, left antenna missing, right fore wing disarticulated. spilomicrus crassiclavis was previously confused with spilomicrus formosus jansson, 1942 by nixon (1980), but this is a different species (notton 1999). spilomicrus annulicornis kieffer, 1911a: 788. valid name spilomicrus annulicornis kieffer, 1911. summary of types syntype ♀, bmnh number 9.642. syntype ♀. primary type data 27-69/ mickleham; type; 249.; cameron coll./ 1910-55.; annulicornis k. (syntype ♀). 275.; spilomicrus/ annulicornis, k.; cameron coll./ 1910-302.; determined by/ dr. kieffer (syntype ♀). type locality scotland and england, mickleham. remarks this species was described from an unspecified number of females. kieffer gave the type locality as scotland; however, one of the syntypes is labelled almost illegibly “mickleham” and the other has no locality label. since the specimens agree with the description, there seems no reason to exclude either syntype from the type series because kieffer is known to have misinterpreted many of the localities on cameron’s specimens. both syntypes are mounted on cards, the one from mickleham has lost the tip of its right hind tarsus, the other has lost the tip of its left hind tarsus. notton d.g., diapriinae types at the natural history museum, london 57 spilomicrus minor kieffer, 1911a: 791. valid name paramesius rufipes (fonscolombe, 1832). summary of types syntype ♀, bmnh number 9.664. syntype ♀. primary type data dalry; type; 242.; cameron coll./ 1910-55; minor k. (syntype ♀). 55-69/ box hill; 241; cameron coll./ 1910-55 (syntype ♀). type locality scotland and england, box hill. remarks this species was described from an unspecified number of females; two females were found which agreed with the description. although the published type locality is scotland, there is no reason to exclude the english specimen from the type series because kieffer is known to have misinterpreted many of the localities on cameron’s specimens. both syntypes are mounted on cards; the one from scotland is entire, the one from box hill has the tip of the left antenna missing. spilomicrus tripartitus kieffer, 1911a: 792. valid name spilomicrus stigmaticalis westwood, 1832. summary of types syntype ♀, bmnh number 9.643. primary type data [illeg.]/ viii; type; 248.; cameron coll./ 1910-55.; tripartitus k. type locality scotland; england; france, mesnil-le-roy and versailles. remarks this species was described from at least four specimens including both sexes. the syntype agrees well with the description and while the notauli are percurrent, it is worth noting they are deep posteriorly and weak anteriorly. the syntype is mounted on a card and is entire. idiotypa nigriceps kieffer, 1911a: 814. valid name idiotypa nigriceps kieffer, 1909. summary of types lectotype ♀, bmnh number 9.746, here designated. european journal of taxonomy 75: 1-123 (2014) 58 primary type data lochawe; type; 230.; cameron coll./ 1910-55.; nigriceps k. type locality scotland, lochawe village. remarks this species was described from an unspecified number of females, apparently including the type series of idiotypa nigriceps kieffer, 1909 with additional material from england. the specimen which is the lectotype of i. nigriceps kieffer, 1909 is also designated here as the lectotype of i. nigriceps kieffer, 1911 to create objective synonymy. the lectotype of i. nigriceps kieffer, 1911 is mounted on a card and is entire. aneurhynchus obliquus kieffer, 1911a: 828. valid name aneurhynchus obliquus kieffer, 1911. summary of types syntype ♂, bmnh number 9.633. primary type data dumf; type; 437.; cameron coll./ 1910-55.; obliquus k. type locality scotland. remarks this species was described from an unspecified number of males. the syntype is mounted on a card and has the left hind tarsus mounted separately. galesus (schizogalesus) mayeti var. hispanicus kieffer, 1911a: 845. valid name coptera hispanica (kieffer, 1911). summary of types syntype ♀, bmnh number 9.651. primary type data giberalter [sic]; type; gibraltar; 401.; cameron coll./ 1910-55.; mayeti k. v./ hispanicus k. type locality gibraltar. remarks this species was described from an unspecified number of females. the syntype is mounted on a card and the left fore tarsus is disarticulated. notton d.g., diapriinae types at the natural history museum, london 59 galesus (g.) parvulus kieffer, 1911a: 853. valid name psilus parvulus (kieffer, 1911). summary of types syntype ♂, bmnh number 9.650. primary type data clober; type; 407.; cameron coll./ 1910-55.; parvulus k. type locality scotland. remarks this species was described from an unspecified number of males. the syntype is mounted on a card and is entire. galesus (g.) foersteri var. nigricornis kieffer, 1911a: 856. valid name psilus fuscipennis (curtis, 1831) syn. nov. summary of types lectotype ♂, bmnh number 9.922, here designated. paralectotypes 2 ♂♂. primary type data glen lyon; type; 408.; cameron coll./ 1910-55.; atricornis. type locality scotland, glen lyon. remarks the original description was based on an unspecified number of males, of which three were found. all the syntypes agree well with the description. kieffer evidently changed the name between labelling the specimens and writing the description; the connection is clear from the names: atricornis and nigricornis are closely related, since ater and niger are both latin words meaning black. the lectotype is mounted on a card and is entire. this specimen belongs to psilus fuscipennis following the interpretation of nixon (1980) and the two species are synonymized here. galesus (g.) bispinosus kieffer, 1911a: 865. valid name psilus fuscipennis (curtis, 1831). summary of types syntype ♀, bmnh number 9.920. european journal of taxonomy 75: 1-123 (2014) 60 primary type data scar/ scarbr; type; 406.; cameron coll./ 1910-55.; bispinosus k.; galesus/ bispinosus/ holotype kieff./ g.nixon 1977. type locality probably england, scarborough. remarks this species was described from an unspecified number of females. despite nixon’s label, this specimen cannot be a holotype as kieffer did not designate a holotype. the type locality is most probably scarborough in england, based on the data reported by cameron (1910), rather than scotland. the syntype is mounted on a card, the left antenna missing beyond a4, wings crumpled and right fore wing broken. galesus (g.) cameroni kieffer, 1911a: 865. valid name psilus fuscipennis (curtis, 1831). summary of types syntype ♀, bmnh number 9.649. syntype ♀. primary type data [illeg.]; type; 403.; cameron coll./ 1910-55.; cameroni k. (syntype ♀). mickleham/ 31-76; 404.; cameron coll./ 1910-55 (syntype ♀). type locality probably england. remarks this species was described from an unspecified number of females. the syntypes agree well with the description; although kieffer says the head has five anterior teeth, this probably means four plus the median ocellus which protrudes anteriorly; also, the “arête en avant jusque entre les antennes” is weak but observable from the side. although the published type locality is scotland, it appears from the locality (caterham) given by cameron (1910) that kieffer saw english specimens, so english specimens have not been excluded from the type series. in any case, kieffer is known to have misinterpreted many of the localities on cameron’s specimens. both syntypes are mounted on cards; the syntype numbered 403 has the metasoma missing; syntype 404 is entire. phaenopria halterata kieffer, 1911a: 880. valid name trichopria halterata (kieffer, 1909). summary of types lectotype ♂, bmnh number 9.715, here designated. primary type data galloway; type; 478; cameron coll./ 1910-55.; halterata k. notton d.g., diapriinae types at the natural history museum, london 61 type locality scotland, galloway. remarks this species was described from an unspecified number of males, apparently from the same type series as for p. halterata kieffer, 1909. the type agrees well with the description, although it has a weak scutellar pit. a lectotype is designated here; this is the same specimen as the lectotype of p. halterata kieffer, 1909, creating objective synonymy between the two names. the lectotype is mounted on a card, left fore leg mounted separately, and most tarsi broken. phaenopria subimpressa kieffer, 1911a: 880. valid name trichopria subimpressa (kieffer, 1911). summary of types lectotype ♀, bmnh number 9.716, designated by notton (1995). primary type data 13-69/ hounslow; type; diapria tritoma; 232.; cameron coll./ 1910-55.; subimpressa k. type locality england, hounslow. remarks this species was described from an unspecified number of females. although the published type locality is scotland, cameron (1910) says he has no definite locality for this species, and the specimen label says hounslow in england. there is no reason to exclude this specimen from the type series because kieffer is known to have misinterpreted many of the localities on cameron’s specimens. the lectotype is mounted on a card, with its head disarticulated, right antenna with a4-10 missing, mesosoma broken, propodeum and metasoma disarticulated from the rest of the mesosoma, left fore wing folded and left hind wing missing. phaenopria cameroni kieffer, 1911a: 881. valid name trichopria cameroni (kieffer, 1909). summary of types lectotype ♀, bmnh number 9.717, here designated. primary type data m; type; suspecta nees; 231; cameron coll./ 1910-55.; cameroni k. type locality probably england, mickleham. european journal of taxonomy 75: 1-123 (2014) 62 remarks this species was described from an unspecified number of females and apparently from the same type series as p. cameroni kieffer, 1909 q.v. a single syntype was located and is designated here as a lectotype. the lectotype of p. cameroni kieffer, 1911 is also the lectotype of p. cameroni kieffer, 1909, creating objective synonymy. type locality and condition of lectotype as for p. cameroni kieffer, 1909. tetramopria donisthorpei kieffer, 1911a: 891. valid name tetramopria cincticollis wasmann, 1899. summary of types lectotype ♂, bmnh number 9.892, designated by notton (1995). primary type data ♂/ tetramopria/ donisthorpei/ whitsand/ bay/ 14.iv.1909; type; ex coll./ donisthorpe/ b.m.1934-4; assoc. with/ tetramorium/ caespitum; tetramopria ♂/ donisthorpei kief.; lectotype ♂/ tetramopria/ donisthorpei/ kieffer, 1911/ det. d.g.notton, 1992; ♂ tetramopria/ cincticollis/ det. d.notton, 1993. type locality england, cornwall, whitsand bay. remarks this species was described from an unspecified number of females and males from whitsand bay and two females and a male from hohscheid. the lectotype is mounted on a card and is entire. an additional female labelled as a cotype is not syntypic since it was collected in june; the original date can be seen on the mount. tetramopria donisthorpei var. femoralis kieffer, 1911a: 892. valid name tetramopria cincticollis wasmann, 1899. summary of types lectotype ♂, bmnh number 9.893, designated by notton (1995). paralectotypes 2♀♀. primary type data ♂/ tetramopria/ femoralis/ whitsand/ bay/ 15.iv.1909; type; ex coll./ donisthorpe/ b.m.1934-4; assoc. with/ tetramorium/ caespitum.; tetramopria/ donisthorpei var./ femoralis kief.; lectotype ♂/ tetramopria donisthorpei var./ femoralis kieffer, 1911/ det. d.g.notton, 1992; ♂ tetramopria/ cincticollis/ det. d.notton, 1993. type locality england, cornwall, whitsand bay. remarks this species was described from an unspecified number of females and males. the lectotype is mounted on a card and the gaster, right hind wing and left mid leg are missing. notton d.g., diapriinae types at the natural history museum, london 63 planopria pedestris kieffer, 1911a: 896. valid name platymischus dilatatus westwood, 1832. summary of types syntype ♀, bmnh number 9.764. primary type data planopria/ pedestris/ luccombe/ chine/ 1.viii.1909; ex coll./ donisthorpe/ b.m.1934-4; assoc. with/ lasius niger; planopria/ pedestris kief.; type. type locality england. remarks this species was described from an unspecified number of females. the syntype is mounted on a card and is entire. basalys collaris kieffer, 1911a: 903. valid name basalys collaris kieffer, 1911. summary of types syntype ♀, bmnh number 9.748. primary type data york; type; 225.; cameron coll./ 1910-55.; collaris k. type locality probably england, york. remarks this species was described from an unspecified number of females. one syntype was found which agrees well with the description; however, the collar is not red edged whitish as stated by kieffer “collier de feutrage roux bordé de blanchâtre”, but the nucha and pronotal cuticle underneath it is reddish and the hairs of the collar are whitish, the overall effect resembling kieffer’s description. although scotland is the published type locality, cameron (1910) noted a specimen from york and so this specimen is not excluded from the type series. the syntype is mounted on a card and is entire. basalys rufiscapus kieffer, 1911a: 909. valid name basalys formicarum (kieffer, 1911). summary of types syntype ♂, bmnh number 9.743. european journal of taxonomy 75: 1-123 (2014) 64 primary type data mull; type; 243.; cameron coll./ 1910-55.; rufiscapus k. type locality scotland. remarks this species was described from an unspecified number of males. the syntype is mounted on a card, and is missing its right fore wing and right antenna beyond a9. loxotropa cursitans kieffer, 1911b: 916. valid name basalys pedisequa (kieffer, 1911) syn. nov. summary of types syntype ♀, bmnh number 9.921. primary type data lade at/ cadder/ 2-8; 209.; cameron coll./ 1910-55. type locality scotland. remarks this species was described from an unspecified number of females. the specimen formerly regarded as the type of l. cursitans by masner (1965) and nixon (1980) is actually a syntype of l. cursitans var. alacris kieffer q.v. a different specimen was found which is considered here to be a true syntype of the nominate l. cursitans. this type has much shorter wings (kieffer says “remplacées par une écaille”), and a shorter head, and agrees better with kieffer’s description. this syntype is mounted on a card and is entire. reinterpretation of the true syntype of l. cursitans shows that it is the same as nixon’s (1980) interpretation of basalys pedisequa. consequently it is removed from synonymy with b. parvus thomson, 1858. loxotropa cursitans var. alacris kieffer, 1911b: 916. valid name basalys parvus thomson, 1858. summary of types syntype ♀, bmnh number 9.729. syntypes 2♀♀. primary type data eccles [illeg.]; type; 208.; cameron coll./ 1910-55.; cursitans/ kieff. (syntype ♀). cladich; 210.; cameron coll./ 1910-55 (syntype ♀). 212; cameron coll./ 1909-182 (syntype ♀). type locality scotland. notton d.g., diapriinae types at the natural history museum, london 65 remarks this variety was described from an unspecified number of females. three syntypes were found, of which the one numbered as type 9.729 had erroneously been regarded as the type of l. cursitans q.v. by masner (1965) and nixon (1980). all syntypes have the wings reaching the basal quarter of the metasoma, a gradually expanded triangular marginal vein, and the head not higher than long and thus agree better with kieffer’s description of l. cursitans var. alacris. all three syntypes are mounted on cards; syntype number 212 is missing part of the left antenna and part of the left fore tarsus and the other two syntypes are entire. the synonymy of this species with basalys parvus thomson, 1858, following nixon’s (1980) interpretation of b. parvus, is confirmed here. loxotropa macroptera kieffer, 1911b: 929. valid name basalys macroptera (kieffer, 1911). summary of types syntype ♀, bmnh number 9.730. primary type data loch awe; type; 227.; cameron coll./ 1910-55.; macroptera k. type locality scotland. remarks this species was described from an unspecified number of females. the syntype is mounted on a card and is entire. loxotropa sulcata kieffer, 1911b: 931. valid name basalys macroptera (kieffer, 1911). summary of types syntype ♀, bmnh number 9.731. syntype ♂. primary type data cladich; type; 456.; cameron coll./ 1910-55.; sulcata k. (syntype ♀). mugdock; 457.; loxotropa/ sulcata, k.; cameron coll./ 1910-302.; determined by/ dr kieffer; basalys/ iphicla nix./ 1979/ g.nixon det. 1978 (syntype ♂). type locality scotland. remarks this species was described from an unspecified number of females and males. both syntypes are mounted on cards and are entire. european journal of taxonomy 75: 1-123 (2014) 66 loxotropa convexa kieffer, 1911b: 932. valid name basalys abruptus thomson, 1858. summary of types syntype ♀, bmnh number 9.732. syntype ♀. primary type data type; 237.; cameron coll./ 1910-55.; convexa k. (syntype ♀). clober/ 87; 240; cameron coll./ 191055 (syntype ♀). type locality scotland. remarks this species was described from an unspecified number of females. two females were found matching the description. both syntypes are mounted on cards; the syntype from clober is entire and the other is missing the right antenna beyond a8. this species was synonymized with basalys abruptus by nixon (1980); however, only one syntype agrees with his concept of b. abruptus. a lectotype needs to be designated for loxotropa convexa, but it is recommended that thomson’s type of basalys abruptus is examined first to ensure that the synonymy established by nixon (1980) is maintained if at all possible. loxotropa longipennis kieffer, 1911b: 932. valid name basalys longipennis (kieffer, 1911). summary of types syntype ♀, bmnh number 9.734. primary type data mull; type; 211.; cameron coll./ 1910-55.; longipennis k. type locality scotland. remarks this species was described from an unspecified number of females. the syntype is mounted on a card and is entire. loxotropa nigricornis var. subterranea kieffer, 1911b: 939. valid name trichopria nigricornis (marshall, 1868). summary of types lectotype ♂, bmnh number 9.785, designated by notton (1995). notton d.g., diapriinae types at the natural history museum, london 67 primary type data loxotropa/ subterranea/ blackgang chine/ aug. 17/ 1908; type; ex coll./ donisthorpe/ b.m.19344.; assoc. with/ lasius flavus; loxotropa/ subterranea kief.; lectotype ♂/ loxotropa/ nigricornis/ subterranea/ det. d.g.notton. type locality england, isle of wight, blackgang chine. remarks this species was described from an unspecified number of males. the lectotype is mounted on a card and is entire. loxotropa bifoveata kieffer, 1911b: 940. valid name basalys bifoveatus (kieffer, 1911). summary of types syntype ♂, bmnh number 9.735. primary type data cladich; type; 467.; cameron coll./ 1910-55.; bifoveata k. type locality scotland. remarks this species was described from an unspecified number of males. the syntype is mounted on a card and is entire; it has both wings twisted. loxotropa luctuosa kieffer, 1911b: 941. valid name basalys luctuosus (kieffer, 1911). summary of types syntype ♂, bmnh number 9.733. primary type data dumfries; type; 470; cameron coll./ 1910-55.; luctuosa k. type locality scotland. remarks this species was described from an unspecified number of males. this syntype agrees well with the description, although the basal scutellar pit is not clearly bifoveate; however, this part of the specimen european journal of taxonomy 75: 1-123 (2014) 68 is dirty, so probably kieffer was mistaken. the syntype is mounted on a card and has three of the tarsi broken. loxotropa atricrus kieffer, 1911b: 941. valid name basalys fumipennis westwood, 1833. summary of types syntype ♂, bmnh number 9.749. syntype ♂. primary type data [illeg.]; type; 458.; cameron coll./ 1910-55; atricrus k. (syntype ♂). [illeg.]’ 461.; loxotropa/ atricrus, k.; cameron coll./ 1910-302.; determined by/ dr kieffer (syntype ♂). type locality scotland. remarks this species was described from an unspecified number of males. both syntypes are mounted on cards: the specimen numbered “9.749” has one tarsus broken; the other is entire. the specific epithet is derived from ater, latin for black, and crus, latin for leg, and is indeclinable. loxotropa scotica kieffer, 1911b: 942. valid name basalys scoticus (kieffer, 1911). summary of types syntype ♂, bmnh number 9.750. syntype ♂. primary type data [illeg.]; type; 462.; cameron coll./ 1910-55; scotica k. (syntype ♂). dumfries; 463.; cameron coll./ 1910-55; basalys/ scotica k./ g.nixon det. 1978 (syntype ♂). type locality scotland. remarks this species was described from an unspecified number of males. both syntypes are mounted on cards: the one numbered 9.730 is missing its left fore wing; the other is entire. loxotropa ciliata kieffer, 1911b: 943. valid name basalys ciliatus (kieffer, 1911). notton d.g., diapriinae types at the natural history museum, london 69 summary of types syntype ♂, bmnh number 9.737. primary type data dumfries; dumfries; type/ g.n.; 269.; loxotropa/ ciliata k.; cameron coll./ 1910-302; selected as/ type of/ loxotropa/ ciliata k./ by g.e.j.n.1934. type locality scotland. remarks this species was described from an unspecified number of males, of which one was found. masner (1965) notes that nixon labelled this specimen as a lectotype, but nixon did not publish a designation so it is still a syntype. the syntype is mounted on a card and has one tarsus broken. loxotropa unifoveata kieffer, 1911b: 943. valid name basalys bifoveatus (kieffer, 1911). summary of types syntype ♂, bmnh number 9.736. primary type data [illeg.]; type; 527.; cameron coll./ 1910-55.; unifoveata k. type locality scotland. remarks this species was described from an unspecified number of males. the syntype is mounted on a card and is entire. loxotropa formicarum kieffer, 1911b: 944. valid name basalys formicarum (kieffer, 1911). summary of types not seen. primary type data according to masner (1965): “loxotropa formicarum ♂; darenth wood, 24.ix.1909; bmnh typelabel”. type locality england, london. european journal of taxonomy 75: 1-123 (2014) 70 remarks this species was described from an unspecified number of males. although one male type was listed for the bmnh by masner (1965), no material was found; nor was this specimen noted by nixon (1980). diapria (tropidopria) inaequalis kieffer, 1911b: 956. valid name trichopria aequata (thomson, 1858). summary of types lectotype ♀, bmnh number 9.693, designated by notton (1995). primary type data type; 477.; cameron coll./ 1910-55.; inaequalis k.; lectotype ♀/ diapria (tropidopria)/ inaequalis kieffer/ des. notton, 1995. type locality scotland. remarks described from an unspecified number of females. the lectotype agrees well with the description apart from “article 11e…plus de deux fois aussi gros que le 12e…”, which does not make sense and is most probably a slip of the pen. the lectotype is mounted on a card and the right antenna is missing beyond a6. diapria (tropidopria) tetratoma kieffer, 1911b: 958. valid name trichopria hyalinipennis (thomson, 1858). summary of types lectotype ♀, bmnh number 9.694, designated by notton (1995). primary type data gloster; type; 228.; cameron coll./ 1910-55.; tetratoma k.; trichopria/ hyalinipennis thoms./ det. l.huggert-89; holo-/ type; lectotype ♀/ diapria tropidopria/ tetratoma kieffer/ des. notton, 1995. type locality england, gloucester. remarks this species was described from an unspecified number of females. although the published type locality is scotland, cameron (1910) records it from gloucester, so this specimen is not excluded from the type series. the lectotype is mounted on a card and is entire. diapria (tropidopria) conotoma kieffer, 1911b: 966. valid name trichopria conotoma (kieffer, 1911). notton d.g., diapriinae types at the natural history museum, london 71 summary of types lectotype ♀, bmnh number 9.725, designated by notton (1995). primary type data alsasua/ spain 30-6; 475.; spain; cameron coll./ 1910-55.; conotoma k.; type; holo-/ type; lectotype ♀/ diapria tropidopria/ conotoma kieffer, 1911: 966/ des. notton, 1995. type locality spain, alsasua. remarks this species was described from an unspecified number of females. the lectotype is mounted on a card and is missing its left fore wing. although the type locality was recorded as scotland, cameron (1910) records it from spain, so this specimen is not excluded from the type series. a spanish locality is also more likely because this is a largely southern european species which only just extends into south-east england. british material of both sexes of t. conotoma has been examined (specimens in bmnh); the male is identical with the description of oxypria bouceki masner, 1959 syn. nov. diapria (tropidopria) variipes kieffer, 1911b: 967. valid name trichopria aequata (thomson, 1858). summary of types lectotype ♀, bmnh number 9.696, designated by notton (1995). primary type data type; 218.; cameron coll./ 1910-55; variipes k.; lectotype ♀/ diapria tropidopria/ variipes kieffer/ des. notton, 1995. type locality scotland. remarks this species was described from an unspecified number of females. the lectotype is mounted on a card and has both antennae and two legs broken. diapria (tropidopria) melanopa kieffer, 1911b: 968. valid name trichopria subimpressa (kieffer, 1911). summary of types lectotype ♀, bmnh number 9.697, designated by notton (1995). paralectotypes ♀, ♂. primary type data mugd./ 26-7; type; 471.; cameron coll./ 1910-55.; melanopa k.; lectotype ♀/ diapria tropidopria/ melanopa kieffer/ des. notton, 1995. european journal of taxonomy 75: 1-123 (2014) 72 type locality scotland, mugdock. remarks this species was described from an unspecified number of females and males. the lectotype is mounted on a card and is missing its right mid leg. according to the publication dates on page 1015 of kieffer (1911b), fascicules 107-8 (including trichopria subimpressa) predate fascicules 109-110 (including t. melanopa, which is here confirmed to be a junior synonym). diapria (tropidopria) nocticolor kieffer, 1911b: 977. valid name trichopria oogaster (thomson, 1858). summary of types lectotype ♂, bmnh number 9.698, designated by notton (1995). primary type data york; type; 266.; cameron coll./ 1910-55.; nocticolor k.; holo-/ type; lectotype ♂/ diapria tropidopria/ nocticolor kieffer/ des. notton, 1995. type locality england, york. remarks this species was described from an unspecified number of males. the lectotype is mounted on a card, the right antenna is broken and the right fore wing and both hind wings are missing. diapria (tropidopria) clavatipes kieffer, 1911b: 978. valid name trichopria subimpressa (kieffer, 1911). summary of types lectotype ♂, bmnh number 9.695, designated by notton (1995). primary type data bishop.; type; 267.; cameron coll./ 1910-55.; clavatipes k.; holo-/ type; lectotype ♂/ diapria tropidopria/ clavatipes kieffer/ des. notton, 1995. type locality scotland, bishopton. remarks this species was described from an unspecified number of males. by comparison with other specimens, the type locality “bishop.” almost certainly means bishopton in dumfries and galloway. the lectotype is mounted on a card and has four tarsi broken and the right hind wing lost. notton d.g., diapriinae types at the natural history museum, london 73 trichopria fimbriata kieffer, 1911b: 995. valid name trichopria nigra (nees, 1834). summary of types lectotype ♂, bmnh number 9.703, designated by notton (1995). primary type data craigton/ fungus 22-9; type; 270.; cameron coll./ 1910-55.; fimbriata k.; holo-/ type; lectotype ♂/ trichopria/ fimbriata kieffer/ des. notton, 1995. type locality scotland, craigton. remarks this species was described from an unspecified number of males. the type locality is most probably craigton in dumfries and galloway, but might instead by craigton in glasgow. the lectotype is mounted on a card, has both antennae and two tarsi broken, and both fore wings are disarticulated. trichopria ciliaris kieffer, 1911b: 1000. valid name trichopria basalis (thomson, 1858). summary of types lectotype ♀, bmnh number 9.704, designated by notton (1995). paralectotypes 2 ♀♀. primary type data clober 1-6; type; 229.; diapria/ ciliaris; cameron coll./ 1910-55.; ciliaris k.; lectotype ♀/ trichopria/ ciliaris kieffer/ des. notton, 1995. type locality scotland, clober. remarks this species was described from at least two females, since a range of lengths is given. the lectotype is mounted on a card and is entire. trichopria formicaria kieffer, 1911c: 385. valid name trichopria sociabilis masner, 1965. summary of types lectotype ♀, bmnh number 9.768, designated by notton (1995). european journal of taxonomy 75: 1-123 (2014) 74 primary type data trichopria/ sociabilis/ box/ hill/ 20.v.1910; type; ex coll. donisthorpe/ b.m.1934-4; assoc. with/ formica fusca; trichopria/ formicaria kief.; holo-/ type; ♀ lectotype/ trichopria/ formicaria/ kieffer, 1911: 385/ des. notton, 1995/ =trichopria/ sociabilis masner, 1965. type locality england, box hill. remarks this species was described from an unspecified number of females. the lectotype is mounted on a card and is entire. schizopria fallax kieffer, 1912a: 68, pl. 3, fig. 11. valid name entomacis fallax (kieffer, 1912). summary of types syntype ♀, bmnh number 9.683. syntypes 2 ♀♀. primary type data 103; silhouette, ’08-9,/ seychelles exp.; schizopria/ fallax/ j.j.kieffer/ type; figured specimen; type; type/ h.t.; seychelle islands./ percy sladen trust/ expedition./ 1913-170 (syntype ♀). 97; mahe, ’089,/ seychelles exp.; schizopria/ fallax/ j.j.kieffer/ paratype; seychelle islands./ percy sladen trust/ expedition./ 1913-170 (syntype ♀). 84; mahe, ’08-9,/ seychelles exp.; schizopria/ fallax/ j.j.kieffer/ paratype; seychelle islands./ percy sladen trust/ expedition./ 1913-170 (syntype ♀). type locality seychelles, silhouette, near mont pot-à-eau; silhouette (as mahé), mare aux cochons dist.; mahé, cascade estate. remarks three syntypes were found which agree with the description except for having 13-segmented antenna; although kieffer says the antenna is 12-segmented, this may be explained by a12-13 being closely joined, giving the appearance of 12 segments. the holotype label and paratype labels are erroneous, as kieffer did not designate a holotype; consequently, they are all syntypes. the specimen numbered “103” is mounted on a card and is entire; “97” is mounted on a point and has the head and one antenna mounted separately; “84” is mounted on a point and is entire. all three syntypes belong to entomacis. schizopria flaviclava kieffer, 1912a: 68, 69. valid name entomacis flaviclava (kieffer, 1912). summary of types holotype ♀, bmnh number 9.684, by monotypy. notton d.g., diapriinae types at the natural history museum, london 75 primary type data high jungle/ 126; silhouette, ’08-9./ seychelles exp.; schizopria/ flaviclava/ j.j.kieffer/ type.; type; type/ h.t.; seychelle islands./ percy sladen trust/ expedition./ 1913-170. type locality seychelles, silhouette, mare aux cochons. remarks the holotype is mounted on a card and is entire. the antenna is sub-12 segmented, the club apparently formed of two closely joined segments. entomacis longicornis kieffer, 1912a: 69. valid name entomacis longicornis kieffer, 1912. summary of types syntype ♀, bmnh number 9.685. primary type data 97; mahe, ’08-9./ seychelles exp.; entomacis/ longicornis/ j.j.kieffer/ type; type; type/ h.t.; seychelle islands./ percy sladen trust/ expedition./ 1913-170. type locality seychelles, mahé, cascade estate. remarks one syntype was found standing over this name. the holotype label is erroneous, as kieffer did not designate a holotype. the syntype is mounted on a card point and is entire. entomacis curticornis kieffer, 1912a: 69, 70. valid name entomacis curticornis kieffer, 1912. summary of types syntype ♂, bmnh number 9.686. syntype ♀. primary type data 97; mahe, ’08-9./ seychelles exp.; entomacis/ curticornis/ j.j.kieffer/ type.; type; type/ h.t.; seychelle islands./ percy sladen trust/ expedition./ 1913-170. (syntype ♂). 146; mahe, ’08-9./ seychelles exp.; entomacis/ curticornis/ j.j.kieffer/ paratype.; seychelle islands./ percy sladen trust/ expedition./ 1913-170 (syntype ♀). type locality seychelles, mahé, cascade estate; mahé, anse aux pins and anse royal. european journal of taxonomy 75: 1-123 (2014) 76 remarks two syntypes were found. the holotype and paratype labels are erroneous, as kieffer did not designate a holotype. the male syntype is mounted on a card point and is entire; the female syntype is mounted on a card and is entire. loxotropa exsul kieffer, 1912a: 70. valid name basalys exsul (kieffer, 1912). summary of types syntype ♀ bmnh number 9.738. syntype ♂. primary type data 22; silhouette, ‘08/ seychelles exp.; loxotropa/ exsul/ j.j.kieffer/ type; type; type/ h.t.; seychelle islands./ percy sladen trust/ expedition./ 1913-170 (syntype ♀). 20; 20/ silhouette, ‘08/ seychelles exp.; loxotropa/ exsul/ j.j.kieffer/ paratype; seychelle islands./ percy sladen trust/ expedition./ 1913170 (syntype ♂). type locality seychelles, silhouette, mare aux cochons; mahé, slopes of morne seychellois. remarks two syntypes were found. the holotype and paratype labels are erroneous, as kieffer did not designate a holotype. the male syntype is mounted on a card point and is entire; the female syntype is mounted on a card point and the left flagellum, gaster and tip of the right hind wing are missing. loxotropa semirufa kieffer, 1912a: 70. valid name basalys semirufus (kieffer, 1912). summary of types holotype ♂, bmnh number 9.739, by monotypy. primary type data 19/ silhouette, ‘08/ seychelles exp.; loxotropa/ semirufa/ j.j.kieffer/ type; type; type/ h.t.; seychelle islands./ percy sladen trust/ expedition./ 1913-170. type locality seychelles, silhouette, mare aux cochons. remarks the holotype agrees well with the description, although it is somewhat faded. it is mounted on a card and is missing its right wings. notton d.g., diapriinae types at the natural history museum, london 77 rhopalopria vulgaris kieffer, 1912a: 71, pl. 3, fig. 12. valid name trichopria vulgaris (kieffer, 1912). summary of types lectotype ♀, bmnh number 9.689, designated by huggert (1979). paralectotypes ♀, 3 ♂♂. primary type data 99/ mahe, ’08-9./ seychelles exp.; paratype; rhopalopria/ vulgaris/ j.j.kieffer/ paratype; seychelle islands./ percy sladen trust/ expedition./ 1913-170; lectotypus/ rhopalopria/ vulgaris/ kieff./ design. ♀/ l.huggert-77; trichopria ♀/ vulgaris kieff./ det. l.huggert-77. type locality seychelles, mahé. remarks five syntypes were found. the holotype and paratype labels are erroneous as kieffer did not designate a holotype. the syntype series was analysed by huggert (1979) who designated a lectotype. the lectotype is mounted on a cork block and has the left antenna and left fore wing mounted on a microslide and the left hind wing missing. huggert considered one paralectotype male of r. vulgaris to be a different species and designated it the holotype of trichopria euvulgaris huggert q.v. diapria seychellensis kieffer, 1912a: 71, 72, fig. 13. valid name trichopria seychellensis (kieffer, 1912). summary of types lectotype ♂, bmnh number 9.699, designated by huggert (1979). paralectotypes 2 ♀♀, ♂. primary type data 22/ silhouette, ‘08/ seychelles exp./ ♂/ diapria/ seychellensis; type; type/ h.t.; diapria/ seychellensis/ j.j.kieffer/ type; figured specimen; seychelle islands./ percy sladen trust/ expedition./ 1913-170; lectotypus/ diapria ♂/ seychellensis/ kieff./ design./ l.huggert-77; trichopria (a.) ♂/ seychellensis (kieff.)/ det. l.huggert-77. type locality seychelles, silhouette, mare aux cochons. remarks this species was described from three females and five males. two females and two males were located by huggert (1979), of which he designated one as lectotype of d. seychellensis. the lectotype is mounted on a papered cork block; the left antenna and left wings are mounted on a microslide on the same pin. huggert considered two of the paralectotypes were not conspecific with t. seychellensis and designated one the holotype of trichopria ensia q.v. and another the holotype of trichopria percyi q.v. european journal of taxonomy 75: 1-123 (2014) 78 diapria scotti kieffer, 1912a: 71, 72. valid name trichopria scotti (kieffer, 1912). summary of types lectotype ♂, bmnh number 9.700, designated by huggert (1979). paralectotype ♀. primary type data 67; diapria/ scotti/ j.j.kieffer/ type; mahe, ‘08-9./ seychelles exp.; type; type/ h.t.; seychelle islands./ percy sladen trust/ expedition./ 1913-170; lectotypus/ diapria ♂/ scotti kieff./ design./ l.huggert-76. type locality seychelles, mahé, nr. morne blanc. remarks the holotype label on the lectotype is erroneous as kieffer did not designate a holotype. the lectotype is mounted on a card and has the right hind wing missing and the left antenna and left wings on a microslide on the same pin. diapria mahensis kieffer, 1912a: 71, 73. valid name trichopria mahensis (kieffer, 1912). summary of types holotype ♀, bmnh number 9.701, by monotypy. primary type data diapria/ mahensis/ j.j.kieffer/ type; mahe, ’08-9./ seychelles exp.; type; type/ h.t.; holotypus ♀/ diapria/ mahensis kieffer/ l.huggert-76; trichopria ♀/ mahensis kieff./ det. l.huggert-76. type locality seychelles, mahé. remarks the holotype is mounted on a card with the left fore wing and left antenna on a microslide mounted on the same pin. diapria saxatilis kieffer, 1912a: 71, 73. valid name trichopria saxatilis (kieffer, 1912). summary of types lectotype ♂, bmnh number 9.702, designated by huggert (1977). paralectotypes ♀, ♂. notton d.g., diapriinae types at the natural history museum, london 79 primary type data 48; diapria/ saxatilis/ j.j.kieffer/ type; praslin, ’08./ seychelles exp.; type; lectotypus/ diapria ♂/ saxatilis kieff./ design./ l.huggert-76; trichopria ♂/ saxatilis kieff./ det. huggert-76. type locality seychelles, praslin, côtes d’or estate, vallée de mai. remarks this species was described from two males and three females. huggert (1977) found two males and one female syntype, designating one male as the lectotype of d. saxatilis. the lectotype is mounted on a card; the left wings and left antenna are mounted on a microslide on the same pin. huggert considered that two of the paralectotypes were not conspecific with t. saxatilis and designated one the holotype of trichopria pseudosaxatilis q.v. and the other the holotype of t. nirva q.v. calogalesus parvulus kieffer, 1912a: 73. valid name calogalesus parvulus kieffer, 1912. summary of types holotype ♀, bmnh number 9.675, by monotypy. primary type data type/ h.t.; mahe ’08-9/ seychelles exp.; calogalesus/ parvulus/ j.j.kieffer/ type; type. type locality seychelles, mahé, cascade estate. remarks kieffer based his description on one specimen, hence a holotype by monotypy. the holotype is mounted on a card point; the left fore wing, right hind wing and some legs are lost. microgalesus quadridens kieffer, 1912a: 74. valid name basalys quadridens (kieffer, 1912) comb. nov. summary of types holotype ♀, bmnh number 9.674, by monotypy. primary type data type/ h.t.; mahe, ’08-9/ seychelles exp.; microgalesus/ quadridens/ j.j.kieffer/ type. type locality seychelles, mahé, cascade estate. european journal of taxonomy 75: 1-123 (2014) 80 remarks kieffer based his description on one specimen, hence a holotype by monotypy. the holotype is mounted on a card point; the head, right hind wing and parts of the legs are lost; the right fore wing is disarticulated. the triangular stigmalis, three-segmented antennal club, notauli represented by a weak trace, and form of the propodeum are typical of basalys. the absence of the basal vein is not surprising since the specimen is so small. the beak-like mandibles, anteriorly indented head with teeth, and the transverse petiole are known in other representatives of this genus. microgalesus quadridens is here transferred to basalys. since m. quadridens is the type species of microgalesus, microgalesus is now a junior synonym of basalys syn. nov. galesus (schizogalesus) silvestrii kieffer, 1913a: 91. valid name coptera silvestrii (kieffer, 1913). summary of types paralectotypes 3 ♂♂. remarks three male syntypes were found. these are now paralectotypes following the lectotype designation of yoder in yoder & wharton (2002). loxotropa donisthorpei kieffer, 1913b: 176. remarks the specimen with bmnh number 9.763 is not a type; the lectotype is in the collection of the university museum of natural history, oxford (notton 1995). aneurhynchus nodicornis marshall, 1867: 225. remarks a male of this distinctive species was found, originally from the claude morley collection and labelled as a metatype, i.e., a specimen that had been compared with the type by the author of the species. this specimen has no formal type status within the iczn code; however, it was determined by marshall and may be of use in the interpretation of this species should the type not be located. mimopria campbellorum masner, 1976: 124, 126, figs 1-6. valid name mimopria campbellorum masner, 1976. summary of types paratypes 3 ♀♀. spilomicrus boweni masner, 1991: 118, 124, figs 26, 28, 83-85, 108, 123-125. valid name spilomicrus boweni masner, 1991. notton d.g., diapriinae types at the natural history museum, london 81 summary of types paratypes ♀, ♂. spilomicrus distinctus masner, 1991: 118, 128, figs 27, 81-82, 107, 114, 122. valid name spilomicrus distinctus masner, 1991. summary of types paratypes ♀, ♂. spilomicrus exul masner, 1991: 117, 131, figs 13, 77-79, 109, 112, 119. valid name spilomicrus exul masner, 1991. summary of types paratype ♂. spilomicrus inornatus masner, 1991: 119, 136, figs 25, 65-67, 102, 115. valid name spilomicrus inornatus masner, 1991. summary of types paratypes ♀, ♂. spilomicrus palustris masner, 1991: 118, 139, figs 86-86a, 110. valid name spilomicrus palustris masner, 1991. summary of types paratype ♂. spilomicrus pumilio masner, 1991: 119, 140, figs 4, 46-48, 94. valid name spilomicrus pumilio masner, 1991. summary of types paratype ♀. spilomicrus sanbornei masner, 1991: 117, 145, figs 5-7, 40-42, 95. valid name spilomicrus sanbornei masner, 1991. european journal of taxonomy 75: 1-123 (2014) 82 summary of types paratype ♂. spilomicrus stepheni masner, 1991: 118, 147, figs 19, 74-76, 106. valid name spilomicrus stepheni masner, 1991. summary of types paratype ♀. spilomicrus superbus masner, 1991: 117, 151, figs 3, 43-45, 93, 120. valid name spilomicrus lubomiri özdikmen, 2010. summary of types paratype ♂. spilomicrus sylvicola masner, 1991: 118, 153, figs 11-12, 37-39, 92, 116. valid name spilomicrus sylvicola masner, 1991. summary of types paratype ♀. chilomicrus pecki masner & garcía, 2002: 42, figs 32, 33. valid name chilomicrus pecki masner & garcía, 2002. summary of types paratype ♂. apopria coveri masner & garcía, 2002: 65, figs 8, 9. valid name apopria coveri masner & garcía, 2002. summary of types paratype ♀. eladio cruzi masner & garcía, 2002: 80, figs 45-47. valid name eladio cruzi masner & garcía, 2002. notton d.g., diapriinae types at the natural history museum, london 83 summary of types paratype ♀. hansonia pauli masner & garcía, 2002: 82, fig. 57. valid name hansonia pauli masner & garcía, 2002. summary of types paratype ♀. leucopria cylindricornis masner & garcía, 2002: 86, fig. 61. valid name leucopria cylindricornis masner & garcía, 2002. summary of types paratype ♀. turripria woldai masner & garcía, 2002: 111, fig. 100. valid name turripria woldai masner & garcía, 2002. summary of types paratype ♀. peckidium enigmaticum masner & garcía, 2002: 119, figs 82-83. valid name peckidium enigmaticum masner & garcía, 2002. summary of types paratype ♀. loxotropa morleii morley, 1931: 15. valid name trichopria halterata (kieffer, 1909). summary of types holotype ♀, bmnh number 9.784, by original designation. primary type data 3 ix 97; loxotropa/ morleyi/ ms. chitty; det. a.j.c.; probably paramesius/ ♀ undescribed tam; type/ cm. european journal of taxonomy 75: 1-123 (2014) 84 type locality england, suffolk, sproughton marshes. remarks the name loxotropa morleii was introduced as a nomen nudum by morley (1929) and then made available in 1931. morley credited the name to chitty, and had a manuscript description from him placing this species in trichopria; however, by placing it in loxotropa förster, 1856 and comparing it to other species of loxotropa, it appears that morley has significantly changed chitty’s manuscript and was largely responsible for the text, and so is credited here as the author. the specimen is mounted on a card and is very dirty. termitopria sheasbyi muesebeck, 1965: 190, fig. 1a-e. valid name termitopria sheasbyi muesebeck, 1965. summary of types paratypes 4 ♀♀. psilus aquilonius muesebeck, 1980: 4, 6, 8, figs 1-1a, 4, 23, 34, 43-44, 61. valid name psilus aquilonius muesebeck, 1980. summary of types paratype ♂. psilus dissidens muesebeck, 1980: 3, 4, 10, figs 7, 25, 36, 47, 48, 62. valid name psilus dissidens muesebeck, 1980. summary of types paratype ♂. psilus fortis muesebeck, 1980: 4, 5, 11, figs 10, 27, 37, 51. valid name psilus fortis muesebeck, 1980. summary of types paratype ♂. psilus masneri muesebeck, 1980: 4, 12, figs 11, 38, 52. valid name psilus masneri muesebeck, 1980. notton d.g., diapriinae types at the natural history museum, london 85 summary of types paratype ♀. psilus masoni muesebeck, 1980: 3, 6, 13, figs 12, 28, 39, 53, 54. valid name psilus masoni muesebeck, 1980. summary of types paratype ♂. coptera angulata muesebeck, 1980: 24, 29. valid name coptera angulata muesebeck, 1980. summary of types paratype ♂. coptera distans muesebeck, 1980: 23, 26, 34. valid name coptera distans muesebeck, 1980. summary of types paratype ♂. coptera pomonellae muesebeck, 1980: 23, 26, 44, figs 80, 100, 119. valid name coptera pomonellae muesebeck, 1980. summary of types paratype ♂. coptera strauziae muesebeck, 1980: 22, 25-26, 46, figs 82, 101, 120-123, 133. valid name coptera strauziae muesebeck, 1980. summary of types paratype ♂. galesus (schizogalesus) quadridens nixon, 1930: 400, 401, 402. valid name coptera quadridens (nixon, 1930). european journal of taxonomy 75: 1-123 (2014) 86 summary of types syntype ♂, bmnh number 9.669. syntype ♀, bmnh number 9.669b. syntypes ♀, 8 ♂♂. primary type data type; natal/ van reenen,/ drakensberg/ 1-22.i.1927; africa/ r.e.turner/ brit. mus./ 1927-54; galesus/ quadridens/ 1930 nixon/ type ♀ (syntype ♀). paratype; natal/ kloof/ 1500ft/ sept. 1926; africa/ r.e.turner/ brit. mus./ 1926-404; galesus/ quadridens/ 1930 nixon (syntype ♀). type; natal/ van reenen,/ drakensberg/ 1-22.i.1927; africa/ r.e.turner/ brit. mus./ 1927-54; galesus/ quadridens/ 1930 nixon/ type ♂ (syntype ♂). paratype; natal/ van reenen,/ drakensberg/ 1-22.i.1927; africa/ r.e.turner/ brit. mus./ 1927-54; galesus/ quadridens/ 1930 nixon (syntypes 6♂♂). paratype; orange f. state/ harrismith/ feb. 1927; s.africa/ r.e.turner/ brit mus./ 1927-117; galesus/ quadridens/ 1930 nixon (syntype ♂). paratype; zululand:/ gingindhloyu/ 15.vi.1926; s.africa/ r.e.turner/ brit. mus./ 1926-277; galesus/ quadridens/ 1930 nixon (syntype ♂). type locality south africa, free state, harrismith; kwazulu-natal, gingindlovu, kloof and van reenen. remarks there are two females and nine males agreeing with nixon’s description and labelled by him as galesus quadridens. of these, two are labelled as “type” and the rest as “paratype“; however, since neither a unique holotype or lectotype has yet been fixed, all are still syntypes (article 73.2). all are mounted on card points, the females are entire, the male labelled by nixon as “type” is missing its head, and three other males are missing parts of their antennae. galesus (schizogalesus) bidens nixon, 1930: 400, 401, 403. valid name coptera bidens (nixon, 1930). summary of types syntype ♀, bmnh number 9.668. syntype ♂, bmnh number 9.688b. syntypes 4 ♀♀, 5 ♂♂. primary type data type; mossel bay/ cape province/ october, 1921; s.africa/ r.e.turner/ brit. mus./ 1921-450; galesus/ bidens/ 1930 nixon/ type ♀ (syntype ♀). paratype; mossel bay/ cape province/ june, 1921; s.africa/ r.e.turner/ brit. mus./ 1921-294; galesus/ bidens/ 1930 nixon (syntype ♀). paratype; mossel bay/ cape province/ october, 1921; s.africa/ r.e.turner/ brit. mus./ 1921-450; galesus/ bidens/ 1930 nixon (syntype ♀). paratype; mossel bay/ cape province/ april, 1921; s.africa/ r.e.turner/ brit. mus./ 1921-210; galesus/ bidens/ 1930 nixon (syntype ♀). paratype; mossel bay/ cape province/ aug. 1921; s.africa/ r.e.turner/ brit. mus./ 1921-353; galesus/ bidens/ 1930 nixon (syntype ♀). type; mossel bay/ cape province/ april, 1921; s.africa/ r.e.turner/ brit. mus./ 1921-210; galesus/ bidens/ 1930 nixon/ type ♂ (syntype ♂). paratype; mossel bay/ cape province/ febr. 1922; s.africa/ r.e.turner/ brit. mus./ 1922-97; galesus/ bidens/ 1930 nixon (syntype ♂). paratype; mossel bay/ cape province/ 18-30.xi.1921; s.africa/ r.e.turner/ brit. mus./ 1922-2; galesus/ bidens/ 1930 nixon (syntype ♂). paratype; mossel bay/ cape province/ 5-31.vii.1921; s.africa/ r.e.turner/ brit. mus./ 1921-315; galesus/ bidens/ 1930 nixon (syntype ♂). paratype; mossel bay/ cape province/ 1-13. iii.1922; s.africa/ r.e.turner/ brit. mus./ 1922-129; galesus/ bidens/ 1930 nixon (syntypes 2♂♂). notton d.g., diapriinae types at the natural history museum, london 87 type locality south africa, western cape province, mossel bay. remarks there are five females and six males agreeing with nixon’s description and labelled by him as galesus bidens. of these, two are labelled as “type” and the rest as “paratype”; however, since neither a unique holotype or lectotype has yet been fixed, all of the type series is syntypic (article 73.2). all the syntypes are mounted on cards and one female has an antenna mounted separately; two males are missing parts of their antennae. galesus (schizogalesus) inquisitor nixon, 1930: 400, 402, 404. valid name coptera inquisitor (nixon, 1930). summary of types syntype ♀, bmnh number 9.659. syntype ♂, bmnh number 9.659b. syntypes 12 ♀♀, 12 ♂♂. primary type data type; mossel bay/ cape province/ 5-31.vii.1921; s.africa/ r.e.turner/ brit. mus./ 1921-315; galesus/ inquisitor/ 1930 nixon/ type ♀ (syntype ♀). paratype; mossel bay/ cape province/ 15-28.iii.1922; s.africa/ r.e.turner/ brit. mus./ 1922-153; galesus/ inquisitor/ 1930 nixon (syntype ♀). paratype; mossel bay/ cape province/ 18-30.xi.1921; s.africa/ r.e.turner/ brit. mus./ 1922-2; galesus/ inquisitor/ 1930 nixon (syntype ♀). paratype; mossel bay/ cape province/ aug. 1921; s.africa/ r.e.turner/ brit. mus./ 1921-353; galesus/ inquisitor/ 1930 nixon (syntypes ♀, 4 ♂♂). paratype; mossel bay/ cape province/ may, 1921; s.africa/ r.e.turner/ brit. mus./ 1921-248; galesus/ inquisitor/ 1930 nixon (syntype ♀). paratype; mossel bay/ cape province/ october, 1921; s.africa/ r.e.turner/ brit. mus./ 1921-450; galesus/ inquisitor/ 1930 nixon (syntypes ♀, 4 ♂♂). paratype; mossel bay/ cape province/ 5-31.vii.1921; s.africa/ r.e.turner/ brit. mus./ 1921-315; galesus/ inquisitor/ 1930 nixon (syntypes ♀, 2 ♂♂). paratype; aliwal north/ cape province/ 4350ft/ 1-13.i.1923; s.africa/ r.e.turner/ brit. mus./ 1923-70; galesus/ inquisitor/ 1930 nixon (syntype ♀). paratype; port st. john/ pondoland/ april 5-30. 1923; s.africa/ r.e.turner/ brit. mus./ 1923-286; galesus/ inquisitor/ 1930 nixon (syntype ♀). paratype; natal/ kloof/ 1500ft/ sept. 1926; s.africa/ r.e.turner/ brit. mus./ 1926-404; galesus/ inquisitor/ 1930 nixon (syntype ♀). paratype; natal/ van reenen/ drakensberg/ nov. 1926; s.africa/ r.e.turner/ brit. mus./ 1926-499; galesus/ inquisitor/ 1930 nixon (syntypes 2 ♀♀). paratype; orange f. state/ harrismith/ march 1-20. 1927; s.africa/ r.e.turner/ brit. mus./ 1927-147; galesus/ inquisitor/ 1930 nixon (syntype ♀). type; cape province/ ceres/ march 1925; s.africa/ r.e.turner/ brit. mus./ 1925-161; galesus/ inquisitor/ 1930 nixon/ type ♂ (syntype ♂). paratype; mossel bay/ cape province/ dec. 1921; s.africa/ r.e.turner/ brit. mus./ 1922-25; galesus/ inquisitor/ 1930 nixon (syntype ♂). paratype; port st. john/ pondoland/ may. 1924; s.africa/ r.e.turner/ brit. mus./ 1924-289; galesus/ inquisitor/ 1930 nixon (syntype ♂). type locality south africa, eastern cape province, aliwal north and port st johns; free state, harrismith; kwazulunatal, van reenen and kloof; western cape province, ceres and mossel bay. european journal of taxonomy 75: 1-123 (2014) 88 remarks there are thirteen females and thirteen males agreeing with nixon’s description and labelled by him as galesus inquisitor. of these, two are labelled as “type” and the rest as “paratype”; however, since neither a unique holotype or lectotype has yet been fixed, all of the type series is syntypic (article 73.2). all syntypes are mounted on cards, except the four from natal and orange free state, which are mounted on card points. galesus (schizogalesus) distinctus nixon, 1930: 400, 405. valid name coptera distincta (nixon, 1930). summary of types holotype ♀, bmnh number 9.655, by monotypy. primary type data type; s.africa/ r.e.turner/ brit. mus./ 1921-315; mossel bay/ cape province/ 5-31.vii.1921; galesus/ distinctus/ nixon/ ♀ type 1930. type locality south africa, western cape province, mossel bay. remarks the holotype is mounted on a card and is entire. galesus (schizogalesus) rectangularis nixon, 1930: 400, 402, 406. valid name coptera rectangularis (nixon, 1930). summary of types syntype ♀, bmnh number 9.656. syntype ♂, bmnh number 9.656b. syntypes 3 ♀♀. primary type data type; port st. john/ pondoland/ nov. 1923; s.africa/ r.e.turner/ brit. mus./ 1924-6; galesus/ rectangularis/ nixon/ ♀ type 1930 (syntype ♀). paratype; port st. john/ pondoland/ aug. 15-31 1923; s.africa/ r.e.turner/ brit. mus./ 1923-463; galesus/ rectangularis/ nixon/ ♀ paratype 1930 (syntype ♀). paratype; port st. john/ pondoland/ jan. 1924; s.africa/ r.e.turner/ brit. mus./ 1924-97; galesus/ rectangularis/ nixon/ ♀ paratype 1930 (syntype ♀). paratype; mossel bay/ cape province/ dec. 1921; s.africa/ r.e.turner/ brit. mus./ 1922-25; galesus/ rectangularis/ nixon/ ♀ paratype 1930 (syntype ♀). type; mossel bay/ cape province/ june 1921; s.africa/ r.e.turner/ brit. mus./ 1921-294; galesus/ rectangularis/ nixon/ ♂ type 1930 (syntype ♂). type locality south africa, eastern cape province, port st johns; western cape province, mossel bay. notton d.g., diapriinae types at the natural history museum, london 89 remarks there are four females and one male agreeing with nixon’s description and labelled by him as galesus rectangularis. of these, two are labelled as “type” and the rest as “paratype”; however, since neither a unique holotype or lectotype has yet been fixed, all of the type series is syntypic (article 73.2). all the syntypes are mounted on card points and are entire. galesus (schizogalesus) abdominalis nixon, 1930: 400, 402, 406. valid name coptera abdominalis (nixon, 1930). summary of types syntype ♀, bmnh number 9.667. syntype ♂, bmnh number 9.667b. syntypes 2 ♀♀, ♂. primary type data type; port st. john/ pondoland/ july 10-31, 1923; s.africa/ r.e.turner/ brit. mus./ 1923-398; galesus/ abdominalis/ nixon; ♀ type, 1930 (syntype ♀). paratype; port st. john/ pondoland/ sept. 1923; s.africa/ r.e.turner/ brit. mus./ 1923-510; galesus/ abdominalis/ nixon; ♀ paratype, 1930 (syntype ♀). paratype; port st. john/ pondoland/ 6-25 feb. 1923; s.africa/ r.e.turner/ brit. mus./ 1924-136; galesus/ abdominalis/ nixon; ♀ paratype, 1930 (syntype ♀). type; port st. john/ pondoland/ july 10-31, 1923; s.africa/ r.e.turner/ brit. mus./ 1923-398; galesus/ abdominalis/ nixon; ♂ type, 1930 (syntype ♂). paratype; port st. john/ pondoland/ july 10-31, 1923; s.africa/ r.e.turner/ brit. mus./ 1923-398; galesus/ abdominalis/ nixon; ♂ paratype, 1930 (syntype ♂). type locality south africa, eastern cape province, port st johns. remarks there are three females and two males agreeing with nixon’s description and labelled by him as galesus abdominalis. of these, two are labelled as “type” and the rest as “paratype”; however, since neither a unique holotype or lectotype has yet been fixed, all of the type series is syntypic (article 73.2). all the syntypes are mounted on cards and are entire. galesus (schizogalesus) magnificus nixon, 1930: 401, 408. valid name coptera robustior (silvestri, 1913). summary of types syntype ♀, bmnh number 9.660. syntype ♀. primary type data type; natal/ kloof/ 1500ft./ sept. 1926; s.africa/ r.e.turner/ brit. mus./ 1926-404; galesus/ magnificus/ ♀ type nixon/ 1930 (syntype ♀). paratype; port st. john/ pondoland/ 6-25.feb.1924; s.africa/ r.e.turner/ brit. mus./ 1924-136.; galesus/ magnificus/ ♀ paratype nixon/ 1930 (syntype ♀). type locality south africa, eastern cape province, port st johns; kwazulu-natal, kloof. european journal of taxonomy 75: 1-123 (2014) 90 remarks there are two females agreeing with nixon’s description and labelled by him as galesus magnificus. of these, one is labelled as “type” and the other “paratype”; however, since neither a unique holotype or lectotype has yet been fixed, all of the type series is syntypic (article 73.2). both syntypes are mounted on card points and are entire. galesus (schizogalesus) modestus nixon, 1930: 401, 408. valid name coptera modesta (nixon, 1930). summary of types syntype ♀, bmnh number 9.665. syntype ♀. primary type data type; port st. john/ pondoland/ july 10-31, 1923; s.africa/ r.e.turner/ brit. mus./ 1923-398; galesus/ modestus/ nixon ♀ type 1930 (syntype ♀). type; port st. john/ pondoland/ jan. 1924; s.africa/ r.e.turner/ brit. mus./ 1924-97; galesus/ modestus/ nixon ♀ paratype 1930 (syntype ♀). type locality south africa, eastern cape province, port st johns. remarks there are two females agreeing with nixon’s description and labelled by him as galesus modestus. of these, one is labelled as “type” and the other “paratype”; however, since neither a unique holotype or lectotype has yet been fixed, all of the type series is syntypic (article 73.2). both syntypes are mounted on card points and are entire. galesus (schizogalesus) turneri nixon, 1930: 401, 402, 409. valid name coptera turneri (nixon, 1930). summary of types syntype ♀, bmnh number 9.670. syntype ♂, bmnh number 9.670b. syntypes 2 ♀♀, ♂. primary type data type; port st. john/ pondoland/ oct. 1923; s.africa/ r.e.turner/ brit. mus./ 1923-547; galesus/ turneri/ nixon/ ♀ type 1930 (syntype ♀). paratype; port st. john/ pondoland/ 18-31 mar. 1924; s.africa/ r.e.turner/ brit. mus./ 1924-191; galesus/ turneri/ nixon/ ♀ paratype 1930 (syntype ♀). paratype; mossel bay/ cape province/ dec. 1921; s.africa/ r.e.turner/ brit. mus./ 1922-25; galesus/ turneri/ nixon/ ♀ paratype 1930 (syntype ♀). type; mossel bay/ cape province/ dec. 1921; s.africa/ r.e.turner/ brit. mus./ 1922-25; galesus/ turneri/ nixon/ ♂ type 1930 (syntype ♂). paratype; mossel bay/ cape province/ 18-30.xi.1921; s.africa/ r.e.turner/ brit. mus./ 1922-2; galesus/ turneri/ nixon/ ♂ paratype 1930 (syntype ♂). type locality south africa, eastern cape province, port st johns; western cape province, mossel bay. notton d.g., diapriinae types at the natural history museum, london 91 remarks there are three females and two males agreeing with nixon’s description and labelled by him as galesus turneri. of these, two are labelled as “type” and the rest as “paratype”; however, since neither a unique holotype or lectotype has yet been fixed, all of the type series is syntypic (article 73.2). the two syntypes from port st john are mounted on card points, the three syntypes from mossel bay are mounted on cards; all are entire except for the male labelled as a paratype, which has the metasoma mounted separately. galesus (schizogalesus) turneri var. carinaticeps nixon, 1930: 401, 402, 410. valid name coptera carinaticeps (nixon, 1930). summary of types syntype ♀, bmnh number 9.671. syntype ♂, bmnh number 9.671b. syntype ♂. primary type data type; mossel bay/ cape province/ october 1921; s.africa/ r.e.turner/ brit. mus./ 1921-450; galesus/ turneri nixon/ v. carinaticeps/ nixon/ ♀ type 1930 (syntype ♀). type; mossel bay/ cape province/ 15-31.vii.1921; s.africa/ r.e.turner/ brit. mus./ 1921-315; galesus/ turneri nixon/ v. carinaticeps/ nixon/ ♂ type 1930 (syntype ♂). paratype; port st. john/ pondoland/ sept. 1923; s.africa/ r.e.turner/ brit. mus./ 1923-510; galesus/ turneri nixon/ v. carinaticeps/ nixon/ ♂ paratype 1930/ g.nixon det. (syntype ♂). type locality south africa, western cape province, mossel bay, but evidently including eastern cape province, port st johns. remarks there are one female and two males agreeing with nixon’s description and labelled by him as galesus turneri v. carinaticeps. of these, two are labelled as “type” and the third as “paratype”; however, since neither a unique holotype or lectotype has yet been fixed, all of the type series is syntypic (article 73.2). the specimen labelled as “paratype” is from port st john, not mossel bay; however, this appears to be a lapsus by nixon. it appears he included it in the description since he referred to two males, so it is not excluded here from the type series. the two syntypes labelled as “type” are mounted on card points, the other is mounted on a card. galesus (schizogalesus) longiceps nixon, 1930: 400, 402, 410. valid name coptera longiceps (nixon, 1930). summary of types syntype ♀, bmnh number 9.666. syntype ♂, bmnh number 9.666b. primary type data type; port st. john/ pondoland/ sept. 1923; s.africa/ r.e.turner/ brit. mus./ 1923-510; galesus/ longiceps/ nixon/ ♀ type 1930 (syntype ♀). type; mossel bay/ cape province/ april 1921; s.africa/ r.e.turner/ brit. mus./ 1923-510; galesus/ longiceps/ nixon/ ♂ type 1930 (syntype ♂). european journal of taxonomy 75: 1-123 (2014) 92 type locality south africa, eastern cape province, port st johns; western cape province, mossel bay. remarks there are one female and one male agreeing with nixon’s description and labelled by him as galesus longiceps, both of which are labelled as “type”. since neither a unique holotype or lectotype has yet been fixed, both specimens are syntypes (article 73.2). the female syntype is mounted on a card point, the male syntype is mounted on a card, and both are entire. galesus (schizogalesus) silvestrii var. nigricornis nixon, 1930: 401, 412. valid name coptera mosselensis nom. nov. summary of types syntype ♂, bmnh number 9.658. syntypes 3 ♂♂. primary type data type; mossel bay/ cape province/ dec. 1921; s.africa/ r.e.turner/ brit. mus./ 1922-25; galesus/ silvestrii kieff./ var. nigricornis/ nixon (syntype ♂). paratype; mossel bay/ cape province/ 18-30. xi.1921; s.africa/ r.e.turner/ brit. mus./ 1922-2; galesus/ silvestrii kieff./ var. nigricornis/ nixon (syntypes 2♂♂). paratype; mossel bay/ cape province/ 5-31.vii.1921; s.africa/ r.e.turner/ brit. mus./ 1921-315; galesus/ silvestrii kieff./ var. nigricornis/ nixon (syntype ♂). type locality south africa, western cape province, mossel bay. remarks there are four males agreeing with nixon’s description and labelled by him as galesus silvestrii v. nigricornis. of these, one is labelled as “type” and rest as “paratype”; however, since neither a unique holotype or lectotype has yet been fixed, all of the type series is syntypic (article 73.2). all syntypes are mounted on cards and are entire. this taxon is not conspecific with coptera silvestrii; as well as the characters given by nixon 1930, it differs also in the shape of the frontal projections (outer angle forming a longer, more acute projection in silvestrii) and in the size of the lateral fovea on the scutellar disc (larger in nigricornis, approaching the size of the basal scutellar fovea) and should be given the rank of species. galesus (schizogalesus) silvestrii var. nigricornis nixon, 1930 is permanently invalid as a junior primary homonym of galesus (g.) foersteri var. nigricornis kieffer, 1911 and no other name appears to be available for it, so it is given a replacement name here based on the name of the type locality, mossel bay. galesus (schizogalesus) difficilis nixon, 1930: 401, 402, 412. valid name coptera difficilis (nixon, 1930). summary of types syntype ♀, bmnh number 9.657. syntype ♂, bmnh number 9.657b. syntypes 2 ♀♀, 12 ♂♂. notton d.g., diapriinae types at the natural history museum, london 93 primary type data type; mossel bay/ cape province/ febr. 1922; s.africa/ r.e.turner/ brit. mus./ 1922-97; galesus/ difficilis/ 1930 nixon/ type ♀ (syntype ♀). paratype; mossel bay/ cape province/ febr. 1922; s.africa/ r.e.turner/ brit. mus./ 1922-97; galesus/ difficilis/ 1930 nixon (syntype ♀). paratype; zululand/ mtunzini/ 7.vi.1926; s.africa/ r.e.turner/ brit. mus./ 1926-277; galesus/ difficilis/ 1930 nixon (syntype ♀). type; s.africa/ r.e.turner/ brit. mus./ 1922-129; mossel bay/ cape province/ 1-13. iii.1922; galesus/ difficilis/ 1930 nixon/ type ♂ (syntype ♂). paratype; mossel bay/ cape province/ 1-13.iii.1922; s.africa/ r.e.turner/ brit. mus./ 1922-129; galesus/ difficilis/ 1930 nixon (syntypes 4♂♂). paratype; mossel bay/ cape province/ febr. 1922; s.africa/ r.e.turner/ brit. mus./ 1922-97; galesus/ difficilis/ 1930 nixon (syntypes 4♂♂). paratype; mossel bay/ cape province/ 15-28.iii.1922; s.africa/ r.e.turner/ brit. mus./ 1922-158; galesus/ difficilis/ 1930 nixon (syntypes 3♂♂). paratype; umatata/ transkei/ 18.ii-18.iii.1923; s.africa/ r.e.turner/ brit. mus./ 1923-189; galesus/ difficilis/ 1930 nixon (syntype ♂). type locality south africa, eastern cape province, transkei; kwazulu-natal, mtunzini; western cape province, mossel bay. remarks there are three females and thirteen males agreeing with nixon’s description and labelled by him as galesus difficilis. of these, two are labelled as “type” and the rest as “paratype”; however, since neither a unique holotype or lectotype has yet been fixed, all of the type series is syntypic (article 73.2). all the male syntypes and the female syntype from mossel bay labelled as “paratype” are mounted on cards and are entire. the other female syntypes are mounted on card points and are entire. trichopria lewisi nixon, 1940: 59, fig. 1a-d. valid name trichopria lewisi nixon, 1940. summary of types holotype ♀, bmnh number 9.706, by original designation. paratypes 24♀♀, ♂. primary type data type; kenya colony/ kabete/ ex glossina/ brevipalpis &/ g. fuscipleuris/ dr e.a.lewis; trichopria/ lewisi nixon/ ♀ type 1940; pres. by/ imp. inst. ent./ b.m.1940-181. type locality kenya, kabete. remarks although nixon mentioned 24 ♀♀ and 1 ♂ as paratypes, there are 63 ♀♀ and 3 ♂♂ specimens labelled as paratypes, so it is not clear which are nixon’s original paratypes. while the error in labelling paratypes is unfortunate, it is not likely to be a problem, as the holotype is certainly recognised and all the type specimens appear to be the same species. nixon’s preparation of the male genitalia is missing. the holotype is mounted on a card point and is entire. it is worth noting a singular feature of the male of this species which was not recorded by nixon: the male head is truncate behind, and the genal pubescence is long, and, unlike other trichopria, turned back to form a row from the top to the bottom of the gena, european journal of taxonomy 75: 1-123 (2014) 94 defining a vertical channel behind each gena, between the back of the head and the pronotum. by analogy with the observed mating behaviour of other trichopria it is predicted that this channel is used to constrain the female antenna during courtship. spilomicrus myrmecophilus nixon, 1947: 787, figs 1-3. valid name spilomicrus myrmecophilus nixon, 1947. summary of types holotype ♀, bmnh number 9.646, by original designation. paratypes 3 ♀♀. primary type data type; 93/ mauritius/ 26.x.1945/ r.mamet; in nest of solenopsis/ mameti/ donis.; spilomicrus/ myrmecophilus/ nixon/ type ♀, 1947; pres. by/ imp. inst. ent./ b.m.1947-417. type locality mauritius, corps de garde mountain. remarks the holotype is on a card point and is entire. entomacis penelope nixon, 1980: 12. valid name entomacis penelope nixon, 1980. summary of types holotype ♀, bmnh number 9.809, by original designation. primary type data eire/ co. sligo/ bundoran/ vii.1933/ g.e.j.nixon; entomacis/ penelope/ nixon/ type ♀, 197. type locality ireland, county sligo, bundoran. remarks the holotype is mounted on a card point and is entire. entomacis laertes nixon, 1980: 13. valid name entomacis graeffei kieffer, 1909. summary of types holotype ♀, bmnh number 9.810, by original designation. paratypes 5 ♀♀. notton d.g., diapriinae types at the natural history museum, london 95 primary type data sweden, sk./ ring sjö/ 27.vi.1938/ d.m.s.p. & j.f.p./ b.m.1938-414; entomacis/ laertes nixon/ type ♀, 197. type locality sweden, skåne county, ring sjö. remarks the holotype is mounted on a card point and is entire. spilomicrus pelion nixon, 1980: 17, figs 33, 40. valid name spilomicrus crassiclavis kieffer, 1911. summary of types holotype ♂, bmnh number 9.811, by original designation. paratype ♂. primary type data surrey/ horsley/ 14.vi.1930/ g.nixon; spilomicrus/ pelion nixon/ type ♂, 197. type locality england, surrey, horsley. remarks the holotype is mounted on a card point and is entire. aneurhynchus fragilis nixon, 1980: 23, fig. 60. valid name aneurhynchus fragilis nixon, 1980. summary of types holotype ♀, bmnh number 9.814, by original designation. primary type data england:/ glos. e./ badgeworth/ field mushroom/ 18.ix.1963; pres. by/ a.muncaster/ b.m.1963-697; aneurhynchus/ fragilis nixon/ type ♀ 1969. type locality england, gloucestershire, badgeworth. remarks the holotype is mounted on a card point, one antenna and the left mid leg are disarticulated, and the tips of three tarsi are lost. european journal of taxonomy 75: 1-123 (2014) 96 aneurhynchus ariadne nixon, 1980: 23, 24, fig. 67. valid name aneurhynchus ariadne nixon, 1980. summary of types holotype ♀, bmnh number 9.815, by original designation. paratypes 3 ♀♀. primary type data france, fi./ morlaix/ 10.vi.1954/ j.f.perkins/ b.m.1954-396; aneurhynchus/ ariadne/ nixon/ holotype ♀, 197. type locality france, morlaix. remarks the holotype is mounted on a card point and is entire. aneurhynchus mese nixon, 1980: 23, fig. 70. valid name aneurhynchus mese nixon, 1980. summary of types holotype ♀, bmnh number 9.816, by original designation. paratypes 3 ♀♀. primary type data eng: northants/ spratton/ viii. 1975/ i. & p.gauld; aneurhynchus/ mese nixon/ type ♀, 197. type locality england, northamptonshire, spratton. remarks the holotype is mounted on a card point, with the right antenna disarticulated. aneurhynchus coryphe nixon, 1980: 25, fig. 69. valid name aneurhynchus coryphe nixon, 1980. summary of types holotype ♂, bmnh number 9.817, by original designation. paratypes 3 ♂♂. primary type data france, fi./ huelgoat/ 3.vi.1954/ j.f.perkins/ b.m.1954-396; aneurhynchus/ coryphe nixon/ type ♂, 197. notton d.g., diapriinae types at the natural history museum, london 97 type locality france, huelgoat. remarks the holotype is mounted on a card point and is entire. basalys orion nixon, 1980: 26, fig. 81. valid name basalys orion nixon, 1980. summary of types holotype ♀, bmnh number 9.818, by original designation. paratype ♀. primary type data berks:/ windsor forest/ ix.1931/ h.st.j.donisthorpe; basalys/ orion nixon/ type ♀, 197. type locality england, berkshire, windsor forest. remarks the holotype is mounted on a card point and is entire. basalys cymocles nixon, 1980: 27, 29, fig. 74. valid name basalys cymocles nixon, 1980. summary of types holotype ♀, bmnh number 9.819, by original designation. paratypes 3 ♀♀. primary type data surrey/ ashtead/ 30.vii.1932/ g.nixon; basalys/ cymocles/ nixon/ type ♀, 197. type locality england, surrey, ashtead. remarks the holotype is mounted on a card point and is entire. basalys semele nixon, 1980: 27. valid name basalys semele nixon, 1980. summary of types holotype ♀, bmnh number 9.820, by original designation. european journal of taxonomy 75: 1-123 (2014) 98 primary type data white downs, near/ dorking surrey/ england/ lg. bouček 29.8.1970; basalys/ semele nixon/ type ♀, 197. type locality england, surrey, white downs near dorking. remarks the type is mounted on a card and is entire. basalys insignificans nixon, 1980: 28. valid name basalys insignificans nixon, 1980. summary of types holotype ♀, bmnh number 9.821, by original designation. paratype ♀. primary type data wales: glamorgan/ pontypridd/ 25.v.1975; grassland/ j.s.noyes/ b.m.1975-265; basalys/ insignificans/ nixon/ type ♀, 197. type locality wales, glamorgan, pontypridd. remarks the holotype is mounted on a card point and is entire. basalys singularis nixon, 1980: 28. valid name basalys singularis nixon, 1980. summary of types holotype ♀, bmnh number 9.823, by original designation. paratype ♀. primary type data furry glen/ co. du., a.w.s./ 26.9.37; ireland/ a.w.stelfox coll./ bmnh 1949-110; basalys/ singularis/ nixon/ type ♀, 197. type locality ireland, county dublin, furry glen. remarks the holotype is mounted on a card and is entire. notton d.g., diapriinae types at the natural history museum, london 99 basalys euterpe nixon, 1980: 30. valid name basalys semele nixon, 1980. summary of types holotype ♂, bmnh number 9.822, by original designation. paratypes 4 ♂♂. primary type data co. sligo/ trawallua/ 24-29.7.1933/ g.nixon; basalys/ euterpe/ nixon/ type ♂, 197. type locality ireland, county sligo, trawallua. remarks the holotype is mounted on a card point and is entire. basalys iphicla nixon, 1980: 30. valid name basalys macroptera (kieffer, 1911) syn. nov. summary of types holotype ♂, bmnh number 9.824, by original designation. paratype ♂. primary type data inverness:/ loch morlich/ 24.vi.1934/ r.b. & j.e.benson; pres. by the/ trustees of the/ godman fund./ b.m.1934-429; basalys/ iphicla/ nixon/ type ♀, 197. type locality scotland, inverness, loch morlich. remarks the holotype is mounted on a card point and has a12-14 and both hind tarsi missing. this species is evidently the same as basalys macroptera, with which it shares a number of characters, most notably its relatively large size, hairs on anterior margin of large tergite, straight carina just below apex of antennal shelf, dark colour and northern distribution. corynopria aphrodite nixon, 1980: 32, fig. 9. valid name monelata aphrodite (nixon, 1980). summary of types holotype ♀, bmnh number 9.825, by original designation. paratypes 3 ♀♀. primary type data s.e.london/ norwood/ vii.1929/ g.nixon; corynopria/ aphrodite/ nixon/ type ♀, 1979. european journal of taxonomy 75: 1-123 (2014) 100 type locality england, london, norwood (as tulse hill in original description). remarks the type series is from norwood, not tulse hill as nixon stated, although these places are very close; evidently this was just an error in recording the locality by nixon and should not prevent them from being types. the holotype is mounted on a card point and is entire. trichopria isis nixon, 1980: 33. valid name trichopria aequata (thomson, 1858). summary of types holotype ♀, bmnh number 9.826, by original designation. paratypes 2 ♀♀. primary type data farley, s.h./ 12.vi.1938/ r.b.benson/ b.m.1938-190; trichopria/ isis nixon/ type ♀, 1977. type locality england, hants, farley. remarks the holotype is mounted on a card point and is entire. trichopria prema nixon, 1980: 34. valid name trichopria prema nixon, 1980. summary of types holotype ♀, bmnh number 9.827, by original designation. primary type data boston manor/ mx., ix.1938; trichopria/ prema/ nixon/ type ♀, 197. type locality england, middlesex, boston manor. remarks the holotype is mounted on a card point, and is entire. trichopria bifoveata nixon, 1980: 34, 38, figs 92, 102, 104, 124. valid name trichopria nixoni notton, 1995. notton d.g., diapriinae types at the natural history museum, london 101 summary of types lectotype ♀, bmnh number 9.894, designated by notton (1995). paralectotypes 7 ♀♀, 2 ♂♂. primary type data devon/ torquay dist./ aug. 1929/ g.nixon; trichopria/ bifoveata k./ g.nixon det. 1975; lectotype ♀/ trichopria/ bifoveata/ nixon, 1980/ det. d.g.notton,1992. type locality england, devon, torquay district. remarks although nixon attributed the name to kieffer, there is no species of that name described by kieffer and nixon (1980) must be considered the author, the description provided in the key being sufficient to make the name available (notton 1995). the lectotype is mounted on a card point, has the right fore wing folded and the left hind tarsus broken off and stuck to the right fore wing. trichopria credne nixon, 1980: 34, fig. 119. valid name trichopria credne nixon, 1980. summary of types holotype ♀, bmnh number 9.828, by original designation. paratypes 2 ♀♀. primary type data surrey/ ashtead/ 6.viii.1932/ g.nixon; trichopria/ credne nixon/ type ♀, 197. type locality england, surrey, ashtead. remarks the holotype is mounted on a card point and is entire. trichopria alifera nixon, 1980: 34, fig. 118. valid name trichopria incrassata (jansson, 1955). summary of types holotype ♀, bmnh number 9.829, by original designation. paratypes 2 ♀♀. primary type data england/ surrey/ weybridge/ viii.1944/ g.e.j.nixon; trichopria/ alifera nixon/ type ♀, 197. type locality england, surrey, weybridge. european journal of taxonomy 75: 1-123 (2014) 102 remarks the holotype is mounted on a card point, the left fore leg, right hind leg and apex of the left hind tarsus are lost and the right fore wing is chipped. trichopria nana nixon, 1980: 36. valid name trichopria paludicola notton, 1995. summary of types holotype ♀, bmnh number 9.830, by original designation. primary type data cambs:/ wicken fen/ 10.ix.1934/ h.st.j.donisthorpe; trichopria/ nana nixon/ type ♀, 197. type locality england, cambridgeshire, wicken fen. remarks the holotype is mounted on a card point and is entire. trichopria sequester nixon, 1980: 37, fig. 97. valid name trichopria subimpressa (kieffer, 1911). summary of types holotype ♂, bmnh number 9.831, by original designation. paratypes 4 ♂♂. primary type data england, se:/ epping forest/ 14-15.viii.1954/ j.a. & d.j.clark/ b.m.1954-521; trichopria/ sequester/ nixon/ type ♂, 197. type locality england, essex, epping forest. remarks the holotype is mounted on a card point and is entire. trichopria crassifemur nixon, 1980: 38, fig. 91. valid name trichopria crassifemur nixon, 1980. summary of types holotype ♂, bmnh number 9.832, by original designation. paratypes 18 ♂♂. notton d.g., diapriinae types at the natural history museum, london 103 primary type data surrey:/ ashtead/ 10.viii.1930/ g.nixon; trichopria/ crassifemur/ nixon/ type ♂, 197. type locality england, surrey, ashtead. remarks the holotype is mounted on a card point and its right antenna is lost. trichopria stelenes nixon, 1980: 38. valid name trichopria credne nixon, 1980. summary of types holotype ♂, bmnh number 9.833, by original designation. paratype ♂. primary type data burnham beeches/ bucks., england/ bouček 14.ix.74; trichopria/ stelenes nixon/ type ♂, 197. type locality england, buckinghamshire, burnham beeches. remarks the holotype is mounted on a card and is entire. the synonymy of notton (1995) is confirmed here based on similarities in the morphology of the head, scutellum and petiole, and the coincident distribution and simultaneous collection of males and females. phaenopria miron nixon, 1980: 39: 40, fig. 7. valid name trichopria fucicola (walker, 1834). summary of types holotype ♀, bmnh number 9.834, by original designation. paratypes 7 ♀♀. primary type data s.e.london/ norwood/ ix.1929/ g.nixon; from rotten/ fungus; phaenopria/ miron nixon/ holotype ♀, 1979. type locality england, london, west norwood. remarks the holotype is mounted on a card point and is entire. nixon says there were nine paratypes; however, only seven were found. european journal of taxonomy 75: 1-123 (2014) 104 trichopria polita notton, 1993: 139, figs 1-13. valid name trichopria polita notton, 1993. summary of types holotype ♀, bmnh number 9.884, by original designation. primary type data uk: cambs: chippenham/ fen: carr-reedbed:/ tl650693: malaise/ 22.8-5.9.1985/ j.field; holotype ♀/ trichopria/ polita/ d.g.notton. type locality england, cambridgeshire, chippenham fen. remarks the holotype is mounted on a card and is entire. trichopria striata notton, 1993: 141, figs 14-21. valid name trichopria striata notton, 1993. summary of types holotype ♀, bmnh number 9.885, by original designation. paratype ♂. primary type data ncc fen survey/ water/ trap; uk: w.norfolk/ brancaster/ tf768443/ 2-16.9.1988: a.foster; ♀ holotype/ trichopria striata/ notton, d.g., 1992. type locality england, norfolk, brancaster. remarks the holotype is mounted on a card and is entire. trichopria melanopa scilloniensis notton, 1993: 143, figs 22-28. valid name trichopria subimpressa scilloniensis notton, 1993. summary of types holotype ♀, bmnh number 9.889, by original designation. paratypes 9 ♀♀, 8 ♂♂. primary type data england: scilly/ tresco/ 21.ix.1975; j.s.noyes/ brit. mus./ 1975-479; ♀ holotype/ trichopria/ melanopa/ scilloniensis n. ssp./ det. d.g.notton 1991. notton d.g., diapriinae types at the natural history museum, london 105 type locality england, isles of scilly, tresco. remarks the holotype is mounted on a card and is entire. diapria luteipes notton, 1993: 145, figs 33-42. valid name diapria luteipes notton, 1993. summary of types holotype ♀, bmnh number 9.886, by original designation. primary type data ncc survey/ pitfall/ trap; uk: w.norfolk: middle/ haring: tl989856/ 1-15.9.1988/ a.foster; holotype ♀/ diapria/ luteipes/ d.g.notton. type locality england, norfolk, middle harling. remarks the holotype is mounted on a card and is entire. diapria cava notton, 1993: 145, 147, figs 43-47. valid name diapria cava notton, 1993. summary of types holotype ♂, bmnh number 9.887, by original designation. primary type data ncc code – water/ trap: rspb 3.a/ open pits; uk: east kent/ dungeness: tr0718/ 22.8-5.9.1988/ r.k.a.morris ncc. type locality england, kent, dungeness. remarks the holotype is mounted on a card and is entire. trichopria quadrifida notton, 1994: 201, fig. 1a. valid name trichopria quadrifida notton, 1994. european journal of taxonomy 75: 1-123 (2014) 106 summary of types holotype ♀, bmnh number 9.890, by original designation. paratype ♀. primary type data kent/ eynsford/ 31.viii.1929/ g.nixon; phaenopria/ incrassata janss./ g.nixon det. 1978; ♀ holotype/ trichopria/ quadrifida/ n. sp./ d.g.notton, 1992. type locality england, kent, eynsford. remarks the holotype is mounted on a card point, and the left hind leg is missing. diapria drosophilae perkins, 1910: 629. valid name trichopria drosophilae (perkins, 1910). summary of types syntype ♂. primary type data honolulu/ oahu 6-19-05; diapria/ foreign; r.c.l.perkins coll./ b.m.1942-95/ hawaii. type locality hawaii, oahu, honolulu. remarks the syntype is mounted on a card point with a small fly and is entire. diapria modesta ratzeburg, 1848: 144. valid name trichopria modesta (ratzeburg, 1848). summary of types neotype ♀, bmnh number 9.891, designated by notton (1995). primary type data 46-16; diapria/ modesta/ foerst./ aachen; neotype; neotype/ diapria/ modesta/ ratzeburg 1848/ des./ d.g.notton, 1994. type locality germany, aachen. notton d.g., diapriinae types at the natural history museum, london 107 remarks this species was described by ratzeburg from an unspecified number of specimens, probably female, sent by a. förster, most probably collected in germany. the original type(s) is lost and a neotype was designated by notton (1995). the neotype is mounted on a card point and the point has been mounted on a card. the tip of the left hind tarsus is disarticulated. spilomicrus quadriceps smith, 1878: 6. valid name spilomicrus quadriceps smith, 1878 stat. rev. summary of types syntype ♂, bmnh number 9.682. primary type data new/ zeal.; type; new/ zeal./ 77-100; spilomicrus/ quadriceps/ type sm. type locality new zealand, otago. remarks this species was described from an unspecified number of males, of which one was found. the syntype is mounted on a card, generally dirty, the right a9-13 are disarticulated, and the tip of the left hind tarsus is missing. this species was placed in malvina by masner (1965); however, malvina is here considered a derived species group of spilomicrus and the original combination is reinstated. psilus fucicola walker, 1834: 117. valid name trichopria fucicola (walker, 1834). summary of types lectotype ♂, bmnh number 9.799, designated by masner (1965). paralectotypes 2 ♂♂. primary type data brit. mus./ walker coll.; type; psilus?/ fucicola/ see ms/ stood under this/ name in old b.m./ coll.; fucicola; lecto-/ type/ l.m.; selected as/ lectotype by/ l.masner 28.xi.61; lectotype ♂/ psilus fucicola/ walker, 1834: 117/ des. l.masner, 1965. type locality england, devon, torquay. remarks three syntypes were found, of which one had been designated lectotype by masner (1965). nine more syntypes (now paralectotypes) are known to be in dublin (notton & o’connor 2004). the lectotype is mounted on a card and is entire. european journal of taxonomy 75: 1-123 (2014) 108 diapria apicalis walker, 1860: 359. valid name trichopria walkeri (dalla torre, 1890) comb. nov. summary of types syntype ♀, bmnh number 9.895. primary type data 63-52; apicalis; diapria. type locality sri lanka (as ceylon). remarks the syntype is mounted on a card and is entire. the number 63-52 refers to natural history museum entomology register number 1863-52, which includes a group of insects described as “type specimens described by mr walker in the annuals & magazine for 1858-59-60”. diapria apicalis was renamed diapria walkeri by dalla torre (1890) because of homonymy with diapria apicalis (say 1836). the type of d. apicalis is a trichopria so trichopria walkeri is a new combination. monelata nigra whittaker, 1930: 133. valid name monelata solida (thomson, 1858). summary of types holotype ♀, bmnh number 9.688, by original designation. paratypes 3 ♀♀. primary type data type; type; hollyburn/ b.c. 8.ix.29/ coll. o.w.; canada:/ o.whittaker coll./ per w.h.storey/ b.m.1947212; 4379/ monelata/ nigra, whitt./ ♀/ det. o.whittaker. type locality canada, british columbia, hollyburn. remarks the description was based on 12 ♀♀. whittaker referred to paratypes, implying that there was a holotype. the holotype is mounted on a card and is entire. galesus fissus wollaston, 1858: 25. valid name coptera fissa (wollaston, 1858). summary of types syntype ♀, bmnh number 9.652. syntypes 3 ♀♀. notton d.g., diapriinae types at the natural history museum, london 109 primary type data type; madeira/ wollaston/ 55.7; galesus/ fissus w. (syntype ♀). rib. da/ janella; madeira/ wollaston/ 55.7 (syntype ♀). 1462; madeira/ wollaston/ 55.7 (syntypes 2 ♀♀). type locality madeira. remarks this species was described from at least five syntypes, since wollaston mentions five different localities: s. antonio da serra; ribeira da janella; feijãa de córte; another locality at intermediate altitude; and another locality at high altitude. not all the syntypes could be assigned to these localities with certainty; although two of them have the locality code 1462, a key to localities codes for wollaston’s hymenoptera has not been found. the four syntypes are mounted on cards: one is entire; one has the right hind telotarsus disarticulated; one has the right a12 missing; and the last one has the wings much torn. synonymic list subfamily diapriinae haliday, 1833 tribe psilini fallén, 1812 aneurhynchus westwood, 1832 a. ariadne nixon, 1980 a. coryphe nixon, 1980 a. fragilis nixon, 1980 a. indicus dodd, 1920 a. mese nixon, 1980 a. nodicornis marshall, 1867 a. obliquus kieffer, 1911 aneuropria kieffer, 1905 = mantara dodd, 1920 syn. nov. a. bifurcata (dodd, 1920) comb. nov. coptera say, 1836 c. abdominalis (nixon, 1930) c. angulata muesebeck, 1980 c. bidens (nixon, 1930) c. carinaticeps (nixon, 1930) c. cratocerus (cameron, 1912) comb. nov. c. difficilis (nixon, 1930) c. distans muesebeck, 1980 c. distincta (nixon, 1930) c. fissa (wollaston, 1858) c. hispanica (kieffer, 1911) c. inquisitor (nixon, 1930) c. longiceps (nixon, 1930) c. modesta (nixon, 1930) c. mosselensis nom. nov. = c. nigricornis (nixon, 1930) preocc. c. muscidorum (dodd, 1920) european journal of taxonomy 75: 1-123 (2014) 110 c. pijiguaorum notton, montilla & garcía nom. nov. = c. sexpunctata montilla & garcía, 2008 preocc. c. pomonellae muesebeck, 1980 c. punctulata (kieffer, 1907) c. quadridens (nixon, 1930) c. rectangularis (nixon, 1930) c. robustior (silvestri, 1913) = c. magnifica (nixon, 1930) c. sexpunctata (ashmead, 1893) comb. nov. = c. bipunctata (ashmead in riley, ashmead & howard, 1894) syn. nov. c. silvestrii (kieffer, 1913) c. strauziae muesebeck, 1980 c. turneri (nixon, 1930) labolips förster, 1856 l. innupta haliday, 1857 psilus panzer, 1801 p. aquilonius muesebeck, 1980 p. cornutus panzer, 1801 = p. gracilipes (kieffer, 1907) p. dissidens muesebeck, 1980 p. fortis muesebeck, 1980 p. fuscipennis (curtis, 1831) = p. walkeri (kieffer, 1907) syn. nov. = p. nigricornis (kieffer, 1911) syn. nov. = p. bispinosus (kieffer, 1911) = p. cameroni (kieffer, 1911) p. masneri muesebeck, 1980 p. masoni muesebeck, 1980 p. parvulus (kieffer, 1911) tribe spilomicrini ashmead, 1893 bruchopria kieffer, 1921 = aulatopria brèthes, 1927 b. tucumana (brèthes, 1927) chilomicrus masner & garcía, 2002 c. pecki masner & garcía, 2002 doddius masner & garcía, 2002 d. rugosus (dodd, 1920) entomacis förster, 1856 e. curticornis kieffer, 1912 e. fallax (kieffer, 1912) e. filiformis (ashmead in riley, ashmead & howard, 1894) e. flaviclava (kieffer, 1912) e. graeffei kieffer, 1909 = e. laertes nixon, 1980 notton d.g., diapriinae types at the natural history museum, london 111 e. latipennis (ashmead in riley, ashmead & howard, 1894) e. longicornis kieffer, 1912 e. penelope nixon, 1980 e. subaptera early, 1980 hemilexomyia dodd, 1920 h. spinosa early, 1980 idiotypa förster, 1856 i. nigriceps kieffer, 1909 = i. nigriceps kieffer, 1911 preocc., syn. nov. i. pallida ashmead, 1893 = i. pallida ashmead in riley, ashmead & howard, 1894 preocc., syn. nov. i. tinctipennis (cameron, 1888) neurogalesus kieffer, 1907 n. carinatus kieffer, 1907 paramesius westwood, 1832 p. brachypterus (thomson, 1858) = p. angustipennis kieffer, 1911 = p. cameroni kieffer, 1911 p. laetus fouts, 1927 p. longior dodd, 1920 p. rufipes (fonscolombe, 1832) = p. inchoatus kieffer, 1911 = p. subinermis kieffer, 1911 = p. nigricornis kieffer, 1911 = p. dentatus kieffer, 1911 = p. minor (kieffer, 1911) p. thoracicus ashmead in riley, ashmead & howard, 1894 pentapria kieffer, 1905 p. chiriquensis (cameron, 1888) rostropria early & naumann, 1990 r. simplex early & naumann, 1990 r. spiniventris early & naumann, 1990 spilomicrus westwood, 1832 = malvina cameron, 1889 syn. nov. = antarctopria brues in tillyard, 1920 syn. nov. s. affinis (dodd, 1920) s. aneurus ashmead in riley, ashmead & howard, 1894 s. annulicornis kieffer, 1911 s. aterrimus (dodd, 1920) comb. nov. s. atriclavus ashmead, 1893 = s. kiefferi fouts, 1925 s. barnesi early & horning, 1978 s. boweni masner, 1991 s. campbellanus (yoshimoto, 1964) comb. nov. european journal of taxonomy 75: 1-123 (2014) 112 s. canaliculatus (cameron, 1888) s. coelopae (early, 1978) comb. nov. s. crassiclavis kieffer, 1911 = s. pelion nixon, 1980 s. curvispina (cameron, 1913) s. diomedeae (early, 1978) comb. nov. s. distinctus masner, 1991 s. exul masner, 1991 s. fasciatipennis (cameron, 1888) s. helosciomyzae (early & horning, 1978) comb. nov. s. hemipterus marshall, 1868 = s. pedissequus kieffer, 1911 = s. pedisequus kieffer, 1916 unjustified emend. s. inornatus masner, 1991 s. insulae (early, 1980) comb. nov. s. integer thomson, 1858 = s. variicornis kieffer, 1911 s. kozlovi nom. nov. = s. punctatus kozlov, 1978 preocc. s. latigaster (brues in tillyard, 1920) comb. nov. s. lubomiri özdikmen, 2010 = s. superbus masner, 1991 preocc. = s. masneri özdikmen, 2011 s. maculipennis (cameron, 1888) s. myrmecophilus nixon, 1947 s. obsoletus (dodd, 1920) s. palustris masner, 1991 s. paucisetis (dodd, 1920) s. picicornis (cameron, 1913) s. pilgrimi early, 1978 s. pumilio masner, 1991 s. punctatus (cameron, 1889) comb. nov. s. quadriceps smith, 1878 stat. rev. s. rekohua (early, 1978) comb. nov. s. sanbornei masner, 1991 s. stepheni masner, 1991 s. stigmaticalis westwood, 1832 = s. tripartitus kieffer, 1911 s. sylvicola masner, 1991 s. vulgaris ashmead in riley, ashmead & howard, 1894 tribe diapriini haliday, 1833 acanthopria ashmead, 1896 a. crassicornis ashmead, 1896 a. triangularis (ashmead in riley, ashmead & howard, 1894) apopria masner & garcía, 2002 a. coveri masner & garcía, 2002 notton d.g., diapriinae types at the natural history museum, london 113 asolenopsia kieffer, 1921 a. schwarzmaieri borgmeier, 1939 basalys westwood, 1833 = microgalesus kieffer, 1912 syn. nov. b. abruptus thomson, 1858 = b. convexus (kieffer, 1911) b. bifoveatus (kieffer, 1911) = b. unifoveatus (kieffer, 1911) b. ciliatus (kieffer, 1911) b. clavatus (ashmead, 1893) = b. clavatus (ashmead in riley, ashmead & howard, 1894) b. collaris kieffer, 1911 b. cymocles nixon, 1980 b. exsul (kieffer, 1912) b. flavidipes (kieffer, 1912) = b. flavipes (ashmead, 1896) preocc. b. formicarum (kieffer, 1911) = b. rufiscapus kieffer, 1911 preocc. = b. mullensis (masner, 1965) b. fumipennis westwood, 1833 = b. atricrus (kieffer, 1911) b. grenadae (kieffer, 1912) = b. grenadensis (ashmead, 1896: 807) preocc. b. grenadensis (ashmead, 1896: 803) b. insignificans nixon, 1980 b. longipennis (kieffer, 1911) b. luctuosus (kieffer, 1911) b. macroptera (kieffer, 1911) = b. sulcatus (kieffer, 1911) = b. iphicla nixon, 1980 syn. nov. b. nigrescens (fouts, 1927) b. orion nixon, 1980 b. parvus thomson, 1858 = b. pedester (kieffer, 1907) = b. alacer (kieffer, 1911) b. pedisequa (kieffer, 1911) = b. cursitans (kieffer, 1911) syn. nov. = b. fuliginosi (box, 1921) b. pleuralis (ashmead, 1896) b. quadridens (kieffer, 1912) comb. nov. b. scoticus (kieffer, 1911) b. semele nixon, 1980 = b. euterpe nixon, 1980 b. semirufus (kieffer, 1912) b. singularis nixon, 1980 b. thoracicus (ashmead in riley, ashmead & howard, 1894) diapria latreille, 1796 d. cava notton, 1993 european journal of taxonomy 75: 1-123 (2014) 114 d. luteipes notton, 1993 doliopria kieffer, 1910 d. magniclavata (ashmead, 1896) comb. nov. eladio masner & garcía, 2002 e. cruzi masner & garcía, 2002 hansonia masner & garcía, 2002 h. pauli masner & garcía, 2002 lepidopria kieffer, 1911 l. lloydi (ferrière, 1935) leucopria masner & garcía, 2002 l. cylindricornis masner & garcía, 2002 mimopria holmgren, 1908 m. barbata borgmeier, 1939 m. campbellorum masner, 1976 m. comes borgmeier, 1939 mimopriella masner & garcía, 2002 m. splendens (borgmeier, 1939) monelata förster, 1856 m. aphrodite (nixon, 1980) m. solida (thomson, 1858) = m. nigra whittaker, 1930 neivapria borgmeier, 1939 n. penicillata borgmeier, 1939 notoxoides ashmead, 1903 n. pedissequa (borgmeier, 1939) n. pronotalis (borgmeier, 1939) platymischoides ashmead, 1901 p. molokaiensis ashmead, 1901 platymischus westwood, 1832 p. dilatatus westwood, 1832 = p. pedester (kieffer, 1911) termitopria muesebeck, 1965 t. sheasbyi muesebeck, 1965 tetramopria wasmann, 1899 t. cincticollis wasmann, 1899 = t. donisthorpei kieffer, 1911 = t. femoralis kieffer, 1911 notton d.g., diapriinae types at the natural history museum, london 115 townesella huggert & masner, 1983 t. marjoriae huggert & masner, 1983 trichopria ashmead, 1893 = xyalopria kieffer, 1907 syn. nov. t. aequata (thomson, 1858) = t. inaequalis (kieffer, 1911) = t. variipes (kieffer, 1911) = t. isis nixon, 1980 t. angulifera (ashmead, 1896) t. ashmeadi kieffer, 1912 = t. grenadensis ashmead, 1896: 808 preocc. t. atriceps ashmead in riley, ashmead & howard, 1894 t. balthazari (ashmead, 1896) t. basalis (thomson, 1858) = t. ciliaris kieffer, 1911 t. bifoveata ashmead, 1896 t. cameroni (kieffer, 1909) = t. cameroni (kieffer, 1911) t. capensis robustior ferrière, 1935 t. confusa masner, 1965 = t. grenadensis (ashmead, 1896: 809, 810) preocc. t. conotoma (kieffer, 1911) = t. bouceki (masner, 1959) comb. nov., syn. nov. t. crassifemur nixon, 1980 t. credne nixon, 1980 = t. stelenes nixon, 1980 t. drosophilae (perkins, 1910) t. ensia huggert, 1979 t. euvulgaris huggert, 1979 t. fluminis nom. nov. = t. nigriceps (kieffer, 1907) comb. nov., preocc. t. fucicola (walker, 1834) = t. miron (nixon, 1980) t. grenadensis (ashmead, 1896: 805) = t. grenadicola kieffer, 1916 t. halterata (kieffer, 1909) = t. halterata (kieffer, 1911) preocc., syn. nov. = t. atricornis kieffer, 1911 = t. morleii (morley, 1931) t. hawaiiensis (ashmead, 1901) t. hyalinipennis (thomson, 1858) = t. tetratoma (kieffer, 1911) preocc. t. incrassata (jansson, 1955) = t. alifera nixon, 1980 t. insularis ashmead in riley, ashmead & howard, 1894 t. lewisi nixon, 1940 t. mahensis (kieffer, 1912) t. melanopleura (ashmead, 1896) stat. rev. t. mellea (ashmead in riley, ashmead & howard, 1894) european journal of taxonomy 75: 1-123 (2014) 116 t. modesta (ratzeburg, 1848) t. myrmicae huggert & masner, 1983 t. neotropica masner, 1965 = t. affinis ashmead, 1896 preocc. t. nigra (nees, 1834) = t. inermis kieffer, 1909 = t. fimbriata kieffer, 1911 t. nigriceps (ashmead in riley, ashmead & howard, 1894) comb. nov. t. nigriclavata (ashmead, 1896) t. nigricornis (marshall, 1868) = t. subterranea (kieffer, 1911) = t. donisthorpei (kieffer, 1913) t. nirva huggert, 1977 t. nixoni notton, 1995 = t. bifoveata nixon, 1980 preocc. t. obscura masner, 1965 = t. nigricornis (ashmead, 1896) preocc. t. occidentalis (fouts, 1927) t. oogaster (thomson, 1858) = t. nocticolor (kieffer, 1911) t. pallida (ashmead in riley, ashmead & howard, 1894) t. paludicola notton, 1995 = t. nana nixon, 1980 preocc. t. peraffinis (ashmead, 1896) t. percyi huggert, 1979 t. pleuralis ashmead in riley, ashmead & howard, 1894 t. polita notton, 1993 t. prema nixon, 1980 t. pseudosaxatilis huggert, 1977 t. quadrifida notton, 1994 t. saxatilis (kieffer, 1912) t. scotti (kieffer, 1912) t. seychellensis (kieffer, 1912) t. simillima (ashmead in riley, ashmead & howard, 1894) t. smithii (ashmead, 1896) t. sociabilis masner, 1965 = t. formicaria kieffer, 1911 preocc. t. spinosiceps (dodd, 1920) comb. nov. t. stomoxydis huggert, 1977 t. striata notton, 1993 t. subclavata (ashmead in riley, ashmead & howard, 1894) t. subimpressa (kieffer, 1911) = t. melanopa (kieffer, 1911) = t. clavatipes (kieffer, 1911) = t. sequester nixon, 1980 t. subimpressa scilloniensis notton, 1993 t. tabanivora fouts in cameron, 1926 t. tachinidarum ferrière, 1933 t. thermarum nom. nov. = t. nigriceps (kieffer, 1913) preocc. notton d.g., diapriinae types at the natural history museum, london 117 t. tricarinata (cameron, 1912) t. unicolor (ashmead, 1896) t. vulgaris (kieffer, 1912) t. walkeri (dalla torre, 1890) comb. nov. = t. apicalis (walker, 1860) preocc. turripria masner & garcía, 2002 t. woldai masner & garcía, 2002 zacranium ashmead, 1901 z. oahuense ashmead, 1901 subfamily uncertain calogalesus kieffer, 1912 c. parvulus kieffer, 1912 peckidium masner & garcía, 2002 p. enigmaticum masner & garcía, 2002 discussion the nature of this paper is such as not to require a lengthy closing discussion, except to say that the location and recognition of types and their placement in currently valid genera takes the study of diapriinae further towards a modern understanding and is intended as a basis for future revisionary studies by all. the discovery and 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species of serphoidea (hymenoptera) from british columbia. proceedings of the entomological society of washington 32: 129-135. wollaston t.v. 1858. brief diagnostic characters of undescribed madeiran insects. annals and magazine of natural history, series 3 1: 18-125. yoder m.j. & wharton r.a. 2002. nomenclature of african psilini (hymenoptera: diapriidae) and status of coptera robustior, a parasitoid of mediterranean fruit fly (diptera: tephritidae). the canadian entomologist 134 (5): 561-576. http://dx.doi.org/10.4039/ent134561-5 manuscript received: 13 february 2013 manuscript accepted: 24 october 2013 published on: 26 february 2014 topic editor: koen martens desk editor: charlotte thionois printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; national botanic garden of belgium, meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark. http://biostor.org/reference/104162 http://dx.doi.org/10.4039/ent134561-5 the nutty world of hazel names – a critical taxonomic checklist of the genus corylus (betulaceae) norbert holstein 1,*, sarah el tamer 2 & maximilian weigend 3 1,2,3 nees institut für biodiversität der pflanzen, universität bonn, bonn, nordrhein-westfalen, germany. * corresponding author: holstein@uni-bonn.de 2 email: saraheltamer@hotmail.com 3 email: mweigend@uni-bonn.de abstract. hazelnuts (corylus l.) are the source of one of the globally most important nut crops. despite their economic and cultural importance, taxonomic knowledge is poor, even the number of species is equivocal. weak morphological differentiation, the inconsistent taxonomic treatment of horticultural selections and cultivars, and uncritical regional treatments generated a multitude of names. the situation is further complicated by an ancient history of use (at least 10 400 years), trade (at least 4000 years) and domestication (at least 2000 years). here, we present an annotated checklist of the taxa in the genus corylus based on an extensive literature review, electronic database research, and visits to some european herbaria. full citations are given for all names, typifications are provided for the majority of them. cultivars are listed if described under the rules of the icn. we designate lectotypes and neotypes for 28 names, and discuss the identity of enigmatic c. maxima mill., a taxon not known from the wild. keywords. herbarium digitization, taxon inventory, taxonomy, typification. holstein n., tamer s. el & weigend m. 2018. the nutty world of hazel names – a critical taxonomic checklist of the genus corylus l. (betulaceae). european journal of taxonomy 409: 1‒45. https://doi.org/10.5852/ejt.2018.409 introduction species of the genus corylus l. (betulaceae gray) are distributed in the northern hemisphere and grow as shrubs or trees. the genus is readily recognized by its characteristic fruits, with a large, oil-rich seed enclosed in a brown pericarp, which in turn is enveloped by a large, tubular or foliose involucre. the detailed morphology of the involucre, the shape and size of the nut, and the leaf indument are the most important morphological traits in species delimitation. human use of hazelnuts has an ancient history across its distribution range, and dates back at least to 8400 years bc in western and central europe (holst 2010; zohary et al. 2012; antolín & jacomet 2015), north america (abrams & nowacki 2008; fine et al. 2013) and china (keightley 1983; liu & chen 2012). there is evidence that hazelnuts have been traded at least regionally as early as 4000 years ago (fairbairn et al. 2014) and cultivation and domestication probably developed independently in the western and the eastern mediterranean at least 2000 years ago (pliny the elder 77–79; boccacci & botta 2009). today, hazelnuts are one of the most important nut crops world-wide with more than 850 000 tons produced in 2013 (faostat 2013), with european journal of taxonomy 409: 1–45 issn 2118-9773 https://doi.org/10.5852/ejt.2018.409 www.europeanjournaloftaxonomy.eu 2018 · holstein et al. this work is licensed under a creative commons attribution 3.0 license. r e s e a r c h a r t i c l e 1 mailto:holstein%40uni-bonn.de?subject= mailto:saraheltamer%40hotmail.com?subject= mailto:mweigend%40uni-bonn.de?subject= https://doi.org/10.5852/ejt.2018.409 http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/3.0/ most of the commercial production in turkey. in spite of the economic importance, the taxonomy of corylus is poorly understood and even the wild ancestors of the cultivars are still enigmatic. one of the most mysterious species in this context is c. maxima mill. it has been variously reported as a wild species occurring in lombardy (hence the name “lamberts nut”) (gesner in cordus 1561: 249r) and in the balkans (bobrov 1936a; săvulescu 1952; tutin & walters 1993), yet this name is neither typified nor are unequivocal wild populations known. this is especially surprising because c. maxima is the species name employed for a wide range of commercial cultivars and is commonly regarded as (one of the) progenitor(s) of the cultivated hazelnut in europe (kosenko 2002). also, in the context of wild crop relatives, for example, in attempts to use resistant strains for breeding against the various diseases, such as eastern filbert blight or hazel dieback “moria”, a clear understanding of the systematics of corylus would be highly desirable. the delimitation of corylus is undisputed due to the easily recognizable nut. corylus is sister to the other genera of the coryloideae: carpinus l., ostrya scop., and ostryopsis decne. (forest et al. 2005; li 2008; grimm & renner 2013). the infrageneric classification varies, however, depending on whether tree-like growth is regarded as an important character (schneider 1916) or if involucre morphology is considered more important (spach 1841; candolle 1864; bobrov 1936a; li & cheng 1979). undisputed, however, is the acceptance of the group of hazels with spiny bracts (c. ferox wall., c. ferox var. tibetica (batalin) franch.) as c. sect. acanthochlamys spach (spach 1841). whitcher & wen (2001) place the remainder into a single second section, c. sect. corylus, based on its data with three subsections: tree hazels (colurnae c.k.schneid.), hazels with tubular bracts (c. subsect. siphonochlamys (bobr.) p.c.li), and corylus (shrubs with leafy bracts). these relationships are also supported by crossing experiments: erdoğan (1999) and erdoğan & mehlenbacher (2000) found incompatibility or only unilateral compatibility especially between species of different subsections, while c. americana walter and c. avellana l. (c. subsect. corylus), and c. sieboldiana blume, c. cornuta marshall, and c. californica (a.dc.) rose (c. subsect. siphonochlamys) were found to be fully interfertile. this study aims at clarifying the published names in the genus corylus as another step towards a taxonomic re-evaluation of betulaceae subfam. coryloideae (holstein & weigend 2016a, 2017). we aim at a complete listing of the published species and infraspecific names under the provisions of the icn (mcneill et al. 2012) for corylus, providing typifications where required and where possible. since many cultivars have been described as biological taxa, we also list cultivars. however, this paper does not intend to list all cultivars or to evaluate which cultivars are to be accepted. it intends to provide their correct names according to the rules of the icn or icncp (brickell et al. 2009) respectively if the cultivars are to be accepted. material and methods we adopted the list of names from the world checklist of selected plant families (wcsp 2016) and supplemented it with overlooked names by an extensive internet search and names given on herbarium specimens. the acceptance status was compared with those on the world checklist and bibliography of fagales (govaerts & frodin 1998) and the relevant floristic treatments: flora of china (li & skvortsov 1999), flora sichuanica (editorial board of the flora sichuanica 2012), flora europaea (tutin & walters 1993), flora of japan (ohwi 1965), flora of north america (furlow 1997), the flora of the ussr (bobrov 1936a), and the last genus-wide revision (bobrov 1936b). for further names, we surveyed the plant list (http://www.theplantlist.org), ipni (http://www.ipni.org), tropicos (http://www.tropicos.org), and ifpni (http://fossilplants.info/). synonyms are listed here according to wcsp (2016), or if not listed there, then according to the species the taxon is currently assigned to. we also list cultivar names if published under the provisions of the icn (mcneill et al. 2012). the names of dochnahl (1860) applicable to corylus are ignored, as the work was added to the opera utique oppressa (cf. greuter & holstein 2016; holstein & greuter 2016; wilson 2017). european journal of taxonomy 409: 1–75 (2018) 2 http://www.theplantlist.org http://www.ipni.org http://www.tropicos.org http://fossilplants.info/ for typification, the first author searched the following herbaria for type specimens in situ: b, bm, br, e, goet, k, liv, m, manch, msb, w, and wu. additional collections were obtained via personal communication (g, le, mt, neu, tbi) or in silico: chinese virtual herbarium (cvh, http://www.cvh.org.cn/), nhn naturalis (l, u, wag), and homepages of the following herbaria: a, blh, gh, le, linn, mo, mw, ny, p, ph, uc, and us. results currently, 22 taxa in 17 species are accepted in corylus (wcsp 2016), and 10 legitimate basionyms can be accepted for cultivars. overall, we located 144 legitimate basionyms, with 85 synonyms for c. avellana var. avellana alone. sixty-one of these were described by a single author, henriksson (1915, 1918, 1923, 1927, 1930, 1931), who meticulously described all differences he found in scandinavian collections of c. avellana. in addition to the legitimate basionyms, 17 illegitimate and 40 invalid names were found for the different corylus species. two names of arguably new taxa, c. spiculata c.c.yang and c. heteromorpha c.c.yang (or c. heleromorpha) were found on chinese herbarium specimens, but no publication was found. checklist of corylus order fagales engl. family betulaceae gray subfamily coryloideae hook.f. tribe coryleae a.dc. corylus l. species plantarum: 998 (linnaeus 1753). – c. sect. avellana spach, annales des sciences naturelles, seconde série – botanique 16: 101 (spach 1841) nom. inval. (art. 22.2, mcneill et al. 2012). – c. subsect. euavellanae c.k.schneid., plantae wilsonianae 2: 447 (sargent 1916) nom. inval. (art. 21.3, mcneill et al. 2012). – c. sect. diphyllon bobrov, sovietskaia botanica 1936 (1): 28 (bobrov 1936b) nom. inval. – c. ser. lacerae bobrov, sovietskaia botanica 1936 (1): 28 (bobrov 1936b) nom. inval. – type species: c. avellana l., lectotypified by britton & brown (1913: 607). c. sect. acanthochlamys spach, annales des sciences naturelles, seconde série – botanique 16: 108 (spach 1841). – c. subg. acanthochlamys (spach) bobr., sovietskaia botanica 1936 (1): 27 (bobrov 1936b). – c. subsect. acanthochlamys (spach) c.j.wang, flora yunnanica 5: 173 (academia sinica kunming institute of botany 1991) nom. inval. [no basionym reference]. – type species: c. ferox wall. c. sect. tuboavellana spach, annales des sciences naturelles, seconde série – botanique 16: 106 (spach 1841). [“tubo-avellana”]. – type species: not designated (the section included c. tubulosa willd. and c. rostrata aiton). c. sect. avellana subsect. colurnae c.k.schneid., plantae wilsonianae 2: 447 (sargent 1916). – type species: c. colurna l. c. subg. phyllochlamys bobrov, sovietskaia botanica 1936 (1): 28 (bobrov 1936b). – type species: not designated. c. subg. phyllochlamys sect. diphyllon ser. incisae bobrov, sovietskaia botanica 1936 (1): 29 (bobrov 1936b). – c. subsect. phyllochlamys (bobr.) c.j.wang, flora yunnanica 5: 174 (academia sinica kunming institute of botany 1991) nom. inval. (no basionym reference). – type: not designated (the series included c. americana walter, c. heterophylla fisch. ex trautv. and c. potaninii bobrov). holstein n. et al., taxonomic checklist of corylus 3 http://www.cvh.org.cn/ c. subg. phyllochlamys sect. monophyllon bobrov, sovietskaia botanica 1936 (1): 32 (bobrov 1936b). – type: not designated (the section included c. colchica albov, c. maxima mill. and c. pontica k.koch). c. subg. siphonochlamys bobrov, sovietskaia botanica 1936 (1): 32 (bobrov 1936b). – c. sect. siphonochlamys (bobr.) c.j.wang, flora yunnanica 5: 174 (academia sinica kunming institute of botany 1991) nom. inval. (no basionym reference). – c. sect. siphonochlamys (bobr.) c.c.yang, flora sichuanica 21: 7 (editorial board of the flora sichuanica 2012). – c. subsect. siphonochlamys (bobr.) p.c.li, flora republicae popularis sinicae 21: 51 (li & cheng 1979). – type species: not designated (the subgenus included c. brevituba kom., c. californica (a.dc.) rose, c. chinensis franch., c. cornuta marshall, c. fargesii (franch.) c.k.schneid., c. mandshurica maxim., c. papyracea hickel, c. sieboldiana blume and c. yunnanensis (franch.) a.camus). 1. corylus americana walter flora caroliniana: 236 (walter 1788). – type: usa, georgia, taccoa, herbersham co., 1200 ft., 17–20 aug. 1895, j.k. small s.n. (neo-, designated by ward (2007: 1099) and here corrected: a00247927!; isoneo-: ny01185413!). – note: walter’s herbarium was in the hands of john fraser’s family and later sold to bm (stafleu & cowan 1976–1988), but apparently lost. ward therefore designated the neotype at the gray herbarium (gh), but the specimen is actually part of the collection of arnold arboretum (a). as both collections are housed in the same building, we correct ward’s minor mistake. there is no type locality mentioned in the protologue. the name is recognized as a species in the flora of north america (furlow 1997) and world checklist and bibliography of fagales (govaerts & frodin 1998). c. humilis willd., enumeratio plantarum horti regii berolinensis: 983 (willdenow 1809). – c. nana bosc, encyclopédie méthodique, agriculture 7: 597 (tessier et al. 1821) nom. nud. (though it would also be illegitimate due to c. nana tourrette & rozier). – c. pumila spach, annales des sciences naturelles, seconde série – botanique 16: 104 (spach 1841) nom. nud. – c. americana [unranked] humilis (willd.) dippel, handbuch der laubholzkunde 2: 132 (dippel 1891) [“1892”]. – c. americana f. humilis (willd.) schelle, handbuch der laubholzbenennung: 50 (beissner et al. 1903). – type: usa, “noveboraco” (new york), s. coll. in herb. c.l. von willdenow s.n. (lecto-, designated here: b-w 17705!). – note: willdenow cited a description by wangenheim (1787: 88), in which wangenheim mentioned that the american hazel was growing rather smallish (8 to 10 feet). initially, willdenow (1805: 471) provided the phrase name corylus americana humilis, but without recognizing it as a distinct taxon. in 1809, willdenow recognized it as a separate species, c. humilis, which he considered as close to c. americana, but with leaves like those of c. avellana. corylus pumila mentioned by spach (1841) is supposed to be based on willdenow. however, to our knowledge there is no willdenow name “corylus pumila”. possibly, spach’s epithet was supposed to be a nom. nov. another name was published by bosc in the agricultural vol. 7 of lamarck’s encyclopédie méthodique (tessier et al. 1821). there, bosc mentioned a c. nana from eastern north america, supposedly based on willdenow. more likely, the name was confused, because willdenow called his dwarf hazel c. humilis, besides that “c. nana” had been published before for a low-growing c. colurna (la tourrette & rozier 1787). the lectotype is taken from willdenow’s own collection. additional original material might exist in wangenheim’s collection, which was at least partly handed over to schreber (zuccarini 1833), and might be housed in m. c. americana var. missouriensis a.dc., prodromus systematis naturalis regni vegetabilis 16 (2): 132 (candolle 1864). – c. americana f. missouriensis (a.dc.) fernald, rhodora 34: 96 (fernald 1932). – type: usa, missouri, saint-louis, n. riehl 91 (holo-: g00418162!; iso-: br0000013516888!, mo1181611!). c. serotina g.kirchn., arboretum muscaviense: 672 (petzold & kirchner 1864). – type: not localized, from cultivation from königliche landesbaumschule sanssouci [potsdam, germany]. – note: european journal of taxonomy 409: 1–75 (2018) 4 petzold was in contact to schlechtendal in hal (stafleu & cowan 1976–1988), possibly original material may be found there. c. virginiana hort., die haselnuss: 90 (goeschke 1887) nom. nud. – c. virginiana hort. ex dippel, handbuch der laubholzkunde 2: 132 (dippel 1891) [“1892”] nom. inval. pro syn. c. calyculata dippel, handbuch der laubholzkunde 2: 132 (dippel 1891) [“1892”]. – c. americana [unranked] calyculata (dippel) schelle, handbuch der laubholzbenennung: 50 (beissner et al. 1903). – c. americana var. calyculata (dippel) h.j.p.winkl., das pflanzenreich iv 61 (heft 19): 48 (winkler 1904). – original citation: “diese von göschke, dem wir unsere pflanze verdanken (...)” – type: not localized [from cultivation by goeschke]. – note: this is supposed to be a weakly growing plant in contrast to the larger c. americana, very likely the same that willdenow described as c. humilis willd. c. americana var. altior farw., report of the michigan academy of science 17: 168 (farwell 1915). – type: usa, michigan, algonac, 7 jul. 1912, o.a. farwell 2822 (syn-: blh0000405); ibid., 8 sep. 1912, o.a. farwell 3154 (syn-: blh0000408, blh0000409, blh0000410); canada, ontario, stoney creek, 27 oct. 1912, o.a. farwell 3266 (syn-: blh0000406, blh0000411). c. americana var. indehiscens e.j.palmer & steyerm., annals of the missouri botanical garden 22: 516 (palmer & steyermark 1935). – type: usa, missouri, greenwood, jackson co., 5 sep. 1924, b.f. bush 10332 (holo-: a00033774!; iso-: k!, min1000722). – additional type material: ibid., e.j. palmer 26035 (para-: a); ibid., arkansas, martinsville, faulkner co., 13 oct. 1924, e.j. palmer 26537 (para-: a); ibid., hot springs, garland co., 11 oct. 1925, e.j. palmer 29160 (para-: a); ibid., north carolina, 19 sep. 1897, biltmore herb. 1244b (isopara-: wu!). 2. corylus avellana l. species plantarum: 998 (linnaeus 1753). – original citation: “habitat in europae sepibus”. – type: “avellana nux sylvestris” (lecto-, designated by sell (jarvis et al. 1993: 38): fuchs (1542: 397, 398) [icon]). – note: the lectotypification by browicz (1972: 97) in rechinger’s flora iranica (linn 1132.1) is superseded because: 1) the chosen specimen featured the note “byzantinus”, which sell interpreted to be in conflict with the protologue, and 2) it may be post 1753 material. however, it would have been a better match for interpretation, because it actually shows the one character linnaeus uses to separate the species from c. colurna, this being the ovate, obtuse stipules. the lectotype chosen by sell, on the other hand does not show stipules, but instead a short cupula, a character that is accepted as typical for the wild c. avellana. as the type of the genus, c. avellana l. is to be accepted as long as the genus itself is accepted. 2a. corylus avellana l. var. avellana c. avellana var. sylvestris ehrh., hannoverisches magazin 18: 234 (ehrhart 1780) nom. nud. c. avellana var. grandis aiton, hortus kewensis 3: 363 (aiton 1789). – c. grandis (aiton) nois., manuel complet du jardinier, maraicher, pépiniériste, botaniste, fleuriste et paysagiste 2: 464 (noisette 1825) (recombination, art. 41.4, mcneill et al. 2012). – c. avellana var. cylindricochlamys spach, annales des sciences naturelles, seconde série – botanique 16: 106 (spach 1841) nom. illegit. superfl. – original citation: only a reference to “corylus sativa fructu rotundo maximo” (bauhin 1623: 418), which does not contain figures. – type: not localized (no original specimen found in bm or k). – note: bauhin only cited “avellana urbana rotunda circa lugdunum, ges.hor.” being an abbreviation for “gesnerus in libro de hortis germaniae and avellana major lugdunensis, cam.” (cordus 1561: 249r; camerarius the younger 1588: 24). both citations lack figures, but describe a large and round fruited hazel from the area of savoy, france and lausanne, switzerland (gesner in cordus 1561: 249r). aiton published the name grandis for the “cob nut”, apparently a popular group of cultivars with large round nuts back then and still widely cultivated, for example, holstein n. et al., taxonomic checklist of corylus 5 in southern italy and northern spain. the form is morphologically close to the extant cultivar c. avellana ‘barcelona’. loudon (1838: 2018) even synonymized c. avellana var. grandis under his c. avellana var. barcelonensis, creating a superfluous name. thus, c. avellana var. grandis would be the botanical name for the cultivated forms of corylus with a large, round nut. they appear to have been in cultivation in italy, france and spain since ancient times and likely represent large-fruited selections of the wild c. avellana. all names cited under c. avellana var. cylindricochlamys spach ultimately refer to bauhin’s corylus sativa fructu rotundo maximo (bauhin 1623), later published as c. avellana var. grandis aiton. therefore, spach’s name is a homotypic synonym. c. avellana var. sylvestris aiton, hortus kewensis 3: 363 (aiton 1789) nom. inval. – c. sylvestris (aiton) salisb., prodromus stirpium in horto ad chapel allerton vigentium: 392 (salisbury 1796) nom. inval. – note: the citation of the name c. avellana var. sylvestris includes the type of c. avellana, and also salisbury cites c. avellana. c. avellana var. striata willd., species plantarum ed. 4 4 (1): 470 (willdenow 1805). – c. avellana var. striata risso, histoire naturelle des principales productions de l’europe méridionale et particulièrement de celles des environs de nice et des alpes maritimes 2: 332 (risso 1826) nom. illegit. (later homonym; art. 53.4, mcneill et al. 2012). – type: not localized. – note: risso most likely used willdenow’s name without citing him. c. avellana var. ovata willd., species plantarum ed. 4 4 (1): 470 (willdenow 1805). – c. avellana var. ovata risso, histoire naturelle des principales productions de l’europe méridionale et particulièrement de celles des environs de nice et des alpes maritimes 2: 331 (risso 1826) nom. illegit. (later homonym). – c. ovata lam. ex steud., nomenclator botanicus ed. 2 1: 424 (steudel 1840) nom. inval. pro syn. – type: iconotype, lamarck (1796–1797: pl. 780n). – note: steudel’s name is often cited as lam. ex steud., but the author of the name is willdenow. willdenow described this variety based on a drawing in lamarck’s illustration des genres, which is more commonly cited as tableau encyclopédique et méthodique (stafleu & cowan 1976–1988). steudel most likely misinterpreted willdenow’s reference to lamarck’s figure and attributed the name to lamarck. as steudel cited the name in synonymy, it is invalid anyway. c. avellana var. maxima audib., catalogue des arbres, arbrisseaux, arbustes, et plantes: 15 (audibert 1817). –type: not localized (cult.). – note: the term “maxima” is misapplied many times. in his c. maxima mill., miller (1768) explicitly refers to a corylus with oblong fruits, whereas the fruits of c. avellana var. maxima audib. are large and rounded, matching the morphology of c. avellana var. grandis. however, both names share long bracts that surpass the fruit, resulting in a similar external appearance. c. avellana var. racemosa risso, histoire naturelle des principales productions de l’europe méridionale et particulièrement de celles des environs de nice et des alpes maritimes 2: 331 (risso 1826). – type: not localized. c. avellana var. crispa loudon, encyclopedia of plants: 792 (loudon 1829). – type: not localized. – note: loudon mentions c. avellana var. crispa briefly in the list (loudon 1829) explaining that it is frizzled (most likely the bracts) in contrast to the other hazels. the diagnosis is short, not an automatic translation of the epithet and distinguishing, and therefore meeting the diagnostic criterion as set in art. 38.2 (mcneill et al. 2012). there is a contemporary description without a latin name by lindley (1829, pl. 70), mentioning that this cultivar originated in a garden at hoveton, near norwich (england). loudon, however, cited lindley only in his arboretum et fruticetum britannicum (loudon 1838: 2018). the fruit is characterized by an elliptical nut enclosed in a flaring, laciniate involucre about twice its length (loudon 1838, fig. 1943). c. avellana var. ardua poit. & turpin, traité des arbres fruitiers 6: 46 (poiteau & turpin 1835). – type: not localized (from wild collection from normandy, france). – note: poiteau & turpin (1835) described this taxon, but they were not clear about its rank. although the name in the title points to the rank of a species, being in the same style as it is done for c. avellana, they wrote in the first sentence that this is supposed to be a variety. we hence follow this designation. in a later publication european journal of taxonomy 409: 1–75 (2018) 6 (poiteau 1846: 214), they added the information that this taxon was first cultivated by hericard ferrand in rue sainte-catherine, paris, in 1812. c. avellana var. microphylla lej., compendium florae belgicae 3: 256 (lejeune & courtois 1836). – original citation: “in ardenna.” – type: [belgium, région wallonne, liège] à monganbroux [mangombroux, today a quarter in verviers], 1832, a.l.s. lejeune s.n. (lecto-, designated here: br!). c. avellana var. barcelonensis loudon, arboretum et fruticetum britannicum 3: 2018 (loudon 1838) nom. illegit. superfl. (superfluous as loudon cited c. avellana var. grandis aiton). c. avellana var. lambertii loudon, arboretum et fruticetum britannicum 3: 2018 (loudon 1838). – type: not localized. – note: this name is used for a cultivated variety from loddiges garden catalogue (loddiges 1836). however, in contrast to the lambert’s nut in the sense of c. maxima (where the name may be derived from lombardy), this name for a cultivar is supposedly derived from a plant breeder named lambert, most likely for what is today widely called “kentish cob” (bunyard 1920). it has an oblong nut with a rather rounded apex and the involucre barely exceeding the nut in length. c. avellana var. pumila loudon, arboretum et fruticetum britannicum 3: 2017 (loudon 1838). – c. pumila lodd., catalogue of plants ed. 16: [page unknown] (loddiges 1836) nom. nud. – type: not localized. c. avellana var. glandulosa godet, mémoires de la société des sciences naturelles de neuchâtel 2: 15 [“dicotyledones”], 3 [“additions et corrections en 1838”] (godet 1839). – c. glandulosa (godet) shuttleworth ex godet, flore du jura: 639 (godet 1853). – original citation: “au dessus de nods et de lignières, au pied de chasseral.” – type: switzerland, in dumetis ad pedem m. chasseral prope lignières, et versus nods, et in vall. lignières frequens, 5 jul. 1835, r.j. shuttleworth s.n. (lecto-, designated here: neu! [acc. no. 204561]). – original material: switzerland, in dumetis ad pedem m. chasseral versus nods et in vall. lignières ubique, 6 jul. 1835, r.j. shuttleworth s.n. (bm!). – note: initially, godet’s name lacks a description that he adds in “additions et corrections en 1838” lateron (submitted on 31 dec. 1838), so only then the publication can be regarded as valid. the name c. glandulosa also appears on a specimen collected by e. perrier in p (p06810712) and is referred to be by ehrhart, however, the present authors are unaware of any ehrhart publication of this name. c. arborea hort. ex steud., nomenclator botanicus ed. 2 1: 424 (steudel 1840) nom. inval. pro syn. c. avellana var. brachychlamys spach, annales des sciences naturelles, seconde série – botanique 16: 106 (spach 1841) nom. illegit. superfl. – note: spach cited c. avellana var. ovata willd. and a “corylus avellana var. sylvestris”, which is most likely the name of aiton, therefore this name is superfluous. c. avellana var. macrochlamys spach, annales des sciences naturelles, seconde série – botanique 16: 106 (spach 1841) nom. illegit. superfl. – note: spach cited c. avellana var. glomerata hort. (kew), which was published by aiton, but also c. avellana var. maxima audib., which is regarded as synonymous to c. avellana var. grandis. c. avellana var. schizochlamys spach, annales des sciences naturelles, seconde série – botanique 16: 106 (spach 1841). – original citation: “commune dans les bois aux environs de paris.” – type: not localized. – note: based on the name chosen, spach referred to the very common form of c. avellana with deeply dissected involucre. c. avellana var. sphaerocarpa rchb., icones florae germanicae et helveticae 12: 5, pl. dxxxviii: 1302 (reichenbach 1850). – type: not localized (the iconotype would be in loc. cit.). – note: reichenbach mentioned that this variety was intermediate between c. avellana s.str. and “c. grandis” (c. avellana var. grandis aiton). c. spicata g.kirchn., arboretum muscaviense: 673 (petzold & kirchner 1864). – type: not localized (cult. in the state nursery near sanssouci, potsdam, germany). c. avellana var. latifolia hort. ex a.dc., prodromus systematis naturalis regni vegetabilis 16 (2): 131 (candolle 1864). – c. filicifolia hort. ex a.dc., prodromus systematis naturalis regni vegetabilis holstein n. et al., taxonomic checklist of corylus 7 16 (2): 131 (candolle 1864) nom. inval. pro syn. – type: not localized. – note: de candolle mentioned a large-leaved variety latifolia that has been in gardens as “latifolia” and “filicifolia”. c. hispanica k.koch, die deutschen obstgehölze: 607 (koch 1867). – type: not localized (from cultivation). – note: koch discussed the early hazel names and mentioned that the taxon behind his c. hispanica was already mentioned by gesner (cordus 1561: 249r). it is therefore evident that this is yet another name for what had already been described as c. avellana var. grandis aiton. c. avellana var. mediopurpurea späth, baumschule. haupt-preis-verzeichniss, 1883 57: 2 (späth 1883). – type: not localized. – note: the present authors did not have the protologue at hand for verification of the validity. späth (1885) described the cultivar as having a red marking in the leaf center. c. avellana var. gloiotricha beck, fl. nieder-oesterreich 1: 267 (beck von mannagetta 1890). – original citation: “in voralpen.” – type: not localized. c. avellana var. glaucescens borbás, c. baenitz herbarium europaeum no. 8333 (1895–1896) nom. nud. in sched. – associate material: hungaria centr., budae-pestini, in declivibus nemoris zugliget, aug. 1895, v. de borbás s.n. (m!, p06809854!, p06809903!, p06810667!, p06810668!, wu!). – note: to the present authors’ knowledge, this is an unpublished name for a hazel with a glaucous lower leaf lamina. c. avellana var. zimmermannii hahne, mitteilungen der deutschen dendrologischen gesellschaft 12: 127 (beissner 1903). – original citation: “zu ende september 1903 machte mich herr lehrer e. zimmermann in schwelm [northrhine-westphalia, germany; …] eine alte hecke im süden der stadt, zu führen.” – type: not localized (if in bonn, then destroyed). – note: this is a formal description for a mutation in which the leaf bases are connected to form a cone. hahne lived in bonn-poppelsdorf, so there may have been a specimen at bonn, but the herbarium was largely destroyed in 1944 and there is no extant material. also, the author’s intention to graft the mutation for further study was apparently not followed. krüssmann (1960: 351) lists this name as a cultivar, but mixing up the birth place and year of e. zimmermann with the original collection data in his reference. as an aberrant individual, it does not deserve taxonomic recognition. c. avellana f. peltata buser, société pour l’étude de la flore franco-helvétique no. 1544 (buser 1904) in sched. – type: switzerland, geneva, haies à lancy, 30 sep. 1903, r. buser s.n. (syn-: p05591949!, p06809904!, p06810696!). – note: the name was also published in the bulletin of the société pour l’étude de la flore franco-helvétique, which is a short list in the bulletin de l’herbier boissier (camus et al. 1905: 974, 976). c. avellana f. grandifolia zapał., conspectus florae galiciae criticus 2: 10 (zapałowicz 1908). – original citation: “ojców (rehman).” – type: not localized (likely in kram, cf. stafleu & cowan 1976–1988). c. avellana var. acuminata henriksson, botaniska notiser 1915 (5): 247 (henriksson 1915). – type: sweden, dalia, paroecia gunnarsnäs ad östevatten, j. henriksson s.n. (holo-: not localized). c. avellana var. acutiuscula henriksson, botaniska notiser 1915 (5): 244 (henriksson 1915). – type: sweden, in helsingia, paroecia segerstad ad elftorp, c. strandberg s.n. (holo-: not localized). c. avellana var. amblyocarpa henriksson, botaniska notiser 1915 (5): 244 (henriksson 1915). – type: sweden, dalia, paroecia holm ad hålsungebyn, p.j. örtengren s.n. (syn-: not localized); sweden, dalia, paroecia gunnarsväs ad mörtviken, j. henriksson s.n. (syn-: not localized). c. avellana var. apiculata henriksson, botaniska notiser 1915 (5): 241 (henriksson 1915). – type: sweden, helsingia, paroecia segerstad ad elftorp, c. strandberg s.n. (syn-: not localized); sweden, dalia, paroecia holm ad hålsungebyn, p.j. örtengren s.n. (syn-: not localized). c. avellana var. arcuata henriksson, botaniska notiser 1915 (5): 239 (henriksson 1915). – type: sweden, dalia, paroecia gunnarsväs ad hällan, backerud et östevatten, j. henriksson s.n. (syn-: not localized); sweden, dalia, paroecia holm ad hålsungebyn, p.j. örtengren s.n. (syn-: not localized). c. avellana var. cervispina henriksson, botaniska notiser 1915 (5): 244 (henriksson 1915). – type: sweden, in oelandia, paroecia bröttorp et hålsungebyn, p.j. örtengren s.n. (holo-: not localized). european journal of taxonomy 409: 1–75 (2018) 8 c. avellana var. compressa henriksson, botaniska notiser 1915 (5): 241 (henriksson 1915). – type: sweden, dalia, paroecia gunnarsnäs, ad mörtviken, j. henriksson s.n. (syn-: not localized); sweden, dalia, paroecia ertemark ad bengtsfors, j. henriksson s.n. (syn-: not localized). c. avellana var. cryptomeria henriksson, botaniska notiser 1915 (5): 241 (henriksson 1915). – type: sweden, dalia, paroecia holm ad bröttorp, p.j. örtengren s.n. (syn-: not localized); sweden, dalia, paroecia ertemark ad bengtsfors, 1 aug. 1914, j. henriksson s.n. (syn-: not localized; isosyn-: ld1780253); sweden, dalia, paroecia gunnarsnäs ad hjulsängen et östevatten, paroecia ör ad långeerud, j. henriksson s.n. (syn-: not localized). c. avellana var. distans henriksson, botaniska notiser 1915 (5): 239 (henriksson 1915). – type: sweden, dalia, paroecia gunnarsnäs ad backerud, 160 m a.s.l., 16 sep. 1915, j. henriksson s.n. (syn-: not localized; isosyn-: ld1798219); sweden, in oelandia, paroecia vickleby ad vicklevy, j. ekström s.n. (syn-: not localized). c. avellana var. ellipsoidea henriksson, botaniska notiser 1915 (5): 246 (henriksson 1915). – type: sweden, dalia, paroecia ertemark ad bengtsfors, j. henriksson s.n. (syn-: not localized); sweden, dalia, paroecia ertemark ad huvudgingen, 1 aug. 1914, j. henriksson s.n. (syn-: not localized; isosyn-: ld1793482). c. avellana var. elongata henriksson, botaniska notiser 1915 (5): 246 (henriksson 1915). – type: sweden, in smolandia, paroecia femsjö ad olastorp, h. persson s.n. (syn-: not localized); sweden, dalia, paroecia holm ad bröttorp, p.j. örtengren s.n. (syn-: not localized). c. avellana var. fusiformis henriksson, botaniska notiser 1915 (5): 246 (henriksson 1915). – type: sweden, in oelandia, paroecia vickleby ad vickleby, j. ekström s.n. (syn-: not localized); sweden, dalia, paroecia holm ad bröttorp, p.j. örtengren s.n. (syn-: not localized); sweden, dalia, paroecia gunnarsnäs ad hjulsängen, j. henriksson s.n. (syn-: not localized). c. avellana var. gibbosa henriksson, botaniska notiser 1915 (5): 245 (henriksson 1915). – type: sweden, in smolandia, paroecia femsjö ad olastorp, h. persson s.n. (syn-: not localized); sweden, dalia, paroecia holm ad hålsungebyn, p.j. örtengren s.n. syn-: not localized); sweden, dalia, paroecia ärtemark ad bengtsfors et huvudgingen, 1 aug. 1914, j. henriksson s.n. (syn-: not localized; isosyn-: ld1793418). c. avellana var. glabrata henriksson, botaniska notiser 1915 (5): 246 (henriksson 1915). – type: sweden, dalia, paroecia holm ad hålsungebyn, p.j. örtengren s.n. (holo-: not localized). c. avellana var. integra henriksson, botaniska notiser 1915 (5): 241 (henriksson 1915). – type: sweden, in smolandia, paroecia femsjö ad olastorp, h. persson s.n. (syn-: not localized); sweden, oelandia, paroecia vickleby as vickleby, j. ekström s.n. (syn-: not localized). c. avellana var. involucrata henriksson, botaniska notiser 1915 (5): 245 (henriksson 1915). – type: sweden, in oelandia, paroecia böda ad böda, o. carlsson s.n. (syn-: not localized); sweden, dalia, paroecia holm ad hålsungebyn, p.j. örtengren s.n. (syn-: not localized). c. avellana var. lacerata henriksson, botaniska notiser 1915 (5): 244 (henriksson 1915). – type: sweden, in smolandia, paroecia femsjö ad olstorp, h. persson s.n. (syn-: not localized); sweden, bahusia ad rödbo prope oppidum kungelv, k. gustafson s.n. (syn-: not localized); sweden, dalia, paroecia holm ad bröttorp, p.j. örtengren s.n. (syn-: not localized); sweden, dalia, paroecia gunnarsnäs, ad stakelund, j. henriksson s.n. (syn-: not localized). c. avellana var. lata henriksson, botaniska notiser 1915 (5): 240 (henriksson 1915). – type: sweden, in gotlandia, paroecia hejde, s.j. stengård s.n. (syn-: not localized); sweden, helsingia, paroecia segerstad ad elftorp, c. strandberg s.n. (syn-: not localized); sweden, dalia, paroecia gunnarsnäs as östevatten, j. henriksson s.n. (syn-: not localized). c. avellana var. limbata henriksson, botaniska notiser 1915 (5): 240 (henriksson 1915). – type: sweden, dalia, paroecia gunnarsnäs, ad mörtviken, 120 m a.s.l., 14 sep. 1915, j. henriksson s.n. (syn-: not localized; isosyn-: br!, ld1797982, ph00006756!); sweden, dalia, paroecia gunnarsnäs ad backerud, hjulsängen, östevatten, j. henriksson s.n. (syn-: not localized); sweden, in oelandia, par vickleby as vickleby, j. ekström s.n. (syn-: not localized); sweden, bahusia, paroecia ryr ad holstein n. et al., taxonomic checklist of corylus 9 heljebol, j. henriksson s.n. (syn-: not localized); sweden, dalia, paroecia ertemark ad bengtsfors, j. henriksson s.n. (syn-: not localized). c. avellana var. menthoides henriksson, botaniska notiser 1915 (5): 241 (henriksson 1915). – type: sweden, in oelandia, paroecia böda ad böda, o. carlsson s.n. (holo-: not localized). c. avellana var. oblonga g.andersson, botaniska notiser 1915 (5): 245 (henriksson 1915). – type: sweden, in dalia, paroecia gunnarsnäs ad östevatten et hjulsängen, paroecia ertemark ad bengtsfors, j. henriksson s.n. (syn-: not localized); sweden, smolandia, paroecia femsjö ad olastorp, h. persson s.n. (syn-: not localized). c. avellana var. obovata henriksson, botaniska notiser 1915 (5): 245 (henriksson 1915). – type: sweden, in bahusia ad rödbo prope kungelv, k. gustafson s.n. (syn-: not localized). c. avellana var. pectinata henriksson, botaniska notiser 1915 (5): 246 (henriksson 1915). – type: sweden, dalia, paroecia gunnarsnäs, ad östevatten, j. henriksson s.n. (holo-: not localized). c. avellana var. sulcata henriksson, botaniska notiser 1915 (5): 241 (henriksson 1915). – type: sweden, dalia, paroecia gunnarsnäs ad östevatten, j. henriksson s.n. (holo-: not localized). c. avellana var. teretiuscula henriksson, botaniska notiser 1915 (5): 239 (henriksson 1915). – type: sweden, in smolandia, paroecia femsjö ad olastorp, h. persson s.n. (syn-: not localized); ibid., oelandia, paroecia vickleby ad vickleby, j. ekström s.n. (syn-: not localized). c. avellana var. truncata henriksson, botaniska notiser 1915 (5): 247 (henriksson 1915). – type: sweden, in smolandia, paroecia femsjö ad olastorp, h. persson s.n. (syn-: not localized); sweden, dalia, paroecia holm ad bröttorp, p.j. örtengren s.n. (syn-: not localized). c. avellana var. turgida henriksson, botaniska notiser 1915 (5): 240 (henriksson 1915). – type: sweden, in gotlandia, paroecia hejde, s.j. stengård s.n. (syn-: not localized); sweden, bahusia paroecia ryr ad heljebol, j. henriksson s.n. (syn-: not localized); sweden, dalia, paroecia gunnarsnäs, ad östevatten, j. henriksson s.n. (syn-: not localized). c. avellana var. velutina henriksson, botaniska notiser 1915 (5): 240 (henriksson 1915). – type: sweden, dalia, paroecia holm ad hålsungebyn, p.j. örtengren s.n. (syn-: not localized); sweden, paroecia gunnarsnäs ad östevatten, j. henriksson s.n. (syn-: not localized [not ld1798302]). c. avellana var. bulbosa henriksson, botaniska notiser 1918 (6): 299 (henriksson 1918). – type: denmark, nærum railway station, e. petersen s.n. (holo-: not localized). c. avellana var. cuneata henriksson, botaniska notiser 1918 (6): 299 (henriksson 1918). – type: denmark, nærum railway station, e. petersen s.n. (holo-: not localized). c. avellana var. cupulifera henriksson, botaniska notiser 1918 (6): 299 (henriksson 1918). – type: denmark, nærum railway station, e. petersen s.n. (holo-: not localized). c. avellana var. orbicularia henriksson, botaniska notiser 1918 (6): 299 (henriksson 1918). – type: norway, southern bergenhus amt, störd [stord], t. bredal s.n. (holo-: not localized). c. avellana var. praefracta henriksson, botaniska notiser 1918 (6): 297 (henriksson 1918). – type: sweden, dalia, paroecia gunnarsnäs in silva hällan ad mörtviken, j. henriksson s.n. (holo-: not localized). c. avellana var. annulata henriksson, botaniska notiser 1923 (3): 283 (henriksson 1923). – type: sweden, dalia, paroecia gunnarsnäs ad östevatten, p.j. örtengren s.n. (syn-: not localized); sweden, dalia, paroecia gunnarsnäs ad östevatten, j. henriksson s.n. (syn-: not localized). c. avellana var. attenuata henriksson, botaniska notiser 1923 (3): 280 (henriksson 1923). – type: sweden, dalia, paroecia gunnarsnäs, ad mörtviken, j. henriksson s.n. (syn-: not localized); sweden, bahusia, paroecia tanum ad rörvik haud procul a grebbestad, v. andersson s.n. (syn-: not localized). c. avellana var. coelata henriksson, botaniska notiser 1923 (3): 280 (henriksson 1923). – type: sweden, dalia, paroecia holm ad bröttorp, p.j. örtengren s.n. (syn-: not localized); sweden, dalia, paroecia gunnarsnäs ad sanatorium kroppefjäll, j. henriksson s.n. (syn-: not localized); sweden, ostrogothia, paroecia landeryd ad uvberget, p.h. johansson s.n. (syn-: not localized). european journal of taxonomy 409: 1–75 (2018) 10 c. avellana var. complanata henriksson, botaniska notiser 1923 (3): 281 (henriksson 1923). – type: sweden, in blekingia haud procul a ronneby, e. essén s.n. (syn-: not localized); sweden, dalia, paroecia gunnarsnäs ad östevatten, p.j. örtengren s.n. (syn-: not localized); sweden, vestrogothia, paroecia häggesled as stinggården, g. brandström s.n. (syn-: not localized); sweden, vestrogothia, paroecia tun ad gammaltorp, g. brandström s.n. (syn-: not localized). c. avellana var. extensa henriksson, botaniska notiser 1923 (3): 281 (henriksson 1923). – type: sweden, dalia, paroecia holm ad bröttorp et hålsungebyn, p.j. örtengren s.n. (syn-: not localized); sweden, dalia, gunnarsnäs ad mörtviken et östevatten, j. henriksson s.n. (syn-: not localized); sweden, uplandia, gustavsberg ad farsta, p.h. johansson s.n. (syn-: not localized); sweden, vestrogothia, paroecia tun ad gammaltorp, g. brandström s.n. (syn-: not localized). c. avellana var. glandulosa henriksson, botaniska notiser 1923 (3): 281 (henriksson 1923) nom. illegit. (later homonym). – type: sweden, in blekingia prope ronneby, e. essén s.n. (syn-: not localized); sweden, dalia, paroecia gunnarsnäs ad mörtviken, j. henriksson s.n. (syn-: not localized). c. avellana var. ovalis henriksson, botaniska notiser 1923 (3): 283 (henriksson 1923). – type: sweden, in blekingia, bustorp ad ronnyby, e. essén s.n. (syn-: not localized); sweden, dalia, paroecia gunnarsnäs ad östevatten, j. henriksson s.n. (syn-: not localized); sweden, ostrogothia, paroecia landeryd ad uvberget, p.h. johansson s.n. (syn-: not localized). c. avellana var. subtecta henriksson, botaniska notiser 1923 (3): 283 (henriksson 1923). – type: sweden, dalia, paroecia gunnarsnäs ad östevatten, j. henriksson s.n. (holo-: not localized). c. avellana var. contracta henriksson, botaniska notiser 1927 (2): 157 (henriksson 1927). – type: sweden, in blekingia ad persborg prope ronneby, e. essén s.n. (syn-: not localized); sweden, dalia, paroecia gunnarsnäs ad sanatorium kroppefjäll, j. henriksson s.n. (syn-: not localized); sweden, sudermannia, paroecia husby rekarne ad hedskolan, g. pettersson s.n. (syn-: not localized). c. avellana var. nitens henriksson, botaniska notiser 1927 (2): 157 (henriksson 1927). – type: sweden, paroecia gunnarsnäs ad sanatorium kroppefjäll, j. henriksson s.n. (holo-: not localized). c. avellana var. prominens henriksson, botaniska notiser 1927 (2): 156 (henriksson 1927). – type: sweden, in bahusia, paroecia tanum ad vik prope grebbestad, v. andersson s.n. (syn-: not localized); sweden, paroecia gunnarsnäs ad sanatorium kroppefjäll, j. henriksson s.n. (syn-: not localized). c. avellana var. retusa henriksson, botaniska notiser 1927 (2): 157 (henriksson 1927). – type: sweden, dalia, paroecia gunnarsnäs ad sanatorium kroppefjäll, j. henriksson s.n. (holo-: not localized). c. avellana var. secalina henriksson, botaniska notiser 1927 (2): 156 (henriksson 1927). – type: sweden, dalia, paroecia laxarby ad gårdsjö, n.i. svensson s.n. (holo-: not localized). c. avellana var. stylifera henriksson, botaniska notiser 1927 (2): 159 (henriksson 1927). – type: sweden, in vermelandia, paroecia sillerud ad vicum sillerud, h. danielsson s.n. (holo-: not localized). c. memorabilis sennen, plantes d’espagne 1928, no. 7082 (1928–1929) in sched. – type: spain [catalonia], barcelone, massif de tibidabo, e loco, 28 may 1928, frère sennen s.n. (syn-: not localized; isosyn-: bm!, ph00006758, p06811312!); ibid., barranco du rabassalet, 28 may 1928, frère sennen s.n. in exsicc. ch. duffour no. 6745 (syn-: not localized; isosyn-: bm!, p06809387!, p06809876!, p06809877!, p06809878!, p06809879!); ibid., 14 aug. 1928 or 1929, frère sennen s.n. in plantes d’espagne no. 7082 (syn-: not localized; isosyn-: bm!, ph00006758, p06811312!). – note: the description is effectively published on printed labels accompanying the type specimens (art. 30.7, mcneill et al. 2012). the name c. memorabilis appeared on labels before [frère sennen in plantes d’espagne no. 4805 (bm!, l!, l!) and frère sennen s.n. (bm!)], but was lacking a description. the sterile collections from 28 may 1928 were also distributed by the société française in 1932 (exsicc. ch. duffour) and are also type material. c. avellana var. acuta henriksson, botaniska notiser 1930 (2): 149 (henriksson 1930). – type: sweden, in gotlandia, paroecia lokrum ad haltarve, g. nilsson s.n. (holo-: not localized). c. avellana var. glandifera henriksson, botaniska notiser 1930 (2): 149 (henriksson 1930). – type: sweden, in gotlandia, paroecia lokrum ad haltarve, g. nilsson s.n. (holo-: not localized). holstein n. et al., taxonomic checklist of corylus 11 c. avellana var. mucronata henriksson, botaniska notiser 1930 (2): 149 (henriksson 1930). – type: sweden, vestrogothia, paroecia järpås ad simmenebol, e. brandström s.n. (holo-: not localized). c. avellana var. pilosa henriksson, botaniska notiser 1930 (2): 149 (henriksson 1930). – type: sweden, vestrogothia, paroecia järpås ad simmenebol, e. brandström s.n. (holo-: not localized). c. avellana var. taeniensis henriksson, botaniska notiser 1930 (2): 149 (henriksson 1930). – type: sweden, vestrogothia, paroecia järpås ad simmenebol, e. brandström s.n. (holo-: not localized). c. avellana var. velutina f. monstrosa henriksson, botaniska notiser 1930 (2): 150 (henriksson 1930). – type: sweden, vestrogothia, paroecia järpås ad simmenebol, e. brandström s.n. (holo-: not localized). c. avellana var. brevilaciniata henriksson, botaniska notiser 1931 (2): 213 (henriksson 1931). – type: sweden, in hallandia, paroecia knäred ad ekernahult, e. bengtsson s.n. (holo-: not localized). c. avellana var. coniformis henriksson, botaniska notiser 1931 (2): 213 (henriksson 1931). – type: sweden, dalia, paroecia gunnarsnäs ad lottsbyn, m. magnusson s.n. (holo-: not localized). c. avellana var. divergens henriksson, botaniska notiser 1931 (2): 213 (henriksson 1931). – type: sweden, dalia, paroecia gunnarsnäs ad lottsbyn, m. magnusson s.n. (holo-: not localized). c. avellana var. gracilescens henriksson, botaniska notiser 1931 (2): 214 (henriksson 1931). – type: sweden, dalia, paroecia gunnarsnäs ad lottsbyn, m. magnusson s.n. (holo-: not localized). c. avellana var. indivisa henriksson, botaniska notiser 1931 (2): 214 (henriksson 1931). – type: sweden, in bahusia, paroecia brastad ad rixö, v. nilsson s.n. (holo-: not localized). c. avellana var. lenticularis henriksson, botaniska notiser 1931 (2): 214 (henriksson 1931). – type: sweden, in bahusia, paroecia sanne ad prästbol, a. johansson s.n. (holo-: not localized). c. avellana var. porrecta henriksson, botaniska notiser 1931 (2): 214 (henriksson 1931). – type: sweden, in bahusia, paroecia brastad ad rixö, v. nilsson s.n. (holo-: not localized). c. avellana var. pyramidata henriksson, botaniska notiser 1931 (2): 214 (henriksson 1931). – type: sweden, in bahusia, paroecia brastad ad rixö, v. nilsson s.n. (holo-: not localized). c. avellana var. simlicinerva henriksson, botaniska notiser 1931 (2): 216 (henriksson 1931). – type: sweden, in bahusia, paroecia brastad ad rixö, v. nilsson s.n. (holo-: not localized). c. avellana var. tiliifolia murr, veröffentlichungen des tiroler landesmuseums ferdinandeum 11: 40 (murr 1931). – original citation: “silz-mötz, am höttinger bild.” – type: not localized. c. mirabilis sennen ex bobrov, sovietskaia botanica 1936 (1): 28 (bobrov 1936b) nom. inval. pro syn. – note: the present authors did not see any specimen with this label, but it may be the name was effectively and validly published on printed exsiccata labels just like c. memorabilis. c. avellana f. monstrosa h.lindb., plantæ finlandiæ exsiccatae no. 1123 (lindberg 1944) nom. illegit. superfl. (later homonym) in sched., non henriksson (1930). – type: finland, usimaa/nyland, paroecia karis, hållsnäs, 25 aug. 1920, h. lindberg s.n. (syn-: k!, wu!). – note: this is a mutation with branched male catkins. the description is effectively published on printed labels accompanying the type specimens (art. 30.7, mcneill et al. 2012), but illegitimate as it is a later homonym. c. avellana f. peltata de langhe, bulletin de la société royale de botanique de belgique 106: 70 (langhe 1973) nom. illegit. (later homonym), non buser (1904). – type: belgium, district mosan, 1961, j.-e. de langhe 544/1961 (holo-: herbarium de langhe). c. avellana f. microphylla gajić & korac, glasnik prirodnaučkog museja u beogradu. serija b biološke nauke. belgrade [bulletin du museum d’histoire naturelle belgrade. serie b. sciences biologiques. belgrade] 33: 60 (gajić & korać 1979) nom. illegit. (later homonym, art. 53.4, mcneill et al. 2012), non c. avellana var. microphylla lej. (lejeune & courtois 1836). – type: serbia/kosovo, m. ošljak, 1450 m a.s.l., 10 jul. 1977, m. gajić and m. korać s.n. (holo-: not localized). c. hispanica mill. ex d.rivera, obón, s.ríos, selma, f.méndez, verde & f.cano, las variedades tradicionales de frutales de la cuenca del río segura. catálogo etnobotánico (1): frutos secos, oleaginosos, frutales de hueso, almendros y frutales de pepita: 37 (rivera núñez et al. 1997) nom. illegit. superfl. (later homonym), non k.koch (koch 1867). – type: spain, in castillae veterae, prope coenobio sancta clarae, in ditione medina de pomar (burgos), 5 aug. 1996, d. rivera and european journal of taxonomy 409: 1–75 (2018) 12 c. obón 47689 (holo-: mub; iso-: mub). – note: rivera núñez et al. (1997) describe this long cultivated form as a new species, synonymizing several valid names below the species rank. corylus avellana var. grandis aiton. is however, already elevated to species rank by noisette (1825), making this the oldest name for the taxon at species rank, if recognized. futhermore, there is already a c. hispanica k.koch, who described the same morphology. in all, c. hispanica mill. ex d.rivera, obón, s.ríos, selma, f.méndez, verde & f.cano is a later homonym. c. avellana var. macrotruncus zernov, определитель сосудистых растений севера российского причерноморья: 94 (zernov 2002). – type: russia, krasnodar, novorossijsk, pylae topolnaja-2, in valle rivuli, 18 aug. 1999, a.s. zernov 557 (holo-: mw; iso-: mosp). – note: this was described as a tree, but synonymized under c. avellana in the plant list (2013). c. domestica kosenko & opalko, інтродукція рослин на початку ххі століття: досягнення і перспективи (до 120-річчяз дня народження академіка м.і.вавилова): матеріали міжнародної наукової конференції (2–4 жовтня 2007 р.): 70–74 (kosenko & opalko 2007) nom. nud. – note: kosenko & opalko (2007) published this name to accommodate the cultivated hazelnut, here considered as of hybrid origin from c. avellana, c. maxima, and c. avellana var. pontica. the protologue, however, lacks a type and a formal diagnosis (i. kosenko, pers. com.), and has been used in a few ukrainian publications mainly on hazel cultivation since then. however, the name refers to the same entity as c. hispanica and c. grandis, and the latter name would have priority. however, as a putative cultigen, the taxon should probably be better named following the code for cultivated plants (icncp, brickell et al. 2009). c. avellana var. alternata henriksson nom. nud. in sched. – associate material: sweden, dalia, paroecia gunnarsnäs ad mörtvkien, 100 m a.s.l., 30 aug. 1926, j. henriksson s.n. (ld1780125). c. avellana f. coronata nom. nud. in sched. – associate material: sweden, vestrogothia, paroecia toarp, skinnarhult, 14 sep. 1921, o. olson s.n. (ld1793794). c. avellana f. sublobulata j.erikson nom. nud. in sched. – associate material: sweden, blekinge, rödeby, 5 sep. 1914, j. erikson s.n. (ld1064362). 2aa. corylus avellana ‘atlas’ c. algeriensis oct.thomas, guide pratique de l’amateur de fruits: 36, 213 (thomas 1876). – original citation: “d’angleterre.” – type: not localized. – note: this is a latin name given for an english cultivar called ‘atlas’. the diagnosis is highly rudimentary, referring only to the clustered fruits. 2ab. corylus avellana ‘aurea’ c. avellana var. aurea g.kirchn., arboretum muscaviense: 670 (petzold & kirchner 1864). – c. avellana f. aurea (g.kirchn.) dippel, handbuch der laubholzkunde 2: 128 (dippel 1891) [“1892”]. – type: not localized. – note: petzold was in contact to schlechtendal in hal (stafleu & cowan 1976– 1988), possibly original material is found there. this is a yellow-leaved garden cultivar, best called c. avellana ‘aurea’. 2ac. corylus avellana ‘contorta’ c. avellana var. contorta bean, the garden 54: 349 (bean 1898). – c. avellana f. contorta (bean) rehder, bibliography of cultivated trees and shrubs hardy in the cooler temperate regions of the northern hemisphere: 111 (rehder 1949). – type: not localized. – note: this is a still popular garden variety with cork-screw shaped branches that was originally discovered near gloucester, england around the year 1850. holstein n. et al., taxonomic checklist of corylus 13 2ad. corylus avellana ‘cosford’ c. avellana var. tenuis loudon, arboretum et fruticetum britannicum 3: 2018, fig. 1943 (loudon 1838). – type: iconotype (lecto-, designated here: lindley 1829: pl. 55). – note: corylus avellana var. tenuis was created as a scientific name for the cultivar ‘cosford’ (lindley 1829). we argue that the cultivar name should be used for what is clearly the result of a horticultural selection process. 2ae. corylus avellana ‘fuscorubra’ c. avellana f. fuscorubra dippel, handbuch der laubholzkunde 2: 128 (dippel 1891) [“1892”]. – type: iconotype, goeschke (1887: tab. 4). – note: dippel cites a german description of this cultivar ‘rotblättrige waldnuss’ by goeschke (1887: 46), who found it in the arboretum of reinhardt behnsch in dürrgoy (tarnogaj) near breslau (wrocław, poland). this is a red-leaved cultivar whose leaf coloration fades soon, and which has an involucre shorter than half of the nut. 2af. corylus avellana ‘impériale de trébizonde’ c. macrocarpa oct.thomas, guide pratique de l’amateur de fruits: 36, 213 (thomas 1876). – type: not localized. – note: corylus macrocarpa is a latin name given for a cultivar called ‘impériale de trébizonde’. the name has been published in garden catalogs before, but the present authors are unaware of an accompanying description. the cultivar is still widely grown and offered for sale. 2ag. corylus avellana ‘pendula’ c. avellana var. pendula nestel, nestel’s rosengarten 2: 17 (nestel 1869). – c. avellana f. pendula (nestel) dippel, handbuch der laubholzkunde 2: 128 (dippel 1891) [“1892”]. – original citation: “wurde vor einer reihe von herrn niessing in einem walde seiner heimat entdeckt, und später als diese merkwürdige, natürliche spielart für ihn als gärtner erhöhtes interesse erhielt, durch uebertragung von pfropfreisen und durch ableger festgehalten.” – type: not localized. – note: corylus avellana var. pendula is a formal taxonomical name for a cultivar with over-hanging branches, best treated as c. avellana ‘pendula’. this garden variety was brought into cultivation by niessing (nestel 1869) and presented for the first time at the world exhibition in paris in 1867 (goeschke 1887: 49). 2ah. corylus avellana ‘urticifolia’ c. avellana [unranked] urticifolia nois., manuel complet du jardinier, maraicher, pépiniériste, botaniste, fleuriste et paysagiste 2: 464 (noisette 1825) [“urticaefolia”]. – c. urticifolia hort. ex loudon, arboretum et fruticetum britannicum 3: 2017 (loudon 1838) nom. inval. pro syn. – c. avellana var. urticifolia (nois.) a.dc., prodromus systematis naturalis regni vegetabilis 16(2): 131 (candolle 1864). – c. urticifolia (nois.) dippel, handbuch der laubholzkunde 2: 128 (dippel 1891) [“1892”]. – original citation: “nous l’avons vu chez m. le comte montmorenci à montgeron [france]”. – type: not localized. – note: none of the later users of the epithet “urticifolia” refer to noisette, who called it “espèce” and “variété” with the name “c. urticaefolia” (thus unranked). however, this leaf shape mutation apparently occurs spontaneously in the wild (franz 1992). mehlenbacher & smith (1995) found the trait to be encoded by a recessive gene. it is thus likely that there are multiple origins of cultivars with the variety name ‘urticifolia’ (and c. avellana var. heterophylla loudon, c. laciniata hort. ex burchardt, and c. avellana var. quercifolia booth et al. ex g.kirchn.). nevertheless, they all refer to a horticultural variety with the same mutation (therefore art. 41.4 applies, mcneill et al. 2012), which is to be treated as c. avellana ‘urticifolia’ as the earliest available name (art. 29.1, brickell et al. 2009). european journal of taxonomy 409: 1–75 (2018) 14 c. avellana var. heterophylla loudon, arboretum et fruticetum britannicum 3: 2017 (loudon 1838). – c. heterophylla lodd., catalogue of plants ed. 16: [page unknown] (loddiges 1836) nom. nud. – c. avellana f. heterophylla (loudon) rehder, bibliography of cultivated trees and shrubs hardy in the cooler temperate regions of the northern hemisphere: 110 (rehder 1949). – type: not localized. – note: the epithet heterophylla is problematic in corylus. at the rank of a variety (and forma), it was formally used for a garden cultivar (c. avellana ‘urticifolia’), whereas in the rank of a species, it was only used as a mere name without description for that cultivar (nom. nud.). however, there is a species, c. heterophylla fisch. ex. trautv., from east asia that must not be confused with with ornamental plant with narrow, deeply incised and double-serrate leaves. as neither trautvetter nor fischer referred to a cultivar, but a siberian wild species, their name is not to be treated as a new combination (art. 41.4, mcneill et al. 2012). the oldest epithet for the deeply incised doubleserrate c. avellana is urticifolia, c. avellana [unranked] urticifolia nois. (noisette 1825), hence this cultivar should be called c. avellana ‘urticifolia’. c. laciniata hort. ex burchardt, verhandlungen des vereins zu beförderung des gartenbaues in den königlich preußischen staaten 15: 84 (burchardt 1841). – c. avellana f. laciniata (burchardt) döll, rheinische flora: 273 (döll 1843). – c. avellana var. laciniata (burchardt) g.kirchn., arboretum muscaviense: 670 (petzold & kirchner 1864). – original citation: “ich habe meinen stamm von den herren gebrüdern baumann, die ihn von einem ihrer reisenden aus dem südlichen frankreich als betula laciniata ohne angabe des orts erhalten hatten.” – type: most likely destroyed. – note: there is a report that a nut collection (‘nußkabinet’) existed, but it was most likely destroyed (anonymous 1854) by burchardt’s elder son (müller 1861). therefore, upon publication, there was original material, but was subsequently lost. the name laciniata is a garden name that was taken up several times independently (see c. avellana f. laciniata and c. avellana var. laciniata). the oldest publication is by theodor heinrich otto burchardt (13 dec. 1771–6 feb. 1853, cf. müller 1861), justice councellor (‘justizrat’) from landsberg a. d. warthe (now gorzów wielkopolski, poland), who described it as a new species. he mentioned that he had the stock from the pépinières baumann from bollwiller (alsace, france), a then famous seed seller, who had received the material from southern france under the name betula laciniata. burchardt mentioned that the plant is also to be found in the catalog of the frères audibert (audibert 1831–1832), where it is listed without a description. the name, however, had already been used in seed catalogs before (dailledouze & dailledouze 1828). although neither döll nor kirchner explicitly referred to burchardt, they referred to the same mutation, and their names are thus to be treated as new combinations (art. 41.4, mcneill et al. 2012). c. serenyiana pluskal, oesterreichisches botanisches wochenblatt 2: 394 (pluskal 1852). – c. avellana var. serenyiana (pluskal) pluskal, verhandlungen des zoologisch-botanischen vereins in wien 3 (2): 22 (pluskal 1853). – original citation: “an dem ufer eines baches bei lomnitz [lomnice u tišnova, czech republic]”. – type: not localized. – note: pluskal mentioned that his plan to describe his name was highly disputed since it closely corresponded to c. avellana var. laciniata, but he insisted on describing it as a new species as he found it in the wild and reported it as new to moravia. c. avellana var. quercifolia booth et al. ex g.kirchn., arboretum muscaviense: 670 (petzold & kirchner 1864). – c. quercifolia hort. ex a.dc., prodromus systematis naturalis regni vegetabilis 16 (2): 131 (candolle 1864) nom. inval. pro syn. – note: rehder (1949) listed this garden variety as a synonym to his c. avellana f. heterophylla. c. avellana var. incisa andré, l’art des jardins: 574 (andré 1879). – type: not localized. c. avellana var. pinnatifida plan. nom. nud. in sched. – associate material: sweden, östergötland, regna, s. coll., s.n. (l1559661!) [n.j. andersson ex s, pers. com.]; sweden, östergötland, regna, [misit jul. 1867], s. coll, s.n. (k!, manch!) [f. ahlberg, pers. com.]; sweden, prope regna ostergothia, 12 jul. 1867, e. rathsman s.n. (bm! k!, k!, k!, ld1790934, manch!, ohn135122); sweden, og, regna, 1 jun. 1865, e. rathsman s.n. (bm!, ld1798977); sweden, ostergothia, regna, aug. 1873, a. magnusson s.n. (p06809852!). – note: this name is found on swedish specimens of c. avellana holstein n. et al., taxonomic checklist of corylus 15 ‘urticifolia’. the name is apparently by j.e. planander, but the present authors are not aware of a valid publication. 2ai. corylus avellana ‘variegata’ c. avellana var. variegata a.dc., prodromus systematis naturalis regni vegetabilis 16 (2): 131 (candolle 1864). – c. avellana f. variegata (a.dc.) dippel, handbuch der laubholzkunde 2: 128 (dippel 1891) [“1892”]. – type: not localized. – note: candolle summarized with this name c. avellana cultivars with white, yellow or red-variegated leaves. 2b. corylus avellana var. pontica (k.koch) h.j.p.winkl. das pflanzenreich iv 61 (heft 19): 47 (winkler 1904). – c. pontica k.koch, linnaea 22: 329 (koch 1849). – original citation: “auf der nordseite des pontischen gebirges auf augitporphyr bis 2500’ höhe.” – type: not localized. – note: corylus avellana var. pontica is recognized by flora of the ussr (bobrov 1936a) and world checklist and bibliography of fagales (govaerts & frodin 1998). it apparently differs from c. avellana var. avellana by double-serrate bracts that surpass the nut and are apically prominently veined. this morphology is also widely found elsewhere and particularly common in cultivars. therefore kosenko (2002) hypothesized that the domesticated hazel might be a hybrid involving or directly selected from c. avellana var. pontica. the eastern origin of hazel cultivation seems to find support in the ancient roman name “nux pontica”, supposedly first introduced from pontus (pliny the elder 77–79). koch neither mentions the romans, nor any cultivars. koch mentions that the nuts resemble those of c. colurna and refers to a wild collection. however, given that hazels freely intercross within the subsections (erdoğan 1999; erdoğan & mehlenbacher 2000) and that hazels have been bred for centuries, the persistence of a “pure” genotype “var. pontica” in the wild (if it ever existed) is more than doubtful. c. pontica var. glandulifera k.koch, linnaea 22: 330 (koch 1849). – c. colurna var. glandulifera (k.koch) a.dc., prodromus systematis naturalis regni vegetabilis 16 (2): 132 (candolle 1864). – original citation: no collections cited. – type: not localized. c. imeretica kem.-nath., trudy tbilisskogo botaniceskogo instituta 1: 111 (kemularia-nathadze 1933). – type: georgia, prov. kutais, distr. schorapani, fauces fl. kvirila, prope urb. tschiathura ad rupes calcareas, 26. jul. 1930, l.m. kemularia-nathadze s.n. (syn-: le [photo at e, k], tbi1024989!); georgia, prov. kutais, distr. letschchum, fauc. fl. rioni, prope pag. atscharis-chidi ad rupes calcareas, 29 jul. 1929, l.m. kemularia-nathadze s.n. (syn-: not localized). – note: corylus imeretica was synonymized under c. avellana var. pontica by rehder (1949). 3. corylus chinensis franch. journal de botanique (morot) 13: 197 (franchet 1899). – c. colurna var. chinensis (franch.) burkill, the journal of the linnean society. botany 26: 503 (burkill 1899). – original citation: “china occidentalis, provincia yunnan, in silvis montanis faucium ad san thang kiou, in via e tali ad ho kin, alt. 2200 m (delavey, nn. 197 et 212)”. – type: china, yunnan, san tchang kiou, ho kin, 2200 m a.s.l., 27 aug. 1884, j.m. delavay 197 [p212] (lecto-, designated here: p06811315!; isolecto-: b 10 0464027!, p06811316!, p06811317!, p06811318!, p06811319!, p06811320!, p06811321!, p06811335!, p06881735!). – note: corylus chinensis is recognized in the flora of china (li & skvortsov 1999), flora sichuanica (editorial board of the flora sichuanica 2012), and world checklist and bibliography of fagales (govaerts & frodin 1998). the specimen chosen as a lectotype here bears two numbers: “no. 197” (with the number circled) and “no. 212.” there are different numbering systems in delavay’s collections (holstein & weigend 2016a), and the 197 is delavay’s original number. the number “212” is apparently the specimen number placed on the sheet at the paris herbarium, which is also found on specimen p06811321. european journal of taxonomy 409: 1–75 (2018) 16 c. papyracea hickel, bulletin de la société dendrologique de france 68: 94 (hickel 1928). – type: cult. in frutic. vilmorinianum olim. cult., r. hickel s.n. (lecto-, designated here: p06811311!). – additional original material: cult. in frutic. vilmorinian., sep. 1910, r. hickel s.n. (p06811310!). – note: corylus papyracea is listed as a synonym of c. chinensis in the flora of china (li & skvortsov 1999), in the flora sichuanica (editorial board of the flora sichuanica 2012), and in the world checklist and bibliography of fagales (govaerts & frodin 1998). the material for the description of this “species” was only observed in cultivation and according to the author, it already had disappeared 10 years before publication. hickel mentions that vilmorin believed that the seeds were from china and had been originally collected by p.g. farges in 1904. the lectotype was chosen because the upper right leaf is the one shown in the illustration accompanying the protologue. the other specimen bears the unpublished name “corylus papyrifera”, but does not differ from the lectotype. possibly, that name was a working name. c. chinensis var. macrocarpa hu, bulletin of the fan memorial institute of biology, botany 8: 32 (hu 1937). – type: china, yunnan, wai-si hsien, yeh chih wa-rung, 3400 m a.s.l., aug. 1935, c.w. wang 68308 (holo-: pe00802441!; iso-: a00033779!, pe00802422!, pe00802446!, pe00802445!, pe00802439!, pe00802430!). – note: corylus chinensis var. macrocarpa is listed as a synonym of c. chinensis in the flora of china (li & skvortsov 1999), flora sichuanica (editorial board of the flora sichuanica 2012), and world checklist and bibliography of fagales (govaerts & frodin 1998). 4. corylus colchica albov prodromus florae colchicae: 219 (albov 1895). – original citation: “hab. in mingrelia: ad rupes calcareas montinum dzhwari et migaria (supra pagum kurdzu), 1900–2000 m, (n. a. 1893, nos 500, 501).” – type: georgia, [samegrelo-zemo svaneti], mingrélie, mt. djwari [jvari], frontière de la forêt, 25 aug. 1893, n.m. alboff 501 (lecto-, designated here: g00418165!). – additional type material: georgia, [samegrelo-zemo svaneti], mingrélie, mt. migaria, 2000 m a.s.l., aug. 1893, n.m. alboff 500 (syn-: g00418164!). – note: corylus colchica is recognized in the flora of the ussr (bobrov 1936a) and in the world checklist and bibliography of fagales (govaerts & frodin 1998). according to bobrov (1936a), the type is in g, whereas a duplicate is found in le. however, it is not mentioned, which of the two syntype specimens is to be regarded as the lectotype (bobrov 1936a: 267), therefore, we designate one. the existence of a le duplicate could not be confirmed (le, pers. com.). 5. corylus colurna l. species plantarum: 999 (linnaeus 1753). – c. tubulosa var. colurna (l.) risso, histoire naturelle des principales productions de l’europe méridionale et particulièrement de celles des environs de nice et des alpes maritimes 2: 333 (risso 1826) nom. illegit. superfl. – orignal citation: “habitat byzantii.” – type: herb. linn. no. 1132.4 (lecto-, designated by browicz 1972: 5: linn!). – note: corylus colurna is recognized in the flora of the ussr (bobrov 1936a), in the flora europaea (tutin & walters 1993), and in the world checklist and bibliography of fagales (govaerts & frodin 1998). c. arborescens münchh., hannover’sche gelehrte anzeigen 3 (61): 929 (münchhausen 1754). – original citation: “es ist bekannt, daß es an einigen orten, als in frankfurt” [am main, germany, in the garden of the merchant du fay (burchardt 1833)] (...) “ich habe dergleichen selber bey mir”. – type: not localized. – note: the frankfurt hazel tree was very well known back then, and it was speculated that it was the same one that was introduced by clusius in the end of the 16th century that he obtained from the austrian baron von ungnad from constantinople (goeschke 1887). the authorship of the name c. arborescens is not obvious, as the article is only cited as “s. im jul 1753. holstein n. et al., taxonomic checklist of corylus 17 m.”. however, the name c. arborescens is usually attributed to münchhausen, the author of the hortiand agricultural book “der hausvater” who owned an estate called schwöbber near hameln in lower saxony, germany. given that the protologue is published in a journal from lower saxony, it is reasonable to assume münchhausen’s authorship of c. arborescens. c. byzantina p.herm. ex mill., the gardeners dictionary ed. 7 [unnumbered] (miller 1759) nom. inval. pro syn. – original citation: “byzantine.” – type: not localized. – note: miller used this name in synonymy, as he did not adopt binomials until the 8th edition of his the gardeners dictionary (miller 1768). there, miller synonymizes c. byzantina under c. colurna. originally, miller cited linnaeus’ hortus cliffordianus (linnaeus 1737: 448) and hermann (1687: 121). the latter refers to caspard bauhin’s avellana peregrina humilis (bauhin 1623: 418) and jean bauhin et al.’s avellana byzantina (bauhin et al. 1650: 270), both of which were also cited by linnaeus (1737). a cluster of nuts with deeply laciniate bracts is depicted in bauhin et al. (1650), clearly what is nowadays regarded as c. colurna, and consistent to his later synonymization. c. nana tourrette & rozier, démonstrations élémentaires de botanique 3: 448 (la tourrette & rozier 1787). – type: not localized. – note: the authors just gave description and the species’ origin, but do not refer to any earlier publication. however, given the origin and description, they mean the lowgrowing form of the byzantine hazel (c. colurna). original material might be found in claret de la tourrette’s or in rozier’s herbarium, both held at lyjb. c. macedonica v.cordus & gesner ex strelin, realwörterbuch für kameralisten und oekonomen 4: 528 (strelin 1788). – type: germany, [cult.] in schwöbber. [originally from macedonia, cult. by münchhausen in schwöbber estate, lower saxony, germany]. – type: not localized. – note: it was described as a tree in contrast to c. colurna that was, incorrectly, considered to be a shrub. basically, this is another name for münchhausen’s c. arborescens. c. avellana var. glomerata aiton, hortus kewensis 3: 363 (aiton 1789). – c. glomerata (aiton) nois., manuel complet du jardinier, maraicher, pépiniériste, botaniste, fleuriste et paysagiste 2: 464 (noisette 1825). – original citation: “corylus nucibus in racemum congestis” (bauhin 1623). – note: apparently a hazel from macedonia [balkan peninsula] growing as a tree with clustered fruits. bauhin’s citations lead to a short description by cordus (1561: 25r) without a figure. according to the descriptions, this is c. colurna. c. colurna var. avellanoides spach, annales des sciences naturelles, seconde série – botanique 16: 104 (spach 1841). – type: cult. in h[ortus] p[arisiensis], (e), 1834, s. coll., s.n. in herb. spach (lecto-, designated here: p06809956!). c. colurna var. brachycarpa spach, annales des sciences naturelles, seconde série – botanique 16: 103 (spach 1841). – type: iconotype, watson (1823‒1825: tab. 99). c. colurna var. leptochlamys spach, annales des sciences naturelles, seconde série – botanique 16: 104 (spach 1841). – type: cult. in h[ortus] p[arisiensis], 1841, s. coll., s.n. in herb. spach (lecto-, designated here: p06809951!). c. colurna var. macrochlamys spach, annales des sciences naturelles, seconde série – botanique 16: 104 (spach 1841). – type: cult. in h[ortus] p[arisiensis], 1841, s. coll., s.n. in herb. spach (lecto-, designated here: p06809954!). c. colurna var. trichochlamys spach, annales des sciences naturelles, seconde série – botanique 16: 104 (spach 1841). – type: cult. in h[ortus] p[arisiensis], ménagerie, 1835, s. coll., s.n. in herb. spach (lecto-, designated here: p06809953!). – additional original material: cult. in h[ortus] p[arisiensis], 1827, s. coll., s.n. in herb. spach (p06809955!); cult. in h[ortus] p[arisiensis], ménagerie, 1835, s. coll., s.n. in herb. spach (p06809955!); cult. in hort[us] par[isiensis], 1841, s. coll., s.n. in herb. spach (p06809955!); s.d., s. coll., s.n. (p06810646!, p06810647!). c. cervorum petrov, byulleten’ moskovskogo obshchestva ispytatelei prirody, otdel biologicheskii n.s., 45: 121 (petrov 1936). – type: azerbaijan, karabagh, querceto-coryleta in decliviis vallis fl. khoross [kores] superiori, alt ci[rc]a 1200‒1400 m supra nivem maris, i. petrov s.n. (holo-: not localized). european journal of taxonomy 409: 1–75 (2018) 18 c. iberica wittm. ex kem.-nath., trudy tbilisskogo botaniceskogo instituta 6: 11 (kemularia-nathadze 1938). – type: georgia, iberia, in sylvis montanis, sep. 1843, wittman 351 (syn-: not localized; isosyn-: bm!, k000859912!); georgia, prov. kuthaisi, ratscha, tschelischskaya datscha, 25 aug. 1910, fijenko s.n. (syn-: not localized); georgia, ratscha, teschelishis udabno prope pag. nikortzminda in calcar., 16 sep. 1929, l.m. kemularia-nathadze s.n. (syn-: not localized); georgia, [no locality given], 20 oct. 1937, l.m. kemularia-nathadze s.n. (syn-: not localized); georgia, лес около сеп. горолеша [in silva gorolesha], известн [limestone], 20 oct. 1937, l.m. kemularianathadze s.n. (syn-: not localized); georgia, prov. tiflis, distr. duschethi, fauces gudamakhari, loc. thchemliani, 1903, a. fomin s.n. (syn-: not localized); georgia, prov. et distr. tiflis, saguramo, 25 jul. 1917, l.m. kemularia-nathadze s.n. (syn-: not localized); georgia, udzo, prope tblisi, 16 aug. 1937, l.m. kemularia-nathadze s.n. (syn-: not localized); georgia, distr. manglis, prope pag. prijut, 9 jul. 1903, a. fomin s.n. (syn-: tbi1006417!); georgia, [without locality], 28 mar. 1936, l.m. kemularia-nathadze s.n. (syn-: not localized); georgia, in faucibus fl. algeti, 22 aug. 1924, l.m. kemularia-nathadze s.n. (syn-: not localized); georgia, distr. aghbulach in silva gudarethi (in calcar.), 28 aug. 1937, l.m. kemularia-nathadze s.n. (syn-: not localized); georgia, distr. thionethi, chisso, 29 aug. 1935, i.i. tumadjanov s.n. (syn-: not localized); georgia, distr. signach, prope pag. lagodechi, 29 sep. 1905, e. koenig s.n. (syn-: not localized). – note: the l.m. kemularianathadze’s specimen from 28 mar. 1936 was not listed in the specimen list in russian. corylus iberica is regarded as a synonym of c. colchica (the plant list 2013), but as kosenko pointed out, it is more likely synonymous to c. colurna (kosenko 2002) in agreement to bobrov (1936b). corylus iberica and c. colurna are both supposed to growth as trees and share the clustered nuts with laciniate bract margins. c. iberica var. glandulosa kem.-nath., trudy tbilisskogo botaniceskogo instituta 6: 15 (kemularianathadze 1938). – original citation: “вместе с типичной формой, часто.” [together with typical form, common.]. no specimens cited. – type: not localized. c. abchasica kem.-nath., zametki po sistematike i geografii rastenii. tiflis 35: 3 (kemularia-nathadze 1978). – c. colchica var. abchasica kem.-nath., flora gruzii 3: 40 (ketskhoveli et al. 1975) nom. inval. (without latin description). – type: georgia, abkhazia, abchazia occid., ad pedem merid. montis arabica, in fruticetis subalpinis, ca. 6000 ft., g. woronow 139 (holo-: not localized; iso-: tbi1005852!). – additional type material: georgia, abkhazia, mons karschlara inter montem mamdzischcha et kutysch ad rupes calcareas; jugum bzibi, m. japsu, 28 mar. 1948, n. purtzchvanidze s.n. (para-: tbi1005851!, tbi1005853!); georgia, abkhazia, tscherkesskaja polyana, 29 jul. 1934, a.g. doluchanov s.n. (para-: tbi1005849!); georgia, distr. gudauty, in jugo chitsma, in monte “lyssaja gora”, 27 jul. 1929, m. gubbis s.n. (para-: tbi1005856!); georgia, abkhazia, suchum, in monte apianča [apiantcha], 29 may 1927, s. petjaev s.n. (para-: tbi1005858!). – note: there is a typographical error in the latin translation of the specimen list of c. abchasica: the paratype collection of n. purtzchvanidze is listed in the latin section as being from march, but the month is written in roman numerals on the sheets and in the russian text, and so the forgotten “v” changes the original august (viii) to march (iii). c. egrissiensis kem.-nath., zametki po sistematike i geografii rastenii. tiflis 35: 6 (kemularianathadze 1978). – type: georgia, megrelia, in silva montis quibia prope pag. salchino, 21 jul. 1953, l.m. kemularia-nathadze s.n. (lecto-, designated here: tbi1006350!; isolecto-: tbi1006351!). – additional type material: georgia, abkhasia, prope novi-athoni [novy afon], angustiae psychra, 300 m a.s.l., 24 jul. 1934, a. doluchanov s.n. (para-: tbi1006337!); georgia, abkhasia, gagri ad ripas flum. djovekwara, 16 oct. 1939, w. sherozia s.n. (para-: not localized); georgia, abkhasia, vicinis gagri, 29 jul. 1912, j. gusev s.n. (para-: tbi1006334!); georgia, abkhasia, pag. otradnoe inter bzipi et gegi [in hortis otradnoe], 3 jul. 1929, s. saakov s.n. (para-: tbi1006339!); georgia, abkhasia, distr. suchum, in m. byrtsch, 4 jul. 1927, s. coll.?, s.n. (para-: not localized); georgia, abkhasia, gudauthi et kaldachvara, 24 jun. 1927, s. coll.?, s.n. (para-: not localized); georgia, abkhasia, prope som. kamany, 29 jun. 1930, s. petjaev s.n. (para-: tbi1006336!); georgia, holstein n. et al., taxonomic checklist of corylus 19 abkhasia, in fauces fl.september gumista, 26 jun. 1927, nikolaev s.n. (para-: not localized); georgia, svanetia, prope pag. chaischi [khashi] in angust. fl. tcheischi [tkheishtshi] et chaischura [khaishuru], 23 jul. 1937, l.m. kemularia-nathadze and a. kharadze s.n. (para-: tbi1006345!); georgia, megrelia, vicinis pag. salchino in angustis fl. tzatschcuris, 20 jul. 1953, l.m. kemularianathadze s.n. (para-: not localized); georgia, adjaria, distr. batumi, angustiae prope pag. macrial, 1908, i. woronov and e. bobrov s.n. (para-: not localized). – note: the lectotype of c. egrissiensis was designated because the protologue does not indicate a herbarium, and there are two specimens in tbi. c. kachetica kem.-nath., zametki po sistematike i geografii rastenii. tiflis 35: 5 (kemularia-nathadze 1978). – c. kachetica kem.-nath., flora gruzii 3: 40 (ketskhoveli et al. 1975) nom. inval. (without latin description). – type: georgia, pshavi: angustiae gometsari, 28 jul. 1935, n. ketzkoveli s.n. (lecto-, designated here: tbi1006504!; isolecto-: tbi1006505!). – additional type material: georgia, kachetia, лагодехи ущелье [lagodekhi gorge] р. апцал [aptsal river], 22 aug. 1938, a.g. doluchanov s.n. (para-: not localized); georgia, ущ. р. качал [gorge of kakhal river], 10 may 1939, l.m. kemularia-nathadze s.n. (spara-: not localized); georgia, kachetia, напареули, дидихеви [napareuli, didikhebu], 12 jul. [aug.?] 1936, p.p. zagareli s.n. (para-: tbi1006501!?); georgia, телавский район [telavi district], около деревии цагаури [near zagauri forest], долина турдо [turdo valley], 1 sep. 1958, s. kutatenadze et al. s.n. (para-: tbi1006506!). – note: the zagareli specimen (tbi1006501) matches the locality and date except for the month (august instead of july as in the protologue). possibly, there was a typographical error in the publication. 6. corylus × colurnoides c.k.schneid. illustriertes handbuch der laubholzkunde 1: 145 (schneider 1904) [“1906”]. – original citation: “in hann. münden, september 1903, c.k. schneider”. – type: not localized. – note: hybrid between c. colurna × c. avellana. recognized by flora of the ussr (bobrov 1936a) and world checklist and bibliography of fagales (govaerts & frodin 1998). c. × fominii kem.-nath., trudy tbilisskogo botaniceskogo instituta 6: 15 (kemularia-nathadze 1938). – type: georgia, hortus botanicus thbilisiensis (sect. silva georgiae orient), 16 aug. 1935, l.m. kemularia-nathadze s.n. (syn-: not localized); ibid., 11 mar. 1936, l.m. kemularia-nathadze s.n. (syn-: tbi1024984!, tbi1024985!); ibid., 16 aug. 1937, l.m. kemularia-nathadze s.n. (syn-: not localized). c. × gudarethica kem.-nath., trudy tbilisskogo botaniceskogo instituta 6: 17 (kemularia-nathadze 1938). – type: georgia, prov. tblisi, distr. aghbulach prope pag. gudarethi in silva (solo calc.), 28 aug. 1937, l.m. kemularia-nathadze and r. zagareli s.n. (holo-: tbi1006382!). – note: although most type material of other names of kemularia-nathadze has not been found in tbi (most likely housed in le), the holotype of c. × gudarethica is localized there. the figure on page 19 is a drawing of the specimen tbi1006382, so this is what the author referred to as c. × gudarethica is supposedly the hybrid of c. iberica (c. colurna) and c. avellana, while in c. × fominii, the parents (parental sexes) are reversed. 7. corylus cornuta marshall arbustrum americanum: 37 (marshall 1785). – type: cult. in hort. marshall., 14 sep. 1835, s. coll., s.n. (neo-, designated here: dwc!). – note: humphry marshall’s original collection is located in dwc with some specimens also in bm (stafleu & cowan 1976–1988). however, there is apparently no extant specimen that can be regarded as original material (s.e. bartholomew-began, pers. com.). the specimen chosen here as neotype is from marshall’s garden, but collected about 50 years later. as it contains a fruit in its striking horn-like involucre, it allows unambiguous identification. european journal of taxonomy 409: 1–75 (2018) 20 7a. corylus cornuta marshall subsp. cornuta note: it has been recognized as having two subspecies by flora of north america (furlow 1997) and world checklist and bibliography of fagales (govaerts & frodin 1998). c. rostrata aiton, hortus kewensis 3: 364 (aiton 1789). – original citation: “cult. [since] 1745 by archibald duke of argyle” [sic!, archibald campbell, third duke of argyll, who laid the groundstock for kew botanical gardens] and “nat. of north america.” material from the cited gronovius (1762): “crescit in dragon swamps” [a river in virginia, usa]. – type: cult., hort. bni [botanici] collinson, mill hill, s. coll., s.n. (neo-, designated here: bm001191197!). – note: the first author did not find any original material in the relevant herbaria. in order to explore which material might be taken into account, we briefly discuss the origin of the name here. as there is no explicit indication of an author, the name is to be attributed to aiton (art 46.8. icn, mcneill et al. 2012). however, the original author of c. rostrata is apparently daniel carlsson solander. although the third volume was mainly edited by dryander (stafleu & cowan 1976–1988), dryander largely used solander’s manuscripts (britten 1912), and in the linnean herbarium, there is a specimen of what is now understood as c. cornuta (linn 1132.2) where j.e. smith writes “c. rostrata mss. 83. 931” and “c. rostrata soland[er] ait. h. kew v.3 364.” as a contemporary of the publication, smith likely knew who the author of the name was, while 81 and 931 might be interpreted as manuscript fragments. therefore, possible original material needs to be linked to solander’s collections. two sources are cited in the protologue, material from duke argyll’s garden and gronovius’ flora virginica ed. 2 (gronovius 1762). material from gronovius is usually found in the clayton herbarium (stafleu & cowan 1976–1988), but there appears to be no original material of this taxon. also, the number cited for clayton’s herbarium in gronovius’ flora virginica ed. 2 is erroneous, because clayton 747 is a juglans sp., which is already cited on the previous page. there is a specimen in bm (bm001191197) by peter collinson (1694–1768) from mill hill, where collinson lived from 1749 on (living & braithwaite 2013). collinson obtained material from gronovius for cultivation (reveal & pringle 1993), and collinson was also in contact with solander. thus, the specimen might be derived from gronovius’ collection. in addition, as a student of linnaeus, solander had access to his herbarium, so the linn specimen (linn 1132.2) might also be taken into consideration. however, neither collinson’s specimen nor the one from the linnean herbarium can unambiguously linked to solander, so they are not original material, making a neotypification necessary. due to be possible link and the completeness of the specimen, we choose the specimen from collinson’s garden as neotype. c. cornuta f. inermis fernald, rhodora 38: 76 (fernald 1936). – original citation: “quebec: abundant in border of woods, east broughton”. – type: canada, quebec, east broughton, 28 aug. 1915, m.l. fernald and h.b. jackson 12073 (holo-: gh). c. cornuta var. megaphylla vict. & rousseau, contributions de l’institut botanique de l’universite de montréal 36: 13 (marie-victorin & rousseau 1940). – type: canada, quebec, matapédia, sur les îles de la restigouche, à l’embouchure de la matapédia (frontière du nouveau-brunswick), 15 aug. 1928, j.l.c. marie-victorin and rolland-germain 28690 (lecto-, designated here: mt00132174!; isolecto-: mt00132175!). – additional type material: canada, quebec, thickets and borders of woods, near mouth of marsouin river (gaspé co.), 26 jul. 1922, m.l. fernald and a.s. pease 25014 (syn-: mt00206842!). – note: marie-victorin & rousseau cited the two collections and mentioned that the type is in the herbarium of the université de montreal (mt). however, there are two specimens of marie-victorin and rolland-germain 28690 in mt, of which one is annotated as a type, and one specimen of fernald and pease 25014 (geoffrey hall, mt, pers. com.). the annotated specimen is here formally designated as lectotype. holstein n. et al., taxonomic checklist of corylus 21 7b. corylus cornuta subsp. californica (a.dc.) a.e.murray kalmia 12: 19 (murray 1982). – c. rostrata var. californica a.dc., prodromus systematis naturalis regni vegetabilis 16 (2): 133 (candolle 1864). – c. californica (a.dc.) rose, garden & forest 8: 263 (rose 1895). – c. cornuta var. californica (a.dc.) sharp, relationships between the floras of california and southeastern united states: 58 (sharp & james 1951). – type: usa, california, 1848, k.t. hartweg 1960 (lecto-, designated here: g-dc! (g00478869); isolecto-: br!, g00418163!, gh00033775!, k000859889!, k!). – note: corylus cornuta subsp. californica is recognized in flora of north america and world checklist and bibliography of fagales (govaerts & frodin 1998). the name c. californica was published twice before rose’s combination. petzold & kirchner (1864: 671) mention a plant in the park of schloss sanssouci garden in potsdam, germany, but neither provide a description, nor explicitly recognize it as something different from c. avellana, therefore their name is invalid. koch (1873: 15) also mentions a c. californica, but also mentions only its status as a cultivated plant without providing a description. neither of the publications refers to candolle, therefore, rose’s combination is the first valid publication of this name. c. rostrata var. tracyi jeps., a manual of the flowering plants of california: 271 (jepson 1923). – original citation: “humboldt co. to siskiyou co. and s. in the sierra nevada.” – type: usa, california, humboldt co., grouse mountain, 5000 ft., 2 sep. 1917, j.p. tracy 4842 (holo-: uc198569!). c. cornuta var. glandulosa b.boivin, phytologia 15: 420 (boivin 1967). – c. cornuta f. glandulosa (b.boivin) t.c.brayshaw, catkin bearing plants of british columbia 18: 157 (brayshaw 1976). – type: canada, british columbia, head of finlayson arm below mt. finlayson, north of victoria, 16 jul. 1961, j.a. calder and k.t. mackay 31517 (holo-: dao). 8. corylus fargesii (franch.) c.k.schneid. illustriertes handbuch der laubholzkunde 2: 896 (schneider 1912). – c. rostrata var. fargesii franch., journal de botanique (morot) 13: 199 (franchet 1899). – c. mandshurica var. fargesii (franch.) burkill, the journal of the linnean society. botany 26: 505 (burkill 1899). – c. chinensis var. fargesii (franch.) hu, iconographia cormophytorum sinicorum 1: 397 (hu 1972). – original citation: “china occidentalis, provincia sutchuen, in silvis as héoupin prope tchen keou tin, alt. 1400 m.” – type: china, sichuan, tchen-keon-tin, 1400 m a.s.l., p.g. farges 1307 (lecto-, designated here: p06811278!; isolecto-: a00033782!, p06810989!, p06810990!, p06810991!, p06811279!, p06811280!, p06811281!, p06811282!). – note: corylus fargesii was recognized in the flora of china (li & skvortsov 1999), flora sichuanica (editorial board of the flora sichuanica 2012), and world checklist and bibliography of fagales (govaerts & frodin 1998). 9. corylus ferox wall. plantae asiaticae rariores 1: 77 (wallich 1830). – type: nepal, ad cacumen sheopore montis, 1821, n. wallich, wallich catalogue no. 2797 (lecto-, designated here: k001117111!; isolecto-: bm001191572!, bm001191573!, e00020277!, k000859893!). 9a. corylus ferox wall. var. ferox note: corylus ferox var. ferox and c. ferox var. tibetica were recognized in the flora of china (li & skvortsov 1999) and world checklist and bibliography of fagales (govaerts & frodin 1998) as two distinct varieties of c. ferox. european journal of taxonomy 409: 1–75 (2018) 22 9b. corylus ferox var. tibetica (batalin) franch. journal de botanique (morot) 13: 201 (franchet 1899). – c. tibetica batalin, trudy imperatorskago s.-peterburgskago botanicheskago sada. acta horti petropolitani 13: 102 (batalin 1893). – type: china, hubei, mar. 1889, a. henry 6778 (syn-: not localized; isosyn-: bm!, k000859906!, k000859907!); china, gansu, 10 jul. 1885, g.n. potanin s.n. (syn-: le; isosyn-: k000859909!, k000859910!, p06810950!, pe00021984!). – note: corylus ferox var. tibetica was recognized as a distinct species in the flora sichuanica (editorial board of the flora sichuanica 2012). 10. corylus heterophylla fisch. ex. trautv. plantarum imagines et descriptiones floram russicam illustrantes 1: 10, fig. 4 (trautvetter 1844). – c. tetraphylla fisch. ex ledeb., denkschriften der königlich-baierischen botanischen gesellschaft in regensburg 3 (2): 58 (ledebour 1841) nom. nud. (sphalm. for c. heterophylla fisch.). – c. avellana var. davurica ledeb., flora rossica 3 (2): 588 (ledebour 1850) [“1850–1851”]. – original citation: “delineavi ad specimina argunensia sicca.” – type: russia, ad fl. argun, s. coll., s.n. (lecto-, designated here: le01012787!). – additional original material: russia, ad fl. argun, [comm.? leg.?] 1836, f.e.l. fischer 5 (br!, p06810938!); russia, ad alterum litteri arguni in dahuria chinensi, 1820, c.f. von ledebour s.n. (p06810936!); russia, dahuria, ad fl. argun, n. turczaninov s.n. (goet!, p06810932!); russia, in montosis ad fl. argun, 1831, s. coll., s.n. (k!, le01012790!, mw0049851!); russia, in montosis dahuriae ad fl. argun, 1831, s. coll., s.n. (le01012785!, le01012788!); china, in dahuriae chinensi, ad fl. argunum, 1830‒1831, n. turczaninov s.n. (le01012784!, le01012789!). – note: the specimen designated as lectotype is from trautvetter’s herbarium, so it is certainly an original. it matches all characters given in the protologue as it is provided with leaves, nuts and young male catkins. the epithet heterophylla was validly published before in corylus as a variety of c. avellana, but it was used for a cultivar with deeply incised leaves (now ‘urticifolia’). however, as neither trautvetter nor fischer dealt with a cultivar, but with a siberian wild species, their name is not to be treated as a new combination (art. 41.4, mcneill et al. 2012). ledebour cited the protologue of c. heterophylla in total under his new name c. avellana var. davurica. as he included the type of c. heterophylla, his name would be superfluous, but the combination to be adopted was not available due to c. avellana var. heterophylla loudon (1838). therefore, c. avellana var. davurica ledeb. is to be regarded as a nom. nov. 10a. corylus heterophylla fisch. ex. trautv. var. heterophylla note: corylus heterophylla var. heterophylla was recognized in the flora of china (li & skvortsov 1999), flora of the ussr (bobrov 1936a), the world checklist and bibliography of fagales (govaerts & frodin 1998), and in the flora sichuanica (editorial board of the flora sichuanica 2012), although in the latter, they were regarded as distinct taxa: c. kweichowensis var. kweichowensis and c. kweichowensis var. sutchuensis. c. mongolica burchardt, verhandlungen des vereins zu beförderung des gartenbaues in den königlich preußischen staaten 9: 108 (burchardt 1833) nom. inval. (name not fully accepted). – c. mongolica k.koch, dendrologie; bäume, sträucher und halbsträucher, welche in mittelund nordeuropa im freien kultivirt werden 2 (2): 13 (koch 1873) nom. inval. pro syn. – note: theodor heinrich otto burchardt (13 dec. 1771–6 feb. 1853, müller 1861) reported that he obtained nuts from “herr von hartwitz” [n.a. von hartwiss] from nikita, crimea, who got his material from f.e.l. fischer from st. petersburg for cultivation. burchardt mentioned in the protologue that he sowed the largest ones in 1829, but none of them germinated. however, burchardt emphasized that the nuts were flattened at their tips, which is typical for what is nowadays referred to as c. heterophylla. fischer had the newly collected central asian material of c.f. von ledebour and turczaninov at hand, and holstein n. et al., taxonomic checklist of corylus 23 apparently also in cultivation in the st. petersburg botanical garden. as there is only one species known from that area, c. heterophylla, and it has flattened nuts, burchardt’s c. mongolica is an earlier name of c. heterophylla. however, burchardt expressed some doubt: “höchst wahrscheinlich werden wir aber die corylus-arten durch eine neue species, mit der corylus mongolica vermehren müssen.” [most likely, we will have to increase the number of corylus species by a new species, corylus mongolica]. due to the explicit doubt expressed, the name does not need to be considered (art. 36, mcneill et al. 2012). c. hasibami siebold, annuaire de la société royale pour l’encouragement de l’horticulture dans les pays-bas: 27 (siebold 1844) nom. nud. – note: the material of this nom. nud. was introduced in leiden botanical garden from material from p.f. von siebold in 1829. two years after publishing this name, siebold himself synonymized the name under c. heterophylla (siebold & zuccarini 1846), while blume (1851: 310) synonymized it under his new c. heterophylla var. thunbergii. c. heterophylla var. thunbergii blume, museum botanicum lugduno-batavum 1: 310 (blume 1851) [“1849–1852”]. – c. thunbergii (blume) k.koch, dendrologie; bäume, sträucher und halbsträucher, welche in mittelund nordeuropa im freien kultivirt werden 2 (2): 13 (koch 1873). – original citation: “in japoniâ.” – original material: japan, e japonia, c.p. thunberg s.n. (ups-thunb 22494!); japan [without locality], h. bürger s.n. (m-0153659!, m-0153660!, m-0153661!). – original material?: japan, japonia, s. coll., s.n. (k00859904!). c. heterophylla var. japonica koidz., the botanical magazine [tokyo] 27: 143 (koidzumi 1913). – original citation: “japonia.” – type: not localized. c. heterophylla var. yezoensis koidz., the botanical magazine [tokyo] 27: 143 (koidzumi 1913). – c. yezoensis (koidz.) nakai (nom. inval. pro syn.), flora sylvatica koreana 2: 9 (nakai 1915). – orinial citation: “yezo: prov. isikari, prope sapporo.” – type: not localized. – note: corylus yezoensis is regarded as a synonym of c. heterophylla var. thunbergii in the flora of japan (ohwi 1965) and the world checklist and bibliography of fagales (govaerts & frodin 1998), where that variety is treated as a distinct variety. c. heterophylla var. shenyangensis l.zhao & d.chen, flora liaoningica 1: 1361 (zhao & chen 1988). – type: china, liaoning, shenyang shi, l. zhao [赵冷] 591 (holo-: ifp03404001a0001). – note: the herbarium where the type is deposited was incorrectly given as “fpi” in the protologue. c. retusa siebold & zucc. nom. nud. in sched. – note: this name solely appears on some specimens collected by siebold (br!, k000859905!, m-0153658!). 10b. corylus heterophylla var. sutchuenensis franch. journal de botanique (morot) 13: 199 (franchet 1899). – c. sutchuensis (franch.) nakai, journal of japanese botany 15 (9): 524 (nakai 1939). – c. kweichowensis var. sutchuenensis (franch.) c.c.yang, flora sichuanica 21: 5 (editorial board of the flora sichuanica 2012) nom. illegit. superfl. – type: china, sichuan, tchen keou tin, 1400 m a.s.l., p.g. farges 805 [originally cited “809”] (lecto-, designated here: p06810915!; isolecto-: b!, b!, p06810916!, p06810917!). – note: corylus heterophylla var. sutchuenensis was recognized in the flora of china (li & skvortsov 1999) and the world checklist and bibliography of fagales (govaerts & frodin 1998) while it was regarded as a variety of a distinct c. kweichowensis in the flora sichuanica (editorial board of the flora sichuanica 2012). c. heterophylla var. crista-galli burkill, the journal of the linnean society. botany 26: 504 (burkill 1899). – type: china, hunan, shimên, mar. 1886, a. henry 7942 (lecto-, designated here: k000859899!). – additional type material: china, hubei, patung, sep. 1886, a. henry 1449 (syn-: k000859898!, k000859900!, k000859903!, p06809382!); china, hubei, ichang, feb. 1887, a. henry 2854 (syn-: k000859903!); china, hubei, ichang, feb. 1887, a. henry 2866 (syn-: k000859900!); china, hubei, tungku, mar. 1889, a. henry 6413 (syn-: bm!, gh00033780, k000859901!, k000859902!, p06809384!); china, yunnan, “gorges of lanho near lankong”, 7 european journal of taxonomy 409: 1–75 (2018) 24 aug. 1883, j.m. delavay 128 (syn-: p06810940!). – additional original material: china, yunnan, les bois à yang in chang près de lankong, 7 aug. 1883, j.m. delavay 128 (k!, p06810941!). – note: this variety is mainly characterized by the long glandular trichomes especially on the petioles. specimens of delavay bearing the delavay’s original number 128 (see holstein & weigend 2016a) were likely collected at different localities. burkill, however, only cited one of these gatherings, being “gorges of lanho near lankong”. the collections from “les bois à yang in chan près lankong” from the same date (k!, p06810941!) are not syntypes, but may be regarded as original material. however, they differ in being somewhat tomentose and lacking glandular trichomes on the petioles and the lower leaf lamina, linking to c. yunnanensis. the specimen chosen as a lectotype here shows the double-serrate bract margin as a name-giving character for this taxon, as well as the young male catkins that typically develop in c. heterophylla as the fruits mature. c. kweichowensis hu, bulletin of the fan memorial institute of biology n.s., 1: 149 (hu 1948). – type: china, kweichow [guizhou], ichaipa, longli county [龙里县], 6 jul. 1930, y. tsiang [蒋英] 8441 (lecto-, designated here: pe00021973!; isolecto-: pe00734547!). – note: the collection number 8411 in the protologue is mistaken, because y. tsiang 8411 is a species of pyrus, conversely y. tsiang 8441 is a corylus and matches the other collection data from the protologue. corylus kweichowensis is regarded by liang (1988) as a distinct species (with a distinct c. heterophylla var. sutchuensis) and treated as such in the flora sichuanica (editorial board of the flora sichuanica 2012). c. kweichowensis var. brevipes w.j.liang, bulletin of botanical research. harbin 8 (4): 117 (liang 1988). – type: china, jiangxi, 990 m a.s.l., 18 sep. 1984, w.j. liang 84001 (holo-: ifp03404999za001!). 11. corylus jacquemontii decne. voyage dans l’inde 4: 160 (jacquemont 1844). – c. colurna var. jacquemontii (decne.) a.dc., prodromus systematis naturalis regni vegetabilis 16 (2): 130 (candolle 1864). – original citation: “frequens à vernague ad jugum banhatte; alt. 2952 m.” – type: laos, inter vernag. et ban hatte, v. jacquemont 784 (lecto-, designated here: p06811323!; isolecto-: p06811322!). c. colurna var. lacera a.dc., prodromus systematis naturalis regni vegetabilis 16 (2): 131 (candolle 1864). – type: india, [uttarakhand], kumaon, r. blinkworth in wallich catalogue no. 2798 (syn-: not localized; isosyn-: bm001191574!, k000859891!, k001117112!). 12. corylus maxima mill. the gardeners dictionary ed. 8 no. 2 (miller 1768). – c. tubulosa willd., species plantarum ed. 4 4 (1): 470 (willdenow 1805) nom. illegit. superfl. – c. avellana var. tubulosa loudon, arboretum et fruticetum britannicum 3: 2018, fig. 1943 (loudon 1838). – original citation: only a reference to bauhin (1623: 418): “corylus sativa fructu oblongo,” which does not contain figures. – type: not localized (no original specimen found in bm or k). – note: corylus maxima was recognized in the flora of the ussr (bobrov 1936a), flora europaea and world checklist and bibliography of fagales (govaerts & frodin 1998). the absence of both a physical specimen and the direct citation of an illustration (art. 7.7, mcneill et al. 2012) makes the interpretation of this name difficult. the rather cryptic description of the stipules being intermediate between c. avellana and c. colurna makes it rather hard to understand the exact intention of the author. miller wrote that “red and white filberts, both which are so well known, as need no description” (and was definitely wrong in that point). solely his citation of bauhin’s descriptive name “corylus sativa fructu oblongo,” gives some indication of what was meant, but the most important character, the involucre of the nut, is not described by him. miller’s idea of c. maxima is clearly connected to the filbert. the word is said to be either derived from “full beard” (loudon 1838; goeschke 1887) or from an old english name (loudon 1838). there are two saints of the name philibert: philibert of jumièges, whose feast day is holstein n. et al., taxonomic checklist of corylus 25 on 20th august, and philibert of toledo, whose feast day is on 22nd august (the benedictine monks of st. augustine’s abbey, ramsgate 1921), both at the time when filberts are supposed to ripen (bunyard 1920). filberts are generally regarded as having a long involucre, much exceeding the ripe nut in length. however, there are two morphs: 1) the involucre is often deeply split and the lobes are striate to reticulate and often bend outwards, and 2) a smooth tubular involucre enclosing the nut and producing a short beak with a laciniate apex, similar to what is found in c. chinensis. brookshaw (1812, pl. 73) illustrates both kinds, the former as “barcelona filbert” (also with an oblong nut) and the latter as “white and scarlet filbert.” the citations under bauhin’s descriptive name all mention that miller’s “filbert” is cultivated and that the fruit is oblong and is either white or red. white and red filberts were described separately as c. avellana var. alba aiton and c. avellana var. rubra aiton later-on, so miller’s species includes different entities. the name c. tubulosa is superfluous, because willdenow cited c. maxima in synonymy. loudon accepted a hazel with a tubular involucre as a distinct and cited c. maxima and c. tubulosa as synonyms, though he adopted the later name. since loudon cited c. maxima mill. fully, c. avellana var. tubulosa is to be interpreted as nomen novum. as there are no types of c. maxima, c. avellana var. tubulosa is not homotypic (illegitimate) with aiton’s varieties c. avellana var. alba and c. avellana var. rubra. however, even loudon did not have a clear image of c. maxima, as loudon’s figure does not correspond to willdenow’s specimens of c. tubulosa (b-w 17704-010!, b-w 17704-020!, b-w 17704–030!). c. avellana var. sativa c.f.ludw., die neuere wilde baumzucht: 14 (ludwig 1783) nom. nud. c. avellana var. alba aiton, hortus kewensis 3: 363 (aiton 1789). – c. alba (aiton) steud., nomenclator botanicus ed. 2 1: 424 (steudel 1840). – c. maxima var. alba (aiton) c.k.schneid., illustriertes handbuch der laubholzkunde 1: 149 (schneider 1904). – original citation: “corylus sativa fructu albo minore sive vulgaris” (bauhin 1623). – type: not localized. – note: this name is only accompanied with the english common name, so it lacks a description itself, but it contains a validating description by bauhin (1623; art. 7.7, mcneill et al. 2012). as there are no corresponding specimens in either kew or bm, nor a figure in bauhin’s pinax theatri botanici, there is no original material. the citations under the name “corylus sativa fructu albo minore sive vulgaris” are ambiguous. they sometimes mention a round nut in contrast to the oblong nut typical of the filbert. this is in contrast to the citation of dodoens’ c. hortensis (dodoens 1757: 515), to whom bauhin also referred. dodoens differentiates and illustrates two hazels, a wild one (a mirror image copy from fuchs (1542: 398), which is the lectotype of c. avellana l.), and the domesticated one (fuchs 1542: 399). dodoens’ cultivated hazel has a laciniate involucre, and he explicitly mentions the red pellicule (testa) of the nut. bauhin also cites fuchs’ avellana domestica, although fuchs clearly states “rubra membrana inclusus”, which appears to be in contrast to the “white” fruit of bauhin’s name. c. avellana var. rubra aiton, hortus kewensis 3: 363 (aiton 1789). – c. rubra (aiton) borkh., theoretisch praktisches handbuch der forstbotanik 1: 723 (borkhausen 1800) nom. illegit. superfl. – c. maxima var. rubra (aiton) c.k.schneid., illustriertes handbuch der laubholzkunde 1: 149 (schneider 1904). – c. maxima var. rubra f. normalis (aiton) c.k.schneid., illustriertes handbuch der laubholzkunde 1: 149 (schneider 1904) nom. inval. (art. 24.3, mcneill et al. 2012). – original citation: “corylus sativa fructu oblongo rubente” (bauhin 1623: 418). – type: not localized. – note: as for c. avellana var. alba, c. avellana var. rubra is only accompanied with the english native name, but it contains a validating description by bauhin (art. 7.7, mcneill et al. 2012). as there are no corresponding specimens in either kew or bm, nor a figure in bauhin’s pinax theatri botanici, there is no original material. according to bauhin’s phrase name, this taxon has oblong reddish nuts, corresponding to the red filbert of miller (1768). rea (1665) already described the “wellknown” filberts and indicates that the “red filbert” has a red testa, but he did not mention red leaves, although he discussed leaf characters for other names. also dodoens (1757: 515) mentioned a red pellicule. therefore, the mention of the red color is thus not referring to the foliage, but the testa. the application and interpretation of borkhausen’s c. rubra is confusing in this context, because european journal of taxonomy 409: 1–75 (2018) 26 the same descriptive epithet was published before, but it was eventually used to describe something different. in all cases, the authors deal with filberts (in the sense of c. maxima mill.) as they refer to the same pre-linnean polynomina by bauhin. however, on the one hand, the epithet in c. avellana var. rubra aiton (1789: 363) describes the red testa. on the other hand, borkhausen understood the epithet to describe a plant with red or brownish leaves, while aiton did not mention the leaves at all. a red-leaved filbert was mentioned by du roi (1771: 178) for the first time, but without giving it a valid name. when du roi dealt with miller’s corylus maxima, he distinguished white filberts and red filberts as did miller and later aiton; he even cited the same polynomina by bauhin. but in contrast to miller and aiton (and any other author to our knowledge), already du roi understood his “red filberts” to have red or brownish leaves. since there is no information on the leaf color of miller’s and aiton’s “red filbert”, it is doubtful whether borkhausen actually meant his name to be something new. even goeschke (1887) mentions that the “rote lambertsnuss”/red filbert has slightly brownish leaves, while he accepted a purple-leaved filbert as something different. corylus rubra is therefore interpreted best a new combination, because borkhausen meant the same “red filbert” that also miller and aiton cited by using bauhin’s polynomen, although aiton and borkhausen referred the epithet to different characters. since borkhausen does not discuss green-leaved filberts or cited any filberts under another name, there is no apparent intent to describe a new species but rather to use a better-fitting name. therefore, art. 41.4 (mcneill et al. 2012) applies. however, borkhausen’s new combination is illegitimate, as he also cites a “corylus maxima münchh.” in synonymy. the present authors are not aware of any publication of this name, but rather interpret this name as an “updated” citation. münchhausen does not use miller’s binomial in the third volume of his der hausvater (münchhausen 1768: 827), but refers to miller’s polynomen (miller 1759) that miller took up later. münchhausen also did not publish that name when he dealt with hazelnuts again in the fifth volume of the der hausvater (münchhausen 1770: 142). he omitted the discussion of the filberts at all by referring to his publication in the third hausvater volume. after miller adopted the binomials in the eigth edition of his the gardeners dictionary and published c. maxima (miller 1768), borkhausen apparently updated münchhausen’s description (who also did not mention red leaves). nevertheless, borkhausen makes an indirect reference to a published name on the same rank without excluding parts of it, rendering his c. rubra superfluous. c. tubulosa var. oblonga risso, histoire naturelle des principales productions de l’europe méridionale et particulièrement de celles des environs de nice et des alpes maritimes 2: 332 (risso 1826). – type: not localized. c. intermedia fingerh., linnaea 4: 384, tab. iii fig. 4 (fingerhuth 1829). – original citation: “wächst an schattigen stellen bei esch im köllner regierungs-bezirk.” – type: not localized. c. sativa poit. & turpin, pomologie française recueil des plus beaux fruits cultivés en france 4: tab.11 (poiteau 1846) nom. illegit. superfl. – note: this name was published with c. tubulosa willd. as a synonym. the plate also depicts the kind of hazel that is here considered to represent c. maxima. poiteau and turpin even show the dark red testa that was mentioned in literature cited by bauhin on which c. maxima relies upon. c. avellana var. viminalis henriksson, botaniska notiser 1918 (6): 297 (henriksson 1918). – type: sweden, dalia, paroecia gunnarsnäs, in silva hällan ad mörtviken, j. henriksson s.n. (holo-: not localized). 12a. corylus maxima ‘purpurea’ c. avellana var. purpurea hort. ex loudon, arboretum et fruticetum britannicum 3: 2017 (loudon 1838). – c. tubulosa [unranked] purpurea audib. ex mouill., traité des arbres & arbrisseaux 2: 1142 (mouillefert 1891–1898) nom. illegit. superfl. – c. maxima var. purpurea (hort. ex loudon) rehder, manual of cultivated trees and shrubs hardy in north america: 154 (rehder 1927). – c. maxima (var. rubra) f. purpurea (hort. ex loudon) c.k.schneid., illustriertes handbuch der holstein n. et al., taxonomic checklist of corylus 27 laubholzkunde 1: 149 (schneider 1904). – c. maxima f. purpurea (hort. ex loudon) rehder, bibliography of cultivated trees and shrubs hardy in the cooler temperate regions of the northern hemisphere: 111 (rehder 1949). – c. purpurea a.dc., prodromus systematis naturalis regni vegetabilis 16(2): 132 (candolle 1864) nom. inval. pro syn. – c. atropurpurea hort. ex loudon, arboretum et fruticetum britannicum 3: 2017 (loudon 1838) nom. inval. pro syn. – c. atropurpurea booth et al., james booth & söhne, eigenthümer der flottbecker baumschulen bei hamburg 1845: 45 (booth et al. 1845).– c. avellana var. atropurpurea (booth et al.) g.kirchn., arboretum muscaviense: 670 (petzold & kirchner 1864). – c. tubulosa var. atropurpurea (booth et al.) a.dc., prodromus systematis naturalis regni vegetabilis 16(2): 132 (candolle 1864). – c. maxima f. atropurpurea (booth et al.) dippel, handbuch der laubholzkunde 2: 130 (dippel 1891) [“1892”]. – type: not localized. – note: this is a cultivar with dark red to purple leaves, but loudon did not give any information on the involucre or the shape of the nut necessary for the differentiation between c. avellana and c. maxima. more complicating, loudon did not differentiate between c. avellana and c. maxima at a species level. loudon cited loddiges’ catalogue of plants ed. 16 (loddiges 1836) as only source. original material therefore came from commercial cultivation, and there is no physical material to our knowledge. goeschke (1887: 79) reported that his “rotblättrige lambertsnuss” (“red-leaved lambert’s hazel or filbert”) was introduced as “corylus atropurpurea” by the pépinières baumann in bollwiller, alsace, france in the 1820s. loudon cited just this name (in synonymy to purpurea), and it is therefore likely that goeschke had the original cultivar in mind in his book. according to goeschke’s description, this cultivar has purple leaves, a smooth tubular involucre, and the nut has a pale pink testa. this cultivar still exists under the name ‘purple aveline’. red/purple foliage is apparently of a single origin in corylus. thompson (1985) reported that the locus for leaf coloration (a for anthocyanin) and the self-incompatibility “gene” were linked in corylus, which is confirmed by ives et al. (2014). the cultivars ‘fuscorubra’ and ‘rode zeller’ have the s6 allele, while ‘purple aveline’ (rotblättrige lambertsnuss/c. atropurpurea hort./c. maxima ‘purpurea’) has the alleles s5s10. the cultivar ‘fuscorubra’ has a short bract husk, while ‘rode zeller’ has a long husk. in ‘rode zeller’, however, the husk is not long tubular, but the bract lamina is striate and presents the nut openly in contrast to c. maxima ‘purpurea’. both cultivars inherit the red leaf coloration (thompson et al. 1996), while c. maxima ‘purpurea’/’purple aveline’ does not and is thus interpreted as graft hybrid on ‘white aveline’, which is c. maxima/c. avellana var. alba. as loudon cited the horticultural epithet atropurpurea in synonymy, it is a pro synonym of purpurea. as a horticultural variety (cultivar) it is best called c. maxima ‘purpurea’, since this epithet is the oldest, unambiguously referable name to the cultivated plant (art. 29.1, brickell et al. 2009). however, it must be noted that the name “c. maxima” has been widely misapplied to a large and round-fruited cultivar of c. avellana, and it was even validly published as c. avellana var. maxima for just that round-fruited cultivar. bean’s c. maxima var. atropurpurea (bean 1898: 350) is such a case, and the described plant likely refers to a cultivar similar to ‘rode zeller’. 13. corylus potaninii bobrov sovietskaia botanica 1936 (1): 29, fig. 3 (bobrov 1936b). – original citation: “hab. china, prov. sutchuen, ta-chien-lu, s. datzzien-lu, tibetice tarsando, 26 vi 1893; 1 vii 1893 (typus) g.n. potanin legit; ut videtur in prov. hupeh, yunnan et kweichow crescit.” – type: not localized. – note: it is unclear from the protologue whether c. potaninii is intended as a new species or a nom. nov. on the one hand, bobrov cited c. heterophylla var. sutchuensis and c. heterophylla var. crista-galli just after the new name in the protologue. however, he did not formally synonymize or accept these names anywhere else, so they might be regarded as synonyms of his new name. however, bobrov expressedly distinguished his new species from these varieties later-on. therefore, c. potaninii is best considered as intended as a new species. corylus potaninii is supposed to be similar to c. heterophylla var. sutchuensis, but differing in rounded leaves and densely pubescent lower leaf european journal of taxonomy 409: 1–75 (2018) 28 laminae and young twigs. bobrov also mentioned similarities to c. heterophylla var. crista-galli most likely referring to the long-stalked glandular trichomes. long-stalked glandular trichomes are also found on an isolectotype specimen of c. heterophylla var. yunnanensis (p06810939), therefore either the type material is mixed or taxon delimitation is obscure. corylus potaninii was recognized by world checklist and bibliography of fagales (govaerts & frodin 1998), but not even mentioned in the flora of china (li & skvortsov 1999) or in the flora sichuanica (editorial board of the flora sichuanica 2012). 14. corylus sieboldiana blume museum botanicum lugduno-batavum 1: 310 (blume 1851) [“1849–1852”]. – c. heterophylla var. sieboldiana (blume) a.dc., prodromus systematis naturalis regni vegetabilis 16 (2): 130 (candolle 1864). – c. rostrata var. sieboldiana (blume) maxim., bulletin de l’académie impériale des sciences de st.-pétersbourg 27: 538 (maximowicz 1881). – original citation: “siki-fasibami s. tsumo-fasibami japonice. sponte in japonice montibus nikoo fide speciminis herb. von siebold.” – type: japan, [locality unknown], p.f. von siebold. 2 or 16 (lecto-, designated here: upper plant of l0366714!). – additional type material: japan, [unclear locality], p.f. von siebold. 2 or 16 (syn-: lower plant of l0366714!). – note: there are actually two collections on the sheet, the upper one with fruits and a lower one without. there are four labels on the specimen. a first label is attached to the upper specimen containing the japanese name in katagana signs シ[shi]キ[ki]ハ[ha]シ[shi]ハ[ha, also to be pronounced as ba] ミ[mi], literally “lucky hazel”. a second label is a leiden label with the latin name c. sieboldiana and transcription of the japanese name “si-ki-hasi-bami”. a third label is siebold’s original in the lower left with the number 16 and the japanese name in katagana signs ツ[tsu]ノ[no]ハ[ha]シ[shi]ハ[ha, also to be pronounced as ba]ミ[mi], literally “horned hazel”. the fourth label bears the number 2, locality as in the protologue, and the latin and the japanese name. it is unclear which label with a number corresponds to which specimen or where the “no. 2” comes from, therefore the lectotype citation here is given rather crudely. the “f” in the transcriptions in the protologue is easily explained as misreading of the “h” of the original label in the kurrent handwriting, which is similar to the “f”. the indument of the tubular involucre varies among the different type specimens of the synonyms. two basic forms can be found in different described varieties of c. sieboldiana. the involucre is glabrous or very finely hairy (c. sieboldiana var. mitis and c. hallaisanensis), while it consists of dense bristly trichomes in c. sieboldiana var. brevirostris, c. sieboldiana var. mandshurica, and c. sieboldiana var. sieboldiana. the type of c. hallaisanensis differs from c. sieboldiana var. mitis in having apically coarsely irregular serrated leaves, like c. sieboldiana var. mandshurica. to the present authors’ knowledge, there is no critical revision or phylogeographic analysis existing that has tested the significance of the involucre indument. 14a. corylus sieboldiana blume var. sieboldiana c. rostrata var. mitis maxim., bulletin de l’académie impériale des sciences de st.-pétersbourg 27: 539 (maximowicz 1881). – c. sieboldiana var. mitis (maxim.) nakai, flora sylvatica koreana 2: 13 (nakai 1915). – original citation: “japonia. nippon. fudzi yama.” – type: japan, fudji yama, 1864, s. tschonoski s.n. (holo-: le; iso-: bm!, br!, gh00033783!, k000859897! k000859896!, p06810985!). – note: corylus rostrata var. mitis was regarded as synonymous to c. sieboldiana in the flora of japan (ohwi 1965) and the world checklist and bibliography of fagales (govaerts & frodin 1998). c. sieboldiana var. brevirostris c.k.schneid., plantae wilsonianae 2: 453 (sargent 1916). – c. brevirostris (c.k.schneid.) miyabe, journal of the faculty of agriculture, hokkaido university 26: 458 (miyabe & kudo 1934). – type: japan, hokkaido, shiribeshi-san, 700 m a.s.l., 27 jul. 1914, e.h. wilson 7268 (holo-: a00033784!). – additional type material: japan, hakodate, 1861, holstein n. et al., taxonomic checklist of corylus 29 c. maximowicz s.n. (para-: bm!); japan, northern honshū, prov. mutsu, hakkoda mountain, aug. 1904, u. faurie 5803 (para-: bm!, e00020189!). – note: corylus brevirostris was recognized as a separate species by world checklist and bibliography of fagales (govaerts & frodin 1998), while it is recognized as c. sieboldiana var. brevirostris in the flora of japan (ohwi 1965). c. hallaisanensis nakai, repertorium specierum novarum regni vegetabilis 13: 250 (nakai 1914). – original citation: “quelpaert: in silvis hallaisan.” – type: korea, hallasan mountain, oct. 1907, e.j. taquet 333 (syn-: not localized; isosyn-: e00020176!, ti); ibid., 12 jun. 1913, t. nakai 335 (syn-: not localized); ibid., 17 may 1913, t. nakai 930 (syn-: not localized). – note: corylus hallaisanensis was recognized as separate species in the world checklist and bibliography of fagales (govaerts & frodin 1998). chang et al. (2004) suggested synonymizing this name to be under c. sieboldiana, but fail to indicate to which of the two varieties it is supposed to belong (c. sieboldiana var. mandshurica or c. sieboldiana var. sieboldiana). 14b. corylus sieboldiana var. mandshurica (maxim.) c.k.schneid. plantae wilsonianae 2: 454 (sargent 1916). – c. rostrata var. mandshurica (maxim.) regel, bulletin de la classe physico-mathématique de l’académie impériale des sciences de saint-pétersbourg 15 (14‒15): 221 (regel 1857). – c. mandshurica maxim., bulletin de la classe physico-mathématique de l’académie impériale des sciences de saint-pétersbourg 15 (8‒9): 137 (maximowicz & ruprecht 1856). – type: russia, amur river, c.j. maximovicz s.n. (lecto-, designated by bobrov 1936a: 268: le; isolecto-: m-0265486!, gh00033781!, k000859911!, p06810979!, p06810986!, pe00021985!). – note: this name was also regarded as a variety in the flora of japan (ohwi 1965), while it was recognized as a distinct species in the flora of the ussr (bobrov 1936a), the flora of china (li & skvortsov 1999), and in the flora sichuanica (editorial board of the flora sichuanica 2012). c. brevituba kom., izvestija glavnogo botaniceskogo sada sssr [bulletin du jardin (botanique) principal de l’urss] 28: 219 (komarov 1929). – c. mandshurica f. brevituba (kom.) kitag., neolineamenta florae manshuricae: 217 (kitagawa 1979). – original citation: “legit o.o. kuzeneva 12 viii 1926 in provincia amurensi ad ripas fl. tomj (fl. zea influentis), in decursu superiore ejus ad pedem montium burejensium: nᵒ 520 et 523. in declvi silvoso ad fontem.” – type: amur area, basin of the zeya river, the tom’ river, upper course, offsets of bureinskii range, slope at the source of spring, 12 aug. 1926, o.o. kuzeneva 523 (lecto-, designated here: le!). – note: corylus brevituba was recognized as separate species by flora of the ussr (bobrov 1936a), although it is noted that is scarcely different from c. mandshurica. the lectotype was placed to le (bobrov 1936a: 268), but without mentioning a specific specimen. c. mandshurica f. glandulosa s.l.tung, bulletin of botanical research. harbin 1 (1‒2): 138 (tung 1981). – type: china, heilongjiang province, shangzhi country, 2 aug. 1956, y.l. chou [周以良] 3902 (holo-: nefi). – additional type material: china, jilin province, linjiang country, 1 jul. 1962, p.h. huang [黄普华] et al. 60222 (para-: not localized); china, heilongjiang province, jingpo lake, [mudanjiang team] 129 (para-: not localized). 15. corylus wangii hu bulletin of the fan memorial institute of biology, botany 8: 31 (hu 1937). – type: china, yunnan, weisi hsein, yeh-chih, 3400 m a.s.l., aug. 1935, c.w. wang 68243 (lecto-, designated by lin et al. 2007: 1249: pe01062681!; isolecto-: a00033785!, k000859895!, p01903232!, pe01062680!). – note: corylus wangii was recognized in the flora of china (li & skvortsov 1999) and the world checklist and bibliography of fagales (govaerts & frodin 1998). hu (1948: 149), on the other hand, regarded his species as synonymous to c. jacquemontii decne. however, c. wangii has an involucre with many glandular trichomes, while c. jaquemontii is eglandular. both c. jacquemontii european journal of taxonomy 409: 1–75 (2018) 30 and c. wangii have long laciniate involucre apices, but these are sometimes divided in the latter. this latter character is very well developed in c. ferox. the involucre of c. wangii are thus strikingly similar to c. × spinescens rehder, an artificial hybrid between c. ferox var. tibetica and c. avellana. 16. corylus wulingensis q.x.liu & c.m.zhang bulletin of botanical research. harbin 10 (1): 35 (liu & zhang 1990). – type: china, hunan, shauangzh xian badagungshan yochang, 1450 m a.s.l., aug. 1988, c.m. zhang [张灿明] 88 08185 (“typus in herb. inst. sylv. hunan conservatur”). – note: corylus wulingensis was recognized by the world checklist and bibliography of fagales (govaerts & frodin 1998), but not mentioned in the flora of china (li & skvortsov 1999). however, according to the protologue and the accompanying figure, this taxon is morphologically intermediate between c. yunnanensis and the type of c. heterophylla var. crista-galli (listed here as a synonym of c. heterophylla var. sutchuensis). it shares the dense indumentum with the former, and the occurrence of glandular trichomes and the short involucre with the latter taxon. 17. corylus yunnanensis (franch.) a.camus bulletin du muséum national d’histoire naturelle ser. 2 1: 438 (camus 1930) [“1929”]. – c. heterophylla var. yunnanensis franch., journal de botanique (morot) 13: 198 (franchet 1899). – original citation: “china occidentalis, in provincia yunnan, in silvis prope collum hee chan men, alt. 3000 m. (r.p. delavay); in monte tsang chan (id., n. 555).” – type: china, yunnan, les bois au col de hee chan men, 3000 m a.s.l., 5 jul. 1889, j.m. delavay s.n. (lecto-, designated here: p01903229!; isolecto-: p06810939!, p06811346!). – additional type material: china, yunnan, hee chan men, feb. 1883, j.m. delavay s.n. (syn-: p06810993!); china, mt. tsang chan, s / tali, 4 jul. 1882, j.m. delavay 555 (syn-: p01903230!, p01903231!). – note: corylus yunnanensis was recognized in the flora of china (li & skvortsov 1999), the flora sichuanica (editorial board of the flora sichuanica 2012), and world checklist and bibliography of fagales (govaerts & frodin 1998). hybrid names c. × vilmorinii rehder, journal of the arnold arboretum 7 (2): 146 (rehder 1926). – type: usa, massachusetts, cult. at arnold arboretum from material by m.l. de vilmorin, sep. 1919 (syn-: not localized); ibid., 18 aug. 1921, j.g. jack bis aa7549 (syn-: a00033778); ibid., 7 apr. 1922 (syn-: not localized); ibid., 23 sep. 1925 (syn-: not localized). – note: artificial hybrid of c. avellana and c. chinensis. c. × spinescens rehder, journal of the arnold arboretum 7 (2): 147 (rehder 1926). – type: usa, massachusetts, cult. by arnold arboretum from material by m.l. de vilmorin, 23 sep. 1925, a. rehder aa19020 (lecto-, designated here: a00033777!; isolecto-: a00033776!). – additional type material: usa, massachusetts, cult. at arnold arboretum from material by m.l. de vilmorin, 6 mar. 1922, a. rehder aa19020 (syn-: not localized); ibid., 5 dec. 1921, a. rehder aa19020 (syn-: not localized). – note: artificial hybrid of c. avellana and c. tibetica. nomina dubia c. avellana var. ölandica fr., summa vegetabilium scandinaviae 1: 55 (fries 1845) nom. nud. c. californica g.kirchn., arboretum muscaviense: 671 (petzold & kirchner 1864) nom. nud. c. mexicana hort. ex g.kirchn., arboretum muscaviense: 672 (petzold & kirchner 1864). – type: not localized (from cultivation from königliche landesbaumschule sanssouci [potsdam, germany]). – holstein n. et al., taxonomic checklist of corylus 31 note: petzold and kirchner received a sapling that is supposed to have come from mexico. the present authors, however, are not aware of any native species of corylus from today’s mexico, however, formerly, mexico included the areas of today’s california, utah, nevada to texas. this name could refer to c. cornuta subsp. cornuta or c. cornuta subsp. californica. c. avellana var. arenata henriksson nom. nud. in sched. – associate material: sweden, dalia, paroecia ärtemark, 2 sep. 1926, j. henriksson s.n. (ld1782996). – note: this voucher contains sterile specimens with branches with leaves with retuse apex, which is common in c. heterophylla and c. mandshurica. c. avellana f. olandica h.j.andersson nom. nud. in sched. – associate material: sweden, öland, borgholm, aug. 1907, h.j. andersson s.n. (bm!, bm!, ld1328264); sweden, borgholms alfvar, jul. 1896, j. erikson s.n. (ld1328384). c. avellana var. silvestris henriksson nom. nud. in sched. – associate material: sweden, dalia, paroecia gunnarsnäs in pascuis silvosis ad hällan, 89 m a.s.l., 10 sep. 1915, j. henriksson s.n. (ld1791305). c. formosana hayata, general index to the flora of formosa as recorded in all literature up to the publication of icones plantarum formosanarum vi: 72 (hayata 1917) nom. nud. c. heteromorpha c.c.yang nom. nud.? in sched. – associate material: china, yunnan, songming county [嵩明县], 邵甸大竹园乡果东照壁山, sparse woodland, shade, wet slopes, 22 apr. 1956, b.j. qiu [邱柄云] 51596 (pe00021981!); ibid., guǒ dōng [果东], 23 aug. 1957, b.-j. qiu [邱柄 云] 54913 (pe00021975!). – note: corylus heteromorpha c.c.yang is listed in chinese virtual herbarium as c. heleromorpha. the bracts are similar to c. jacquemontii, but much shorter, likely only barely surpassing the nuts (fallen out on the specimens). the present authors are not aware of a publication of this name. the seen specimens bear labels by yang from 1990, and later labels with that name again from 2006. c. spiculata c.c.yang nom. nud.? in sched. – associate material: china, s. loc., aug. 1911, s. coll., s.n. (n061030513 n.v.); china, guizhou, hezhang county [赫章县], kunming workstation [昆 明工作站] 1091 (pe00021982!); china, guizhou, qingzhen city [清镇市], dà pō mountain [ 大坡山], 1150 m a.s.l., 17 oct. 1956, x.l. jiang [蒋兴麟] 2124 (pe00021983!); china, guizhou, bijie city [毕节市], 二区保合乡, 19 aug. 1957, p.h. yu [禹平华] 314 (pe00021978!); china, guizhou, hezhang county [赫章县], 财神塘, 1480 m a.s.l., 11 oct. 1957, p.h. yu [禹平华] 1291 (pe00021977!). – note: the present authors are not aware of a publication of this name. the seen specimens bear labels by yang from 1977, and later labels with that name again from 2006. the involucre is leafy with a double-serrate margin, however, it is much longer than the nut. there are similarities to c. heterophylla var. sutchuensis, where the involucre is about are as long as the nut. however, given the large variability of the involucre length, for example in c. avellana, description of this name should be underpinned by other strong evidence than just the involucre morphology. nomina excludenda c. virginiana münchh., der hausvater 3: 830 (münchhausen 1768) nom. utique rejic. (holstein & weigend 2016b; wilson 2017). discussion as for carpinus, ostrya, and ostryopsis (holstein & weigend 2016a, 2017), we found many overlooked basionyms (even if henriksson’s names are ignored). ninety-five basionyms, except for the nomina dubia european journal of taxonomy 409: 1–75 (2018) 32 and excludenda, but including illegitimate and invalid names as far as unambiguously assignable to a valid name were not listed before. the nescience (and rediscovery) of older names, such as c. avellana var. microphylla lej., c. avellana f. peltata buser, and c. avellana f. monstrosa henriksson rendered later names illegitimate. although not of significant consequence in corylus, old names might pose a threat to the stability of currently used names. corylus mongolica burchardt might have challenged the well-established c. heterophylla for priority if burchardt had accepted his species fully. furthermore, some epithets were used on species rank and infraspecific rank despite being based on different types, but not illegitimate according to art. 53.4 (mcneill et al. 2012). corylus avellana var. heterophylla loudon (1838) and c. heterophylla fisch. ex trautv. (trautvetter 1844) and c. avellana var. tenuis loudon (1838) and korallova’s fossil hazel c. tenuis v.v.korallova (korallova & panova 1971) are quite similar; and the present authors did not find validly published combinations of the older names that would render the later name illegitimate. although overlooked names caused the creation of a new epithet for an existing taxon that simply would have needed a status change in ostryopsis (holstein & weigend 2016a) and led to missing four species in a major regional floristic treatment in carpinus (holstein & weigend 2017), the consequences in corylus are minor. but then again, the omission of corylus potaninii, described from sichuan, in the flora of china (li & skvortsov 1999) and the flora sichuanica (editorial board of the flora sichuanica 2012) may be a result of nescience. the vast majority of heterotypic synonyms is found in c. avellana var. avellana (85) followed by c. colurna (14), both from europe, while recognition of the major taxa in north america and east asia is less ambiguous. many of the taxon definitions go back to characters of the involucre and the shape of the nut and combinations of the two. thus, henriksson described 61 legitimate subspecific taxa (plus several illegitimate and invalid names) of c. avellana to record all differences found in scandinavian hazels. while his taxa are hardly of taxonomic value, he 1) followed the early 20th century fashion of splitting, and 2) nicely documented the variability of fruit and involucre characters in what today is treated as a single species without hesitation. one pervasive problem, especially in the horticultural varieties, is their large-scale negligence in botanical collections and especially herbaria. few of the varieties have any herbarium vouchers associated, making their unequivocal identification highly problematic especially if, as in much of the older literature, neither detailed descriptions nor illustrations are provided. this is a problem that not only affects corylus systematics. there is generally an under-documentation of wide-spread and cultivated plants (a. rockinger (m), f. luebert (bonn), pers. com.), with c. maxima, a species widely recognized in floras, but only known from cultivation, being a particularly striking example. the observation that hedera helix l. (linnaeus 1753: 202), a wide-spread cultivated and feral climber, had not been documented for more than 50 years in the vast herbarium of the botanische staatssammlung münchen (f. schuhwerk, pers. com.) is a point in case. while the existence of four major groups in corylus (whitcher & wen 2001; forest et al. 2005; li 2008; grimm & renner 2013) is barely in debate, morphological identification within these groups appears to be more challenging. on the other hand, bassil et al. (2013) found evidence for genetic separation of only some morphologically hardly distinguishable taxa, for example c. cornuta subsp. californica (vs c. cornuta subsp. cornuta) and c. jacquemontii (vs c. colurna). the latest global taxonomic treatment of corylus is by bobrov (1936b), and there has been a major regional update for china (li & skvortsov 1999). much molecular work has been done, but so far with a focus on cultivars (e.g., boccacci et al. 2006; gökirmak et al. 2009; bassil et al. 2013). the present study attempts to provide the basis for further studies on corylus phylogeny and taxonomy by providing a detailed list of taxa recognized at one time or another. there are two overall layers to the taxonomic problems, the natural diversity in the wild, which sometimes leads to the recognition of more or fewer species and subspecies across the range of the genus, and the multiplicity of horticultural forms and selections, which have often been treated as holstein n. et al., taxonomic checklist of corylus 33 varieties and subspecies and even species under the icn in the past. stable and unequivocal naming of cultivars based on latin names is going to be much facilitated by the comprehensive list of names here provided. resolving the remaining taxonomic issues for the wild taxa outside western eurasia might be relatively straightforward, based on critical herbarium and field studies plus modern, molecular tools. the clear definition of the cultivated forms, virtually all from western eurasia, will also find a new basis now, since the clear definition of the wild taxa will help to re-investigate the origins of cultivated forms. there is some overlap between the two, where the identity and origin of cultivated forms remains obscure. the majority of leaf and color variants can be easily assigned to c. avellana in the strict sense, but the large-fruited forms bred for the production of hazelnuts and “filberts” remain highly problematic. fruits of corylus have here been extensively used by humans for at least 10 400 years (holst 2010; zohary et al. 2012; antolín & jacomet 2015), and there is evidence that they have been traded regionally for at least 4000 years (fairbairn et al. 2014). they had been domesticated and were widely traded during the roman empire 2,000 years ago (pliny the elder 77–79; boccacci & botta 2009). as active trade and breeding have such a long tradition in europe and minor asia, different morphs and mutations were most likely taken up independently in different regions. boccacci & botta (2009) argue for an independent domestication of c. avellana in the western and the eastern mediterranean, which would imply that the present-day large fruited forms do not have a single, common origin. the fruit-hazels essentially differ in two sets of characters: nut shape (spherical versus elliptical in outline) and involucres (shorter than or equal to the nut, flaring and about twice as long, or much longer and more or less beaked, i.e., contracted above the nut). these characters are more or less freely combined in different varieties. corylus, and especially the large-fruited forms and the wild c. avellana, have been shown to be fully interfertile except when the parents share self-incompatibility alleles (erdoğan 1999; erdoğan & mehlenbacher 2000), which must have contributed to some of the diversity observed. however, the large fruited forms do not seem to become naturalized anywhere to any extent, as all national and regional floras clearly emphasize (bobrov 1936a; săvulescu 1952), nor is there any clear evidence that wild and domesticated corylus do hybridize noticeably where they co-occur. corylus maxima is a particularly curious case, because the name is widely accepted, but there is neither type material, nor are any wild collections. when miller described the species, he mentioned that it was so widely known, that a detailed description would be superfluous. poiteau & turpin (poiteau 1846) mention and illustrate it (under the illegitimate name c. sativa), yet, herbarium specimens are unknown. we only found the red-leaved c. maxima ‘purpurea’ in herbarium collections, although green-leaved plants are still in cultivation (as ‘aveline blanche longue’, ‘white filbert’, and ‘aveline rouge’ or ‘red filbert’). many “corylus maxima” collections, however, are misidentified and might be treated under the name c. avellana var. grandis in so far as they have large, but round nuts. the application of formal latin names for cultivated plants causes a lot of taxonomic uncertainty, i.e., because they tend not to be associated with the documentation usually provided for new species, such as detailed diagnoses and herbarium vouchers. dochnahl’s singular attempt of naming cultivars separate from biological taxa (greuter & holstein 2016) was not a productive effort to resolve this issue. by tracking the sparse evidence from old literature due to absence of type material, we were able to reconstruct the concept behind miller’s c. maxima. acknowledgements we gratefully acknowledge the great help of the herbarium and literature digitization services, the curators and staff of b, bm, br, dwc, e, g, goet, h, k, le, liv, m, manch, mel, mo, msb, mt, and neu for making the collections available or confirmation of lacking types. thanks are given to john mcneill and federico luebert for discussions on taxonomic problems, eberhard fischer for communicating the collections in tbi and zurab schewardnadse for help with translations from european journal of taxonomy 409: 1–75 (2018) 34 russian and georgian, and tianyun liu for help with translations from chinese language. we also thank genevieve k. walden, tilo henning, and the library staff of b, bm and k for their help to obtain the literature. the reviewers and alejandro quintanar sánchez are thanked for useful comments that helped to improve the manuscript. this research received support (visit to e collection) from the synthesys project 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(ed.) das pflanzenreich. iv 61 (heft 19). wilhelm engelmann, berlin. available from http://biodiversitylibrary.org/item/17309 [accessed 27 jun. 2016]. zapałowicz h. 1908. conspectus florae galiciae criticus. vol. 2. academiae litterarum cracoviensis, kraków. zernov a.s. 2002. определитель сосудистых растений севера российского причерноморья [manual of vascular plants in the north of the russian black sea coast]. kmk, moscow. zhao l. & chen d.y. 1988. flora liaoningica 1: 1988. liaoning science & technology publishing, shenyang. zohary d., hopf m. & weiss e. 2012. domestication of plants in the old world: the origin and spread of domesticated plants in southwest asia, europe, and the mediterranean basin. oxford university press, oxford. zuccarini j.g. 1833. ueber den gegenwärtigen zustand des königlichen herbariums. die bayerischen annalen 1833 (14): 101–103. manuscript received: 13 october 2016 manuscript accepted: 2 march 2017 published on: 28 february 2018 topic editor: koen martens desk editor: alejandro quintanar printed versions of all papers are also deposited in the libraries of the institutes that are members of the ejt consortium: muséum national d’histoire naturelle, paris, france; botanic garden meise, belgium; royal museum for central africa, tervuren, belgium; natural history museum, london, united kingdom; royal belgian institute of natural sciences, brussels, belgium; natural history museum of denmark, copenhagen, denmark; naturalis biodiversity center, leiden, the netherlands; museo nacional de ciencias naturales-csic, madrid, spain; real jardín botánico de madrid csic, madrid, spain. holstein n. et al., taxonomic checklist of corylus 45 http://biodiversitylibrary.org/item/17309 no taxon left behind? – a critical taxonomic checklist of carpinus and ostrya (coryloideae, betulaceae) norbert holstein 1,* & maximilian weigend 2 1,2 rheinische friedrich-wilhelms-universität bonn, bonn, nordrhein-westfalen, germany. * corresponding author: holstein@uni-bonn.de 2 email: mweigend@uni-bonn.de abstract. hornbeams (carpinus) and hop-hornbeams (ostrya) are trees or large shrubs from the northern hemisphere. currently, 43 species of carpinus (58 taxa including subdivisions) and 8 species of ostrya (9 taxa including sudivisions) are recognized. these are based on 175 (plus 16 latin basionyms of cultivars) and 21 legitimate basionyms, respectively. we present an updated checklist with publication details and type information for all accepted names and the vast majority of synonyms of carpinus and ostrya, including the designation of 54 lectotypes and two neotypes. cultivars are listed if validly described under the rules of the icn. furthermore, we consider carpinus hwai hu & w.c.cheng to be a synonym of carpinus fargesiana var. ovalifolia (h.j.p.winkl.) holstein & weigend comb. nov. during the course of our work, we found 30 legitimate basionyms of non-cultivars that have been consistently overlooked since their original descriptions, when compared with the latest checklists and floristic treatments. as regional floras are highly important for taxonomic practice, we investigated the number of overlooked names and found that 78 basionyms were omitted at least once in the eight regional treatments surveyed. more seriously, we found 4 basionyms of accepted species being overlooked in a major floristic treatment. keywords. herbarium digitization, taxon inventory, taxonomy, typification. holstein n. & weigend m. 2017. no taxon left behind? – a critical taxonomic checklist of carpinus and ostrya (coryloideae, betulaceae). european journal of taxonomy 375: 1‒52. https://doi.org/10.5852/ejt.2017.375 introduction in the ongoing discussions on the loss of biodiversity and taxonomy crisis, inventories of taxon names are a crucial step. floristic treatments and annotated checklists for countries, provinces or national parks are widely used as tools for biodiversity assessments, monitoring, and conservation (palmer et al. 1995; funk 2006). regional projects are very important, but regional taxonomic treatments may be of limited use when wide-spread taxa are concerned. the narrow focus of regional treatments carries the risk of overemphasizing local variability and repeatedly describing more wide-spread taxa (scotland & wortley 2003; michelangeli 2005). therefore, taxonomically comprehensive lists as a baseline for taxonomic work are required. this need is addressed by, e.g., the series the world checklist of selected plant families (wcsp 2016) or the plantlist (http://www.theplantlist.org), with names and types available from nomenclatural databases such as ipni (http://www.ipni.org) or tropicos (http://www.tropicos.org). european journal of taxonomy 375: 1–52 issn 2118-9773 https://doi.org/10.5852/ejt.2017.375 www.europeanjournaloftaxonomy.eu 2017 · holstein n. & weigend m. this work is licensed under a creative commons attribution 3.0 license. m o n o g r a p h 1 mailto:holstein%40uni-bonn.de?subject= mailto:mweigend%40uni-bonn.de?subject= https://doi.org/10.5852/ejt.2017.375 http://www.theplantlist.org http://www.ipni.org http://www.tropicos.org http://www.europeanjournaloftaxonomy.eu/index.php/ejt/index https://creativecommons.org/licenses/by/3.0/ the correct and valid application of taxon names (at the rank of family and below), however, is determined by designated taxonomic types (art. 7.1. icn; mcneill et al. 2012). therefore, lists of names are only the initial step, since it is the type specimen that anchors the identity of the taxon. obtaining type information, often deeply hidden in the historical literature, is a tedious process and the physical type specimens may be difficult to locate, but a correct and unequivocal typification is the crucial basis for disentangling taxonomic problems (gaston & mound 1993). only then, as a feedback mechanism, treatments will be improved not only by data density, but also by data quality. almost 20 years ago, the world checklist and bibliography of fagales (govaerts & frodin 1998) provided a baseline for the taxa of this order containing many species that constitute the temperate forests. however, critical taxonomic revisions are still scarce, with a revision of the genus betula l. as a noteworthy exception (ashburner et al. 2013). carpinus l. and ostrya scop. belong to betulaceae subfam. coryloideae hook.f. the species of the two genera are large shrubs or trees in forests nearly across the northern hemisphere. a few species are used for specialty timber or for landscaping and hedges, sometimes also as an ornamental, with carpinus betulus l. and a range of cultivars of that species being by far the most important taxon. carpinus and ostrya are sister to the chinese ostryopsis decne., a genus of three species (holstein & weigend 2016). while some studies find ostrya to be paraphyletic to carpinus (grimm & renner 2013), it is more often found nested in carpinus (yoo & wen 2002, 2007; li 2008). ostrya is easily differentiated morphologically by having its nuts enclosed in pale, papery, inflated tubular bracts with a small blunt distal opening. nuts of carpinus are more or less exposed, at most half-covered by a side wing of the one large abaxial bract. an additional, minute, caducous, adaxial scale bract can be found in some east asian species (carpinus sect. distegocarpus (siebold & zucc.) sarg.) currently, the world checklist of selected plant families (wcsp 2016) lists 42 accepted species of carpinus and 8 of ostrya, with both genera having a center of diversity in china. the revision of carpinus for china by hu (1933) is the latest critical taxonomic treatment on a larger geographical scale, while furlow (1987) worked on the four north american taxa. the latest global key for carpinus and ostrya (rushforth 1987) is an extension of the key from li’s chinese floral treatment (li 1979a). here, we amend the current lists of taxa of carpinus and ostrya with further names based on extensive literature and in silico research, compare the taxonomic status of the names in various floras, and present typification data as far as available. furthermore, we check the number of names not treated in the various treatments despite occurrences in the covered area to analyze the rate of overlooked taxa. material and methods we assembled an initial list from the world checklist of selected plant families (wcsp 2016) and complemented it with names from ipni, the plant list, tropicos and other sources. we took the acceptance status of world checklist of selected plant families as a baseline and compared the status and the basionyms of the corresponding synonyms with the recent literature (furlow 1987; chang et al. 2014a), and the appropriate floras, such as the flora of china (li & skvortsov 1999), flora of taiwan (liao 1996), flora sichuanica (editorial board of the flora sichuanica 2012), flora of japan (ohwi 1965), flora of north america (furlow 1997), and flora of the ussr (bobrov 1936). in order to calculate the number of overlooked basionyms, names relating to cultivars (hanging branches, leaf color and maculation, and leaf serration mutants) were excluded. illegitimate and invalid names were listed in the supplementary file as they are sometimes mentioned and may represent evolutionary significant entities, but they were not included in the calculations either. a name was tagged as “missing” for a specific publication if it was not cited, although it should have been covered as it occurs in the focus area of the publication. type information for the names was identified from the protologues. the following herbaria were searched for type specimens in situ: b, bm, br, e, goet, k, liv, m, manch, msb, w, and wu. european journal of taxonomy 375: 1–52 (2017) 2 www.europeanjournaloftaxonomy.eu/index.php/ejt/editor/downloadfile/500/2456 additional specimens were obtained via personal communication (dwc, le, tbi) or in silico: jstor plant science (http://plants.jstor.org/), chinese virtual herbarium (cdbi, ibk, ibsc, kun, lbg, n, nas, pe, wuk; http://www.cvh.org.cn/), hungaricana (bp; http://gallery.hungaricana.hu/en/), nhn naturalis (l, u, wag), virtuella herbariet (ld, s; http://www.herbarium-ume.se/virtuella_herbariet), and the homepages of the following herbaria: a, f, gh, hk, le, linn, mich, mo, ny, p, ph, tai, taif, and us. types of korean taxa in korean and japanese herbaria are cited from chang et al. (2014b). results we found 175 legitimate basionyms for carpinus (plus 16 for cultivars described under the rules of the icn, two of them not formally mentioned as cultivars) and 21 for ostrya. the list of non-cultivar basionyms with acceptance and taxonomic status, and the digital availability of at least one type specimen per basionym is found in the supplementary file. in total, 78 basionyms were overlooked in at least one of the ten treatments where they should have been covered, plus the recently described c. langaoensis (lu et al. 2017). we found 32 legitimate basionyms not included in the world checklist of selected plant families yet (wcsp 2016). the world checklist of fagales (govaerts & frodin 1998) missed 46 basionyms, plus three basionyms, c. chingiana, c. fargesiana var. tchouana and c. henryana var. chuana (yang 1997). the latter might, however, have been published between the copy deadline and the publication of the checklist. six relevant basionyms have been overlooked in the flora of ussr (bobrov 1936), ten in the flora of japan (ohwi 1965), two in the flora of taiwan (liao 1996), 34 in the flora europaea (tutin & walters 1993), three in the flora of north america (furlow 1997), 14 in the flora of china (li & skvortsov 1999, including c. chingiana, c. fargesiana var. tchouana and c. henryana var. chuana), one in the flora sichuanica (editorial board of the flora sichuanica 2012), and three in the checklist of korea (chang et al. 2014a). the four chinese species c. lipoensis, c. luochengensis, c. mengshanensis, and c. shimenensis are accepted in the world checklist (govaerts & frodin 1998), but not mentioned in the flora of china (li & skvortsov 1999). the acceptance status of 21 names in carpinus and one name in ostrya differs among the different treatments. five basionyms in carpinus and one in ostrya differ in the taxon to which the name is supposed to be synonymous only. one hundred and two taxa of carpinus and 16 of ostrya (including all accepted ones) had original or type material available online, most of them freely. however, almost all herbaria surveyed in situ for type specimens and all herbaria with non-type digitization programs harbored hitherto unrecognized type material to a considerable degree. complete digitization of herbaria, such as in the paris herbarium, naturalis or the china virtual herbarium, was most rewarding for this work, due to the manifold of unrecognized type material and the time-unlimited access. checklist of carpinus order fagales engl. family betulaceae gray subfamily coryloideae hook.f. tribe carpineae a.dc. carpinus l. species plantarum: 998 (linnaeus 1753). ‒ ostrya j.hill, the british herbal: 513 (hill 1757) [“1756”] nom. rejic. – c. sect. eucarpinus sarg., the silva of north america 9: 40 (sargent 1896) nom. inval. [art. 22.2, mcneill et al. (2012)]. – c. subg. eucarpinus (sarg.) nakai, the botanical magazine [tokyo] 29: 37 (nakai 1915) nom. inval. [art. 22.2, mcneill et al. (2012)]. – c. subg. carpinus (l.) hu, acta phytotaxonomica sinica 9: 282 (hu 1964). – c. ser. betulae hu, acta phytotaxonomica sinica 9: 282 (hu 1964), nom. inval. [art. 22.2, 22.6, mcneill et al. (2012)]. – c. subsect. carpinus holstein n. & weigend m., taxonomic checklist of carpinus and ostrya 3 http://plants.jstor.org/ http://www.cvh.org.cn/ http://gallery.hungaricana.hu/en/ http://www.herbarium-ume.se/virtuella_herbariet www.europeanjournaloftaxonomy.eu/index.php/ejt/editor/downloadfile/500/2456 (l.) p.c.li & s.h.cheng, flora reipublicae popularis sinicae 21 (2): 65 (li 1979a). – c. subsect. betulae (hu) c.j.wang, flora yunnanensis 5: 182 (1991) nom. inval. [art. 22.2, 22.6, mcneill et al. (2012)]. ‒ type species: c. betulus l., lectotypified by britton & brown (1913: 606). distegocarpus siebold & zucc., abhandlungen der mathematisch-physikalischen klasse der königlich bayerischen akademie der wissenschaften 4 (3): 226 (siebold & zuccharini 1846). – carpinus sect. distegocarpus (siebold & zucc.) sarg., the silva of north america 9: 40 (sargent 1896). – c. subg. distegocarpus (siebold & zucc.) nakai, the botanical magazine [tokyo] 29: 37 (nakai 1915). ‒ type species: distegocarpus carpinus siebold & zucc. (c. japonica blume). c. sect. brachyspicae nakai, the botanical magazine [tokyo] 29: 37 (nakai 1915). – type species: c. paxii h.j.p.winkl. c. sect. elongatae nakai, the botanical magazine [tokyo] 29: 37 (nakai 1915). – type species: not designated. c. ser. monbeigianae hu, acta phytotaxonomica sinica 9: 285 (hu 1964) [“monberigianae”]. – c. subsect. monbeigianae (hu) p.c.li, flora reipublicae popularis sinicae 21 (2): 70 (li 1979a). – type species: c. monbeigiana hand.-mazz. c. ser. polyneurae hu, acta phytotaxonomica sinica 9: 294 (hu 1964). – c. subsect. polyneurae (hu) p.c.li, flora reipublicae popularis sinicae 21 (2): 84 (li 1979a). – type species: c. polyneura franch. [art. 22.6, mcneill et al. (2012)]. c. ser. pubescentes hu, acta phytotaxonomica sinica 9: 289 (hu 1964). – type species: c. pubescens burkill [art. 22.6, mcneill et al. (2012)]. c. subg. distegocarpus ser. fangianae hu, acta phytotaxonomica sinica 9: 281 (hu 1964). – type species: c. fangiana hu. c. subg. distegocarpus ser. cordatae hu, acta phytotaxonomica sinica 9: 282 (hu 1964). – type species: c. cordata blume. 1. carpinus betulus l. species plantarum: 998 (linnaeus 1753). – original citation: “europe, canada.” – type: sweden, småland, s. coll., s.n. (lecto-, designated by browicz (1972: 3): linn! ex herb. linn. no. 1131.1). – additional original material: sweden, skâne, c. linnaeus s.n. (s09–40045). c. vulgaris mill., the gardeners dictionary ed. 8, no. 1 (miller 1768). – original citation: “in germania copiose, rarius in italia, in scania & inferiore parte smolandiae.” – type: dodoens (1616: 841) [figure]. – additional original material: sweden, småland, s. coll., s.n. (linn! ex herb. linn. no. 1131.1). s. loc., s. coll., s.n. (bm000647417! ex herb. cliff. no. 447). c. sepium lam., flore françoise 2: 212 (lamarck 1779) nom. illegit. superfl. – note: lamarck (1779) cites c. betulus l. (linnaeus 1763: 1416), which includes all citations for c. betulus l. (linnaeus 1753). c. compressus gilib., exercitia phytologica 2: 399 (gilibert 1792) nom. inval., opus utique oppr. c. ulmifolia salisb., prodromus stirpium in horto ad chapel allerton vigentium: 392 (salisbury 1796) nom. illegit. superfl. – note: salisbury (1796) cites c. betulus l. (linnaeus 1763: 1416), which includes all citations for c. betulus l. (linnaeus 1753). c. ulmoides gray, a natural arrangement of british plants 2: 245 (gray 1821) nom. illegit. superfl. – note: gray (1821) cites c. betulus l. (linnaeus 1763: 1416), which includes all citations for c. betulus l. (linnaeus 1753). c. austriaca m.serres, voyage dans le tyrol: 320 (serres 1823) nom. nud. – original citation: “les environs de brixen [...], je distinguai le carpinus austriaca, tout-à-fait étranger à la flore du nord du tyrol.” – associated material: s.d. (p06810513!). c. carpinizza host, flora austriaca 2: 626 (host 1831). – c. betulus [unranked] carpinizza (host) dippel, handbuch der laubholzkunde 2: 140 (dippel 1891) [“1892”]. – c. betulus var. carpinizza (host) c.k.schneid., illustriertes handbuch der laubholzkunde 1: 140 (schneider 1904). – european journal of taxonomy 375: 1–52 (2017) 4 c. betulus subsp. carpinizza (host) o.schwarz, mitteilungen der thüringischen botanischen gesellschaft 1: 96 (schwarz 1949). – original citation: “in sylvis transylvaniae. kitaibel. fl. majo. incolae transylvaniae, valachice loquentes, [...], et carpinizza vocant.” – type: not localized (not in kitaibel’s herbarium in bp or in w). – note: the application of this name is somewhat ambiguous. host mentions in the description the 3-lobed bracts with smooth margins of the side lobes, typical for c. betulus. neilreich (1861: 76), however, mentions that the name “carpinizza” is the vernacular walachian name for c. duinensis scop. (c. orientalis mill.) and blames kitaibel to whom host refers for mixing that up. then again, there is a c. orientalis specimen in kitaibel’s herbarium (xl/84!) with a label that this plant is called “kárpánicza”. it was therefore host, not kitaibel, misapplying the name. schneider (1904) refers the rank variety to neilreich (1861: 76), but neilreich did not publish that combination. therefore, schneider’s combination is the first at this rank. c. betulus var. integriloba spach, annales des sciences naturelles, botanique ser. 2 16: 252 (spach 1841a). – type: ménagerie, 1833, s. coll., s.n. (lecto-, designated here: p06810518!). – note: spach did not cite any locality or specific specimen but this collection from spach’s herbarium bears a note “nob.” (ours). c. betulus var. odontoloba spach, annales des sciences naturelles, botanique ser. 2 16: 252 (spach 1841a). – original citation: no detailed data given. – type: not localized. c. betulus f. serrata schur nom. nud. in sched., non beck (beck von mannagetta 1890). – associated material: romania, siebenbürgen, bergregion bis 3000’, 1848, p.j.f. schur 14164 (p06747296!). romania, siebenbürgen, hermannstadt [sibiu], 1848, p.j.f. schur 14166 (p06747370!). – note: this name is only known from two specimens, but schur published many taxa, and this name might have been published. should schur’s name have been validly published before 1904, then it would have priority over c. betulus var. serrata (beck) c.k.schneid. (c. betulus [unranked] serrata beck, basion.). c. intermedia wierz. ex rchb., icones florae germanicae et helveticae 12: 4, pl. dcxxxiii (reichenbach 1850). – original citation: “in bannatu, am wege von der sägemühle gegen kirscharusch bei steyersdorf [anina in caraș-severin], [p.p]. wierzbicki!”. – type: not localized. c. betulus var. provincialis gren. & godr., flore de france 3 (1): 121 (grenier & godron 1855). – c. provincialis j.gay nom. nud. in sched. – original citation: “forêts, bois et taillis.” – type: france [provence-alpes-côte d’azur: var], pennafort près draguignan, jun. 1830, j.h. perreymond 28 (lecto-, designated here: k!). – additional original material: france, draguignan, j.h. perreymond s.n. (k!); ibid., 1840, j.h. perreymond s.n. (p00543785!). – note: the specimen here designated as lectotype is mounted with the original note of gay from 1835, where he describes his new species c. provincialis. in a note from 1 dec. 1853 —also attached to the sheet— announcing a letter to grenier for the following day, gay rejects his species as a youthful folly (“péché de ma jeunesse”). nevertheless, grenier took up the name and published is as a variety in his flore de france. c. betulus var. obtusifolia g.kirchn., arboretum muscaviense: 667 (petzold & kirchner 1864). – c. betulus f. obtusifolia (g.kirchn.) h.j.p.winkl., das pflanzenreich iv 61 (heft 19): 30 (winkler 1904). – original citation: no detailed data given. – type: not localized. – note: petzold & kirchner accepted their term “spielart” in the sense of “varietas” as only rank below the species (1864: 41). although they discuss other rank-denoting terms, such as “unter-art” (subspecies) and “form” (forma) as being widely used, they only accept “spielarten”. hence, names below the species rank in petzold & kirchner (1864) are to be treated as varieties. c. subcordifolia schur, enumeratio plantarum transsilvaniae: 611 (schur 1866). – original citation: “in wäldern der kalkgebirge: auf dem ecsem-teteje bei sz. – domokos; auf dem kereszthegy bei remete. jul.” – type: not localized. – note: this name is missing in all relevant treatments and current databases surveyed. c. nervata dulac, flore du département des hautes-pyrénées: 141 (dulac 1867). – type: france, hautes-pyrénées, lascazères, j.j. corbin s.n. (syn-: bbf?); ibid., saint-sever-de-rustan, j.j. corbin s.n. (syn-: bbf?). holstein n. & weigend m., taxonomic checklist of carpinus and ostrya 5 c. ulmifolia st.-lag., études des fleurs ed. 8, 2: 742 (cariot 1889) nom. illegit. superfl. – note: saint-lager proposed this name to replace c. betulus l. c. betulus var. haynaldiana borbás, oesterreichische botanische zeitschrift 39 (6): 234 (braun 1889). – original citation: “im auwinkel [zugliget] bei ofen [buda].” – type: hungary [középmagyarország], in silvaticis ad zugliget (auwinkel) budae-pestini, 15 jul. 1888, v. de borbás s.n. (syn-: bm!); ibid., 27 jul. 1888, v. de borbás s.n. (syn-: e!). – note: this name is missing in all relevant treatments and current databases surveyed. c. betulus [unranked] serrata beck, flora von nieder-österreich: 266 (beck von mannagetta 1890). – c. betulus var. serrata (beck) c.k.schneid., illustriertes handbuch der laubholzkunde 1: 140 (schneider 1904). – c. betulus subsp. serrata (beck) o.schwarz, mitteilungen der thüringischen botanischen gesellschaft 1: 96 (schwarz 1949). – c. betulus f. serrata (beck) georgescu, flora republicii populare române 1: 195 (săvulescu 1952). – original citation: no detailed data given. – type: not localized. c. betulus var. typica koehne, deutsche dendrologie: 193 (koehne 1893) nom. inval. [art. 24.3; mcneill et al. (2012)]. c. betulus var. angustifolia błocki ex zapał., conspectus florae galiciae criticus 2: 9 (zapałowicz 1908). – original citation: “lesienice pod lwowem (błocki), strzyżów w jasielskiem (holzer).” – type: [ukraine, lviv] lesienice bei lemberg, 1886, b. błocki s.n. (syn-: not localized; isosyn-: b!, p06747360!, p06747390!, p06809764!, w1985–5965!); [poland, województwo podkarpackie] jasielski, strzyżów, holzer s.n. (syn-: not localized). c. betulus [unranked] angustifolia medw., vestnik tiflisskogo botanicheskogo sada 14: 26 (medwedew 1909) nom. illegit. (later homonym). – c. betulus var. angustifolia (medw.) radde-fom., trudy fizychno-matematychnogo viddilu. kiev [mémoires de la classe des sciences physiques et mathématiques de l’académie des sciences de l’ukraine] 15 (1): 80 (radde-fomina 1929) nom. illegit. (later homonym). – c. betulus f. angustifolia (medw.) radde-fom., trudy fizychnomatematychnogo viddilu. kiev [mémoires de la classe des sciences physiques et mathématiques de l’académie des sciences de l’ukraine] 15 (1): 80 (radde-fomina 1929) nom. inval. pro syn. – original citation: “найдена въ верхней аджаріи (ущ. р. аджарисъ-цкали въ батумской обл.) ло годерзскаго перевала и за нимъ (въ бассейнѣ куры, масальскій, герб. спб. бот. сада) а также въ терской области на хр. салатау, между с. чиркей и хонцагой, на высотѣ 3000’ (алекс., герб. ак. н.)” [found in upper adsharia (acharis-tskali river in batumi province), at goderdzi pass and beyond (in kura basin; massalsky, herb. of the st. petersburg botanical garden). in terek region, in salatau between czirkei and honzagoi, alt. 3000’ (alexeenko, herb. of the academy).]. – type: not localized. – note: medwedew differentiated three forms of c. betulus in the caucasus, but published his c. betulus [unranked] angustifolia explicitly as “var. nov.” nowadays, the use of the two different ranks in a non-successive order is to be interpreted as unranked (art. 37.8; mcneill et al. 2012). radde-fomina (1929: 80) accepted medwedew’s informal use of “form” and cited medwedew’s name in the rank forma and gave it the rank as variety afterward, while citing medwedew’s form in the synonymy. irrespective of this, medwedew’s and radde-fomina’s names are illegitimate due to the priority of zapałowicz’s name from 1908. c. betulus var. acuminata k.reinecke, mitteilungen des thüringischen botanischen vereins 28: 41 (reinecke 1911). – original citation: “im glacis der cyriaksburg b. e. [erfurt, germany].” – type: not localized. c. betulus var. parva radde-fom., trudy fizychno-matematychnogo viddilu. kiev [mémoires de la classe des sciences physiques et mathématiques de l’académie des sciences de l’ukraine] 15 (1): 55, tab. vi f, g, e (radde-fomina 1929). – original citation: “dagestan, agan-kale prope temirchan-schura [former name of buynaksk] 2. x. 1916. budaev [s.n.]!”. – type: not localized. c. betulus f. subacuta domin, flora čechoslovenica exsiccata no. 245 (1931) in sched. – type: czech republic, bohemia centralis, gregarie in valle radotinské údoli prope urbem praha, solo calcareo, altitudine circa 250 m a.s.l., 10 aug. 1931, k. domin and m. deyl s.n. (holo-: not localized; iso-: b10 european journal of taxonomy 375: 1–52 (2017) 6 0752246!, b!, b!, k!, msb-114979!). – note: this name is effectively published on specimen labels accompanying the types (art. 30.7; mcneill et al. 2012). c. betulus var. tuzsoni kárp., botanikai közlemények 34: 195 (kárpáti 1937). – original citation: “während einer excursion unter der leitung des herrn prof. dr. j. v. tuzson machten wir am jánoshegy bei budapest […].”. – type: not localized. – note: this is a variety with fissured bark. c. caucasica grossh., izvestiya azerbaidzhanskogo filiala [akademii nauk sssr] 1940 (5): 34 (grossheim 1940). – type: [russia] sala-tau, czirkei × honzagoi, ± 3000 ft, 28 jul. 1897, t. alexeenko s.n. (holo-: le). – note: the holotype might be the same specimen as the one cited as syntype of c. betulus [unranked] angustifolia medw. 1a. carpinus betulus ‘quercifolia’ c. betulus var. quercifolia hort. ex c.f.ludw. (ludwig 1783). – c. quercifolia (c.f.ludw.) desf., tableau de l’école de botanique: 213 (desfontaines 1804). – c. betulus f. quercifolia hort. ex k.koch, dendrologie 2, 2: 3 (koch 1873) nom. inval. pro syn. – c. betulus f. quercifolia (c.f.ludw.) c.k.schneid., illustriertes handbuch der laubholzkunde 1: 140 (schneider 1904). – c. betulus var. quercifolia (desf.) tzvelev, flora vostochnoĭ evropy 11: 91 (tzvelev 2004) nom. illegit. (later homonym). – original citation: not localized. – type: paris botanical garden [cult.], s. coll., s.n. (neo-, designated here: p06747373!). – note: the oak-leaved hornbeam is an early mentioned cultivar that is usually attributed to desfontaines (1804: 213). however, the mutation was known before 1783 when ludwig published an oak-leaved, c. betulus var. quercifolia. the short description by ludwig (1783) was actually the english name, under which it was supposedly sold in an english garden catalogue. ludwig (1783) cites four english catalogues in the preface and mentions that he did not translate any name himself. the name ‘quercifolia’ does not appear in the loddiges catalogues (loddiges 1779, 1783), while the remaining two could not be examined by the present authors. therefore, it might be possible that the name was published validly even before. as all treatments deal with the same taxon, art. 41.4 (mcneill et al. 2012), applies. the specimen chosen as neotype is likely to represent what was and still is understood under this name. the end of the left branch on the neotype has strong affinities to the cultivar ‘incisa’, showing that these two may not always be readily distinguished. already koch (1873) considered the two names as the same. however, for now, it deems the present authors best to keep these two names separate. c. betulus var. heterophylla hort. ex loudon, arboretum et fruticetum britannicum 3: 2005 (loudon 1838) nom. inval. pro syn. – note: this cultivar with an unusual leaf incision is sometimes regarded as the same as ‘incisa’, but in contrast to ‘incisa’ with acute lobes and shallow teeth, the ones in ‘quercifolia’ are rather obtuse and rounded (cf. nicholson 1883; schneider 1904). koch regarded c. betulus ‘quercifolia’ as synonymous to ‘incisa’ and published the epithet as c. betulus f. incisa (aiton) k.koch (c. betulus var. incisa aiton, basion.). 1b. carpinus betulus ‘incisa’ c. betulus var. incisa aiton, hortus kewensis 3: 362 (aiton 1789). – c. betulus f. incisa (aiton) k.koch, dendrologie 2, 2: 3 (koch 1873). – type: kew [cult.], oct. 1883, s. coll., s.n. (neo-, designated here: k! ex ligneous herb. mr. g. nicholson s.n.). – note: the specimen designated as neotype here is mounted together with another specimen marked as “c. betulus var. asplenifolia”. c. laciniata hort. ex g.nicholson, the garden (london 1871–1927) 24: 419 (nicholson 1883) nom. inval. pro syn. – note: this variety, better to be treated as a cultivar, has leaves with a lobulate margin, triangular lobes, acute apices, and a shallow dentation. holstein n. & weigend m., taxonomic checklist of carpinus and ostrya 7 1c. carpinus betulus ‘variegata’ c. betulus var. variegata lodd. ex loudon, arboretum et fruticetum britannicum 3: 2005 (loudon 1838). – c. betulus f. variegata (lodd. ex loudon) h.j.p.winkl., das pflanzenreich iv 61 (heft 19): 31 (winkler 1904). – original citation: no detailed data given. – type: not localized. – note: this is a cultivar with variegated leaves from loddiges’ plant catalogue (loddiges 1836). winkler cites neither loudon nor loddiges (1836) but dippel (1891: 140), and according to art. 41.4. (mcneill et al. 2012), his name should be considered as a new combination as he is dealing with the same cultivar. the color of the variegation is unknown, as loudon does not mention it. dippel (1891: 149) recognizes only a single variegated cultivar, irrespective from the color of the variegation, while schneider (1904) recognizes a forma albovariegata and a forma aureovariegata. 1d. carpinus betulus ‘heterophylla’ c. betulus var. heterophylla g.kirchn., arboretum muscaviense: 667 (petzold & kirchner 1864). – c. betulus f. heterophylla (g.kirchn.) k.koch, dendrologie 2 (2): 3 (koch 1873). – original citation: no detailed data given. – type: not localized. – note: kirchner gave the name “heterophylla” to plants that have both normal-shaped leaves and deeply serrated leaves, regardless whether the leaf serration is of the ‘incisa’ or ‘quercifolia’ type, which he regarded as synonymous, as did koch. as the mutation occurs spontaneously and may be fixed by cuttings, this name ‘heterophylla’ is complicated to use. kirchner refers to “bth. cat.”, meaning the catalogue of james booth and söhne nursery in klein-flottbek, hamburg, germany. as there is no reference to loudon, kirchner’s name cannot be regarded as a clear recombination of c. betulus var. heterophylla hort. ex loudon, which is an invalid name published as a synonym of c. betulus ‘quercifolia’. therefore, kirchner’s name is not illegitimate, but rather a later homonym. 1e. carpinus betulus ‘pendula’ c. [unranked] pendula massé, revue horticole ser. 4 2: 271 (massé 1853). – c. betulus var. pendula (massé) g.kirchn., arboretum muscaviense: 667 (petzold & kirchner 1864). – c. pendula (massé) k.koch, dendrologie 2 (2): 3 (koch 1873). – c. betulus f. pendula (massé) schelle, handbuch der laubholzbenennung: 48 (beissner et al. 1903). – original citation: no detailed data given. – type: not localized. – note: this is a cultivar with weak, overhanging branches. massé mentions a “carpinus pendula” but immediately called it a variety of the common hornbeam. therefore, the taxonomic status is unclear upon publication, and the name is to be treated as unranked. although neither koch nor beissner et al. (1903) explicitly refer to massé, the names are to be regarded as a new combination (art. 41.4; mcneill et al. 2012). 1f. carpinus betulus ‘purpurea’ c. betulus f. purpurea k.koch, dendrologie 2 (2): 3 (koch 1873). – c. betulus var. purpurea (k.koch) g.nicholson, the garden (london 1871–1927) 24: 419 (nicholson 1883). – original citation: no detailed data given. – type: not localized. – note: this is a cultivar with initially brownish-red leaves that turn greenish soon after. nicholson does not refer to koch, but intended to describe the same cultivar, therefore art. 41.4 (mcneill et al. 2012) applies. 1g. carpinus betulus ‘fastigiata’ c. betulus var. fastigiata hort. ex g.nicholson, the garden (london 1871–1927) 24: 419 (nicholson 1883). – c. betulus f. fastigiata (g.nicholson) schelle, handbuch der laubholzbenennung: 48 (beissner et al. 1903). – original citation: no detailed data given. – type: not localized. – note: the european journal of taxonomy 375: 1–52 (2017) 8 branches of this cultivar are supposed to be “more ascending and the habit altogether more erect.” beissner et al. (1903) do not refer to nicholson, but basically describe the same cultivar, therefore art. 41.4 (mcneill et al. 2012) applies. c. betulus [unranked] pyramidalis dippel, handbuch der laubholzkunde 2: 140 (dippel 1891) [“1892”] nom. illegit. superfl. – c. betulus f. pyramidalis (dippel) h.j.p.winkl., das pflanzenreich iv 61 (heft 19): 31 (winkler 1904) nom. illegit. superfl. – note: dippel (1891) and winkler (1904) both cite c. betulus ‘fastigiata’ as a synonym, therefore their names are superfluous. 1h. carpinus betulus ‘columnaris’ c. betulus [unranked] columnaris beissn., mitteilungen der deutschen dendrologischen gesellschaft 8: 136 (beissner 1899b). – c. betulus f. columnaris (beissn.) schelle, handbuch der laubholzbenennung: 48 (beissner et al. 1903). – c. betulus var. columnaris (beissn.) bean, trees and shrubs, hardy in the british isles 1: 294 (bean 1914). – original citation: “in the gardens of fürst lobkowitz [ferdinand zdenko von lobkowitz] at schloss eisenberg [jezeří castle, czech republic]”. – type: not localized. – note: according to beissner et al. (1903), this cultivar was reported for the first time in a garden catalogue of the späth’sche baumschule in berlin, germany, in 1891, but the present authors could not verify if that publication contains a valid description. although neither beissner et al. (1903) nor bean (1914) refer to beissner (1899b), they all refer to a cultivar with erect branches forming a slender crown, so their names can be considered as new combinations (art. 41.4.; mcneill et al. 2012). 1i. carpinus betulus ‘pyramidalis’ c. betulus var. pyramidalis hort. ex bean, trees and shrubs, hardy in the british isles 1: 294 (bean 1914). – original citation: “a fine specimen grows in the solferino square at rouen.”. – type: not localized. – note: this cultivar has erect branches, but not as slender as those of ‘columnaris’. bean does not refer to dippel (1891) or winkler (1904), whose names are superfluous. 1j. carpinus betulus ‘fastigiata cucullata’ c. betulus f. cucullata hort. ex h.j.p.winkl., das pflanzenreich iv 61 (heft 19): 31 (winkler 1904). – c. betulus fastigiata cucullata g.kirchn., arboretum muscaviense: 667 (petzold & kirchner 1864) nom. inval. [polynomen as it was not published hyphenated, cf. art. 20.3, ex.7, mcneill et al. (2012)]. – original citation: no detailed data given. – type: not localized. – note: this is a garden cultivar with pyramidal growth and hooded leaves. the cultivar name ‘fastigiata cucullata’ is the earliest available one (art. 21.6; brickell et al. 2009), although not valid under the botanical code (art. 23.6c; mcneill et al. 2012), under which it was independently published as c. betulus f. cucullata by winkler (1904). 1k. carpinus betulus ‘albovariegata’ c. betulus f. albovariegata c.k.schneid., illustriertes handbuch der laubholzkunde 1: 140 (schneider 1904). – original citation: no detailed data given. – type: not localized. – note: it was published as “albo-variegata”, corrected according to art. 60.9 (mcneill et al. 2012). 1l. carpinus betulus ‘aureovariegata’ c. betulus f. aureovariegata c.k.schneid., illustriertes handbuch der laubholzkunde 1: 140 (schneider 1904). – original citation: no detailed data given. – type: not localized. – note: it was published as “aureo-variegata”, corrected according to art. 60.9 (mcneill et al. 2012). holstein n. & weigend m., taxonomic checklist of carpinus and ostrya 9 1m. carpinus betulus ‘horizontalis’ c. betulus [unknown rank] horizontalis simon-louis, “preisverw., herbst” 1902–1903 (simonlouis 1902). – c. betulus f. horizontalis (simon-louis) c.k.schneid., illustriertes handbuch der laubholzkunde 1: 140 (schneider 1904) – c. betulus var. horizontalis (simon-louis) bean, trees and shrubs, hardy in the british isles 1: 294 (bean 1914). – original citation: no detailed data given [found by m. jouin on the simon-louis establishments near metz, france (bean 1914)]. – type: not localized. – note: this is a cultivar with a flat crown. the present authors do not have the simon-louis catalogue at hand. depending on whether or not it contains a description and a definite rank, schneider’s name would become valid when he provided both. if the publication by simonlouis constitutes a valid publication with a definite rank, either schneider’s or bean’s combination might become superfluous. 1n. carpinus betulus ‘asplenifolia’ c. betulus var. asplenifolia bean, trees and shrubs, hardy in the british isles 1: 294 (bean 1914). – original citation: no detailed data given. – type: not localized. – note: cultivar with incised leaves. the incision depth is described as “the primary teeth large enough to be termed lobes”. 2. carpinus caroliniana walter flora caroliniana: 236 (walter 1788). – original citation: no detailed data given. – type: not localized. c. ostryoides raf., the medical repository ser. 3 2: 333 (rafinesque 1811) nom. nud., non göppert (1855). – note: göppert’s c. ostryoides is a valid name for a leaf fossil without reference to rafinesque (1811). 2a. carpinus caroliniana walter subsp. caroliniana note: it was accepted as subspecies distinct from c. caroliniana subsp. virginiana (marshall) furlow in the flora of north america (furlow 1997). c. americana michx., flora boreali-americana 2: 201, pl. 7 (michaux 1803). – original citation: “a canadam ad floridam.” – original material: ameriques septentrionales, a. michaux s.n. (p06810890!); ibid., s. coll., s.n. (p06810902!, p06810903!). – note: carpinus americana is regarded as synonymous to c. caroliniana s. lat. in flora of north america (furlow 1997). 2b. carpinus caroliniana subsp. virginiana (marshall) furlow systematic botany 12: 429 (furlow 1987). – c. betulus var. virginiana marshall, arbustrum americanum: 25 (marshall 1785). – c. virginiana (marshall) sudworth, bulletin of the torrey botanical club 20: 43 (sudworth 1893) nom. illegit., non miller (1768). – c. caroliniana var. virginiana (marshall) fernald, rhodora 37: 425 (fernald 1935). – type: s. loc., 16 sep. 1906, r.w. woodward s.n. (neo, designated by furlow (1987: 429): nebc00078970!). – note: original material in marshall herbarium, but it was destroyed (furlow 1987). 3. carpinus chingiana c.c.yang distribution of the woody plants in sichuan [四川树木分布]: 661 (yang 1997). – type (according to ipni): china, sichuan, muli, 2600 m a.s.l., 3 aug. 1978, q.-s. zhao et al. [zhao zhenju, mu ke-hua, yang yabin/赵清盛, 牟克华, 杨亚滨] 6980 (holo-: sz00094298, sz00094299; iso-: cdbi0172181!, cdbi0172182!, cdbi0172183!). – note: this name is accepted in the flora sichuanica (editorial board of the flora sichuanica 2012), but neither listed in the world checklist (govaerts & frodin european journal of taxonomy 375: 1–52 (2017) 10 1998) nor in the flora of china (li & skvortsov 1999). the cdbi specimens were labeled by yang as “carpinus chingii”, and the specimen cdbi0172182 bears an annotation slip by yang with the unpublished name “carpinus muliensis”. 4. carpinus chuniana hu journal of the arnold arboretum 13: 334 (hu 1932). – type: china, kwangtung [guangdong], lokchang, 31 may 1929, c.l. tso [左景烈] 20872 (holo-: pe?; iso-: a00033751!, ibsc0001142, ibsc0001143, ibsc0001144, ny00253858!). 5. carpinus cordata blume museum botanicum lugduno-batavum 1 (20): 309 (blume 1851). – original citation: “fisibami japonice. in japoniâ a cl. viro von siebold detecta.” – type: japan, in japoniâ, s. coll., s.n. (lecto-, designated here: l0040896! ex herb. siebold s.n.; isolecto-?: l0103212). 5a. carpinus cordata blume var. cordata c. erosa blume, museum botanicum lugduno-batavum 1 (20): 308 (blume 1851). – original citation: “in japoniâ.” – type: japan, s. coll. (lecto-, designated here: l0040895! ex herb. siebold s.n.). c. erosa var. microcarpa hayashi, 林業試験場研究報告 [research report of the forestry experiment station] 57: 153 (hayashi 1952). – type: japan, ezo [hokkaido], prov. hiyama, mt. gamushi, 8 aug. 1952, y. hayashi s.n. (holo-: tfm). c. erosa var. velutina hayashi, 林業試験場研究報告 [research report of the forestry experiment station] 57: 153 (hayashi 1955). – type: japan, hondo, prov. rikuchiu, yamagata in kuji, 15 aug. 1952, n. karizumi s.n. (holo-: tfm). ostrya mandshurica budischtschew ex trautv., trudy imperatorskago s.-peterburgskago botaniceskago sada 9: 166 (trautvetter 1884). – type: not localized, likely in le. c. cordata var. pseudojaponica h.j.p.winkl., botanische jahrbücher für systematik, pflanzengeschichte und pflanzengeographie 50 (suppl.): 490 (winkler 1914). – c. erosa var. pseudojaponica miyabe & tatewaki, bulletin of the kyushu university forests 21: 48 (tatewaki 1953) nom. inval. – c. erosa var. pseudojaponica (h.j.p.winkl.) miyabe & kudo ex horikawa, acta phytotaxonomica et geobotanica 15 (1): 13 (horikawa 1953) nom. inval. pro syn. – type: japan, insel yesso, mororan, sep. 1904, u.j. faurie 5775b (holo-: b†) [on u.j. faurie 5775 in b marked with b]. – note: this name is not mentioned in the world checklist (govaerts & frodin 1998) or in the wcsp (2016), but it is synonymized under c. cordata in the flora of japan (ohwi 1965). tatewaki published the combination without reference to older literature or an own description, it is therefore invalid (art. 41.3; mcneill et al. 2012). horikawa referred to the combination as c. erosa var. pseudojaponica (h.j.p.winkl.) miyabe & kudo in synonymy only. the combination may have been published in icones of the essential forest trees of hokkaido (miyabe et al. 1920–1932) though, but the present authors did not have access for verification c. cordata var. faurieana h.j.p.winkl., botanische jahrbücher für systematik, pflanzengeschichte und pflanzengeographie 50 (suppl.): 489 (winkler 1914). – type: japan, insel yesso, sapporo, jun. 1891, u.j. faurie 7111 (lecto-, designated here: p06747141!). – additional type material: japan, iwanai, sep. 1904, u.j. faurie 5777 (syn-: a00033752!, bm000580426!). – note: this name is not mentioned in the world checklist (govaerts & frodin 1998) or the wcsp (2016). based on the subglabrous leaves and the locus classicus being on hokkaido, it may well represent a distinct taxon. c. cordata var. robusta h.j.p.winkl., botanische jahrbücher für systematik, pflanzengeschichte und pflanzengeographie 50 (suppl.): 489 (winkler 1914). – type: japan, insel yesso, mororan, sep. 1904, u.j. faurie 5775 (syn-: bm000580424!, e!). – note: this name is not mentioned in the world checklist (govaerts & frodin 1998) or the wcsp (2016). as for c. cordata var. faurieana holstein n. & weigend m., taxonomic checklist of carpinus and ostrya 11 h.j.p.winkl. it is unlikely that this name is synonymous with the other accepted varieties because of the relatively subglabrous nature and the locus classicus being on hokkaido instead of the asian mainland. c. cordata var. winkleri radde-fom., trudy fizychno-matematychnogo viddilu. kiev [mémoires de la classe des sciences physiques et mathématiques de l’académie des sciences de l’ukraine] 15 (1): 70 (radde-fomina 1929). – original citation: “wladiwostok 1864 budischczev [s.n.]! «suchaja padi» in viciniis pag. rakovka 2. ix. 1915, neczajeva et gordeev! orlinoe gnieso ad marginem sylvae 17. v. 1907. palczevsky [s.n.]! prov primorskaja, flum orlinoje gniesdo ix. 1902 palczevsky [s.n.]!”. – type: not localized. c. cordata var. brevistachyus s.l.tung, bulletin of botanical research, harbin 1 (1‒2): 139 (tung 1981). – type: china, jilin, linjiang [临江], 700 m a.s.l., 22 jun. 1962, y.l. chou [周以良] 60007 (holo-: nefi). 5b. carpinus cordata var. chinensis franch. journal de botanique [morot] 13: 202 (franchet 1899). – c. cordata f. chinensis (franch.) nakai, the journal of the college of science, imperial university of tokyo, japan 31: 205 (nakai 1911). – c. chinensis (franch.) c.pei, botanical bulletin of academia sinica 2: 223 (pei 1948). – type: china, sutchuen [sichuan], in ditione tchen keou tin [chengkou county], 1400 m a.s.l., p.g. (reverend père) farges 14 (lecto-, designated here: p06747063!; isolecto-: l!, p06747058!, p06747059!, p06747060!). – note: carpinus cordata var. chinensis is accepted in the flora of china (li & skvortsov 1999), but it is segregated as c. chinensis in the flora sichuanica (editorial board of the flora sichuanica 2012). 5c. carpinus cordata var. mollis (rehder) w.c.cheng ex y.chen illustrated manual of chinese trees and shrubs: 163 (chen 1937). – c. mollis rehder, journal of the arnold arboretum 11: 154 (rehder 1930). – type: china, sichuan, sungpan hsien, side of stream, 17 aug. 1928, w.p. fang [方文培] 4245 (holo-: a00033760!; iso-: e00275495!, e00275496!, k000859923!, nas00070300, ny00253865!, p01903243!, pe00021928!, pe00021929!). – note: carpinus cordata var. mollis is accepted in flora of china (li & skvortsov 1999), but it is regarded as a synonym of c. chinensis in the flora sichuanica (editorial board of the flora sichuanica 2012). 6. carpinus dayongina k.w.liu & q.z.lin bulletin of botanical research, harbin 6 (2): 143 (liu & lin 1986). – type: china, hunan, dayong [now zhangjiajie], tianmen shan, 1100 m a.s.l., 28 jul. 1985, k.w. liu [liu ke-wang] 33359 (holo-: csfi; iso-: pe). – additional type material: ibid., 25 aug. 1984, y.t. xiao [xiao yu-tan] 40700 (para-: not localized). – note: the epithet is sometimes cited as “dayongiana” (govaerts & frodin 1998; wcsp 2016), but the authors consistently use “dayongina” (liu & lin 1986: 143, 145), and correctly so (art. 60 rec. 60d, mcneill et al. 2012). 7. carpinus faginea lindl. plantae asiaticae rariores 2: 50 (wallich 1831). – type: india [uttar pradesh], gurwal, kamrup [kamroop] s.n. (holo-: cge?). c. latibracteata lindl. nom. nud. in sched. – note: this name is only known from one specimen. it also bears the notes “wallich h.i.” and “lindley 1830”, however, neither wallich nor lindley were in kumaon in 1830. likely, it is material from wallich’s collection (by robert blinkworth?), and the name was written onto the specimen by lindley in 1830 [india, kamaon [kumaon], s. coll., s.n. (k!)], but never published. european journal of taxonomy 375: 1–52 (2017) 12 8. carpinus fangiana hu journal of the arnold arboretum 10: 154 (hu 1929). – type: china, sichuan, nanchuan hsien, in thickets, 1500–1800 m a.s.l., 1 jun. 1928, w.p. fang [方文培] 1351 (lecto-, designated by lin et al. (2007: 1248): pe00021890; isolecto-: a00033753!, e00275500!, e00275501!, ibsc0367917, k000859928!, ny00253860!, p06747103!, pe00021891!, pe00021968!). – additional type material: ibid., w.p. fang 1352 (para-: e!, ibsc0367916, k!, p06747102! pe00021888!, pe00021969!). – note: accepted in the flora of china (li & skvortsov 1999), in the world checklist (govaerts & frodin 1998), and in the flora sichuanica (editorial board of the flora sichuanica 2012). c. wilsoniana hu, journal of the arnold arboretum 10: 154 (hu 1929). – type: china, sichuan, mt. omei, in thickets, 1675–1800 m a.s.l., 9 aug. 1928, w.p. fang 2685 (lecto-, designated here: pe00021889; isolecto-: a00033772!, e00275499!, e00275528!, ibsc0367915, k000859919!, nas00070302!, ny00253869!, p06810764!, pe00021970!). 9. carpinus fargesiana h.j.p.winkl. botanische jahrbücher für systematik, pflanzengeschichte und pflanzengeographie 50 (suppl.): 507 (winkler 1914). – type: china, sutchuen [sichuan], in ditione tchen keou tin [chengkou county] [as “carpinus yedoensis”], p.g. (reverend père) farges s.n. (lecto-, designated here: p06811425!; isolecto-: p06811420!). – additional type material: china, sichuan, a. henry 7053 (syn-: not localized). 9a. carpinus fargesiana h.j.p.winkl. var. fargesiana note: it was accepted with c. fargesiana var. hwai (see c. fargesiana var. ovalifolia) as another variety of c. fargesiana in the flora of china (li & skvortsov 1999). c. daginensis hu, acta phytotaxonomica sinica 9: 293 (hu 1964). – type: china, sichuan, dagin hsien, aning, 2900 m a.s.l., 9 may 1958, x. li [li hsien, 李馨] 77351 (holo-: pe00021885!; iso-: nas00070301, wuk0252019). – note: carpinus daginensis is considered as a synonym of c. fargesiana var. fargesiana in the flora of china (li & skvortsov 1999) and in the flora sichuanica (editorial board of the flora sichuanica 2012: 35). c. fargesiana var. tchouana c.c.yang, distribution of the woody plants in sichuan [四川树木分布]: 662 (yang 1997). – type: not localized. – note: this name is only accepted and mentioned in the flora sichuanica (editorial board of the flora sichuanica 2012), while it is regarded as a synonym of c. fargesiana var. fargesiana in the wcsp (2016). the present authors did not see the protologue. 9b. carpinus fargesiana var. ovalifolia (h.j.p.winkl.) holstein & weigend comb. nov. urn:lsid:ipni.org:names:77167104-1 basionym: c. turczaninovii var. ovalifolia h.j.p.winkl., botanische jahrbücher für systematik, pflanzengeschichte und pflanzengeographie 50 (suppl.): 505 (winkler 1914). – c. hwai hu & w.c.cheng, bulletin of the fan memorial institute of biology n.s. 1: 148 (hu 1948a). – c. fargesiana var. hwai (hu & w.c.cheng) p.c.li, flora reipublicae popularis sinicae 21 (2): 82 (li 1979a) nom. illegit. superfl. – type: china, sichuan, no. [north] wushan, a. henry 7020 (lecto-, designated here: p06811189!; isolecto-: bm!, gh00112538!, k000859934!, k000859935!). – additional type material: china, “n huan tou shan”, g. giraldi s.n. (syn-: not localized); ibid., sichuan, s wushan, a. henry 7019 (syn-: k000859936!); ibid., distr. tchen-keou-tin, p.g. [réverend père] farges 1273 (syn-: p06811190!, p06811191!, p06811192!, p06811194!, p06811503!). ibid., p.g. [réverend père] farges s.n. (syn-: not localized); ibid., sichuan [locality unreadable], 7500 ft, may 1903, e.h. wilson 4489 (syn-, mentioned in appendix to protologue: bm!, ibsc0368215, k000859932!, holstein n. & weigend m., taxonomic checklist of carpinus and ostrya 13 http://www.ipni.org/urn:lsid:ipni.org:names:77167104-1 k000859933!). – note: hu (1948b) cited winkler’s variety for their new species as the only element (thus, c. hwai is a new name, but only legitimate in the rank of a species). when treating this taxon as variety, however, again the name “ovalifolia” has priority, which li apparently overlooked. the lectotype material was mentioned in the appendix of the protologue, and is thus also original material. it bears an annotation of winkler, and duplicates are distributed in several herbaria, so it seems a good choice for a lectotypification. this taxon is recognized both in the flora of china (li & skvortsov 1999) and in the flora sichuanica (editorial board of the flora sichuanica 2012: 35). 10. carpinus firmifolia (h.j.p.winkl.) hu bulletin of the fan memorial institute of biology n.s. 1 (2): 144 (hu 1948a). – c. turczaninovii var. firmifolia h.j.p.winkl., botanische jahrbücher für systematik, pflanzengeschichte und pflanzengeographie 50 (suppl.): 505 (winkler 1914). – c. pubescens var. firmifolia (h.j.p.winkl.) hu ex p.c.li, flora reipublicae popularis sinicae 21 (2): 80 (li 1979a). – type: china, kui-tscheu [guizhou], ma-jo, fr, sep. 1908, j. cavalerie 3135 (holo-: e!; iso-: p04815672!). – note: accepted as c. firmifolia in the flora of china (li & skvortsov 1999), but listed as a variety in the world checklist of fagales (govaerts & frodin 1998). c. pubescens var. bigiehensis hu, acta phytotaxonomica sinica 9: 290 (hu 1964). – type: china, kweichow [guizhou], bigieh hsien, lin-ku’gie-wo, in open woods on rocky cliffs, 1450 m a.s.l., 13 sep. 1957, p.h. yü [yü pin-hwa, 禹平华] 810 (holo-: pe; iso-: ibsc0368211, kun0590865!, lbg00053917, wuk206962). – note: it was listed as a synonym of c. firmifolia in the flora of china (li & skvortsov 1999). 11. carpinus hebestroma yamam. supplementa iconum plantarum formosarum 5: 14, fig. 4 (yamamoto 1932). – type: r. china, taiwan, inter shinjio et batakan, 26 apr. 1917, s. coll. [y. yamamoto?], s.n. (lecto-, designated here: tai [118773]!). – additional type material: r. china, taiwan, inter batakan et naitaroko, apr. 1917, y. yamamoto s.n. (syn-: not localized). 12. carpinus henryana (h.j.p.winkl.) h.j.p.winkl. botanische jahrbücher für systematik, pflanzengeschichte und pflanzengeographie 50 (suppl.): 507 (winkler 1914). – c. tschonoskii var. henryana h.j.p.winkl., das pflanzenreich iv 61 (heft 19): 36 (winkler 1904). – c. hupeana var. henryana (h.j.p.winkl.) p.c.li, flora reipublicae popularis sinicae 21 (2): 83 (li 1979a) nom. illegit. – type: china, sichuan, a. henry 7063 (holo-: b†; lecto-, designated here: k00859926!; isolecto-: bm!, e00275504!, gh00033755!, p06811429!). 12a. carpinus henryana (h.j.p.winkl.) h.j.p.winkl. var. henryana c. henryana var. chuana c.c.yang, distribution of the woody plants in sichuan [四川树木分布]: 661 (yang 1997). – type: not localized. – note: this name is only accepted and mentioned in the flora sichuanica (editorial board of the flora sichuanica 2012), while it is regarded as a synonym of c. henryana var. henryana in the wcsp (2016). the present authors did not see the protologue. 12b. carpinus henryana var. oblongifolia (hu) rushforth plantsman 8: 249 (rushforth 1987). – c. turczaninovii var. oblongifolia hu, sunyatsenia 1 (2‒3): 115 (hu 1933). – c. oblongifolia (hu) hu & w.c.cheng, bulletin of the fan memorial institute of biology n.s. 1: 146 (hu 1948a). – type: china, kiangsu [jiangsu], poa-hwa shan, 450 m a.s.l., 23 european journal of taxonomy 375: 1–52 (2017) 14 jul. 1925, a.n. steward 1476 (holo-: pe00021938!). – note: this taxon is considered as a distinct species in the flora of china (li & skvortsov 1999). c. huana w.c.cheng, contributions from the biological laboratory of the science society of china, botanical series 9: 68 (cheng 1933). – type: china, zhejiang, [lin’an city, west tianmushan], 21 aug. 1924, w.c. cheng 5161 (holo-: pe00021905!). – additional type material: ibid., 15 aug. 1931, k. ling 3519 (para-: not localized). – note: carpinus huana is regarded as a synonym of c. hupeana in the flora of china (li & skvortsov 1999). c. hupeana hu, sunyatsenia 1 (2‒3): 118 (hu 1933). – type: china, hupeh, liang sung kou, 1250 m a.s.l., 9 sep. 1922, w.y. chun et al. 4173 (holo-: pe00021898!). – note: carpinus hupeana is accepted as distinct species in flora of china (li & skvortsov 1999) with c. funiushanensis, c. huana and c. longipes as synonyms. c. longipes hu, acta phytotaxonomica sinica 9: 291 (hu 1964). – type: china, kiangsi [jiangsi], tasiping shan, shih-cha-pu, 15 jul. 1949, y.k. hsiung [hsi un yao-kuo, 熊耀国] 5305 (lecto-, designated by lin et al. (2007: 1248): pe00021917!; isolecto-: pe00021915!, pe00021916!). – note: carpinus longipes is regarded as a synonym of a distinct c. hupeana in the flora of china (li & skvortsov 1999). c. funiushanensis p.c.kuo, flora tsinlingensis 1 (2): 601 (instituto botanico boreali-occidentali academiae sinicae edita 1974). – type: china, henan, 卢氏, 淇河北面山坡 [lu-shih hsien], 920 m a.s.l., 14 sep. 1958, c.c. fu [j.q. fu, 傅竞秋] 656 (holo-: wuk0407353, wuk0114527; iso-: ibsc0367962, nas00070304). – note: carpinus funiushanensis is regarded as a synonym of a distinct c. hupeana in the flora of china (li & skvortsov 1999). kuo did not indicate a herbarium for the holotype. as the flora tsinlingensis is mainly based on material from wuk, the specimens there ought to be regarded as holoand isotype respectively or as one holotype mounted on two sheets if indicated as one gathering. the specimens from other herbaria are considered as duplicates. 12c. carpinus henryana var. simplicidentata (hu) rushforth plantsman 8: 249 (rushforth 1987). – c. simplicidentata hu, bulletin of the fan memorial institute of biology n.s. 1: 143 (hu 1948a). – c. hupeana var. simplicidentata (hu) p.c.li, flora reipublicae popularis sinicae 21 (2): 83 (li 1979a). – type: china, hubei, near chin tai po, 2150 m a.s.l., 2 sep. 1922, w.y. chun 4175 (lecto-, designated by lin et al. (2007: 1249): pe00021955!; isolecto-: pe00021953!, pe00021954!). – additional type original material: china, suon nai ook, 2150 m a.s.l., 2 sep. 1922, w.y. chun 4174 (syn-: not localized). – note: this name is treated in flora of china (li & skvortsov 1999) as a synonym of c. stipulata (listed as synonym of c. turczaninovii here). 13. carpinus insularis n.h.xia, k.s.pang & y.h.tong journal of tropical and subtropical botany 22 (2): 121 (tong et al. 2014). – type: china, hong kong, violet hill, 22°14´25.30´´ n, 114°11´55.04´´ e, 190 m a.s.l., 21 aug. 1913, k.y. tam s.n. (holo-: ibsc0770924; iso-: hk0043873). – note: according to the original authors c. insularis has similarities to c. hebestroma and c. polyneura. 14. carpinus japonica blume museum botanicum lugduno-batavum 1 (20): 308 (blume 1851) – distegocarpus carpinus siebold & zucc., abhandlungen der mathematisch-physikalischen klasse der königlich bayerischen akademie der wissenschaften 4 (3): 227 (siebold & zuccharini 1846). – distegocarpus carpinoidis siebold & zucc., abhandlungen der mathematisch-physikalischen klasse der königlich bayerischen akademie der wissenschaften 4 (3): 240 (siebold & zuccharini 1846) orth. var. (of distegocarpus carpinus). – carpinus carpinus (siebold & zucc.) sarg., garden and forest 6: 364 (sargent 1893) nom. inval. holstein n. & weigend m., taxonomic checklist of carpinus and ostrya 15 (tautonym). – c. carpinoides (siebold & zucc.) makino, the botanical magazine [tokyo] 26: 391 (makino 1912). – c. distegocarpus koidz., the botanical magazine [tokyo] 27: 144 (koidzumi 1913) nom. illegit. superfl. – type: japan, s. loc., h. bürger s.n. (lecto-, designated by akiyama et al. (2013: 349): m-0120911!) – additional original material: japan, s. loc., h. bürger s.n. (m0120912!, m-0120914!, m-0120915!, m-0120916!, m-0120917!); ibid., p.f. von siebold et al. s.n. (br!, br!, gh00033756!, k000859947!, k000859948!, l0175945!, p06811392?!). – original material?: s. loc. [via java], s. coll., s.n. (u1155346!). c. japonica var. caudata h.j.p.winkl., botanische jahrbücher für systematik, pflanzengeschichte und pflanzengeographie 50 (suppl.): 488 (winkler 1914). – type: japan, insel nippon [honshu], in den ontake-bergen, aug. 1905, u.j. faurie 6641 (lecto-, designated here: bm!). c. japonica var. cordifolia h.j.p.winkl., das pflanzenreich iv 61 (heft 19): 26 (winkler 1904). – c. carpinoides var. cordifolia (h.j.p.winkl.) makino, the botanical magazine [tokyo] 26: 391 (makino 1912) nom. illegit. – type: japan, central japan [honshu], im nikko-gebirge, w.f.k. dönitz s.n. (holo-: b†). c. japonica var. pleioneura h.j.p.winkl., botanische jahrbücher für systematik, pflanzengeschichte und pflanzengeographie 50 (suppl.): 488 (winkler 1914). – type: [japan] nikko, prov. schimotsuke, sep. 1887, s. coll., s.n. (lecto-, designated here: p06811395!) – additional type material: japan, miyanoschta, o. warburg 7756 (syn-: b†?); ibid., yokohama, may and oct. 1862, c.j. maximowicz s.n. (syn-: p06811393!). 15. carpinus kawakamii hayata icones plantarum formosanarum 3: 175, pl. xxxiiib (hayata 1913). – type: r. china, taiwan, monte morrison [yushan], oct. 1907, t. kawakami and u. mori 1998 (holo-: ti!). – note: the type cited here is the direct basis for the drawing in plate xxxiiib, and can therefore be interpreted as the holotype. 15a. carpinus kawakamii hayata var. kawakamii note: accepted in the flora of china (li & skvortsov 1999) with a distinct c. kawakamii var. minutiserrata (hayata) s.s.ying, whereas liao (1996) does not accept an infraspecific taxon in the flora of taiwan. c. hogoensis hayata, icones plantarum formosanarum 6: 62 (hayata 1916). – type: r. china, taiwan, musha [wu sha], hōgō [chunyang], 3500 ft, apr. 1916, b. hayata s.n. (holo-: not localized; iso-: taif!). – note: carpinus hogoensis is regarded as a synonym of c. kawakamii var. kawakamii in the flora of china (li & skvortsov 1999). c. sekii yamam., supplementa iconum plantarum formosarum 5: 12 (yamamoto 1932). – type: r. china, taiwan, in monte daibusan, 3000 ft, 24 may 1918, e. matsuda s.n. (holo-: tai [118775]!). – note: carpinus sekii is regarded as a synonym of c. kawakamii var. kawakamii in the flora of china (li & skvortsov 1999). c. auriculifera hayata nom. nud. – note: c. auriculifera is only known from the specimen collected in r. china (ariko-banti, 1 feb. 1917, e. matsuda s.n., tai [035707]!) and is marked as “type”. it might have been published, but the protologue was not localized. the specimen, however, clearly belongs to c. kawakamii. 15b. carpinus kawakamii var. minutiserrata (hayata) s.s.ying coloured illustrated flora of taiwan 3: 271 (ying 1988). – c. minutiserrata hayata, icones plantarum formosanarum 3: 177 (hayata 1913). – type: r. china, taiwan, tandaisha, apr. 1910, u. mori s.n. (lecto-, designated here: ti!; isolecto-: taif007296!, taif007300!). – note: liao (1996) european journal of taxonomy 375: 1–52 (2017) 16 synonymizes this name in the flora of taiwan under c. kawakamii, while it is accepted as separate species in the flora of china (li & skvortsov 1999). 16. carpinus kweichowensis hu sinensia 2: 79, fig. 1 (hu 1931). – type: china, guizhou, chengfeng hsien, pa-na, si-mi-yao, mixed woods, 24 oct. 1930, y. tsiang [蒋英] 4406 (lecto-, designated here: nas00070306!; isolecto-: a00033757!, bm!, e00275503!, k000859925!, n000053496, nas00070305!, ny00253862!, pe00021906!, pe00021908!, us00089310!). – note: the specimen selected here as lectotype is annotated as holotype, but it was not published as such. it is also the specimen depicted in figure 1 in hu (1931). c. austroyunnanensis hu, bulletin of the fan memorial institute of biology n.s. 1 (3): 213 (hu 1949). – type: china, yunnan province, xichou, hsin cheih, 24 oct. 1947, k.m. feng 12609 [erroneously published as 13609] (lecto-, designated here: pe00020175!; isolecto-: pe00818041!, wuk0207172). – note: the type citation of c. austroyunnanensis is erroneous, as k.m. feng 13609 (pe00792657) was collected the 23 nov. 1947 and is a morinda species, while k.m. feng 12609 was annotated by hu as c. austroyunnanensis. 17. carpinus langaoensis z.qiang lu & j.quan liu phytotaxa 295 (2): 191 (lu et al. 2017). – type: china, shaanxi, langao county, hengxi township, 108°48´ e, 32°08´ n, 1170 m a.s.l., 28 jul. 2016, z.q. lu 2016lzq029 (holo-: lzu; iso-: lzu). 18. carpinus laxiflora (siebold & zucc.) blume museum botanicum lugduno-batavum 1 (20): 309 (blume 1851). – distegocarpus laxiflora siebold & zucc., abhandlungen der mathematisch-physikalischen klasse der königlich bayerischen akademie der wissenschaften 4 (3): 228 (siebold & zuccharini 1846). – type: japan, s. loc., h. bürger s.n. (lecto-, designated by akiyama et al. (2013: 349): m-0120909!). – additional original material: japan, s. loc., h. bürger s.n. (m-0120907!, m-0120908!); ibid., s. coll. (l0175947! ex herb. p.f. von siebold s.n., l0175948! ex herb. p.f. von siebold s.n.); ibid., s. coll. (k000859944!, p06811472!, p06811473!, p06811475!). c. laxiflora f. lacera hayashi, journal of geobotany [北陸の植物] 22: 4 (hayashi 1974). – type: not localized. c. laxiflora var. chartacea h.lév., bulletin de la société botanique de france 51: 424 (léveillé 1904). – original citation: without collecting data. – type: not localized. c. laxiflora var. pendula miyoshi, report of the national monument investigation. plants [天然紀念 物調査報告 植物之部 第 8輯] 8: 45 (department of the interior 1928). – c. laxiflora f. pendula (miyoshi) sugim., new keys to japanese trees [日本樹木総検索誌]: 117, 458 (sugimoto 1961). – original citation: “ein einziges exemplar im grundstück von jojuin tempel, tochigi präfektur.” [a single exemplar on the area of the jojuin temple, tochigi prefecture”; 成就院 in niregimachi, kanuma district]. – type: not localized. – note: this is tree with hanging branches, better considered as a cultivar. the tree (most likely a clone) still existed in 2013. c. laxiflora var. macrophylla nakai, the botanical magazine [tokyo] 45: 112 (nakai 1931). – c. laxiflora f. macrophylla (nakai) w.t.lee, lineamenta florae koreae: 165 (lee 1996). – type: south korea [gyeonggi-do], in silvis kôryô prov. keiki, gwangreung, 3 sep. 1930, t. nakai 13201 (holo-: ti). c. laxiflora var. obtusisquama koidz., acta phytotaxonomica et geobotanica 9 (2): 73 (koidzumi 1940). – type: south korea, chiisan [mt. jiri-san], kokiri, 11 jun. 1935, g.s. koidzumi s.n. (holo-: kyo). – note: this name is mentioned neither in the world checklist (govaerts & frodin 1998) nor in the checklist of korea (chang et al. 2014a) nor in the wcsp (2016), synonymization unclear. holstein n. & weigend m., taxonomic checklist of carpinus and ostrya 17 c. laxiflora var. longispica uyeki, suigen gakuho 41: 9 (uyeki & lee 1924). – type: south korea, chyolla australis, mont chokē (holo-: snua†). – type: south korea, jeollanam-do, suncheon city, songkwang-myeon, seungju-eup, jukhak-ri, mt. jogye-san, near the stream in front of songkwang temple, 30 jun. 1993, j.i. jeon and d.j. ha 10005 (neo-, designated by chang & chang (2010: 275): snua; isoneo-: kh). – note: chang et al. (2014a) accept c. laxiflora var. longispica as a distinct taxon with c. laxiflora var. macrothyrsa as its synonym. c. laxiflora var. macrothyrsa koidz., acta phytotaxonomica et geobotanica 9 (2): 73 (koidzumi 1940). – type: south korea, mt. chiisan [jirisan], hannyaho, gawun [gurae-jun], [jeollanam-do], 2 jun. 1935, s. okamoto s.n. (lecto-, designated by chang & chang (2010: 276): kyo). – note: this name is regarded as a synonym of c. laxiflora var. longispica in chang et al. (2014a). c. laxiflora var. gosenensis franch. nom. nud. – note: this name is written on the specimen only: japan, niigata, montagnes de gosen, 28 jul. 1874, u.j. faurie s.n. (p06811469!). c. laxiflora var. onoei nom. nud. – note: this name is written on the specimens only: japan, [chūbu], ono [ōno], l. savatier 2935 (p06811464!); ibid., s. coll., s.n. (p06811462!). 19. carpinus lipoensis y.k.li guizhou science 1983 (2): 20 (li & mang 1983). – type: china, guizhou, lipo, in sylvis montium calcareum, 850 m a.s.l., y.k. li [李永康] 9940 (holo-: hgas? ex “herb. inst. biol. guizhou”). – note: this name is neither listed in the world checklist (govaerts & frodin 1998) nor in the flora of china (li & skvortsov 1999). 20. carpinus londoniana h.j.p.winkl. das pflanzenreich iv 61 (heft 19): 32 (winkler 1904). – type: china, yunnan, 4000 ft, a. henry 11640 (lecto-, designated here: k000859945!; isolecto-: a00033759!, b†, e00275498!, k000859946!, mo-2140881!, mo-2140882!, ny00253864!). 20a. carpinus londoniana h.j.p.winkl. var. londoniana c. poilanei a.camus, bulletin de la société botanique de france 76: 968 (camus 1930). – type: vietnam, annam, lang-bian, entre dran et dung-lo, e. poilane 3983 (lecto-, designated here: p06811169!). – additional type material: vietnam, dalat en face de manline, f. evrard 1168 (syn-: p06811171!). c. poilanei var. chevalieri a.camus, bulletin de la société botanique de france 76: 969 (camus 1930). – type: vietnam, annam, plateau du langbian, entre bellevue et dran, 30 apr. 1919, a. chevalier 40445 (lecto-, designated here: p06747035!; isolecto-: p06747037!, p06811163!). – additional type material: vietnam, belle [bellevue], 17 jun. 1921, b. hayata 707 (syn-: p06811154!); ibid., dalat, chemin de la montagne de l’éléphant ruisseau de prenh, 29 nov. 1924, f. evrard 1918 (syn-: p06811160!). 20b. carpinus londoniana var. lanceolata (hand.-mazz.) p.c.li flora reipublicae popularis sinicae 21 (2): 68 (li 1979a). – c. lanceolata hand.-mazz., oesterreichische botanische zeitschrift 80: 338 (handel-mazzetti 1931). – type: china, hainan, ngitse-leng, g. fenzel 159 (holo-: not localized). – additional type material: china, hainan, secus rivos in media insula, 300–500 m a.s.l., oct.–nov. 1929, g. fenzel 106 (para-: not localized); ibid., ngitse-leng, in glarea, g. fenzel 164 (para-: not localized). european journal of taxonomy 375: 1–52 (2017) 18 20c. carpinus londoniana var. latifolia p.c.li acta phytotaxonomica sinica 17 (1): 87 (li 1979b) [corrected from “latifolius”]. – type: china, chekiang [zhejiang], ning-po [ningbo (宁波)], s. coll. 1018 (holo-: pe). 20d. carpinus londoniana var. xiphobracteata p.c.li acta phytotaxonomica sinica 17 (1): 87 (li 1979b). – type: china, chekiang [zhejiang], yin hsien [yin xian (鄞县)], g.r. chen [chen gun-rung] 2289 (holo-: pe). 21. carpinus luochengensis j.y.liang guihaia 6 (4): 275 (liang 1986). – type: china, guangxi, luocheng, 28 jul. 1983, j.y. liang k1644 (holo-: ibk00190892!). – additional type material: ibid., 5 aug. 1984, f.n. wei [wei fa-nan, 韦 发南] 1743 (para-: ibk00190885!, ibk00190886!). – note: not mentioned in the flora of china (li & skvortsov 1999). 22. carpinus mengshanensis s.b.liang & f.z.zhao bulletin of botanical research, harbin 11 (2): 33 (liang & zhao 1991). – type: china, shandong, pingyi xian, meng shan, 750 m a.s.l., oct. 1984, f.z. zhao 84001 (holo-: sdfs). – note: not mentioned in the flora of china (li & skvortsov 1999). 23. carpinus microphylla z.c.chen ex y.s.wang & j.p.huang guihaia 5 (1): 15 (wang & huang 1985). – type: china, guangxi, tianyang xian, 750 m a.s.l., 16 may 1964, z.c. chen 54089 (holo-: ibk00079486!). – note: the holotype location is given by the authors as “hibg” but according to the index herbariorum (thiers 2016), hibg is the herbarium of hiroshima botanical garden. according to the mandarin text, the holotype is deposited in the “guangxi institute of botany”, which is ibk. 24. carpinus mollicoma hu bulletin of the fan memorial institute of biology n.s. 1 (3): 216 (hu 1949). – c. polyneura var. mollicoma (hu) c.c.yang, distribution of the woody plants in sichuan [四川树木分布]: 42 (yang 1997). – type: china, yunnan, xichou, faa-doou, 25 sep. 1947, k.m. feng 12053 (lecto-, designated here: pe00021946!; isolecto-: ibsc0368097, pe00818159!). – note: carpinus mollicoma is accepted in the flora of china (li & skvortsov 1999) and the world checklist (govaerts & frodin 1998). yang in flora sichuanica (editorial board of the flora sichuanica 2012: 35) notes that his combination c. polyneura var. mollicoma (hu) c.c.yang is a nom. nud. [comb. inval.?] in the 1997 publication, but he treats it as a distinct variety in c. polyneura in 2012. if there was no reference to the basionym in the cited reference, then the new combination was published in 2012. 25. carpinus monbeigiana hand.-mazz. anzeiger der oesterreichischen akademie der wissenschaften mathematisch-naturwissenschaftliche klasse 61: 162 (handel-mazzetti 1925). – original citation: “in silvis dumetisque calide temperatis inter dawan et gwanyilang, legi 3. vii. 1914 (nr. 3431) et supra schidsilu, legi 30. vi. 1914 (nr. 3331) prope urbem yungbe. certe in subtropicis ad fluvium mekong leg. monbeig 1908 ibique observavi infra vicum gangpi eandemque verosimiliter ad fl. salwin circa tschammutong pluries.” – type: china, yunnan, prope urbem yungbei, in regionis calide temperatae silvis inter vicos dawan et swanyilang, 2400–2600 m a.s.l., 3 jul. 1914, h.r.e. von handel-mazzetti 3431 (lecto-, designated holstein n. & weigend m., taxonomic checklist of carpinus and ostrya 19 here: wu030928!; isolecto-: e00275494!, k!). – additional type material: china, yunnan, prope urbem yungbei, in regionis calide temperatae dumetis supra vic. schidsilu, 2400–2700 m a.s.l., 30 jun. 1914, h.r.e. von handel-mazzetti 3324 (syn-: wu030929!); ibid., am mekong, 2400‒2600 m a.s.l., j.t. monbeig 1908 (syn-: not localized). c. densispica hu, acta phytotaxonomica sinica 9: 286 (hu 1964). – type: china, [yunnan], tehchin hsien, tzichung, sila, 2500‒2800 m a.s.l., in mixed forest, 19 [in protologue “18”] jul. 1940, k.m. feng 5618 (lecto-, designated here: pe00021886!; isolecto-: pe01062146!). – additional type material: china, [yunnan], weisi hsien, kangpu, at 2300 m a.s.l., in mixed woods, jul. 1938, c.w. wang [王启无] 64343 (para-: pe01062148!). c. likiangensis hu, acta phytotaxonomica sinica 9: 287 (hu 1964). – type: china, yunnan, likiang hsien, snow range, 2700 m a.s.l., in mixed forest, 18 aug. 1942, k.m. feng 9055 (lecto-, designated by lin et al. (2007: 1248): pe00021913!; isolecto-: pe00021914!). – additional type material: ibid., 2400 m a.s.l., 19 sep. 1955, k.m. feng 21595 (para-: pe00021912!, pe00818189!). c. monbeigiana var. weisiensis hu, acta phytotaxonomica sinica 9: 285 (hu 1964). – type: china, yunnan, weisi hsien [weixi lisu autonomous county], yehchi, 2040 m a.s.l., 24 aug. 1956, p.y. mao [mao ping-yi/mao pin-yih, 毛品一] 197 (lecto-, designated here: pe00021933!; isolecto-: pe00021934!). 26. carpinus omeiensis hu & fang acta phytotaxonomica sinica 9: 296 (hu 1964). – type: china, sichuan, mount omei [éméi shān], dashiaochenfeng, 1900 m a.s.l., 7 oct. 1957, k.h. yang [yang kuang-hway, 杨光辉] 57490 (lecto-, designated here: pe00021939!; isolecto-: ibsc0368104, kun0590785!, nas00070308, pe00818214!). – additional type material: china, guizhou, teking hsien, yaimenko, 1200 m a.s.l., in dense forest on slope, 15 aug. 1959, north kweichow exped. [黔北队] 1631 (para-: kun0590783!, nas00070309!, pe00818217!, pe00818218!). 27. carpinus orientalis mill. the gardeners dictionary ed. 8, no. 3 (miller 1768). – c. nigra moench, verzeichniss ausländischer bäume und stauden: 19 (moench 1785) nom. illegit. superfl. – type: oriens, j.p. de tournefort s.n. (lecto-, designated here: bm!). – additional original material: oriens, j.p. de tournefort s.n. (p [tournefort 5561], linn no. hs1481.2!, linn no. hs1481.3!). – note: the tournefort specimen in bm was designated to be lectotype because 1) miller cited tournefort’s corollarium t. 40 in the protologue, just as given on the label and 2) miller’s material is deposited in bm (stafleu & cowan 1976–1988). the other specimen in bm, dating from that time (bm001041899), is a cultivar from leiden (“hort. bni. [botanici] boerhaave”). miller most likely had material from boerhaave at hand, but he did not mention cultivated material from leiden in the protologue. hence it be or may not be that this specimen could be considered as original material. there are two c. orientalis specimens in the smith herbarium (linn-hs 1481.2 and linn-hs 1481.3). one (linn-hs 1481.3) refers only to tournefort’s institutiones rei herbariae (tournefort 1700) and the other one to both tournefort’s publications (tournefort 1700, 1703) but not to the actual page number in contrast to the specimen in bm. additionally, miller did not refer to the institutiones rei herbariae nor do the linn specimens match the description since they have (sub-)acute leaf apices, while miller explicitely refers to obtuse apices. hence, we do not believe that miller saw the material in the smith herbarium for his description. the lectotypification with the tournefort specimen in paris by olshanskyi (2014: 68) is not effective because the author forgot the phrase “designated here” or an equivalent (art. 7.10, mcneill et al. 2012). we deviate from his suggestion because miller’s herbarium is in fact now located in bm (stafleu & cowan 1976–1988), hence lectotypes of his names should preferably be placed there. european journal of taxonomy 375: 1–52 (2017) 20 27a. carpinus orientalis mill. var. orientalis c. duinensis scop., flora carniolica ed. 2, 2: 243 (scopoli 1772). – original citation: “circa duinum” [duino]. – type: not localized. c. minor pall., bemerkungen auf einer reise in die südlichen statthalterschaften des russischen reichs 2: 95 (pallas 1801). – original citation: “sobald man den belbek [a river], etwa zwölf werste von bachtschisarai [bakhchysarai] durchfahren hat.” [this is a locality on crimea, and the plant must have been collected soon after easter 1793]. – type: not localized. c. betulus var. edentula heuff., verhandlungen der zoologisch-botanischen gesellschaft in wien 8: 196 (heuffel 1858). – c. edentula kit., descriptiones et icones plantarum rariorum hungariae 2: xxxii (waldstein & kitaibel 1803‒1805) nom. nud. – original citation: “smyrmium et banatum inhabitant.” – type: in comitatu syrmiensi et in sylvis banatus atque serviae, p. kitaibel a57 (lecto-, designated here: bp-xl/144!; isolecto-: b-w 17700–030!). – additional original material: s. loc., a. wolny s.n. (bp-xl/86! ex herb. p. kitaibel). – note: carpinus edentula is a nom. nud. appearing first in kitaibel’s diary in 1804 (cf. mólnar 2015). spach (1842: 221) separates this name in a key, and refers to a “carpinus edentula roch. bann.”. rochel (1828: 26) just listed this name as having been communicated by kitaibel. steudel (1840: 300) published this name again, but only mentions the perennial life form and the origin, so his publication cannot be considered as valid, and spach (1842: 221) published it as synonym to an invalid c. betulus var. β. in 1858, heuffel recognized kitaibel’s name as a variety of c. betulus and amended it with a description thus validating the name. c. orientalis f. umbraculifera (beissn.) schelle, handbuch der laubholzbenennung: 49 (beissner et al. 1903). – c. orientalis [unranked] umbraculifera beissn., mitteilungen der deutschen dendrologischen gesellschaft 8: 133 (beissner 1899a). – c. betulus [unranked] umbraculifera hort. ex beissn., mitteilungen der deutschen dendrologischen gesellschaft 8: 133 (beissner 1899a) nom. inval. (not accepted by the author). – original citation: “schloss eisenberg ist herrlich oben am dicht bewaldeten erzgebirge gelegen, im tiefer liegenden park”. – type: not localized. – note: beissner mentioned a cultivar with a spherical crown that has been called c. betulus umbraculifera, but he supposed it to be a c. orientalis. a definite rank was given in 1903. c. orientalis f. calcarea radde-fom., trudy fizychno-matematychnogo viddilu. kiev [mémoires de la classe des sciences physiques et mathématiques de l’académie des sciences de l’ukraine] 15 (1): 95 (radde-fomina 1929). – original citation: “prov. czernomorskaja, distr. novorossijsk, ad capitem dob, prope pag. kabardinka. 30 vi. 1912, [d.i.] sosnowsky!” – type: not localized. c. orientalis var. coronata radde-fom., trudy fizychno-matematychnogo viddilu. kiev [mémoires de la classe des sciences physiques et mathématiques de l’académie des sciences de l’ukraine] 15 (1): 107 (radde-fomina 1929). – c. coronata schischk. nom. nud. – original citation: “anatolien, trapezund, unterer teil des deïrmen-dere-tales; gebüsch westlichen abhängen, 12 juni 1917.” – type: not localized (le?). c. orientalis var. gibbosa radde-fom., trudy fizychno-matematychnogo viddilu. kiev [mémoires de la classe des sciences physiques et mathématiques de l’académie des sciences de l’ukraine] 15 (1): 107 (radde-fomina 1929). – c. ovata schischk. ex radde-fom., trudy fizychno-matematychnogo viddilu. kiev [mémoires de la classe des sciences physiques et mathématiques de l’académie des sciences de l’ukraine] 15 (1): 107 (radde-fomina 1929) nom. inval. pro syn. – original citation: “anatolien, sandshak trapezund, schlucht gadshevara, zwischen platana und fol; abhänge mit gebüsch. 27. juni 1917.” – type: not localized (le?). – note: carpinus ovata was a name suggested by schischkin on a herbarium specimen that he had sent to radde-fomina for revision. she suggested that this plant should be called “c. orientalis var. gibbosa” instead of being described as a new species. c. orientalis f. grandifolia radde-fom., trudy fizychno-matematychnogo viddilu. kiev [mémoires de la classe des sciences physiques et mathématiques de l’académie des sciences de l’ukraine] holstein n. & weigend m., taxonomic checklist of carpinus and ostrya 21 15 (1): 94 (radde-fomina 1929). – original citation: “tauria ad viam e simpheropol in aluschtam” [alushta], viii. 1925, leg. n. dubowik!” – type: not localized. c. paucicostata schischk. ex radde-fom., trudy fizychno-matematychnogo viddilu. kiev [mémoires de la classe des sciences physiques et mathématiques de l’académie des sciences de l’ukraine] 15 (1): 107 (radde-fomina 1929) nom. inval. pro syn. – note: radde-fomina regarded schischkin’s plant as a mere c. orientalis. however, she indicated that he intended to describe the name later on. this name is therefore provisional and invalid here. the present authors were unable to locate a later, validating publication. c. orientalis f. banatica kárp., botanikai közlemények 34: 194 (kárpáti 1937). – original citation: “im banat bei herkulesbad” [băile herculane, romania]. – type: not localized. – note: this is supposed to be a narrow-leaved form. 27b. carpinus orientalis subsp. macrocarpa (willk.) browicz flora iranica 97: 2 (browicz 1972). – c. orientalis var. macrocarpa willk., forstliche flora von deutschland und oesterreich ed. 2: 368 (willkomm 1887). – c. macrocarpa (willk.) h.j.p.winkl., das pflanzenreich iv 61 (heft 19): 38 (winkler 1904). – c. orientalis f. macrocarpa (willk.) raddefom., trudy fizychno-matematychnogo viddilu. kiev [mémoires de la classe des sciences physiques et mathématiques de l’académie des sciences de l’ukraine] 15 (1): 90 (radde-fomina 1929) nom. nud. pro syn. – original citation: “in turkomanien hat [g.s.] karelin”. – type: not localized. – note: the published combination c. orientalis f. macrocarpa as a synonym was a misunderstanding by radde-fomina. she believed that medwedew (1909: 32) regarded the taxon as a forma. however, he used the term “form” informally only, because “form” appears in only the text (medwedew 1909: 32) while “var. macrocarpa” is written explicitly in the appendix (medwedew 1909: 41). medwedew’s name is unranked, but would have been a later homonym anyway. 28. carpinus paohsingensis w.y.hsia contributions from the institute of botany, national academy of peiping 2: 179 (hsia 1934). – type: china, sichuan, paohsing-hsien (mupin) [baoxing], 1960 m a.s.l., 8 jul. 1933, t.h. tu 4356 (lecto-, designated here: pe00021940!; isolecto-: ibk00079502!, ibsc0368268, pe00021941!). – note: carpinus paohsingensis is accepted in the world checklist (govaerts & frodin 1998), but accepted neither in the flora of china (li & skvortsov 1999) nor by chang et al. (2014a) but listed as synonymous with c. tschonoskii. c. falcatibracteata hu, acta phytotaxonomica sinica 9: 297 (hu 1964). – c. tschonoskii var. falcatibracteata (hu) p.c.li, flora reipublicae popularis sinicae 21 (2): 85 (li 1979a). – type: china, sichuan, hunghwa to wali, 1900 m a.s.l., 14 jul. 1959, szechuan econ. bot. liangshan exp. [四川经济植物（凉山）调查队] 1219 (lecto-, designated here: pe00021887!; isolecto-: cdbi0172170!, pe00802589!). – additional type material: china, sichuan, le-po-hsien, on open mountain slope, 1950 m a.s.l., 8 aug. 1934, t.t. yu [俞德浚] 3639 (para-: ibsc0368265, ibsc0368266, ibsc0368269, pe00802576!, wuk0330666); ibid., liang ho-ko, in forest, 1700 m a.s.l., 28 sep. 1930, f.t. wang 22597 (para-: pe00802565!); ibid., kiangsi [jiangxi], shui swe hsien, mofu shan, along gully, 10 sep. 1947, y.k. hsiung [hsiung yao-kuo/熊耀国] 5830 (para-: lbg00001770!, pe00802561!, pe00802562!, pe00802563!). – note: this name is neither accepted in the world checklist (govaerts & frodin 1998) nor in the flora of china (li & skvortsov 1999) nor by chang et al. (2014a). it is regarded in the first treatment as a synonym of c. paohsingensis, which itself is treated as synonymous with c. tschonoskii in the latter two treatments. in the flora sichuanica (editorial board of the flora sichuanica 2012), it is accepted pro parte minore as distinct variety of c. tschonoskii, while the rest ought to be synonymous with c. tschonoskii and with c. paohsingensis as full synonym of c. tschonoskii. european journal of taxonomy 375: 1–52 (2017) 22 c. obovatifolia hu, acta phytotaxonomica sinica 9: 289 (hu 1964). – type: china, yunnan, tsenyih hsien, shiao-ma-la in forest, 1954, y.h. li [li yen-hei, 李延辉] 150 (lecto-, designated by lin et al. (2007: 1249): pe00021937!; isolecto-: pe00021936!). – note: this name is listed as synonymous with c. tschonoskii in the flora of china (li & skvortsov 1999), in the flora sichuanica (editorial board of the flora sichuanica 2012), and by chang et al. (2014a) for the flora of korea, while it is synonymous with c. paohsingensis in the world checklist (govaerts & frodin 1998). 29. carpinus polyneura franch. journal de botanique [morot] 13: 202 (franchet 1899). – c. turczaninovii var. polyneura (franch.) h.j.p.winkl., das pflanzenreich iv 61 (heft 19): 38 (winkler 1904). – type: china, sutchuen [sichuan], in ditione tchen keou tin [chengkou county], p.g. (reverend père) farges s.n. (lecto-, designated here: p06811144!; isolecto-: p06811145!?, p06811146!). – note: carpinus polyneura is accepted as a species, with the two varieties (with c. polyneura var. sungpanensis) that are listed here, in the world checklist (govaerts & frodin 1998) and in the flora sichuanica (editorial board of the flora sichuanica 2012), but they are accepted as two distinct species in the flora of china (li & skvortsov 1999). 29a. carpinus polyneura franch. var. polyneura c. handelii rehder, journal of the arnold arboretum 1: 59 (rehder 1919). – type: china, hunan, inter urbes linling (yungchoufu) e sinning in silvis collium supra vicum tjentiesse, 400 m a.s.l., 14 aug. 1917, h.r.e. von handel-mazzetti 421 (holo-: a00033754!; iso-: wu 30926!). – type: china, hunan, in silva infra tungdjiapai prope minas hsikwangchan, dist. hsinhwa, 550 m a.s.l., 20 may 1918, h.r.e. von handel-mazzetti 11884 [“534”] (para-: e!, p06811414!, wu 30927!). – note: this name is listed as a synonym of c. polyneura in the world checklist (govaerts & frodin 1998), in the flora of china (li & skvortsov 1999), and in the flora sichuanica (editorial board of the flora sichuanica 2012). rehder cites the type as “421”, which is a crossed-out number written with a pencil only on the specimen in a. it is therefore the specimen that rehder explicitly cited and type, as the wu duplicate does not contain that number. c. polyneura var. glandulosopunctata c.j.qi, acta phytotaxonomica sinica 22 (6): 494 (qi 1984). – c. glandulosopunctata (c.j.qi) c.j.qi, bulletin of botanical research, harbin 20 (1): 3 (lin & qi 2000) [“glanduloso-punctata”]. – type: china, hunan, ningxiang, on hillside, 200 m a.s.l., 10 oct. 1982, y.t. xiao [xiao yu-tan] 40004 (holo-: csfi; iso-: pe). – note: this name is listed as a synonym of c. polyneura in the world checklist (govaerts & frodin 1998), but it is not mentioned otherwise. c. polyneura var. wilsoniana h.j.p.winkl., botanische jahrbücher für systematik, pflanzengeschichte und pflanzengeographie 50 (suppl.): 506 (winkler 1914). – type: china, [sichuan,] am berge omi [éméi shān], may 1904, e.h. wilson 5791 (lecto-, designated here: k000859929!; isolecto-: a00033763!, bm!). – additional type material: china, hupeh, e.h. wilson 1170 p.p. (syn-: w). – note: this name is mentioned neither in the world checklist (govaerts & frodin 1998) nor in the flora of china (li & skvortsov 1999) nor in the flora sichuanica (editorial board of the flora sichuanica 2012). 29b. carpinus polyneura var. sungpanensis (w.y.hsia) p.c.li flora reipublicae popularis sinicae 21 (2): 86 (li 1979a). – c. sungpanensis w.y.hsia, contributions from the institute of botany, national academy of peiping 2: 180 (hsia 1934). – c. tschonoskii var. sungpanensis (w.y.hsia) c.c.yang, distribution of the woody plants in sichuan [四川树木分布]: 42 (yang 1997). – type: china, sichuan, sungpan-hsien, 2050 m a.s.l., 16 sep. 1932, t.t. yü 2580 (lecto-, designated here: pe01062157!; isolecto-: ibsc0368164, ibsc0368182, pe01062158!). – note: the taxon is accepted as a distinct species in flora of china (li & skvortsov 1999), but treated holstein n. & weigend m., taxonomic checklist of carpinus and ostrya 23 as a variety of c. polyneura in the world checklist (govaerts & frodin 1998) and in the flora sichuanica (editorial board of the flora sichuanica 2012). 29c. carpinus polyneura var. tsunyihensis (hu) p.c.li flora reipublicae popularis sinicae 21 (2): 88 (li 1979a). – c. tsunyihensis hu, acta phytotaxonomica sinica 9: 296 (hu 1964). – type: china, kweichow [guizhou], tsunyih hsien, nanchien shan, on hilly slope, 900‒1050 m a.s.l., 17 aug. 1956, szechuan-kweichow exp. 973 (holo-: pe00818388!). – note: this taxon is accepted as a distinct species in the flora of china (li & skvortsov 1999), but it is considered as a variety of c. polyneura in the world checklist (govaerts & frodin 1998) and in the flora sichuanica (editorial board of the flora sichuanica 2012). 30. carpinus pubescens burkill journal of the linnean society, botany 26 (178): 502 (forbes & hemsley 1899) [“1890”]. – type: china, yunnan, mi-lê district, a. henry 9929 (holo-: k000859939!; iso-: a00033764!, e00275493!, ny00253866!, pe00934232!). c. seemeniana diels, botanische jahrbücher für systematik, pflanzengeschichte und pflanzengeographie 29: 279 (diels 1901). – c. pubescens var. seemeniana (diels) hu, acta phytotaxonomica sinica 9: 290 (hu 1964). – type: china, s nan ch’uan, shan tzu p’ing, wald, aug., c. bock and a. von rosthorn [“bvr”] 294 (lecto-, designated here: gzu-rosthorn 000261231!; isolecto-: a00033767!, b†, k000859940!). c. pinfaensis h.lév. & vaniot, bulletin de la société botanique de france 52: 142 (léveillé 1905). – type: china, kouy-tchéou, pin-fa, 28 mar. 1903, j. cavalerie 1011 (holo-: e00275492!; iso-: a00077469!). c. austrosinensis hu, sinensia 2: 87, fig. 4 (hu 1931). – type: china, kweichow [guizhou], anlung, in lightly shaded mixed woods, 620 m a.s.l., 22 oct. 1930, y. tsiang [蒋英] 7470 (holo-: not localized; iso-: a00033748!, bm000580520!, e00275505!, ibsc0368184, k000859927!, n000053493, ny00253859!, p06747032!, pe00020176!, pe00020177!, us00089311!). – note: the type shown in hu (1931: fig. 4) is not among the specimens seen. c. kweitingensis hu, sinensia 2: 82, fig. 2 (hu 1931). – c. pubescens var. kweitingensis (hu) hu, acta phytotaxonomica sinica 9: 290 (hu 1964). – type: china, guizhou, guiding/kweiting: pingfa, light shaded woods, 400‒550 m a.s.l., 29 jun. 1930, y. tsiang [蒋英] 5440 (holo-: not localized; iso-: a00077466!, a00077467!, e00275502!, ibsc0001132, m-0265493!, nas00070311!, p06811206!, pe00021904!). – note: the type shown in hu (1931: fig. 2) is not among the specimens seen. c. tsiangiana hu, sinensia 2: 90, fig. 5 (hu 1931). – type: china, guizhou, kwangcheng, open hillsides, 23 jul. 1930, y. tsiang [蒋英] 8610 (lecto-, designated here: nas00070318!; isolecto-: a00033770!, bm000580521!, e00275491!, k000859920!, nas00070312!, nas00070313!, nas00070314!, ny00253867!, pe00021964!, us00089309!). – note: the specimen chosen as lectotype here is the one depicted in hu (1931: fig. 5). c. tungtzeensis hu, sinensia 2: 85, fig. 3 (hu 1931). – type: china, guizhou, tungtze. 450 m a.s.l. (1476.4 ft.), 26 may 1930, y. tsiang [蒋英] 5111 (syn-: nas00070319, nas00070320, nas00070321; isosyn-: bm000580523!, k000859921!, n000053499, ny00253868!, pe00021966!, pe00021967!, us00089308!). c. kweiyangensis hu, bulletin of the fan memorial institute of biology n.s. 1 (2): 187 (hu 1948b). – type: china, kweichow [guizhou], kweiyang [贵阳市, guiyang], chen-lin shan, 10 oct. 1937, p.c. tsoong [钟补勤] 1118 (holo-: pe00021948!). c. lancilimba hu, bulletin of the fan memorial institute of biology n.s. 1 (2): 185 (hu 1948b). – type: china, kweichow [guizhou], tse-ching hsien [zhijin co., 织金县], [dongshan, 县东山], 28 oct. 1938, p.c. tsoong [钟补勤] 1879 (lecto-, designated here: pe00021909!; isolecto-: pe00021910!). european journal of taxonomy 375: 1–52 (2017) 24 c. pilosinucula hu, bulletin of the fan memorial institute of biology n.s. 1 (2): 142 (hu 1948a). – type: china, guizhou, anlong co., 15 jul. 1940, p.c. tsoong [钟补勤] 1536 (holo-: pe00021943!). c. pingpienensis hu, bulletin of the fan memorial institute of biology n.s. 1 (2): 188 (hu 1948b). – type: china, yunnan, ping-pien hsien, 1500 m a.s.l., 18 jul. 1934, h.t. tsai [tsai hsi-tao, 蔡希陶] 60994 (holo-: pe00021944!; iso-: ibsc0368199, lbg00053976!, wuk0046644). c. tsoongiana hu, bulletin of the fan memorial institute of biology n.s. 1 (2): 186 (hu 1948b). – type: china, kweichow [guizhou], yungchun hsien, on rocky slope along river [贵州省, 从江县, guizhou province from river county], 29 jul. 1937, p.c. tsoong [钟补勤] 1084 (holo-: pe00021965!). c. wangii hu & w.c.cheng, bulletin of the fan memorial institute of biology n.s. 1 (2): 147 (hu 1948a). – type: china, yunnan, foo-ning, lung-mai, on rocky hill, 1000 m a.s.l., c.w. wang [王启 无] 89104 (holo-: not localized; iso-: kun0590973!). c. marlipoensis hu, bulletin of the fan memorial institute of biology n.s. 1 (3): 215 (hu 1949). – type: china, yunnan, malipo county, pan-chia-chu, 31 oct. 1947, k.m. feng 12619 (lecto-, designated here: pe00021921!; isolecto-: pe00021922!, wuk0207179). c. marlipoensis var. angustifolia hu, acta phytotaxonomica sinica 9: 288 (hu 1964). – type: china, yunnan, faadou, 1400 m a.s.l., in mixed forests on rocky hills, 24 may 1943, p.c. tsoong and k.z. kuang [钟补勤、匡可任] 426 (lecto-, designated here: pe00021925!; isolecto-: ibsc0368204, pe00021923!, pe00021924!). c. parva hu, acta phytotaxonomica sinica 9: 292 (hu 1964). – type: china, se yunnan: sichour hsien, in shrubbery on cretaceous cliff, 1600 m a.s.l., 17 oct. 1958, h.t. tsai [tsai hsi-tao, 蔡希 陶] 58–8559 (holo-: pe00021942!). c. pingpienensis var. lichuanensis w.c.cheng, 科学 [science] 32 (8): 246 (cheng 1950) nom. nud.? – c. pingpienensis var. lichuanensis w.c.cheng, 湖北植物志 [flora hupehensis] 1: 95 (hubei institute of botany 1976) nom. nud. – c. pingpienensis var. hupehensis w.c.cheng nom. nud. – note: the name “c. pingpienensis var. hupehensis” only appears on the specimens pe00021911!, a00033762 and k000859922! (china, [hubei,] li-chuan, hsio-ho, long-tawn-pu, in forest, 4100 ft, 13 sep. 1948, w.c. cheng and c.t. hwa 950) and appears to be provisional. cheng annotated the pe specimen in 1960 as “carpinus lichuanensis hu” and as “carpinus pingpienensis var. lichuanensis” in 1967. tang (1987) cited the name c. pingpienensis var. lichuanensis as a “n.n.” (nom. nud.) as does chen (2015). the present authors did not find the protologue for validation. synonymization follows tang (2007) as the name is not mentioned in the flora of china (li & skvortsov 1999). c. whilungpaensis w.c.cheng, 科学 [science] 32 (8): 246 (cheng 1950) nom. nud.? – note: tang (2007) cited this name as a “n.n.” [nomen nudum?] found on a specimen of w.c. cheng and c.t. hwa 866 (duplicate without that note: k!). the present authors did not find the protologue, and the name is not cited in the flora of china (li & skvortsov 1999). 31. carpinus purpurinervis hu acta phytotaxonomica sinica 9: 293 (hu 1964). – type: china, kwangsi [guangxi], du-an, in forests, 2 jul. 1957, y.k. li [li yin-kuen/李荫昆] p01567 (holo-: pe; iso-: ibk00190887!, ibk00190888!, ibk00190889!, ibk00190890!, ibk00190891!, ibsc0368220). 32. carpinus putoensis w.c.cheng contributions from the biological laboratory of the science society of china, botanical series 8: 72 (cheng 1932). – type: china, zhejiang, [普陀区, 佛顶山/putuo district, fodingshan], 15 may 1930, k.k. tsoong [钟观光] 94 (lecto-, designated here: pe00021950!; isolecto-: pe00021949!, pe00021951!). holstein n. & weigend m., taxonomic checklist of carpinus and ostrya 25 33. carpinus rankanensis hayata icones plantarum formosanarum 6: 63, fig. 8 (hayata 1916). – type: r. china, [taiwan, yilan county], rankanzan [lan-k’an shan, a mountain range ne of nanao], 4000 ft, 12 may 1916, b. hayata s.n. (lecto-, designated here: ti!; isolecto-: taif007302!, taif007303!, taif007304!). 33a. carpinus rankanensis hayata var. rankanensis 33b. carpinus rankanensis var. matsudae yamam. supplementa iconum plantarum formosarum 5: 15, fig. 5 (yamamoto 1932). – type: r. china, taiwan, ariko-banti, 8 jul. 1917, e. matuda [also called matsuda] 11 (holo-: tai118774!; iso-: ti!). – note: this variety is accepted as a distinct variety of c. rankanensis in flora of china (li & skvortsov 1999) but not by liao (1996). 34. carpinus rupestris a.camus bulletin de la société botanique de france 76: 966 (camus 1930). – type: s. loc., s. coll., eventually j. cavalerie 4560 (lecto-, designated here: p01903242!; isolecto-: p01903240!, p01903241!). – additional type material: china, kouy-tchéou [guizhou], houang-tia-pa [houang-ts’ao-pa], grands rochers, j. cavalerie 4560 (syn-: a00033766!, e00240621!, k!); ibid., [guizhou], hiny-hien [missionary station in sw of guizhou, nw of xingyi city, hsien-i-fu, 興義府], rochers, j. cavalerie s.n. (syn-: p01903239!). – note: the specimen designated here as lectotype bears an annotation by camus, so it is original material, but it lacks (as do the isolectotypes) an original label by cavalerie. the specimen p01903242 bears a written note on an extracted page with the protologue that “cavalerie 4560” is apparently missing in the paris herbarium. possibly, the original label became detached and is lost. the specimens p01903240–2 are not duplicates of the other syntype p01903239, which has male catkins and narrower leaves. in addition, p01903241 and p01903242 contain infrutescences. 35. carpinus × schuschaensis h.j.p.winkl. das pflanzenreich iv 61 (heft 19): 32 (winkler 1904). – c. betulus [unranked] schuschaensis (h.j.p. winkl.) medw., vestnik tiflisskogo botanicheskogo sada 14: 33 (medwedew 1909). – c. betulus f. schuschaensis (h.j.p.winkl.) radde-fom., trudy fizychno-matematychnogo viddilu. kiev [mémoires de la classe des sciences physiques et mathématiques de l’académie des sciences de l’ukraine] 15 (1): 80 (radde-fomina 1929) comb. inval. pro syn. – type: karabagh, schuscha, r.f. hohenacker s.n. (syn-: b†, g-boiss., w). – note: radde-fomina published the combination c. betulus f. schuschaensis in synonymy, because she believed that medwedew understood the name as a forma. however, medwedew used the term “form” informally, as he used form (medwedew 1909: 33) and variety (medwedew 1909: 41) in parallel. his combinations are therefore unranked (art. 37.8; mcneill et al. 2012). biologically, this is a hybrid between c. betulus and c. orientalis. c. × grosseserrata h.j.p.winkl., das pflanzenreich iv 61 (heft 19): 40 (winkler 1904). – type: iran, bei radkan, f.a. buhse 1043 (holo-: b†). c. × hybrida h.j.p.winkl., das pflanzenreich iv 61 (heft 19): 40 (winkler 1904). – type: transkaukasien, r.f. hohenacker (syn-: b† and/or le). karabagh, s. coll., s.n. (syn-: b† and/or le). c. × oxycarpa h.j.p.winkl., das pflanzenreich iv 61 (heft 19): 31 (winkler 1904). – c. betulus [unranked] oxycarpa (h.j.p. winkl.) medw., vestnik tiflisskogo botanicheskogo sada 14: 27 (medwedew 1909). – c. betulus f. oxycarpa (h.j.p. winkl.) radde-fom., trudy fizychno-matematychnogo viddilu. kiev [mémoires de la classe des sciences physiques et mathématiques de l’académie des sciences de l’ukraine] 15 (1): 83 (radde-fomina 1929) comb. inval. pro syn. – type: karabagh, european journal of taxonomy 375: 1–52 (2017) 26 schuscha, r.f. hohenacker, u.i. 1838 (syn-: b†, g-boiss.?, le, herb. moritz winkler, w†). – note: olga radde-fomina’s combination in synonymy is a misunderstanding. she believed that medwedew published a forma, but he used the term “form” informally, which is apparent in the index of his treatment, where he explicitly notes “var. oxycarpa (winkl.) medw.” (medwedew 1909: 41). medwedew’s combination is nevertheless to be regarded as unranked (art. 37.8, mcneill et al. 2012). in contrast to winkler’s note (winkler 1904), duplicates are not present in “breslau” (wrocław; brsl or wrsl, pers. comm. k. świerkosz). c. × oxycarpa nothovar. betuloides h.j.p.winkl., das pflanzenreich iv 61 (heft 19): 32 (winkler 1904). – type: karabagh, bei schuscha, r.f. hohenacker, u.i. 1838 (syn-: b†, g, le). c. × geokczaica radde-fom., trudy fizychno-matematychnogo viddilu. kiev [mémoires de la classe des sciences physiques et mathématiques de l’académie des sciences de l’ukraine] 15 (1): 89 (raddefomina 1929). – original citation: “prov. baku distr. geokczai, in frutecetis siccis in ascensu as pagum vaenk 1700’. 31 vii. 1899. alexeenko (herb. acad. petrop.) prov. baku distr. schemacha, prope pagum müdshi, in decliviis meridionalibus, in papidosis calcareis. 4000’. 30. vii. 1900. alexeenko (herb. acad. petrop.)”. – type: not localized. 36. carpinus shensiensis hu bulletin of the fan memorial institute of biology n.s. 1 (2): 145 (hu 1948a). – type: china, shensi [shaanxi], yuan kwan t’ang, mixed with celtis, 15 oct. 1935, y.y. pai 2891 (lecto-, designated here: pe00021952!; isolecto-: nas00070315, wuk0072726). – additional type material: china, shaanxi, hsing lung sze, 15 oct. 1935, y.y. pai 2860 (para-: pe00802529!, wuk0072723, wuk0170781). c. shensiensis var. paucineura s.c.qu & k.y.wang, bulletin of botanical research, harbin 8 (4): 111 (qu & wang 1988). – type: china, shaanxi, zhenping xian, feihe xiang, laoyachung, 1800 m a.s.l., aug. 1986, k.y. wang 747 (holo-: nwfc). – additional type material: ibid., 1200 m a.s.l., k.y. wang 740 (para-: not localized). 37. carpinus shimenensis c.j.qi journal of nanjing technological college of forest products 1981 (3): 123 (qi 1981). – type: china, hunan, shimen district, huping shan, 2050 m a.s.l., p.c. cai [cai pingcheng] 20241 (holo-: csfi017382; iso-: nf). – note: this name is accepted in the world checklist (govaerts & frodin 1998), but it is not mentioned in the flora of china (li & skvortsov 1999). 38. carpinus tientaiensis w.c.cheng contributions from the biological laboratory of the science society of china, botanical series 8: 135 (cheng 1932). – c. laxiflora var. tientaiensis (w.c.cheng) hu, sunyatsenia 1: 112 (hu 1933). – type: china, zhejiang, tiantai, tien tai shan, open woods, 2800 ft, 12 aug. 1927, y.l. keng [耿 以礼] 1065 (lecto-, designated by lin & sun (2007: 178): pe00021959!; isolecto-: a00033765!, pe00021935!). – additional type material: china, zhejiang, tiantai, tientai shan, in wood partially shaded, 3500 ft, 10 may 1924, r.c. ching 1547 (syn-: pe00021958!). – note: the isolectotype from pe lacks a specimen label, but it has a collection label that matches the lectotype collection label exactly. it is therefore very likely a duplicate. however, it is annotated as “carpinus nobilis hu type”, a name whose publication status is unknown to the present authors. 39. carpinus tropicalis (donn.sm.) lundell lloydia 2: 79 (lundell 1939). – c. americana var. tropicalis donn.sm., botanical gazette 15: 28 (smith 1890). – c. caroliniana var. tropicalis (donn.sm.) standl., contributions from the united states holstein n. & weigend m., taxonomic checklist of carpinus and ostrya 27 national herbarium 23 (1): 169 (standley 1920). – type: guatemala, alta verapaz, chicoyonito, 4300 ft, apr. 1889, j. donnell smith in donn. sm. exs. pl. cit. 1667 (lecto-, designated by furlow (1987: 431): us00089312!; isolecto-: gh00033747!, k000512590!). – additional type material: guatemala, alta verapaz, bei coban tactic purulá, 1250‒1500 m a.s.l., may 1882, f.c. lehmann pl. guatemal., costaric. columbianae 1446 (syn-: bm000956101!, k!); ibid., cumbre de sta. lucia, jan. 1878, k.g. bernoulli and r. cario fl. guatemal. 2606 (syn-: goet!, k!); ibid., las majadas, may 1878, k.g. bernoulli and r. cario fl. guatemal. 2607 (syn-: goet!, k!). 39a. carpinus tropicalis (donn.sm.) lundell var. tropicalis 39b. carpinus tropicalis var. mexicana furlow systematic botany 12 (3): 431 (furlow 1987). – type: mexico [méxico], [jalisco], municipio de gómez farias, tamaulipas, camino al rancho del cielo, 1150 m a.s.l., 8 aug. 1969, h. puig 5159 (holo-: encb). – additional type material: mexico [méxico], cerro de corona, zacualpan, 2‒3 may 1954, e. matuda et al. 30712 (para-: bh); ibid., municipio de ocuilan, cerca del rio chalma, ahuehuete, 19 sep. 1962, may nah am-48 (para-: encb, mich); ibid., 5 km al sw de sultipec, sobre el camino a amatepec, 2350 m a.s.l., 18 feb. 1979, j. rzedowski 36059 (para-: encb, mich); ibid., [michoacán], municipio de pueblo nuevo, ladera s del volcán tancitaro, 2000 m a.s.l., 19 mar. 1977, l.m.v. de puga and s. carvajal h. 10134 (para-: encb); ibid., nuevo león, dulces nombres, 13 jul. 1948, f.g. meyer and d.j. rogers 2752 (para-: bm000956117!, e!, gh); ibid., nuevo león, municipio de montemorelos, trail from la trinidad to sierra de la cebolla, 20 aug. 1939, c.h. muller 2867 (para-: gh). 40. carpinus tsaiana hu bulletin of the fan memorial institute of biology n.s. 1 (2): 141 (hu 1948a). – type: china, yunnan, ping-pien hsien [屏边苗族自治县, 水塘坡, pingbian miao autonomous county], 7 jul. 1934, h.t. tsai 62398 (lecto-, designated here: pe00021960!; isolecto-: pe00021961!, pe00021962!). – additional type material: ibid., 3 jun. 1934, h.t. tsai 62191 (para-: lbg00053971, pe00802549!); ibid., yunnan, si-chou hsien, far-doe [西畴县, 法斗], 12 dec. 1939, c.w. wang [王启无] and y. li [刘瑛] 85686 (para-: pe00021918!, pe00021919!, pe00021920!, pe00802552!) [partly misspelled as 2 dec. and 65686 but all with same label data]. – note: carpinus tsaiana hu is accepted in the flora of china (li & skvortsov 1999). the lectotype designated here is not the unpublished one, labeled as holotype. we deviate from the original intention, because the intended “holotype” has lost the fruits of its infructescence, therefore we choose a more complete specimen. c. sichourensis hu, bulletin of the fan memorial institute of biology n.s. 1 (3): 214 (hu 1949). – type: china, yunnan, xichou [西畴县], ma-chia, 14 oct. 1947, k.m. feng 12516 (holo-: pe00802497!). 41. carpinus tschonoskii maxim. bulletin de l’academie imperiale des sciences de st.-pétersbourg 27: 534 (maximowicz 1881b). – type: japan, [kanagawa], hakone, 1864, s. tschonoski s.n. (syn-: le; isosyn-: bm!, br!, gh00033769 p.p.!, k000859942!, l! m-0265492!, p06809892!, p06811215!, p06809882!?, p06811396!?, p06809866!?, pe00021963 p.p.!, wu!); ibid., fudzi-yama [fuji yama], 1864, s. tschonoski s.n. (syn-: le; isosyn-: bm000580518!, gh00033769 p.p.!, k000859941!, p06811217!, pe00021963 p.p.!). c. yedoensis maxim., bulletin de l’academie imperiale des sciences de st.-pétersbourg 27: 535 (maximowicz 1881b). – type: japan, [tokio,] cult. in yedo [edo, tokio] garden, nov. 1862, c.j. maximowicz s.n. (syn-: not localized); ibid., l. savatier 1172 (syn-: not localized). european journal of taxonomy 375: 1–52 (2017) 28 c. fauriei nakai, the botanical magazine [tokyo] 26: 325 (nakai 1912). – type: south korea, [jeju], quelpaert, in silvis, oct. 1907, e.-j. taquet 587 (syn-: ti, kyo); ibid., aug. 1911, t. mori 32 (syn-: ti); ibid., aug. 1911, t. mori 33 (syn-: ti). c. eximia nakai, repertorium specierum novarum regni vegetabilis 13: 249 (nakai 1914). – c. tschonoskii var. eximia (nakai) hatus., bulletin of the kyushu imperial university forests 5: 48 (hatushima 1934). – type: south korea, [jeollanam-do], secus vias et in silvis circa templum senonji pede montem chirisan, 280 m a.s.l., 15 jul. 1913, t. nakai 11 (holo-: ti). – note: this name is accepted as a distinct species in the world checklist (govaerts & frodin 1998), but listed as a synonym of c. tschonoskii in chang et al. (2014a) and the wcsp (2016). c. tschonoskii var. jablonskyi h.j.p.winkl., botanische jahrbücher für systematik, pflanzengeschichte und pflanzengeographie 50 (suppl.): 500 (winkler 1914). – type: japan, prov. kotzuke, bamba und nogurizawa, 29 jul. 1894, m. shirai s.n. (syn-: not localized); ibid., 1 aug. 1894, m. shirai s.n. (syn-: not localized). – note: this name is mentioned neither in the world checklist (govaerts & frodin 1998) nor in the wcsp (2016) nor in the flora of japan (ohwi 1965). chang et al. (2014a) list it as synonymous with c. tschonoskii. c. tschonoskii var. serratiauriculata h.j.p.winkl., botanische jahrbücher für systematik, pflanzengeschichte und pflanzengeographie 50 (suppl.): 500 (winkler 1914). – type: japan, [tokyo,] jedo, l. savatier 1172 (syn-: not localized); ibid., l. savatier s.n. (syn-: not localized). ibid., 1874, l. savatier s.n. (syn-: p06811219!, p06810761!). – note: this name is mentioned neither in the world checklist (govaerts & frodin 1998) nor in the wcsp (2016) nor in the flora of japan (ohwi 1965). chang et al. (2014a) list it as synonymous with c. tschonoskii. c. tschonoskii var. subintegra h.j.p.winkl., botanische jahrbücher für systematik, pflanzengeschichte und pflanzengeographie 50 (suppl.): 501 (winkler 1914). – type: south korea, [jeju], insel quelpart, hallaisan [hallasan mt.], jul. 1907, u.j. faurie 1535 (holo-: not localized; iso-: a00033735!, bm!, e00388827!, p06809858!). – note: this name is not mentioned in the world checklist (govaerts & frodin 1998), while chang et al. (2014a) list it as synonymous with c. tschonoskii. c. tschonoskii var. brevicalycina nakai ex kawamoto, handbook of korea-manchurian forestry: 92 (kawamoto 1939). – type: not localized. – note: this name is not mentioned in the world checklist (govaerts & frodin 1998) nor in the wcsp (2016). jeon & chang (1997) consider it as a synonymous with c. tschonoskii, chang et al. (2014a) list it as a invalid name. the present authors did not see the protologue, so validity of the name is unclear. c. coreensis koidz., acta phytotaxonomica et geobotanica 9 (2): 73 (koidzumi 1940). – original citation: “carpinus fargesiana (non winkl.) nakai fl. sylv. korea. ii (1915). […] korea: mt. chiisan, hannyaho, kokiri”. – type: not localized. – note: jeon & chang (1997) and chang et al. (2014a) consider this name to be synonymous with c. tschonoskii, as it is listed in the world checklist (govaerts & frodin 1998). c. tschonoskii f. pendula hayashi, 林業試験場研究報告 [research report of the forestry experiment station] 125: 72 (hayashi 1960). – type: japan, hondo, prov. mushi, kokubunji, 10 may 1958, y. hayashi s.n. (holo-: tfm). – note: this name is not mentioned in the world checklist (govaerts & frodin 1998) or in the flora of japan (ohwi 1965). most likely, this is to be regarded as a cultivar with hanging branches rather than as a botanical taxon. c. mianningensis t.p.yi, bulletin of botanical research, harbin 12 (4): 335 (yi 1992). – type: china, sichuan, mianning county, lingshan, 18 jul. 1991, t.p. yi [易同培] 91170 (holo-: sifs; iso-: pe00934231!). – note: this name is accepted in the flora sichuanica (editorial board of the flora sichuanica 2012), but it is listed as synonymous with c. tschonoskii in the flora of china (li & skvortsov 1999). holstein n. & weigend m., taxonomic checklist of carpinus and ostrya 29 41. carpinus turczaninovii hance journal of the linnean society, botany 10 (44): 203 (hance 1868). – type: china, in collibus ad occidentem urbis peking [beijing] sitis, aug. 1865, s.w. williams 12681 (holo-: k000859937!). – additional original material: china, in collinis pekinensibus, aug. 1865, s.w. williams 12681 (bm000580390!). – additional original material?: china, in ditione pekinensis, aug. 1866, s.w. williams 12681 (gh00112537!, le). – note: several gatherings are distributed with the number 12681, all belonging to the same species. winkler (1914: 502) already mentions the admixed nature of the specimens with the same “collection” number. hance cited “collibus ad occidentem urbis peking, m. augusto 1865,” which is only written on the specimen k000859937, which is therefore the holotype. however, the other specimens seen by the present authors were distributed from hance’s collection, so they are original material. c. paxii h.j.p.winkl., das pflanzenreich iv 61 (heft 19): 35 (winkler 1904). – type: china, peking, tsetai-ssu, h. wawra von fernsee 1067 (syn-: b†, w†; isosyn-: a00033761!). – note: winkler (1914: 502) acknowledged c.k. schneider’s comment that his c. paxii is synonymous to c. turczaninovii. this name is not mentioned in the flora of china (li & skvortsov 1999), but it is listed as synonym of c. turczaninovii in the flora of japan (ohwi 1965) and in the flora sichuanica (editorial board of the flora sichuanica 2012). c. stipulata h.j.p.winkl., das pflanzenreich iv 61 (heft 19): 35 (winkler 1904). – c. turczaninovii var. stipulata (h.j.p.winkl.) h.j.p.winkl., botanische jahrbücher für systematik, pflanzengeschichte und pflanzengeographie 50 (suppl.): 505 (winkler 1914). – type: china, schensi [shaanxi], c-china, im tsin-ling-shan-gebirge, 10 jul. 1900, g. [giuseppe, but also called joseph; reverend] giraldi 7267 (lecto-, designated here: k000859930!; isolecto-: a00077468!, b†). – note: this name is listed as a synonym of c. turczaninovii in the flora of japan (ohwi 1965), but it is accepted as a distinct species in the flora of china (li & skvortsov 1999). c. tanakaeana makino, the botanical magazine [tokyo] 28: 32 (makino 1914). – type: japan, tokushima, prov. tosa, mt. yokogura (on limestone), t. makino s.n. (holo-: mak046499!). c. turczaninovii var. makinoi h.j.p.winkl., botanische jahrbücher für systematik, pflanzengeschichte und pflanzengeographie 50 (suppl.): 505 (winkler 1914). – type: japan, prov. tosa, 1889, t. makino 277 (syn-: not localized); ibid., s. loc., m. shirai s.n. (syn-: not localized). – note: this name is not listed in the world checklist (govaerts & frodin 1998) or in the wcsp (2016), but listed as a synonym of c. turczaninovii in the flora of japan (ohwi 1965). c. coreana nakai, the botanical magazine [tokyo] 40: 162 (nakai 1926). – c. turczaninovii var. coreana (nakai) w.t.lee, lineamenta florae koreae 166 (lee 1996). – type: south korea, [jeollanam-do], in monte kannonzan insl. wangto, 20 jun. 1913, t. nakai 581 (holo-: ti). – additional type material: south korea, [incheon], prov. keiki, chemulpo [jemulpo], in horto residentis germanici, sep. 1906, u.j. faurie 202 [cited as e. taquet 202] (para-: bm000580428, e00275514!, p05591545!); ibid., [jeollanam-do,], prov. zennan, monte yudalsan [mt.], mokpo [city near yudalsan], t. nakai 9377 (para-: not localized); ibid., [jeollanam-do], in silvis genkeimen, t. nakai 9378 (para-: not localized); ibid., [jeollanam-do], in monte insl. chito, t. nakai 9379 (para-: not localized); ibid., [jeollanam-do], in silvis montis semsatsuzan insl. chinto [jindo], t. nakai 9381 (para-: not localized); ibid., [jeollanam-do], in silvis insl. baikato, t. ishidoya and chung 3382 (para-: not localized); ibid., [jeollanam-do], in monte monganzan insl. daikokuzanto, t. ishidoya and chung 3384 (para-: not localized); ibid., [jeju,] quelpaert [jeju-do]: in silvis, e. taquet 6220 (para-: not localized); ibid., [jeollanam-do], kantoku, t. nakai 1439 (para-: not localized); japan, [nagasaki], tsusima [tsushima] in monte sumo-shiratake, t. nakai s.n. (para-: not localized). – note: carpinus coreana is cited as a distinct species in the world checklist (govaerts & frodin 1998), but listed as a synonym of c. turczaninovii in wcsp (2016). nakai described his new species by using the differences of some korean specimens that had been discussed by winkler (1914) as being similar to c. turczaninovii, but not typical. sun et al. (2011) used three accessions in their phylogeny and european journal of taxonomy 375: 1–52 (2017) 30 found one to be almost inseparable from c. turczaninovii [lee s.n. (skk); yoo & wen (2002)], and two being closely related to c. fangiana and c. japonica. although the three accessions could not be examined, it is likely that the first one concerns c. coreana. for the two other accessions no specimen is cited, but they were discussed by sun et al. (2011) as differing by “long hairs on the mid vein, inflorescences, and leaf margin veins”. the placement in the phylogeny, however, rather suggests that those specimens belong to the distegocarpus group, and hence might be misidentified c. japonica or c. cordata. c. coreana var. major nakai, the botanical magazine [tokyo] 40: 162 (nakai 1926). – type: south korea, [hwanghae-namdo], yooshin, prov. kokei, t.h. chung [chung-tyai-hyon] s.n. (holo-: ti). – note: according to the diagnosis, the large leaf and bract size rather suggest stronger affinities to c. tschonoskii than to c. turczaninovii, however this remains speculative without having examined the type. jeon & chang (1997), chang et al. (2014a) and the wcsp (2016) consider this to be a synonym of c. turczaninovii. c. chowii hu, journal of the arnold arboretum 13: 333 (hu 1932). – type: china, hopei, fang-shan, 500 m a.s.l., 5 oct. 1931, h.f. chow 41730 (lecto-, designated here: pe00021883!; isolecto-: a00033750!, pe00021884!, pe00802985!). c. coreana var. multiflora nakai ex kawamoto, handbook of korea-manchurian forestry: 92 (kawamoto 1939). – type: not localized. – note: this name is neither mentioned in the world checklist (govaerts & frodin 1998) nor in the flora of china (li & skvortsov 1999) or the wcsp (2016). jeon & chang (1997) consider is to be synonymous with c. turczaninovii, chang et al. (2014a) list it as invalid name. the present authors did not have the protologue at hand for verification. c. turczaninovii var. arguta uyeki, woody-plants and their distributions in tyôsen: 16 (uyeki 1940). – type: not localized. – note: this name is not mentioned in the world checklist (govaerts & frodin 1998) or the wcsp (2016), but chang et al. (2014a) list it as a invalid name. the present authors did not see the protologue to confirm this. c. turczaninovii var. chungnanensis p.c.kuo, flora tsinlingensis 1 (2): 601 (instituto botanico boreali-occidentali academiae sinicae edita 1974). – type: china, shaanxi, chang-an hsien [终南山, chung-nan-shan; zhongnanshan], 1700 m a.s.l., 15 jun. 1933, h.w. kung 2838 (holo-: wuk0000756, wuk0000764). – note: if necessary, the lectotype should be chosen from one of the two cited specimens, because the flora tsinlingensis is mainly based on material from wuk. as there is no image of the specimens available to the present authors, the designation of a lectotype here does not seem to be appropriate. 43. carpinus viminea lindl. plantae asiaticae rariores 2: 4 (wallich 1831). – type: india, in kamaon, r. blinkworth in n. wallich list no. 2800b (lecto-, designated here: k001117116!; isolecto-: bm000580526!). – additional type material: nepal, in montibus napaliae, n. wallich list no. 2800a (syn-: bm000580527!, e00301177!, e00301178!, k001117115!, k000859917!, k000859918!, m!); india, sirmore [sirmaur], w.a. webb s.n. (syn-: not localized). – note: it is accepted in flora of china (li & skvortsov 1999) with c. viminea var. chiukiangensis as a distinct variety. the kew specimen k000859918 bears a note “napalia wallich 1830” but wallich was in nepal in 1821 (wallich list no. 2800a). most likely, the number has to be regarded as the year when the specimen was received at kew. c. laxiflora var. macrostachya oliv., icones plantarum 20 (4): tab. 1989 (oliver 1891). – c. macrostachya (oliv.) koidz., acta phytotaxonomica et geobotanica 9 (2): 73 (koidzumi 1940). – type: china, hupeh, north patung, a. henry 7013 (lecto-, designated here: k000859943!; isolecto-: bm!, e!, p06811517!). – note: this name is treated as a synonym of c. viminea var. viminea in the flora of china (li & skvortsov 1999) and the flora sichuanica (editorial board of the flora sichuanica 2012). holstein n. & weigend m., taxonomic checklist of carpinus and ostrya 31 c. laxiflora var. davidii franch., journal de botanique [morot] 13: 203 (franchet 1899). – c. davidii (franch.) c.k.schneid., illustriertes handbuch der laubholzkunde 2: 893 (schneider 1912). – type: china, china centralis, prov. kiukiang, in montibus ly chan, sep. 1868 [reverend père, j.p.] a. david 750 (lecto-, designated here: p06811502!; isolecto-: p06811505!, p06811506!, p06811507!). – note: this name is treated as a synonym of c. viminea var. viminea in the flora of china (li & skvortsov 1999) and the flora sichuanica (editorial board of the flora sichuanica 2012). c. fargesii franch., journal de botanique [morot] 13: 202 (franchet 1899). – c. laxiflora var. fargesii (franch.) burkill, journal of the linnean society, botany 26 (178): 501 (forbes & hemsley 1899). – type: china, sutchuen [sichuan], in ditione tchen keou tin [chengkou county], p.g. [reverend père] farges 699 (holo-: p06811515!; iso-: ny00253861!). – note: this name is treated as a synonym of c. viminea var. viminea in the flora of china (li & skvortsov 1999) and the flora sichuanica (editorial board of the flora sichuanica 2012). c. kenpukwan koidz., acta phytotaxonomica et geobotanica 9 (2): 74 (koidzumi 1940). – type: [china, china centralis] hu & chun (1927: 15, tab. 15) [tabula, sub c. laxiflora (siebold & zucc.) blume]. – note: this name is mentioned neither in the flora of china (li & skvortsov 1999) nor in the flora sichuanica (editorial board of the flora sichuanica 2012). c. tehchingensis hu, acta phytotaxonomica sinica 9: 283 (hu 1964). – type: china, yunnan, tehchin hsien, sila, in mixed woods, 2700‒2900 m a.s.l., 18 jul. 1944, k.m. feng [冯国楣] 5593 (holo-: pe00021957!). – note: this name is also synonymous with c. viminea var. viminea according to the flora of china (li & skvortsov 1999) and the flora sichuanica (editorial board of the flora sichuanica 2012). c. viminea var. chiukiangensis hu, acta phytotaxonomica sinica 9: 282 (hu 1964). – type: china, yunnan, chiukiang valley, in wood, 1950 m a.s.l., 31 jul. 1938, t.t. yü 19531 (lecto-, designated here: pe00802369!; isolecto-: a00033771!, e!, pe00802338!, pe00802368!). – note: this name is accepted as distinct variety of c. viminea in flora of china (li & skvortsov 1999). c. fargesii var. latifolia s.y.wang & c.l.chang, journal of the henan agricultural college 14 (2): 6 (ding et al. 1980). – type: china, henan, luanchuan (栾川), 12 aug. 1978, s.y. wang et al. [王遂 义等] 78 0346 (holo-: heac). invalid names c. betulus var. marmoratus de vos, beredeneerd woordenboek der voornaamste heesters en coniferen, in nederland gekweekt: 21 (vos 1867) nom. nud. – note: the name by de vos lacks a valid description, so it is to be ignored under the icn rules (mcneill et al. 2012) but available as a cultivar name (art. 21.6. icncb, brickell et al. 2009): c. betulus ‘marmorata’. according to krüssmann (1951), the name was also published in a späth garden catalogue, but these were not at hand to the present authors for the verification of the validity. c. albopilosa hu nom. nud. – note: no publication of this name is known, name only written on the specimen pe00020172! (china, sichuan, leibo county, 1934, w.y. chun 3942). c. calcarea w.c.cheng nom. nud. – note: this name is only written on a specimen (china, metasequoia area, hubei-sichuan, 1948 m a.s.l., w.c. cheng and c.t. hwa 866: a00033749!, k!). a duplicate in hib (hib0156026) is stored as ostrya japonica. the specimen was also cited under the apparently invalid name c. whilungpaensis w.c.cheng. c. cambodica l.pierre nom. nud. – note: apparently an unpublished name. the specimens are stored in p herbarium, p06811149!, p06811167! and p06810780! (vietnam, crescit ad montem chiao shan in cochinchina, feb. 1877, l. pierre 3300), under c. poilanei a.camus (synonymous to c. londoniana h.j.p.winkl. var. londoniana). european journal of taxonomy 375: 1–52 (2017) 32 c. changhwaensis hu nom. nud. – note: no publication of this name known, only written on the specimen pe00020179! (china, zhejiang, linan, 8 may 1929, s. coll. 363). c. japonica var. auricula maxim. nom. nud. – note: this name is only listed on written specimens bm000580481!, br! and k! (japan, yokohama, 1862, c.j. maximowicz s.n.). c. laifengensis wuzhi nom. nud. – note: apparently an unpublished herbarium name, mentioned in chen (2015). c. mexicana griseb. nom. nud. – note: this name is only written on a specimen in goet! (mexico, w. schaffner pl. mexicanae no. 14) and on two specimens in p, p06810806! (mexico, w. schaffner pl. mexicanae no. 14) and p06810805! (mexico, in montanis mexique orizaba, m.b. [m. botteri?]). despite having the same name, but at a different rank, furlow (1987) does not mention the apparently unpublished grisebach name. c. microbracteata hu nom. nud. – note: there is no publication with this name known to the present authors; it is only written on the specimens bm! and pe00818273! (china, sichuan, baoxing, 1900 m a.s.l., 11 sep. 1936, k.l. chu [曲桂龄] 3824). c. tatarinowii hance nom. nud. – note: there is no publication with this name known to the present authors; it is only written on the specimens p06811225! and p06811224! (china, ad pekinum [giulles], p.g. [reverend père] farges s.n.). checklist of ostrya order magnoliales juss. ex bercht. & j.presl family betulaceae gray subfamily coryloideae hook.f. tribe carpineae a.dc. ostrya scop. flora carniolica: 414 (scopoli 1760) nom. cons. – type: ostrya carpinifolia scop. – note: the name was conserved against ostrya j.hill (hill 1757: 513) [“1756”], which is a publication of a superfluous name for carpinus l. 1. ostrya carpinifolia scop. flora carniolica ed. 2, 2: 244 (scopoli 1772). – o. carpinifolia var. genuina fliche, bulletin de la société botanique de france 35: 166 (fliche 1888) nom. inval. (although not explicitly cited but in context including the type, see art. 24.3, mcneill et al. 2012). – o. italica subsp. carpinifolia (scop.) h.j.p.winkl., das pflanzenreich iv 61 (heft 19): 22 (winkler 1904). – carpinus ostrya l., species plantarum: 998 (linnaeus 1753) p.p. – original citation: “italia, virginia.” – type: s. loc., s. coll. (lecto-, designated by iamonico & reveal (2012: 866): linn! ex herb. linn. no. 1131.4). – additional original material: s.d. (bm000647418! ex herb. clifford 447, carpinus 2). o. vulgaris willd., species plantarum ed. 4, 4 (1): 469 (willdenow 1805) nom. illegit. superfl. – note: willdenow cites o. carpinifolia as synonym, so his name is superfluous. o. italica p.micheli ex spach, annales des sciences naturelles, botanique ser. 2 16: 246 (spach 1841b) nom. illegit. superfl. – o. italica scop., flora carniolica: 414 (scopoli 1760) nom. inval. (polynomen), pro syn. – carpinus italica steud., nomenclator botanicus ed. 2, 1: 300 (steudel 1841) nom. inval. pro syn. – o. ostrya var. italica (p.micheli ex spach) c.k.schneid., illustriertes holstein n. & weigend m., taxonomic checklist of carpinus and ostrya 33 handbuch der laubholzkunde 1: 143 (schneider 1904) nom. inval. (tautonym). – note: ostrya italica scop. is rather a misinterpretation as scopoli discusses only genera and as he did not adopt linnaeus’ binomials in this work. scopoli’s name might better be read as polynomial “ostrya italica, carpini folio liongiore &? breviore.” that he took from micheli (1729: 223) as one of the names used for the genus ostrya. spach accepted o. italica in 1841, but he cited o. carpinifolia in synonymy. one year later, spach refers to the species as o. vulgaris willd. steudel refers to an unpublished name, or used by scopoli, “carpinus italica” (or mistaken by steudel) as a synonym of o. vulgaris. o. cylindrica friv., a’magyar tudós társaság évkönyvei 3 (3): 163 (frivaldszky von frivald 1837) nom. nud. – note: the name also appears on a specimen (wu!) from macedonia. o. carpinifolia var. alpestris ces., saggio su la geografia botanica e su la flora della lombardia: 61 (cesati 1844). – type: not localized (ro?). – note: there is a specimen by cesati from varenna, lombardy, italy (p06751764!). it was annotated by him as “ostrya carpinifolia β collina mihi”. eventually, cesati changed his mind about the epithet upon publication of his new variety. o. ladelcii sanguin., flora romanae prodromus alter: 818 (sanguinetti 1864). – original citation: “in sylvaticis latii, frascati, albano.” – type: not localized (ro?). o. carpinifolia var. corsica fliche, bulletin de la société botanique de france 35: 166 (fliche 1888). – type: france, corsica, forêt communale de pietrosa, may 1885, p. fliche s.n. (syn-: not localized). france, corsica, forêt de tova, may 1885, p. fliche s.n. (syn-: not localized); ibid., forêt de bavella, mathieu s.n. (syn-: not localized); ibid., plateau de nebbio, mathieu s.n. (syn-: not localized). o. carpinifolia var. lanceolata kem.-nath., trudy tbilisskogo botaniceskogo instituta 2: 131 (kemularianathadze 1937). – type: georgia, prov. kutais, distr. letschchum, fauces fl. ladjanura loc. orpiri, prope pag. atscharis-chidi ad rupes calcareas, 22 may 1929, l.m. kemularia-nathadze s.n. (syn-: not localized); ibid., 19 oct. 1932, l.m. kemularia-nathadze s.n. (syn-: not localized). 2. ostrya chinensis i.m.turner annales botanici fennici 51: 308 (turner 2014). – o. multinervis rehder, journal of the arnold arboretum 19: 71, pl. 217 (1938) nom. illegit., non ettingshausen (1868) (later homonym). – type: china, hunan, ma-ling-tung, sinning hsien, in mixed forest on slope, 650 m a.s.l., 13 oct. 1935, c.s. fan and y.y. li 605 (holo-: a00033789!; iso-: bm001191567!, nas00286610!, l!, p06811043!). 3. ostrya japonica sarg. the silva of north america 9: 32 (sargent 1896). – o. ostrya var. japonica (sarg.) c.k.schneid., illustriertes handbuch der laubholzkunde 1: 143 (schneider 1904) nom. inval. (tautonym). – type: japan, hokkaido, hill near sapporo, 18 sep. 1892, c.s. sargent s.n. (syn-: gh00055441!); ibid., oiwagi, 1861, c.j. maximowicz s.n. (syn-: bm!, br!, k000859914!, l!, m-0265487!, p06809338!, p06811038!, w1886–1595!); ibid., nambo, 1865, s. tschonoski s.n. (syn-: k0000859913!, p06811037!); ibid., in sylvis frondosis circa lacum konoma (syn-: not localized); cult. in arnold arboretum from seeds sent from japan by h. mayr (syn-: not localized). – note: the name o. japonica sarg. was only accepted by the author in the 1896 publication. before, sargent (1893, 1894) initially expressed certainty on the distinctness “in the forests of yezo i felt no doubt of its specific distinctness […] all, things considered, it is, perhaps, best to consider the japanese tree as specifically distinct”. however, just in the next sentence, he weakened his opinion stating “only after it has been grown here during many years side by side with the american species will it be possible to reach any opinion on this subject worthy of much consideration. if it proves to be distinct it should bear the name of ostrya japonica”. according to art. 36.1. b (mcneill et al. 2012), the two early publications (sargent 1893, 1894) cannot be considered as a valid publication of o. japonica. in 1896, however, sargent accepted the japanese collections as fully distinct and formalized the name. european journal of taxonomy 375: 1–52 (2017) 34 o. carpinifolia f. japonica schelle, handbuch der laubholzbenennung: 49 (beissner et al. 1903) nom. nud. o. liana hu, journal of the arnold arboretum 11: 49 (hu 1930). – type: china, chihli, eastern tomb, 15 sep. 1929, c.f. li timber no. 3 [field no. 3, herbarium number 15] (lecto-, designated here: pe00021986!; isolecto-: a00033788!). o. japonica var. homochaeta honda, the botanical magazine [tokyo] 47: 433 (honda 1933). – type: japan, honshu, nikko, prov. shimotsuke, 1889, m. shirai s.n. (holo-: ti). – additional type material: ibid., 1931, h. itoo s.n. (para-: not localized); ibid., prov. bicchu, takahashi, 1915, g. koidzumi s.n. (para-: not localized); ibid., hokkaido, in monte moiwa, circ. sapporo, 1893 g. yamada (para-: not localized); south korea, [seogwipo,] in monte hallaisan, ins. saishuutoo, 1917, e.j. taquet 4439 (para-: e00388817); ibid., [jeju,] ins. saishuto [jeju-do, quelpaert], 1917, t. nakai 4905 (para-: not localized); ibid., [haenam], in monte daikoji, prov. zenra-nandoo, 1921, t. nakai 9384 (para-: not localized). o. virginica var. japonica sarg., garden and forest 6: 383 (sargent 1893) nom. nud. – note: sargent wrote that maximowicz (1881a) had named the japanese ostrya collections as var. japonica, but there is no such name mentioned in the cited maximowicz paper. 4. ostrya knowltonii coville garden and forest 7: 114 (coville 1894). – o. carpinifolia f. knowltonii (coville) schelle, handbuch der laubholzbenennung: 49 (beissner et al. 1903). – type: usa, arizona, yavapai co., grand canon of the colorado river, 10 jul. 1892, j.w. toumey 272 (holo-: us00130392!; iso-: a00263686!). o. baileyi rose, contributions from the united states national herbarium 8 (4): 293 (rose 1905). – type: usa, texas, guadalupe mountains, 19 aug. 1901, v. bailey 435 (holo-: us00089317!). o. chisosensis correll, wrightia 3 (7): 128 (correll 1965). – o. knowltonii subsp. chisosensis (correll) a.e.murray, kalmia 12: 22 (murray 1982). – o. knowltonii var. chisosensis (correll) a.e.murray, kalmia 13: 10 (murray 1983). – ostrya virginiana var. chisosensis (correll) henrickson ex a.m.powell, trees and shrubs of the trans-pecos and adjacent areas: 75 (powell 1998). – type: usa, texas, brewster co., chisos mts, big bend np, base of north-facing ledges, emory peak, 17 jun. 1964, d.s. correll 29733 (holo-: ll00370396!; iso-: c10009751!, gh00033786!, ll00370395!, mexu00122440!, mexu00711117!, mich1192171!, ncu00000363!, tex, us00089316!). – additional type material: usa, texas, high shady crevice in rock bluff, near nail place, chisos mts., 21 aug. 1915, m.s. young s.n. (para-: srsc3716!, tex00378112); ibid., north of crown mt, chisos mts, 1 jul. 1937, b.h. warnock 21633 (para-: srsc3715!, srsc3721!, tex00378110); ibid., igneous soil on emory-boot trail, big bend np, chisos mts, 7500 ft, 8 jun. 1952, b.h. warnock 10478 (para-: ll00378111, srsc3718!); ibid., along a trail from boot spring and basin, chisos mts, north side of emory peak, big bend np, 15 jul. 1955, b.h. warnock 12726 (para-: ll00378109, srsc3714!). – note: ostrya chisosensis was recognized in flora of north america (furlow 1997). however, its taxonomic status has been challenged recently (mccauley & paces 2015). 5. ostrya rehderiana chun journal of the arnold arboretum 8 (1): 29 (chun 1927). – type: china, tien moh shan, 400 m a.s.l., fairly common in open woods, 2 oct. 1925, r.c. ching 3385 (lecto-, designated here: a00033790!; isolecto-: n, us00089314!). 6. ostrya trichocarpa d.fang & y.s.wang guihaia 3 (3): 189 (fang & wang 1983). – type: china, guangxi, napo county (那坡县), 百都公社, 弄化, 1260 m a.s.l., 石山 [rocky hill], 23 apr. 1981, d. fang, t.-h. tan and z.-g. wang [方鼎, 覃 holstein n. & weigend m., taxonomic checklist of carpinus and ostrya 35 德海, 王振刚] 22412 (holo-: gxmi050301). – additional type material: ibid., d. fang, m.-x. lai et al. 25379 (para-: gxmi); ibid., guanxi zhuang auton. region, jinxi county, 780 m a.s.l., 17 nov. 1962, z.-j. li and h.-q li 1650 (para-: ibk00079836!, ibk00079837!). 7. ostrya virginiana (mill.) k.koch dendrologie 2 (2): 6 (koch 1873). – carpinus virginiana mill., the gardeners dictionary ed. 8, no. 6 (miller 1768). – zugilus virginica raf., florula ludoviciana: 159 (rafinesque 1817) orth. var., nom. illegit. superfl. – o. ostrya var. virginiana (mill.) c.k.schneid., illustriertes handbuch der laubholzkunde 1: 143 (schneider 1904) nom. inval. (tautonym). – type: [cult. garden specimen], 1740, chelsea physic garden 938 (lecto-, designated by reveal (1990): bm001041898!). – note: in his protologue, miller only cited a plukenet (1692) drawing, which does not correspond well to what is today considered as o. virginiana. reveal (1990) found a matching typotype (h.s. 95:143, bmsl) that turned out to be o. carpinifolia scop., and three specimens from that era that miller might have had seen (but without actual evidence) of what is now considered o. virginiana. nevertheless, the proposal to conserve the name with a new type was rejected, and it was decided that one of the specimens should serve as lectotype (brummitt 1993). 7a. ostrya virginiana (mill.) k.koch subsp. virginiana carpinus virginica münchh., der hausvater 5 (1): 120 (münchhausen 1770) orth. var. – o. carpinifolia var. virginica (münchh.) fliche, bulletin de la société botanique de france 35: 166 (fliche 1888). – note: münchhausen misspelled the name given by miller. carpinus triflora moench, methodus: 694 (moench 1794). – type: not localized. – note: the protologue does not state any locality, and moench’s herbarium is destroyed (stafleu & cowan 1976–1988). o. americana f.michx., histoire des arbres forestiers de l’amérique septentrionale 3 (1): 54 (michaux 1812) nom. inval. – associated material: a canada ad carolinam montosam, s. coll., s.n. (p06810885!). – note: while michaux briefly mentions an “ostrya americana”, he apparently does not consider it to be a species different from “carpinus ostrya” (ostrya carpinifolia). the name is not mentioned in the index either, so the name is rather to be interpreted as meaning the plants from america. the associated material by michaux also suggests that. o. virginica var. eglandulosa spach, annales des sciences naturelles, botanique ser. 2 16: 246 (spach 1841b). – type: cult. at trianon garden, 1833, s. coll., s.n. (lecto-, designated here: p06811525!). – note: the specimen designated as lectotype here bears a note by spach as author “nob.” after the name. o. virginica var. glandulosa spach, annales des sciences naturelles, botanique ser. 2 16: 246 (spach 1841b). – o. virginiana var. glandulosa (spach) sarg., botanical gazette 67: 216 (sargent 1919). – o. virginiana f. glandulosa (spach) j.f.macbr., publications of the field museum of natural history, botanical series 4 (7): 192 (macbride 1929). – type: cult. at h. p. [hortus parisiensis], 1833, s. coll., s.n. (lecto-, designated here: p06809319!). – note: the specimen designated as lectotype here bears a note by spach as author “nob.” after the name. o. ostrya (l.) macmill., the metaspermae of the minnesota valley: 287 (macmillan 1892) nom. inval. (tautonym), p.p. – note: macmillan referred only to the virginian part of linnaeus’ carpinus ostrya l., but he produced a tautonym, which is not valid according to art. 23.4 of the icn (mcneill et al. 2012). o. virginiana var. lasia fernald, rhodora 38: 414 (fernald 1936). – o. virginiana subsp. lasia (fernald) a.e.murray, kalmia 12: 22 (murray 1982). – type: usa, florida, columbia co., lake city, 11‒19 jul. 1895, g.v. nash 2158 (holo-: gh00033787!; iso-: e00870598!, k!, p06810598!, p06810607!, us00089313!, wu!). – additional type material: usa, virginia, henrico co., richmond, 5 may 1894, j.r. churchill s.n. (para-: not localized); ibid., princess anne co., rich dry woods, little neck, european journal of taxonomy 375: 1–52 (2017) 36 m.l. fernald et al. 4627 (para-: not localized); ibid., norfolk co., dry rich woods, east of gertie, m.l. fernald et al. 4628 (para-: not localized). 7b. ostrya virginiana subsp. guatemalensis (h.j.p.winkl.) a.e.murray kalmia 13: 10 (murray1983). – o. italica var. guatemalensis h.j.p.winkl., das pflanzenreich iv 61 (heft 19): 22 (winkler 1904). – o. guatemalensis (h.j.p.winkl.) rose, contributions from the united states national herbarium 8 (4): 292 (rose 1905). – o. virginiana var. guatemalensis (h.j.p.winkl.) j.f.macbr., publications of the field museum of natural history, botanical series 4 (7): 193 (macbride 1929). – type: guatemala, volcán de fuego, 6400 ft, 7 aug. 1873, o. salvin s.n. (lecto-, designated here: w1886–8378!; isolecto-: k!). – additional type material: guatemala, j. donnell smith in donn. sm. pl. guatem. 2470 (syn-: k!); ibid., san miguel uspantán, depart. quiché, 6000 ft, apr. 1892, e.t. heyde and e. lux in donn. sm. pl. guatem. 2910 (syn-: k!, m-0265489!, mo-188762!, wu!); “guatemala und costa rica” (winkler 1904), j. von warszewicz 55 (syn-: not localized). mexico, orizaba, aug. 1853, f. müller 1526 (syn-: w0065339!); mexico, jalapa, 23 may 1899, c.g. pringle 8188 (syn-: bm000993542!, br0000005269754!, e00870592!, f, goet!, k!, m-0265488!, manch!, p06810561!, p06810571!, p06810594!, ph00020034!, us00898166, uvmvt221269!, w1900–3036!, wu!). – note: the lectotype was chosen to be the salvin specimen in w because it is the only guatemalan specimen that bears a note by winkler. the pringle 8188 specimen in w also bears a winkler note but rose separated it to his o. mexicana rose. choosing this specimen as lectotype would make plenty of isolectotypes, but a new name would become necessary in case that the mexican and the guatemalan specimens turn out to be different after all. o. mexicana rose, contributions from the united states national herbarium 8 (4): 292 (rose 1905). – type: mexico, guerrero, omilteme, 25 may 1903, e.w. nelson 7050 (lecto-, designated here: us00089315!; isolecto -: ny00253871!, ny00253872!). – additional type material: mexico, orizaba, m. botteri 284 (syn: goet!, us00898164); ibid., jalapa, 23 may 1899, c.g. pringle 8188 (syn-: bm000993542!, br0000005269754!, e00870592!, f, goet!, k!, m-0265488!, manch!, p06810561!, p06810571!, p06810594!, ph00020034!, us00898166, uvmvt221269!, w1900–3036!, wu!). 8. ostrya yunnanensis w.k.hu acta phytotaxonomica sinica 17 (1): 87 (li 1979b). – type: china, yunnan, lu-chuan hsien, in thin forest, 2600 m a.s.l., 5 dec. 1938, p.y. mao [mao pin-yi] 1935 (lecto-, designated here: pe00021988!; isolecto-: kun446238, kun446239, pe00734260!, pe00734261!). discussion our finding that 78 basionyms are missing in major floristic treatments, 46 (plus 3 likely too late to be included) in the world checklist and the bibliography of fagales (govaerts & frodin 1998), and 32 in the wcsp (2016) alone is thought-provoking. most of the basionyms that have not been treated in the checklists are below species level and none was considered as accepted upon publication. floristic treatments draw a different picture though, although it must be noted that full synonymy is often not given for stylistic reasons. however, missing four accepted species in the flora of china should be alarming given its importance as a major tool for species identification. most of the basionyms that have been missed, in general, are likely explained due to the unavailability of the protologues, mainly published in literature from the former soviet union, china, and japan. this is also most likely the explanation for the overlooked species in the flora of china. the fact that ipni did not record infraspecific names before 1971 contributes to the gap. the old-fashioned concept of formalizing minor morphological differences in new taxon names (endersby 2009) led to a multitude of names that needs to be revised. holstein n. & weigend m., taxonomic checklist of carpinus and ostrya 37 page (2016) coined the term “dark texts” for literature that is not digitally available. however, that term can be expanded to all literature that is unavailable, even if the obstacle is formed by language or the nescience or unavailability of the hard-copy literature. limited access to material in collections far away from the natural origin, and limited access to, or knowledge of existing literature were – and still are – pitfalls for taxonomy. with new taxa being described constantly, overlooked names are a source of error contributing to the instability of names. this is demonstrated by the fact that the name ostrya multinervis had already been published for a fossil species (turner 2014) or by the description of a new ostryopsis intermedia b.tian & j.q.liu although a valid basionym, corylus davidii var. cinerascens franch. was already available (holstein & weigend 2016). the immense efforts to digitize literature and herbaria during the last years are an enormous help for taxonomic work (knapp et al. 2002; wheeler 2004, godfray 2007; turner 2014), but there is a need for 1) an expansion to publications in non-latin script, and 2) from a structured review of old literature. the former, then, would profit if the script were not only readable, but also markable for automatic translation. this is of special relevance for texts before 1935 for the question of validity (art. 39.1, mcneill et al. 2012). overlooking names due to limited access to literature or nescience is critical not only for the sake of completeness, but because it may lead to overlooking potentially endangered taxa and underestimating biodiversity. taxon differences are hypotheses that are tested traditionally using morphological characters, and rejection of such hypotheses ends in synonymization of the applied names (gaston & mound 1993; holstein & luebert 2017). uncritical adoption of synonymizations might ease taxonomic and floristic work at first, but may well be the source of incorrect subsequent taxonomical decisions. when synonymizations were made, material availability may have been limited or type material may not have been accessible. critical revision including new data is therefore important (wheeler 2004). furthermore, adopting synonymy and ignoring infraspecific taxa appears to be justified, but this is only true if species are regarded as a homogeneous entity. those who rely on the recognition of heterogeneity within species, such as population geneticists, plant breeders or phytochemists may beg to differ. overlooking infraspecific taxa should therefore not be belittled (nixon & wheeler 1990). the percentage of non-cultivar synonyms in relation to accepted basionyms is 66.8% in carpinus and 62% in ostrya, both being on the lower spectrum of the numbers found by scotland & wortley (2003) for a range of other taxa. however, the number of synonyms between the accepted taxon names greatly differs among the carpinus taxa. while another 14 names represent cultivars of c. betulus, many synonymous basionyms belong to wide-spread species. some of them were likely created due to limited access to material, thus overemphasizing the differences, e.g., in the treatments of winkler (1904, 1914) and partly of hu (1929, 1930, 1931, 1932, 1933). although iwasaki et al. (2010) report only a few instances of hybridization between c. cordata and c. japonica, and between c. tschonoskii and c. laxiflora, some of the synonyms, also in c. turzaninovii, might be explained by hybridization, or introgression. the five names under c. × schuschaensis may represent either natural variability or different degrees of introgressive hybridization between c. betulus and c. orientalis, which had already been suggested in the flora of the ussr (bobrov 1936). a good number of synonyms, however, was described in regional treatments or even outside a wider taxonomic treatment. an exaggerated focus on local taxa ignoring the species outside of the respective province or country borders carries the risk of overemphasizing local variability and describing more wide-spread taxa repeatedly (mabberley 1991; scotland & wortley 2003; michelangeli 2005). the aim to produce floras (target 1 of the gspc), which usually focus on regional to national scales (funk 2006), might thus create problems, since the whole scope of a species is ignored. the iucn red list (international union for conservation of nature and natural resources; status from 29 march 2016) lists 27 species of carpinus with 13 of them classified as “data deficient” and 2, c. putoensis and c. tientaiensis, as “critically endangered”. shaw et al. (2014) also regard c. hebestroma european journal of taxonomy 375: 1–52 (2017) 38 as critically endangered, whereas it is listed as “not assessed” online. for ostrya, the red list gives data for only six species with o. chisosensis and o. rehderiana listed as critically endangered. as there are at least 42 accepted species plus 17 infraspecific taxa of, partly, poorly known distribution in carpinus alone, the conservation status is clearly very insufficiently documented for this genus. changing this situation may not be easy, since the acceptance status differs among the current treatments and checklists in 21 of the carpinus names, with four species missing in the flora of china. the stability of taxon status and recognizability of taxa are, however, crucial for conservation, comparison, and monitoring (mace 2004). the fact that revisionary work is important for conservation has also been shown elsewhere, for example in juncaceae and potamogetonaceae (kirschner & kaplan 2002) or in cotula (powell et al. 2014). target 2 of the global strategy for plant conservation (gspc) of the convention of biological diversity (cbd, https://www.cbd.int) aims at an assessment of the conservation status of all known plant species. meeting target 2, however, not only requires an assessment of the taxonomic status as-is, but also critical revisions. one of the crucial tasks of taxonomic revision, being the localization and identification of type material is tackled here for carpinus and ostrya. this is, however, not always straight-forward, and collections are often wide-spread among different herbaria. the valuable information on where potential type collections are located in taxonomic literature ii (stafleu & cowan 1976–1988) help to prioritize herbarium visits, but may leave many collections being ignored for a long time. we found that many types of most (in case of ostrya even all) taxa are already available in silico, but many of them have not been marked as type. completely (or nearly so) digitized collections, such as the ones of the muséum national d’histoire naturelle (paris), naturalis (l, u, wag), and the chinese virtual herbarium (pe and partly others) were therefore very convenient. digitizing type specimens may be a priority and ease taxonomic work quickly (berents et al. 2010), but complete digitization is inevitable. our finding of the unrecognized type of c. orientalis in bm among the specimens in the general collection may serve as a good example, apart from our result that all surveyed herbaria contained hitherto unrecognized type material. digital collections have been confirmed as a useful low-cost tool (steudel et al. 2012), but they also save time and help preserving specimens thanks to the reduced risks of transportation and handling damage (friis 2017). however, information on the history of the collectors and collections, usually the expertise from curatorial staff, is a not to be underestimated asset. premature decisions, such as inappropriate lectoor neotypifications due to only partly digitized collections may become problematic, just as ongoing taxonomic changes or novelties due to unavailability of (even more modern) literature and physical material. we therefore consider open and easy access to fully-digitized collections to be not only cost-effective in the long-term but also a prerequisite for correct taxonomic work, especially since digital collections are being used more and more. conclusion despite the great efforts of digitization, we still find yet undiscovered taxon names in carpinus and ostrya, and even recent floristic treatments miss taxa. this is especially critical if publications are hardly accessible and in languages that are not widely understood, which then may explain the limited accessibility in libraries. the loss of names, and therefore hypotheses for taxa, due to oversight of littleknown publications, language barriers, and uncritical synonymization, contributes to future taxonomic instability or overlooked biodiversity. regional treatments ignoring taxa from neighbouring countries might do more harm than good by adding more (superfluous) names. the urgency to inventory biodiversity is without dispute, but it should not go without critical revisionary work. digital collections turn out to be very valuable for this work, but with many types not recognized as such, partial digitization might give the dangerous illusion of taxonomic work already completed, where it has not even really started. holstein n. & weigend m., taxonomic checklist of carpinus and ostrya 39 https://www.cbd.int acknowledgements we are grateful to the various literature and herbarium digitization projects that enable the easy access to the resources, and to the curators and staff of b, bm, br, e, goet, k, liv, m, manch, msb, w, and wu for making the collections available. we also thank adam boratynski, hans-joachim esser, tilo henning, qi lin, zurab manvelidze, sergei l. mosyakin, alexander sukhorukov, genevieve k. walden, and the library staff of b, bm, and k for their help in obtaining literature. thanks are given to federico luebert for discussions on taxonomy, tianyun liu for translation help from chinese language, to two anonymous reviewers and to alejandro quintanar sánchez for additions and helpful comments. this research received support (visit to e collection) from the synthesys project (http://www.synthesys.info/), which is financed by the european community research infrastructure action under the fp7 “capacities” program. references aiton w. 1789. hortus kewensis. vol. 3. printed for george nicol, bookseller to his majesty, pall mall, london. https://doi.org/10.5962/bhl.title.4504 akiyama s., thijsse g., esser h.-j. & ohba h. 2013. siebold and zuccarini’s type specimens and original material from japan, part 2. angiosperms. dicotyledoneae 1. journal of japanese botany 88: 346‒377. ashburner k., mcallister h.a. & rix m. 2013. the genus betula: a taxonomic 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