01_Korneyev_01_22.indd UDC 595.773.4(494) A NEW SPECIES OF RHAGOLETIS (DIPTERA, TEPHRITIDAE) FROM SWITZERLAND, WITH DISCUSSION OF ITS RELATIONSHIPS WITHIN THE GENUS S. V. Korneyev1,2,3,4, J. J. Smith2, D. L. Hulbert2, J. E. Frey5, V. A. Korneyev1 1Schmalhausen Institute of Zoology NAS of Ukraine, vul. B. Khmelnytskogo, 15, Kyiv, 01030 Ukraine E-mail: s.v.korneyev@gmail.com; valery.korneyev@gmail.com 2Michigan State University, Department of Entomology, Natural Sciences Building, 288 Farm Lane, East Lansing, MI 48824, USA E-mail: jimsmith@msu.edu; dhulbert87@gmail.com 3University of California, Davis, Department of Entomology and Nematology, College of Agricultural and Environmental Sciences, One Shields Avenue, Davis, California, 95616 USA 4California Department of Food and Agriculture, Plant Pest Diagnostics Branch, 3294 Meadowview Road, Sacramento, California 95832-1448 USA 5Federal Department of Economic Aff airs, Education and Research EAER, Mueller-Th urgau-Strasse 29, 8820 Waedenswil, Switzerland E-mail: juerg.frey@agroscope.admin.ch S. V. Korneyev (https://orcid.org/0000-0001-8599-7695) J. J. Smith (https://orcid.org/0000-0003-1492-3095) J. E. Frey (https://orcid.org/0000-0001-6628-8834) V. A. Korneyev (https://orcid.org/0000-0001-9631-1038) urn:lsid:zoobank.org:pub:F742168B-14A5-4286-BD2D-D68320E6D26F A New Species of Rhagoletis (Diptera, Tephritidae) from Switzerland, with Discussion of its Relationships within the Genus.  Korneyev, S. V., Smith, J. J., Hulbert, D. L., Frey, J. E., Korneyev, V. A. — Rhagoletis merzi sp. n., is described and illustrated based on specimens swept and reared from Juniperus sabina L. in Switzerland. A comparative review of Palaearctic species and a key to Palearctic and Nearctic species similar to R. merzi is provided. Based on DNA sequences from the COI, CAD, 28S, period, and AATS genes (4270 bp) of 92 isolates from two outgroup species (Anastrepha ludens, Euphranta canadensis), one species of Carpomya and 35 species representing most of species groups of Rhagoletis, a MrBayes analysis recovered a monophyletic lineage of Juniper-infesting species within a monophyletic cluster of R. fausta, R. batava, as well as the suavis, cingulata, pomonella, tabellaria and juniperina groups. Th e juniperina group includes both Nearctic (R. juniperina and undescribed forms) and Palaearctic species (R. fl avigenualis and R. merzi). Rhagoletis merzi is more similar to the Nearctic R. juniperina in both morphological characters (wing pattern, occiput, mesonotum and legs coloration, shape of male surstyli) and molecular sequences than to the Palearctic R. fl avigenualis. K e y w o r d s : phylogeny, multigene DNA analysis, juniper. Introduction Th e genus Rhagoletis Loew, 1862 belongs in the tribe Carpomyini and includes more than 75 described species of the true fruit fl ies occurring mostly in the Holarctic and Neotropical Regions and, to the lesser de- gree, in the Oriental Region (Smith & Bush, 1999; Korneyev & Korneyev, 2019). Many Rhagoletis species are economically important, including pests of apples, cherries, blueberries, and walnuts (Boller & Prokopy, 1976). Two pest species, Rhagoletis cingulata (Loew, 1862) (the eastern cherry fruit fl y) and R. completa (Cresson, 1929) (the walnut husk fl y) have been introduced from North America to Europe (Merz, 1991), and R. cerasi (Linnaeus, 1758) (the European cherry fruit fl y) has recently been introduced to North America. Zoodiversity, 56(1): 1–20, 2022 DOI 10.15407/zoo2022.01.001 Fauna and Systematics 2 S. V. Korneyev, J. J. Smith, D. L. Hulbert, J. E. Frey, V. A. Korneyev Larvae of most Rhagoletis species infest fl eshy fruits of various angiosperm plants, except a few species whose larvae feed in gymnosperm fl eshy cones or galbuli (commonly called “juniper berries”). Rhagoletis species are oligo- or monophagous, specialized frugivores, oft en infesting a single host species. Host plant specialization and fi delity are important aspects of Rhagoletis biology (Bush, 1966). Th erefore, studying the patterns of host use and host shift s in Rhagoletis is an important part of fundamental evolutionary research and valuable from an economic perspective. A host shift to an economically important crop from a native host has been documented for Rhagoletis pomonella (Walsh) from its native host plant Crataegus spp. (hawthorn) to introduced commercial apples (Ma- lus domestica) in the last ~170 years (Walsh, 1867; Bush, 1969). Such host plant shift s in Rhagoletis have led to the formation of host races, the hypothesized initial stage of speciation-with-gene-fl ow, and have provided an opportunity to study the fundamental nature of population divergence and the formation of new species, especially in the R. pomonella species complex (Bush, 1969 and many other publications), which has become a textbook example of speciation in action (Schluter, 2000; Coyne & Orr, 2004; Nosil, 2012). Rhagoletis species have been classifi ed taxonomically into defi ned species groups beginning with the semi- nal work of Bush (1966), in which the pomonella, tabellaria, suavis, cingulata and ribicola species groups were proposed for classifi cation of most North American Rhagoletis species based on analyses of their morphological characteristics. Analyses based on morphology (Jenkins, 1996), allozymes (Berlocher & Bush, 1982), morphol- ogy and mitochondrial DNA (Smith et al., 2006) further defi ned the species groups, but did not resolve relation- ships among these groups. Th e phylogenetic relationships among these fi ve Rhagoletis species groups remained unresolved until Hamerlinck et al. (2016) demonstrated that the pomonella and tabellaria species groups are sister groups, and more recently Hulbert (2018) produced a more detailed and well supported phylogeny indi- cating the relationships among all fi ve species groups. Many of the Palearctic Rhagoletis species were grouped by Kandybina (1977) into the cerasi, juniperina, alternata and meigenii species groups, originally based on larval characters (she also provided refi ned diagnoses of the suavis, cingulata, and pomonella species groups); see Kandybina (1961, 1962, 1972), Kandybina & Richter (1976), and Richter & Kandybina (1997) for larval descriptions. Th ese Palearctic Rhagoletis species were alter- natively placed into the cerasi, fl avicincta, meigenii, and zernyi species groups by Smith et al. (2006) based on mitochondrial COII and adult morphological characters. Th e Palearctic Rhagoletis species (including those originally described in Zonosema, Megarrhagoletis, and Microrhagoletis, now considered synonyms of Rhagoletis) comprise more than 30 described taxa, most of which were reviewed and keyed by Rohdendorf (1961). Later, additional species were described by Jermy (1961), Kandybina (1972), Richter (1974), Kandybina & Richter (1976), Richter & Kandybina (1997), and Korneyev & Merz (1997). Th e Middle Asian, Far East Palearctic, and Oriental species of Rhagoletis and the closely related Carpomya were keyed by Korneyev & Merz (1997), Korneyev & Ovchinnikova (2004), Ito (2011), whereas Fre- idberg & Kugler (1989), Merz (1994), Korneyev (1997), and Mohamadzade Namin & Rasoulian (2009) treated the European, Middle Eastern, and Caucasian species. Korneyev et al. (2018 a) provided a key for the species of Europe, Asia Minor, Caucasus, and Near East, including Iran, along with references to each species, data on known host plants and distribution, as well as some taxonomic remarks. Recently, Korneyev & Korneyev (2019) provided a key to the Asian (except eastern) species. Although not all non-Nearctic species were included, recent work using a Bayesian phylogenetic and Maximum Likelihood analysis based on DNA sequences at fi ve loci (Hulbert, 2018) recovered a well-supported monophyletic group containing all of the Nearctic Rhagoletis species that belong to the fi ve species groups named by Bush (1966), plus two unplaced Nearctic species (R. fausta and R. juniperina), as well as the Palearctic species R. batava and R. fl avigenualis (Hulbert, 2018). Mitochondrial COII sequences had earlier suggested that R. batava and R. fl avigenualis might have taxonomic affi nity to the “core Nearctic taxa” (Smith et al., 2006). Additionally, Hulbert (2018) supported the monophyly of a group within this larger clade consisting of the two juniper-infesting species, the Nearctic R. juniperina Marcovitch and the Palearctic R. fl avigenualis Hering. Th us, our working hypothesis is that all of the Juniperus-infesting species (R. juniperina, R. fl avigenualis, R mongolica Kandybina, and R. zernyi Hering) comprise a monophyletic group, based on these initial DNA sequences and the very similar morphological characters and host plant preferences of these species. At this point in time, however, DNA sequences from R. mongolica and R. zernyi are lacking to test this hypothesis. In this paper, we determined the position of a newly described species from Switzerland, R. merzi sp. n., and its relationships with the other Rhagoletis species. We provided comparison of its morphological characters with the species occurring in the Palaearctic Region, and DNA sequences at fi ve genetic loci using a wide sample of other representatives of the genus. Material and methods Th e following acronyms refer to collections housing specimens: MHNG: Muséum d’Histoire Naturelle, Geneva, Switzerland; MSU: Michigan State University, East Lansing, MI, USA; SIZK: I. I. Schmalhausen Institute of Zoology, National Academy of Sciences of Ukraine, Kyiv, Ukraine; SMNS: Staatliches Museum für Naturkunde Stuttgart, Germany. 3A New Species of Rhagoletis (Diptera, Tephritidae) from Switzerland… Morphological terminology generally follows White et al. (1999) except for the wing venation, which fol- lows Cumming & Wood (2017). Wing bands are labeled on fi gure 1 a. Measurements are given in mm. Body length of females includes the oviscape. Genitalia were prepared for study using the following procedure: the abdomen was excised from a relaxed specimen, cleared in NaOH solution (10 %) for 2 hours at 90–95 °C, and then washed in distilled water. Genitalia were examined in a drop of glycerine on a microscope slide with a depression or under a glass cover slip. Detached parts are stored in polypropylene microvials containing glycerine pinned together with a specimen. Structures were measured with an ocular micrometer. Wing and habitus photographs were taken using a Canon PowerShot A640 camera connected to a Zeiss Stemi C-2000 (SIZK) microscope or using a Leica DFC 490 camera mounted on a Leica Z16 APO A microscope (MNKB). Photographs of genitalia were taken using a Nikon Coolpix P50 camera through the eyepiece of a Wild M11 light microscope. Digitized photographs were stacked using CombineZM® (Hadley, 2007) and Helicon Focus®. Some photos of R. merzi were taken at Michigan State University using a Visionary Digital Passport II system (Dun, Inc., Palmyra, Virginia, USA). Photographic stacks were taken with a Canon EOS 5D Mark II equipped with a 65 mm Canon Macro lens (Canon Inc., Tokyo, Japan) and Dynalite MH2015 Road Flash Heads (Dynalite Flash Equipment, Union, New Jersey, USA) mounted onto a Stack Shot Macro Rail system (Cognisys, Inc, Traverse City, Michigan, USA) and montaged in Helicon Focus 7.5.6 (Helicon Soft , Kharkiv, Ukraine). Postprocessing was performed with Adobe Photoshop v 19.1.9 (Adobe Inc., San Jose, California, USA). D N A I s o l a t i o n a n d P C R a m p l i f i c a t i o n , s e q u e n c i n g a n d e d i t i n g DNA extraction for all specimens except R. merzi sp. n. was done at Michigan State University, Depart- ment of Entomology in Smith’s lab. DNA was extracted using the DNeasy Blood & Tissue Kit protocol (Qiagen, Valencia, CA). For most specimens a single leg or head was taken for analysis. Five loci, with total length of 4270 base pairs, previously shown to be useful in systematic analyses of insects, especially tephritids, were analyzed (Han et al., 2002; Hebert et al., 2004; Moulton & Wiegmann, 2004; Barr et al., 2005; Smith & Brown, 2008, Ha- merlinck et al., 2016); see also Hulbert (2018: Table 2.2) for details. Th e mitochondrial protein coding gene cytochrome oxidase I (COI) (684 bp), the nuclear protein coding carbamoyl-phosphate synthase (CPS) domain of carbamoyl-phosphate synthetase 2, aspartate transcarbamy- lase and dihydroorotase gene (CAD) (990 bp), the large subunit of the nuclear ribosomal gene (28S) (1359 bp), the nuclear protein coding period gene (614 bp) and the nuclear protein coding alanyl t-RNA synthetase gene (AATS) (623 bp) were the PCR-amplifi ed regions. Primers and thermocycler conditions used for each of the fi ve genes amplifi ed were listed by Hulbert (2018: Table 2.2). Amplifi cations of COI, period, and AATS employed a single primer pair. For 28S, two primer pairs were used to amplify two non-overlapping fragments separated by 58 bp (1445 bp total). For CAD, we used two primer pairs to amplify two overlapping fragments (1003 bp total). Verifi cation of successful amplifi cation for all PCR products was confi rmed electrophoretically via agarose gels (1 % w/v) prior to purifi cation of PCR products using a QIAquick PCR Purifi cation Kit (Qiagen, Valencia, CA) according to the manufacturer’s specifi cations. Sanger sequencing was performed at the Michigan State University Research Technology Support Facility via BigDye Terminator Sequencing on an Applied Biosystems 3730xl DNA Analyzer (Foster City, CA, USA) using the PCR primers as sequencing primers. All sequences were edited manually by visual comparison of the automatic base calls to the original elec- tropherogram traces using MEGA (version 7.0.14) (Tamura et al., 2011) and deposited in GenBank (accession numbers MG825190-MG825320) (see Hulbert, 2018: Table 2.3) except for the new sequences obtained and uploaded in 2022 (table 1). Alignments of DNA sequences were constructed and edited in MEGA. We used the default parameters in MUSCLE (Edgar, 2004) as implemented in MEGA to align DNA sequences for all loci except 28S. For the 28S alignment, we used MAFFT with default parameters (Katoh et al., 2002). Th e alignment is available as a supplementary fi le at https://doi.org/10.5281/zenodo.6111737. Th e names of terminal taxa correspond to those in table 1 and Hulbert (2018: Table 2.3), except R. bushi Hulbert & Smith, 2018 sometimes mentioned as “buff aloberry_fl y” by Hulbert (2018). S a m p l i n g a n d D N A e x t r a c t i o n o f R h a g o l e t i s m e r z i s p . n . Infested fl eshy cones of Juniperus sabina L. were sampled near Visperterminen, Switzerland in October 2016 by JJS. Th ey were transferred to plastic boxes containing sterile sand to enable larvae to emerge from the fl eshy cones and pupate. DNA of the pupae was extracted with the GenElute™ Mammalian Genomic DNA Miniprep Kit (Merck Sigma-Aldrich Chemie, Buchs, Switzerland) according to the manufacturer’s recommendations. S e q u e n c i n g L i b r a r y p r e p a r a t i o n a n d D N A s e q u e n c i n g For library preparation, the Nextera® XT DNA Library Prep kit (Illumina, San Diego, CA, USA) was used with 1ng of DNA according to the manufacturer’s recommendations. As the library was to be sequenced together with other samples on a MiSeq (Illumina, USA), the DNA had to be tagged by a barcoding PCR performed using primer pairs from the Nextera XT Index kit (Illumina, USA). Th e amplifi cation reaction was conducted with the same reagents and the thermocycler was set to the following parameters: 95 °C for 3 min, 4 S. V. Korneyev, J. J. Smith, D. L. Hulbert, J. E. Frey, V. A. Korneyev 20 cycles of 95 °C (30 s), 55 °C (30 s), 72 °C (30 s), one additional step at 72 °C (5 min) and a fi nal step at 4 °C until further processing. Amplifi ed products were cleaned with the protocol of the AMPure XP beads (Beckman-Coulter, Fullerton, CA, USA). Th e quality of the product was checked on a 1 % agarose gel and quantifi ed on a Qubit 3.0 Fluorimeter (Th ermo Fisher Scientifi c, Waltham, MA, USA) using the corresponding High-sensitivity dsDNA HS Assay Kit. Th e Rhagoletis merzi library was then pooled with the other samples prepared for this run in an equimolar way and loaded on a cartridge on the Illumina MiSeq sequencing system (Illumina, USA). MiSeq Reagent Kit V3 (2 × 300 bp) sequencing reagents (Illumina, USA) were used for this experiment together with 1 % of PhiX Control v.3 (Illumina, USA) spiking for quality control. Th e collection and voucher information for the Rhagoletis and outgroup species sequenced are given in Hulbert (2018) except those given in the table 1 and the accession numbers in the table 2. We used the dataset of Hulbert (2018) in its entirety, adding to it three specimens of the new species R. merzi, and three specimens collected from J. horizontalis in 2015, which we designate as R. sp. nr. juniperina for the analyses here. P h y l o g e n e t i c a n a l y s e s We used a Bayesian framework for our phylogenetic analyses of the fi ve-locus alignment. We predefi ned the following biologically relevant partitions in the alignments per the recommendation of the PARTITION- T a b l e 1 . Specimen collection records (compatible with Table 2.1 in Hulbert, 2018) Species group No Taxon Locality Host plant Date Collector juniperina (= zernyi) 1 R. sp. ex J. horizontalis 15GHB2 Centerville Township, Michigan. USA 44.94, -85.81 Cupressaceae: Juniperus horizontalis 8.10.2015 J. J. Smith juniperina (= zernyi) 2 R. sp. ex J. horizontalis 15gab3 Centerville Town- ship, Michigan. USA 44.94, -85.81 Cupressaceae: Juniperus horizontalis 8.10.2015 J. J. Smith juniperina (= zernyi) 3 R. sp. ex J. horizontalis 15BetsieA Lake Township, Michigan. USA 44.69, -86.25 Cupressaceae: Juniperus horizontalis 8.10.2015 J. J. Smith juniperina (= zernyi) 4 R. merzi sp. n. 2016 1 Switzerland, Valais, Visperterminen 46.262, 7.899 Cupressaceae: Juniperus sabina 17.10.2016 J. J. Smith juniperina (= zernyi) 5 R. merzi sp. n. 2016 2 Switzerland, Valais, Visperterminen 46.262, 7.899 Cupressaceae: Juniperus sabina 17.10.2016 J. J. Smith juniperina (= zernyi) 6 R. merzi sp. n. 2018 1 Switzerland, Valais, Visperterminen 46.262, 7.899 Cupressaceae: Juniperus sabina 17.10.2016 J. J. Smith T a b l e 2 . GenBank accession numbers of the DNA sequences (compatible with Table 2.3 in Hulbert, 2018) Taxon designation COI CAD period AATS 28S part A 28S part B R. sp. ex J. horizontalis 15GHB2 OM541942 OM718809 OM718815 OM718821 OM569803 OM569801 R. sp. ex J. horizontalis 15gab3 OM541943 OM718810 OM718816 OM718822 OM569805 OM569802 R. sp. ex J. horizontalis 15BetsieA OM541944 OM718811 OM718817 OM718823 OM569806 OM569803 R. merzi sp. n. 2016 1 OM632706 OM718812 OM718818 OM718824 OM649845 R. merzi sp. n. 2016 2 OM632707 OM718813 OM718819 OM718825 OM649846 R. merzi sp. n. 2018 1 OM632708 OM718814 OM718820 OM718826 OM649847 FINDER soft ware user-guide (Lanfear et al., 2017): 28S (fragments were concatenated and considered a single partition for phylogenetic analysis), and separate partitions for each nucleotide position (1st, 2nd, 3rd codon position) for COI, CAD, period and AATS. Next, we used PARTITIONFINDER v2.1.1 (Lanfear et al., 2017) to determine combinability of partitions and nucleotide substitution models. We then ran PARTITIONFINDER implementing PhyML (Guindon et al., 2010) with the “greedy” algorithm (Lanfear et al., 2012) using the cor- rected Akaike Information Criterion (AICc) to assess model and partition quality. We conducted runs of PAR- TITIONFINDER restricted to MRBAYES models. Th e Bayesian analysis used four independent runs each with four Metropolis-coupled chains with default heating parameters (one cold and three heated) in MRBAYES. Th e chains were sampled once every thousand generations for 5 million generations and the fi rst 25 % of samples were discarded as burn-in. All analyses con- verged to an average standard deviation of split frequencies below 0.02 (Ronquist et al., 2012). 5A New Species of Rhagoletis (Diptera, Tephritidae) from Switzerland… A maximum-likelihood (ML) analysis was also perfo med in MEGA 11 (Tamura et al., 2021) using Tamu- ra-Nei model (Nei & Kumar, 2000) with rates among sites Gamma distributed with invariant sites (G+I), NNI heuristic method, with the initial tree automatically made (NJ/BioNJ). Th e reliability of clustering patterns in the ML tree was determined by the bootstrap test (500 replications) in 4 threads. Mesquite 3.20 (Maddison & Maddison, 2021) was used to visualize the phylogenetic trees and to estimate genetic distances using Kimura’s two-parameter (K2P) model of nucleotide substitution. Results Rhagoletis Loew, 1862 Rhagoletis Loew, 1862 b: 44. T y p e s p e c i e s : Musca cerasi Linnaeus, 1758 (by monotypy). Zonosema Loew, 1862 b: 43. T y p e s p e c i e s : Tephritis alternata Fallén, 1814 (by subsequent designation of Rondani, 1870: 6). Microrrhagoletis Rohdendorf, 1961: 187. T y p e s p e c i e s : Microrrhagoletis samojlovitshae Rohdendorf, 1961 (by original designation). Megarrhagoletis Rohdendorf, 1961: 196. T y p e s p e c i e s : Megarrhagoletis magniterebra Rohdendorf, 1961 (by original designation). D i a g n o s i s . Medium-sized (3.0–8.0 mm) fruit fl ies with 3 frontal and 2 (rarely 1) or- bital setae, pale or dark postocellar seta, short head, fi rst fl agellomere usually with pointed apex (rarely rounded); thorax entirely orange to entirely black; postpronotal lobe usually pale yellow; scutellum orange to black with creamy white or yellow disc (black in R. psalida); surstylus of male with long and variously shaped, usually acute, posterior lobe; oviscape with T-shaped desclerotized posteromedial area ventrally; aculeus usually uniformly ta- pered apically. Th ird instar larva with variable number (from 3 to 20) of oral ridges and stomal sensory organ with or without preoral teeth. R e m a r k s . Th e morphological diagnosis of Rhagoletis almost entirely overlaps with that of Carpomya, which sometimes cannot be undoubtedly diff erentiated (except the mesonotum pattern and number of frontal setae; both variable). Current concepts of these genera need revision based on a sound multi-locus DNA reconstruction of phylogenetic relationships among the genera of Carpomyini. Key to the Rhagoletis species similar to R. merzi sp. n. Th is key includes R. merzi and the species similar to it, having a black body with postpronotal lobe and major part of scutellum yellow or white, the wing with 4 bands, of which the apical band is connected to the subapical band and separated from the apical wing margin by a crescentic marginal hyaline area: Rhagoletis juniperina Marcovitch, 1915, R. zernyi Hendel, 1927, R. fl avigenualis Hering, 1958 (the juniperina group), R. tabellaria (Fitch, 1855), R. persimilis Bush, 1966, R. electromorpha Berlocher, 1982, R. bushi Hulbert et al., 2018 (tabellaria group), R. ebbettsi Bush, 1966, R. ribicola Doane, 1899 (ribicola group), and the following species unassigned to groups, R. scutellata Zia, 1938, R. batava Hering, 1958, R. mongolica Kandybina, 1972), and R. bagheera Richter & Kandybina, 1997. 1. Dark subbasal and discal bands widely connected in posterior part of wing (see Foote et al., 1993: fi gs 378–379). Nearctic Region only. .............................................................................. tabellaria group, part 2 — Dark subbasal and discal bands widely separated or connected by a pale grey, indistinctly darkened area (fi g. 1). Palearctic and Nearctic Regions. ......................................................................................................... 4 2. Apical crescentic hyaline area (cr) shorter, reaching at most to posterior 1/3 of costa in cell r4+5; subapi- cal band uniformly dark, crossvein dm-m pale emarginated (see: Foote et al., 1993: fi g. 379). ................. ................................................................................................................................... R. electromorpha Berlocher — Apical crescentic hyaline area longer, reaching vein M1; subapical band uniformly dark, crossvein dm-m not pale emarginated (see: Foote et al., 1993: 359). ........................................................................... 3 3. Aculeus shorter than 0.7 mm. Larvae in fruits of Cornus (Cornaceae) and Vaccinum (Ericaceae). ......... .................................................................................................................................................R. tabellaria (Fitch) — Aculeus longer than 0.8 mm. Larvae in fruits of Prosartes hookeri (Liliaceae). ............................................ ................................................................................................................................................... R. persimilis Bush 4. Occiput completely yellow. Wing pattern usually yellowish brown, with bands laterally emarginated with brown (fi gs 1, a–b). Femora mostly or entirely yellow. Larvae in Juniperus fl eshy cones. ................. ......................................................................................................................................... juniperina group, part 5 6 S. V. Korneyev, J. J. Smith, D. L. Hulbert, J. E. Frey, V. A. Korneyev — Occiput with at least medial sclerite above occipital foramen, or oft en lateral of occipital suture black or brown. Wing pattern with blackish brown bands (fi gs 1, c–e). Femora coloration and host plants variable. .............. 6 5. Discal and subapical bands widely connected at least in cell r2+3 (fi g. 1, a). Spain. ..........R. zernyi Hendel — Discal and subapical bands entirely separated (fi g. 1, b). Asia: Turkey, Caucasus, Iran, Middle Asia. ..... .......................................................................................................................................... R. fl avigenualis Hering 6. Discal and subapical bands connected in cell r4+5 (see: Bush, 1966: fi g. 200; Foote et al., 1993: fi g. 381). USA: California. ............................................................................................................... R. ebbettsi Bush, 1966 — Discal and subapical bands widely isolated. .................................................................................................... 7 7. Femora yellow. .................................................................................................................................................... 8 — At least mid and hind femora black. ................................................................................................................ 9 8. Only dorsal third of occiput with dark transverse spot. Larvae in Juniperus sabina. Asia: Mongolia, (?) Kyrgyzstan. Genitalia not examined. ...................................................................... R. mongolica Kandybina — Occiput with wide black horseshoe-shaped pattern reaching its lower half. Lateral surstylus with short posterior lobe. Spermatheca narrow and long, worm-like (see Bush, 1966: fi g. 167). Larvae in Ribes spp. North America. ........................................................................................................................ R. ribicola Doane 9. Occiput with wide black horseshoe-shaped pattern reaching its lower half. Lateral surstylus with long and narrow posterior lobe (fi g. 2, a). Spermatheca rounded, small, < 0.05 mm in diameter, with neck longer than spermatheca itself (fi g. 2, d). .......................................................................................................10 — Occiput black only across upper 1/3. Lateral surstylus with short posterior lobe (fi g. 2, c) (not known for R. scutellata). Spermatheca globose, larger, > 0.05 mm in diameter, with neck shorter than sper- matheca itself (fi g. 2 f) (not known in R. scutellata). Associated with Juniperus (host not known for R. scutellata). .......................................................................................................................................................... 12 10. Nearctic Region. Scutellum laterally and fore coxa usually black. Larvae in Shepherdia argentea (Pursh) Nutt. (Eleagnaceae). ................................................................................................ R. bushi Hulbert & Smith — Palearctic Region. Scutellum laterally and fore coxa usually yellow. Host plants diff erent. .................. 11 11. Smaller: wing length less than 2.8 mm (in { 2.0–2.4, rarely up to 2.7 mm), in } 2.2–2.5 mm). Larvae in Rhamnus pallasii (Rhamnaceae). Asia: Armenia, Georgia. .................. R. bagheera Richter & Kandybina — Larger: wing length greater than 2.9 mm (in { 3.0–3.6 mm, in } 3.3–4.2 mm). Europe, Asia (Caucasus, Middle Asia, Siberia, China). Larvae in Hippophae rhamnoides L. (Elaeagnaceae). ..... R. batava Hering 12. Abdominal tergites 2–4 uniformly brown or black, without pale bands on posterior margins. Genital characters not examined. Host plants unknown. China (Gansu). .................................... R. scutellata Zia — Abdominal tergites 2–4 with pale bands on posterior margins. ................................................................ 13 Fig. 1. Palearctic species of Rhagoletis species similar to R. merzi, wings: a — R. zernyi; b — R. fl avigenualis; c — R. merzi, sp. n.; d — R. bagheera; e — R. batava. Bands are marked as follows: A — apical, D — discal, SA — subapical, SB — subbasal. Red arrow shows connection of D and SA; cyan arrow shows cr — crescentic hyaline area. Scale: 1 mm. 7A New Species of Rhagoletis (Diptera, Tephritidae) from Switzerland… 13. North America. Associated with various Juniperus spp. ................................................................................. ............................................................ R. juniperina Marcovitch(possibly an assemblage of cryptic species) — Europe: Switzerland. Associated with Juniperus sabina. ....................................................... R. merzi sp. n. Overview of Palearctic species similar to R. merzi Rhagoletis bagheera Richter & Kandybina, 1997 (fi g. 3) Rhagoletis bagheera Richter & Kandybina, 1997: 915 (description, biology); Korneyev et al., 2018 a: 462 (key); Korneyev & Korneyev, 2019: 93 (key). T y p e m a t e r i a l . Holotype {: Armenia: “Erevan, ex fruits of Rhamnus pallasii, em. 28.VI.1971” (G. Ariutunan) (ZISP). P a r a t y p e s . Armenia: Erevan, ex fruits of Rhamnus pallasii, em. 28.06.1971, 1 {, 1 } (G. Arutiunan); Asni, Vedi Distr., 5.08.1965, 37 {, 20 } (V. Richter), 1 {, 1 }, same labels (SIZK); Georgia: Vashlovan Nature Reserve, ex fruits of Rhamnus pallasii, 2 }, em. 24.06.1974 (I. Hodzevanishvili); idem, em. 4–18.05.1981 (I. Hodzhevanishvili) (ZISP). D i a g n o s i s . Head yellow, antenna and frontal vitta dark yellow; ventral half of median occipital sclerite and occiput lateral of it widely black or brown, widely yellow emarginated. Flagellomere 1 pointed apically. Scutum black, with 4 gray microtrichose vittae. Wing pattern with four brown bands, without intercalary band; apical band separated from costa by hyaline area in cells r2+3 and r4+5 (fi g. 1, d). Femora dark brown to black. Abdomen black, tergites with yellow posterior margins. Th is species is similar to R. batava and R. merzi in general appearance and femora coloration, but is conspicuously smaller (body length = 2.7–2.9 mm and wing length = 2.2–2.36 mm for R. bagheera, > 3 mm in R. batava and Fig. 2. Palearctic species of Rhagoletis species similar to R. merzi, epandrium and surstyli, posterior view (a–c) and spermatheca (d–f): a, d — R. batava; b, e — R. fl avigenualis; c, f — R. merzi sp. n. 8 S. V. Korneyev, J. J. Smith, D. L. Hulbert, J. E. Frey, V. A. Korneyev R. merzi) and has diff erent host plants: Rhamnus (Rhamnaceae) for R. bagheera, and Hippophae (Elaeagnaceae) for R. batava and Juniperus (Cupressaceae) for R. merzi. M e a s u r e m e n t s . Wing length { = 2.0–2.4 mm; wing length } = 2.2–2.5; costal cell length = 0.7; aculeus length = 0.71 mm; aculeus length / costal cell length = 1 (Richter & Kandybina, 1997). H o s t p l a n t . Rhamnus pallasii Fisch. & C.A. Mey (Richter & Kandybina, 1997). D i s t r i b u t i o n . Armenia, Georgia. Rhagoletis batava Hering, 1958 (fi g. 4) Rhagoletis batava Hering, 1958: 2 (description); Kandybina, 1977: 145 (larvae); Norrbom et al., 1999 (catalogue); Korneyev et al., 2018 a: 462, 2018 b: 43 (key; distribution); Korneyev & Korneyev, 2019: 93 (key). Fig. 3. Rhagoletis bagheera paratype male (a, с–e) and female (b, f–h): a — habitus left (abdomen dissected), b — same, dorsal; c, d — epandrium, hypandrium and surstyli (c — left , d — posterior), e — phallus glans; f — aculeus apex, g — ovipositor, h — spermatheca. 9A New Species of Rhagoletis (Diptera, Tephritidae) from Switzerland… T y p e m a t e r i a l . Holotype }: Th e Netherlands: Terschelling I., Boschplaat (Th eowald) (SMNS). N o n - t y p e m a t e r i a l . Kyrgyzstan: Tien Shan, 1500 m, “Pristan-Przewalsk”, near Karakol (= Przewalsk), 42.5756º  N, 78.3011º E, 28.07.1986, 4 { (Korneyev); Karakol, on Hippophae, 15.08.1994, 1 } (Korneyev); Terskei Alatau, Karakol ravin, h = 2050–2850 m, 42.4431º N, 78.4129º E, 12–13.08.1998, 3 {; 2 } (Korneyev & Kameneva); Yssyk-Kol Region, Chong-Kyzyl-Suu, 42.250º N 78.130º E, 16–17.08.1998, 3 {, 1 } (Korneyev & Kameneva); Alai , 45 km S of Kyzyl-Kiya, Kichik-Alai Ridge, Isfairam-Sai basin, Langar, h = 1800–1900 m, 39.8264º N, 72.1133º E, 30.07.1999, 4 {; 3  } (Korneyev & Kameneva) (SIZK), idem, 3 {, 3 } in alcohol (Korneyev & Kameneva) (MSU); Russia: Altay, Chikhachev Ridge, reared ex fruits Hippophae rhamnoídes 09.1966–17.03.1967, 3 {, 2 } (Litvinchuk) (SIZK); Th e Netherlands: Hompelvoet Z.H. 10–18.07.2000, 2 {, 3 } (B. V. Aartsen) (SIZK); Tajikistan: Peter First Range, 39.14382º N,71.56161º E, 3 km S Muk, 2320 m asl, swept from Hippophae rhamnoides, 26.07.2018, 3 {, 7 }; 39.07035º N,70.79778º E, Mirazyon, 1950 m asl, swept from Hippophae rhamnoides, 27.07.2018, 1 {; Turkestan Range, N slope, 39.520714º N, 68.925904º E, 25 km SE Dzharkutan, 2840 m asl, 6.08.2018, 3 } (V. Korneyev) (SIZK). Fig. 4. Rhagoletis batava male (a, с–e) and female (b, f–j): a — habitus dorsal, b — same, left ; c, d — epandrium, hypandrium and surstyli (c — left , d — posterior), e — phallus glans; f — aculeus apex, g — ovipositor, h — spermatheca; i — eversible membrane, ventral. Scale: f — 0.1 mm, g — 0.5 mm. 10 S. V. Korneyev, J. J. Smith, D. L. Hulbert, J. E. Frey, V. A. Korneyev D i a g n o s i s . Rhagoletis batava is similar to R. bagheera and R. merzi in general appearance and in having black femora, diff ering from R. bagheera by its larger size (wing length = or > 3 mm in R. batava vs. < 2.5 mm in R. bagheera) and from R. merzi by the conspicuously longer posterior lobe of the lateral surstylus (1.3 times as long as surstylus basal of prensisetae (fi g. 4, d) vs. only 0.6–0.7 times as long as surstylus basal of prensisetae in R. merzi (fi g. 8, b)) in the male and by spermatheca size and shape in the female (oval, 0.03 mm in diameter, and with the neck as long as the spermatheca in R. batava (fi g. 4, i) vs. spherical, 0.09 mm in diameter, with the neck at most 0.8 times as long as the spermatheca itself in R. merzi (fi gs 2, f; 8 f)), as well as by the diff erent host plants. M e a s u r e m e n t s . Body length {  =  3.64 mm; body length } = 3.9 mm; wing length { = 3.0–3.6 (m  =  3.38) mm; wing length } = 3.3–4.2 (m = 3.8); costal cell length = 0.9; aculeus length = 0.78 mm; aculeus length /costal cell length = 0.9. H o s t p l a n t . Hippophae rhamnoides L. (Elaeagnaceae) (Kandybina, 1962). D i s t r i b u t i o n . Europe (Korneyev et al., 2018 b); south of West and East Siberia; Middle Asia. Rhagoletis fl avigenualis Hering, 1958 (fi gs 5–6) Rhagoletis zernyi: Zaitzev, 1947: 6 (misidentifi cation; records from Georgia; host plants). Rhagoletis fl avigenualis Hering, 1958 (description); Rohdendorf, 1961 (key, description); Kandybina, 1977 (larva; distribution; host plants); Korneyev & Merz, 1997 (key, distribution); Norrbom et al., 1999 (catalogue); Gilasian & Merz, 2008; Mohamadzade & Rasoulian, 2009: 84; Korneyev et al., 2018 a: 466; 2018 b: 32 (key, distribution). T y p e m a t e r i a l . Holotype {: Turkey: S. Anatolia, Antalya-Kas, Katrandag, 1100 m (SMNS). N o n - t y p e m a t e r i a l . Kazakhstan: Aksu-Djabagly, on Juniperus zeravshanica, 17.08.1964, 2 {, 2 }; Fig. 5. Rhagoletis fl avigenualis male (a) and female (b–e): a–b — habitus left , c — abdomen dorsal; d — occiput and mesonotum, posterodorsally. 11A New Species of Rhagoletis (Diptera, Tephritidae) from Switzerland… 15.09.1964 (Fisechko); Kyrgyzstan: Kyrghyz Alatau, 30 km S of Bishkek, 42°35.9΄ N 73°52.1΄ E, h = 1950– 2100 m, 5–7.08.1998, 3 {, 1 } (Korneyev & Kameneva) (SIZK); idem, 1 {, 1 }, in alcohol (MSU); Yssyk-Kol Re- gion, Terskey Alatau, from Juniperus sabina, 5.08.1972 (Kandybina) (SIZK); Tajikistan: Ghissar Range, 2.5 km E Iskanderkul, 39.08530º N, 68.40226º E, 2360 m asl, swept from Juniperus, 7–8.07.2018, 3 { (V. Korneyev) (SIZK); Turkmenistan: [Kopet-Dagh, between Firuza & state border], 23.09.1930, 1 } (L. Bianchi) (SIZK). D i a g n o s i s . Rhagoletis fl avigenualis can be diff erentiated from other species of the Rhagoletis juniperina group by having the medial occipital sclerite entirely yellow, entirely or predominantly yellow femora (at most hind femur brown basally), wing with basicostal cell clearly tinged with brown, wing bands partially yellowish-brown, with discal and subapical bands separated. R. fl avigenualis is similar to R. bagheera, R. batava, R. juniperina, R. mongolica, and R. merzi in general appearance, diff ering from them also by having the occiput entirely yellow (or at most the occipital sutures tinged with brown) (vs. with the median occipital sclerite widely black at least on the ventral half in the other species); it also diff ers from the juniper-associated R. juniperina and R. merzi by having a conspicuously longer male posterior lobe of the lateral surstylus (1.3–1.5 times as long as surstylus basal of prensisetae (fi g. 6, a) vs. 0.6–0.75 times as long as surstylus basal of prensisetae in R. merzi (fi g. 8, b) and R. juniperina (Bush, 1966: fi g. 83)). Abdomen black with posterior margins of tergites yellow. It is also similar to Rhagoletis zernyi in having the occiput and femora yellow and the wing bands partially yellowish-brown, diff ering from it by the entirely separated discal and subapical bands (in R. zernyi, the discal, subapical, and apical bands are connected at the anterior margin of the wing). Wing length 3.8–4.0 mm. Fig. 6. Rhagoletis fl avigenualis male (a–с) and female (d–g): a, b — epandrium, hypandrium and surstyli (a — left , b — posterior), c — phallus glans; d — aculeus apex, e — aculeus, f — spermatheca; g — eversible mem- brane, ventral. Scale: d, f — 0.1 mm, e, g — 0.5 mm. 12 S. V. Korneyev, J. J. Smith, D. L. Hulbert, J. E. Frey, V. A. Korneyev H o s t p l a n t s . Juniperus isophyllos (as “isocellos”) C. Koch; J. foetidissima Willd. (Zaitzev, 1947); J. excelsa M. Bieb., J. seravschanica Kom. (Kandybina, 1977). D i s t r i b u t i o n . Georgia (Zaitzev, 1947: as “R. zernyi” — misidentifi cation); Kazakh- stan, Kyrgyzstan, Tajikistan, S Turkmenistan (Kandybina, 1977), Turkey, Iran (Gilasian & Merz, 2008; Mohamadzade Namin & Rasoulian, 2009). R e m a r k s . Th is species is widespread in the Asia Minor, Caucasus and Central Asia. Rhagoletis merzi sp. n. (fi gs 1, c; 7–8) urn:lsid:zoobank.org:act:ADCCE1F4-7DB8-4EB6-9B5C-339A7F8F6ED9 Rhagoletis batava: Merz, 1994: 108 (misidentifi cation); Rhagoletis fl avigenualis: V. Korneyev in: Merz, 2006: 8 (misidentifi cation); Rhagoletis sp. near fl avigenualis: Korneyev et al., 2018 a: 466. T y p e m a t e r i a l . Holotype {: Switzerland: Visperterminen, VS, 1400 m, 26.07.1990 (Merz) (MNHG ENTO 00012822) (MHNG). Paratypes: Switzerland: 1 }, Visperterminen, h = 1400 m, 17.07.1995 (Merz) (MNHG ENTO 00012824); Visperterminen, VS, 1400 m: 1 {, 18.07.1993 (Merz) (MNHG ENTO 0001825); 1 {, idem, 1520 m, 20.07.1993 Wald (Merz) (MNHG ENTO 0001828); 1 {, idem, 17.07.1995 (Merz) (MNHG ENTO 0001823); 1 {, Visper- terminen, Kreuz, h = 1500 m, 21.07.2004 (Merz) (MNHG ENTO 0001827) (MHNG); Visperterminen, [Kreuz,] h = 1300–1900 m [swept from Juniperus sabina], 21.07.2004, 1 {, 1 } (S. & V. Korneyev) (SIZK). Fig. 7. Rhagoletis merzi sp. n. paratypes (MNHG): male (a–b) and female (c–d): a, c — habitus left , b, d  — same, dorsal (photos by Bernard Landry). 13A New Species of Rhagoletis (Diptera, Tephritidae) from Switzerland… N o n - t y p e s p e c i m e n s . Switzerland: Visperterminen, h = 1300–1900 m, reared from Juniperus sabina fl eshy cones, 3 puparia [used for DNA extraction completely], 17.10.2016 (J. Smith). D i a g n o s i s . Rhagoletis merzi is similar to all other species having the wing pattern with four dark bands, apical band joined to subapical band and separated by a crescent hyaline area from the costal vein anteroapically. It is most similar to, and in fact to our knowledge morphologically indistinguishable from, the Nearctic R. juniperina. Both species have the occiput widely black or brown on the upper 1/3, wing bands uniformly brown to blackish, mid and hind femora black, male lateral surstylus with the posterior lobe relatively short, 0.6–0.75 times as long as surstylus basal of prensisetae (fi g. 8, b), and female spermathecae large, 0.09 mm in diameter, with short neck (fi g. 2, f). We recognize R. merzi as a distinct species from R. juniperina based on the signifi cant genetic distance between their COI sequences (K2P = 0.071). Rhagoletis merzi is also very similar to the Central Asian R. mongolica and R. scutellata (both known only from their holotypes, not examined for potential genitalic diff erences) in general appearance, including the wing pattern and having the occiput widely black on the upper 1/3. Rhagoletis mongolica is also associated with J. sabina, like R. merzi, whereas the host for R. scutellata is unknown. Rhagoletis merzi diff ers from R. mongolica by having black rather than yellow femora and from R. scutellata by abdominal tergites 2–4 having whitish or yellowish posterior margins and the basicostal cell brownish (in R. scutellata, basicostal cell entirely hyaline and abdominal tergites uniformly black or brown). Fig. 8. Rhagoletis merzi sp. n. paratypes (SIZK): male (a–с) and female (d–g): a, b — epandrium, hypandrium and surstyli (a — left , b — posterior), c — phallus glans; d — aculeus apex, e — aculeus, f — spermatheca; g — eversible membrane, ventral. Scale: d, f — 0.1 mm, e, g — 0.5 mm. 14 S. V. Korneyev, J. J. Smith, D. L. Hulbert, J. E. Frey, V. A. Korneyev Th is species readily diff ers from the West Palearctic R. fl avigenualis and R. zernyi by having the widely black or brown median occipital sclerite, black mid and hind femora, and uniformly brown wing bands (in R. fl avigenualis and R. zernyi median occipital sclerite and all femora uniformly yellow (very rarely only hind femur partly brown), and the wing bands at least partly yellow with brownish borders; R. zernyi diff ers also by having the discal and subapical bands widely fused). Th e genetic distance between R. merzi and R. fl avigenualis is also signifi cant (K2P = 0.063–0.066). Rhagoletis merzi is similar to the Palearctic species R. bagheera and R. batava, and the Nearctic R. bushi in having the wing bands uniformly brown to blackish, and mid and hind femora black, diff ering from them by having the male lateral surstylus with the posterior lobe conspicuously shorter, 0.6–0.75 times as long as the surstylus basal of the prensisetae (fi g. 8, b) vs. 1.3–1.4 times as long as the surstylus basal of the prensisetae in R. bagheera (fi g. 3, c) and R. batava (fi g. 6, a), and female spermathecae larger, 0.09 mm in diameter, with a short neck (fi g. 2, f) vs. 0.02–0.03 mm in diameter, with the neck longer than the spermatheca itself in R. bagheera and R. batava. Rhagoletis merzi also has a diff erent host plant, Juniperus sabina L., vs. Hippophae rhamnoides (Elaeagnaceae) for R. batava and Rhamnus palasii (Rhamnaceae) for R. bagheera. Th e genetic distance between R. merzi and R. batava is K2P = 0.064–0.068, and between R. merzi and R. bushi K2P = 0.078–0.079. D e s c r i p t i o n . Head. Orange-yellow, ocellar triangle, ventral part of median occipital sclerite and oft en occiput lateral of it black or brown. Antennal arista pubescent. Setae black except postocellar, posterior genal, and some occipital setae white. Paravertical seta short, about as long as black acuminate postocular setae. — Th orax. Scutum black, yellowish setulose, with microtrichia pattern with two pairs of partly fused matte grayish vittae separated by subshining darker areas. Postpronotal lobe and notopleural stripe creamy white to yellow; scutellum pale yellow, black on anterior margin dorsally and laterally. All thoracic setae black; basal scutellar seta inserted into black area. Halter yellow to creamy white. — Legs. Fore coxa yellow, mid and hind coxae black or brown; fore and mid trochanters yellow; hind trochanter brown or black; fore femur yellow anteroventrally, black posterodorsally; mid and hind femora black except apices yellow; hind femur somewhat thickened in male, with 2–3 longer subapical anterodorsal and 2–3 longer subapical anteroventral setae; tibiae and tarsi yellow (fi g. 7). — Wing (fi g. 1, c). 2.3 times as long as wide, with pattern consisting of basicostal cell with brownish tinge and four dark brown bands; subbasal band from humeral crossvein over basal half of costal cell through cells br, bm and cua (= anal cell auctt.) slightly into cell cup, discal band from pterostigma over crossvein r-m to posterior margin between veins M4 (= CuA1) and CuA + CuP (= CuA1+A1), subapical band from middle of cell r1 over crossvein dm-m (= dm-cu) and apical band from middle of cell r1 into apex of cell m4; discal band separated from both subbasal and subapical bands (fi gs. 1, c; 7, a) or at most narrowly fused with subapical band at posterior margin (fi g. 7, c); subapical and subapical bands fused in cells r1 and r2+3; apical band separated from costa between apex of cell r1 and vein M1; no intercalary band; vein R4+5 dorsally with 1 seta at node. — A b d o m e n . All segments mostly black, posterior margin of tergites 2–4 in male, and 2–5 in female narrowly creamy yellow (fi gs. 9, b, d). Oviscape shining black, as long as tergite 5; setae and setulae black. — G e n i t a l i a . M a l e . Epandrium black. Proctiger as long as epandrium (fi g. 10, b). Surstylus dark yellow, lateral susrtylus with posterior lobe short, 0.6–0.75 times as long as surstylus basal of prensisetae (fi g. 9, b). Phallus with moderately large glans (fi g. 8, c) having membranous, narrow, fi nger-like apicodorsal process, large prepuce with smooth walls, and acrophallus with pair of semitubular fi laments, very similar to that of R. bagheera (Richter & Kandybina, 1997: fi g. 5), R. fl avigenualis (fi g. 6, c) and R. juniperina (Bush, 1966: fi g. 125); preglans short and simple, without eversible caecum. F e m a l e . Eversible membrane with two pairs of taeniae 0.5 × as long as membrane itself, ventral side of membrane with scales of diff erent size, medial ones larger than lateral ones and moderately pointed (fi g. 9, g). Two globular spermathecae, 0.09 mm in diameter, with 15A New Species of Rhagoletis (Diptera, Tephritidae) from Switzerland… long scale-like papillae on surface (fi g. 9, f). Aculeus brown, 5.5 × as long as wide, with acute apex (fi gs. 9, d–e). M e a s u r e m e n t s . Body length { = 3.8–4.2 mm; wing length { = 4.1–4.2 mm. Body length } = 4.0–4.4 mm; wing length { = 3.0, wing length } = 3.6 mm, costal cell length = 0.9; aculeus length = 0.85 mm; aculeus length /costal cell length = 0.9. H o s t p l a n t . Juniperus sabina L. Th e pupae for DNA analysis were reared from the same plants and in the same locality as the type specimens were swept. D i s t r i b u t i o n . Switzerland. E t y m o l o g y . Th is species is named in honor of the eminent Swiss dipterist Dr. Bernhard Merz, who collected most of the type specimens, in recognition of his contributions to the study of fruit fl ies. R e m a r k s . Kandybina (1977) reported specimens of “R. mongolica” with entirely black femora and partly black tibiae reared from Juniperus sabina in Kyrgyzstan, which need re-examination to determine whether they are conspecifi c with R. merzi. Rhagoletis mongolica Kandybina, 1972 Kandybina, 1972: 913 (description), 1977 (larva; distribution; host plants; new records); Korneyev & Merz, 1997: 63 (key); Norrbom et al., 1999: 201 (catalogue); Korneyev & Ovchinnikova, 2004: 482 (key). C o m m e n t s . Th is species was originally described based on a single female and a third instar larva reared from Juniperus sabina L. in Mongolia (Kandybina, 1972). Later, the larva was redescribed based on material from the same host plant in Kyrgyzstan (Kandybina, 1977). Th e adult specimens from Kyrgyzstan were briefl y described as having completely black femora and partly black tibiae; neither their male nor female genitalia have been described. We therefore consider R. mongolica to have entirely yellow femora as in the holotype, and the Kyrgyz specimens to be likely non-conspecifi c. Th e geographically “intermediate form” from Kyrgyzstan must be analyzed to clarify its taxonomic position and molecular diff erences from both R. mongolica and R. merzi. Rhagoletis scutellata Zia, 1938 Zia in: Zia & Chen, 1938: 34 (description); Wang, 1998: 124 (redescription); Norrbom et al., 1999: 202 (catalogue); Korneyev & Ovchinnikova, 2004: 482 (key). R e m a r k s . Th is species is known from the holotype male from Gansu, China (IZAS), unavailable in this study. According to the original description, it possesses black femora, uniformly dark brown bands on the wing, and entirely black abdominal tergites; body length { = 3.7; wing length { = 4.0. Additional study of the type and topotypic material from China, including morphology of the male and female genitalia, DNA sequences, and host plants, is needed. Rhagoletis zernyi Hendel, 1927 (fi g. 1, b) Hendel, 1927: 76 (description, key); Merz & Blasco-Zumeta, 1995: 132 (host plant, distribution); Merz, 2001: 92 (checklist); Norrbom et al., 1999: 202 (catalogue); Korneyev et al., 2018 a: 468 (key, distribution). M a t e r i a l . Spain: Monegros, Pina-de-Negro, 13.08.1992, 1 { (Blasco-Zumeta) (Merz det., 1994) (SIZK). D i a g n o s i s . Rhagoletis zernyi can be diff erentiated from most species of Rhagoletis by the characters given for the juniperina group. It diff ers from the other species of that group by having yellow femora and the wing pattern with partially yellowish-brown discal and subapical bands that are broadly fused at anterior margin (other species have the wing pattern uniformly brown except R. fl avigenualis, which has the discal and subapical bands separated (fi g. 1, b)); male and female genitalia not examined. Abdomen black with posterior halves of tergites creamy or yellow. Body length 4.0–4.8 mm; wing length { = 3.5–3.6 mm. 16 S. V. Korneyev, J. J. Smith, D. L. Hulbert, J. E. Frey, V. A. Korneyev D i s t r i b u t i o n . Spain. H o s t p l a n t s . Juniperus thurifera L. (Merz & Blasco-Zumeta, 1995). Phylogenetic analysis Hulbert’s (2018) multigene molecular analysis grouped the included Rhagoletis spe- cies into ten lineages corresponding to 12 previously established species groups (alternata, cerasi, meigenii, ferruginea, nova, striatella, juniperina, ribicola, cingulata, suavis, tabel- laria, and pomonella), which mainly correspond to the groups proposed by Bush (1966), Kandybina (1977), and Smith & Bush (1999) plus the unplaced R. fausta and R. batava. Our analyses here is based on the 85 individuals from Hulbert (2018) plus three speci- mens of R. merzi and three specimens of R. sp. nr. juniperina, using the same 4270 bp from the COI, CAD, 28S, period, and AATS genes used in Hulbert (2018). Th e taxon sample includes two outgroup species (Anastrepha ludens and Euphranta canadensis), one species of Carpomya and 35 species representing most of the species groups of Rhagoletis. Th e Bayesian analysis recovered a monophyletic cluster with high support, containing the suavis, cingulata, pomonella, tabellaria, and juniperina groups plus the unplaced species R. fausta and R. batava (fi g. 9). All the juniper-associated species form a well-supported monophyletic lineage corresponding to the juniperina group within the clade described above. Th is includes both Nearctic (R. juniperina and undescribed species R. sp. nr. juni- perina) and Palaearctic species (R. fl avigenualis and R. merzi). It is interesting that R. merzi is more similar to the Nearctic taxa (R. juniperina, R. sp. near juniperina) in both morpho- logical characters (wing pattern, occiput, mesonotum and leg coloration) and molecular sequences than to the Palearctic R. fl avigenualis. Th e Maximum Likelihood tree has a similar topology to that obtained with the Bayes- ian analysis, yet with some notable diff erences that mostly concern position of some basal groups (fi g. 9). Th e juniperina group is recovered in the ML analysis with a bootstrap value of 58 and, within this group, R. merzi is nested within the North American lineage as sister to R. sp. near juniperina (bootstrap 75). Conclusions As a result of molecular and careful morphological study (including the structure of male and female genitalia), the population of fruit fl ies found to infest Juniperus sabina in the Swiss Alps, previously misidentifi ed either as R. batava (Merz, 1994) or R. fl avigenualis (V. Korneyev in: Merz, 2006) was found to represent a new species, R. merzi. Rhagoletis merzi is a case of discovery of a previously unknown animal species in the very heart of Europe and in the part of the continent with the most thoroughly studied tephritid fauna. It remained dubiously identifi ed for decades, though it diff ers morphologically from known west Palearctic species, and is genetically distinct from all other species of the genus. Both morphological characteristics and DNA-based phylogenies show that R. merzi is related to three other juniper-infesting species of Rhagoletis from both the Nearctic and Palaearctic Regions: R. juniperina, R. fl avigenualis, and R. zernyi. Morphological characters can be used to diagnose R. merzi from some other juniper- infesting species. Rhagoletis merzi has a combination of black colored femora, round spermathecae and relatively short surstyli, strongly resembling the Nearctic R. juniperina and R. sp. near juniperina, but R. merzi is quite distant genetically from both of these species. Th e new species has been already confused with R. batava, which feeds on sea buckthorn and has black femora as well. Th e use of a particular host plant can be a proxy for species identifi cation in Rhagoletis, and is usually reliable in species identifi cation within the genus, but caution must be exercised as there are rare exceptions where host-specifi c fl ies have been reared from ‘non-natal’ hosts that likely do not represent established populations (Bush, 1966; Yee & Goughnour, 2008; Hood et al., 2012; Yee et al., 2015). 17A New Species of Rhagoletis (Diptera, Tephritidae) from Switzerland… Fig. 9. Bayesian phylogeny of Rhagoletis inferred from an alignment of 4270 bp of fi ve genes (COI, CAD, ribo- somal 28S, period and AATS) using MrBayes, and Max- imum-Likelihood using MEGA 11. Th e fi rst number on each branch is the bootstrap support from ML analysis; the second number represents posterior probability from Bayesian inference (BI). Asterisks (*) over branch- es indicate a Bayesian posterior probability of 1.0 and 100 % bootstrap support for the clade. Dash (- /) means that the clade inferred by MrBayes was not recovered by the ML analysis. Grey background shows position of the “core Nearctic taxa” (Smith et al., 2006). Th e bold numbers and color rectangles indicate species groups as follows: 1 — alternata group, 2 — cerasi group, 3 — clus- ter of ferruginea +nova + striatella groups, 4 — meigenii group, 5 — cingulata group, 6 — suavis group, 7 — ribi- cola group (sensu Bush, 1966), 8  — pomonella group, 9 — tabellaria group, 10 — juniperina group. Abbrevia- tions: NA — Nearctic Region, PA — Palaearctic Region. Inlay shows relationships within the juniperina group. 18 S. V. Korneyev, J. J. Smith, D. L. Hulbert, J. E. Frey, V. A. Korneyev Unfortunately, we lack data about such peculiar species as R. zernyi and the morphologically similar species R. scutellata and R. mongolica from Central Asia, making the relationships of the Nearctic and Palearctic juniper-infesting Rhagoletis not fully resolved. Additional studies are needed to clarify the diff erences between R. merzi and R. mongolica, for which no molecular data are available. It will be important to obtain samples from the type locality of R. mongolica morphologically identical to its holotype, because the specimens infesting Juniperus sabina recorded by Kandybina (1977) from Kyrgyzstan as “R. mongolica” may be misidentifi ed R. merzi as they diff er from the R. mongolica holotype in the coloration details. Th ese tasks are forthcoming, as are descriptions of a number of previously unknown new juniper-infesting species in North America (Hulbert et al., in prep.). Rhagoletis merzi has a unique COI haplotype that shows essential diff erences from similar sequences from both the juniper-infesting R. fl avigenualis (K2P = 0.063–0.066) and Nearctic R juniperina (K2P = 0.71), as well as from the superfi cially similar R. batava (K2P = 0.064–0.068). Despite intensive studies in the genus Rhagoletis, mainly restricted to the pest and model species, the Juniperus-associated species remain one of the most poorly examined groups. Both sweeping and rearing from juniper fl eshy cones oft en give very poor output; a full day of sweeping over juniper trees usually brings 1–2 specimens, and rearing, which is more productive, requires collecting cones during exact short time periods late in summer or in autumn. Numerous superfi cially similar species associated with various hosts and diff ering by a few coloration and genitalic characters form a paraphyletic formation in the base of a monophyletic lineage represented mostly by Nearctic species. Both Bayesian and ML analyses show that the species of the tabellaria group, as well as R. ribicola and R. batava, do not form a monophyletic clade with the juniperina group (fi g. 9) despite the strong morphological similarities of these fl ies. It is believed that the purely Nearctic species groups, such as the pomonella, cingulata and suavis groups are derivatives from the forms superfi cially similar to R. juniperina, R. batava or R. ebbettsi. Authors’ responsibilities All authors collected material in the fi eld. DLH, JEF, JJS and SVK extracted and sequenced DNA. All morphological dissections and descriptions, photographs, and trees were produced mostly by SVK and VAK, and analysis provided by DLH, SVK, JJS and VAK. Th e text was mostly written by DLH, SVK, VAK and JJS. Acknowledgements Th is work was fi nancially supported by IIE, Fulbright Ukraine, Research and Development Program, 2017–2018 (IIE Participant ID: PS00246781; Grantee ID: E0579598) fellowship for S. Korneyev and the Smith laboratory, Department of Entomology at Michigan State University. We thank Bernard Landry and Emmanuel F.A. Toussaint for help in examination of specimens located at Muséum d’histoire naturelle Geneva (MHNG) and Rachel Osborn for her help at MSU. We also thank Nicole E. Wonderlin, Courtney Larson, Alexa R. Warwick, Inna Barysh, Anton Senenko, Kristina Kernytska, Elena P. Kameneva and the late Marta Kolomayets for their valuable help. We also thank two anonymous reviewers for their valuable comments. Mention of trade names or commercial products in this publication is solely for the purpose of providing specifi c information and does not imply recommendation or endorsement. References Barr, N. B., Cui, L., McPheron, B. A. 2005. Molecular systematics of nuclear gene period in genus Anastrepha (Tephritidae). Annals of the Entomological Society of America 98, 173–180. Berlocher, S. H., Bush, G. L. 1982. An electrophoretic analysis of Rhagoletis (Diptera: Tephritidae) phylogeny. Systematic Zoology, 31, 136–155. Boller, E. F., Prokopy, R. J. 1976. Bionomics and management of Rhagoletis. 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