Open access journal: http://periodicos.uefs.br/ojs/index.php/sociobiology ISSN: 0361-6525 DOI: 10.13102/sociobiology.v68i4.7276Sociobiology 68(4): e7276 (December, 2021) Social parasitism has evolved several times and in different forms among ants, showing amazing morpho- functional, chemical and behavioral adaptations to this peculiar habit (Visicchio et al., 2001; Buschinger, 2009). Inquilinism is considered the most extreme of such forms: inquiline social parasites produce few to no workers while they concentrate their efforts on the production of reproducers which their hosts raise (Hölldobler & Wilson, 1990; Buschinger, 2009; Degueldre et al., 2021). As a result, inquiline ant species are rarely detected during field surveys. Abstract We conducted a survey on the Alpine fauna of one of the largest Natural Park of the Italian Alps (Stelvio National Park) in the framework of a broad ecological monitoring of Alpine biodiversity. A two-years standardized sampling employing pitfall traps along a 1200 m altitudinal gradient led to the discovery of the inquiline social parasite ants Myrmica myrmicoxena Forel, 1895 and M. microrubra Seifert, 1993. Myrmica myrmicoxena, which is classified as Vulnerable according to the IUCN Red List, was so far known from only three sites across a narrow geographic range between Italy and Switzerland. Our data support the previous hypothesis over its ecology and host association. Myrmica microrubra is considered an incipient species of high evolutionary interest, sometimes regarded as an intraspecific form of M. rubra. While having a wide distribution in Europe, its presence in Italy was hitherto known only from a single site. Our record extends its altitudinal distribution limit in Europe upwards by about 600m. Sociobiology An international journal on social insects Enrico Schifani1, Cristina Castracani1, Fiorenza A Spotti1, Daniele Giannetti1, Martina Ghizzoni1, Mauro Gobbi2, Valeria Lencioni2, Luca Pedrotti3, Donato A Grasso1, Alessandra Mori1 Article History Edited by Evandro Nascimento Silva, UEFS, Brazil Received 18 June 2021 Initial acceptance 25 October 2021 Final acceptance 14 November 2021 Publication date 24 November 2021 Keywords Inquiline social parasitism, rare species, Alpine insects. Corresponding authors Enrico Schifani Department of Chemistry, Life Sciences & Environmental Sustainability, University of Parma, Parco Area delle Scienze 11/a, 43124 Parma, Italy. E-Mail: enrico.schifani@unipr.it Cristina Castracani Department of Chemistry, Life Sciences & Environmental Sustainability, University of Parma, Parco Area delle Scienze 11/a, 43124 Parma, Italy. E-Mail: cristina.castracani@unipr.it There is an underdeveloped understanding of their biology, ecology, and conservation status (López-Soria, 1991; Schifani, 2017). However, a large number of the European inquiline ants were classified as Vulnerable on the IUCN Red List (Social Insects Specialists Group, 1996). The genus Myrmica Latreille, 1804 counts about 190 species distributed across the Holarctic region (Bolton, 2021), and it includes a remarkable variety of inquiline social parasites: in the West-Palearctic region, over 30 Myrmica species are recognized (Radchenko & Elmes, 2010; Seifert, 2018), 1 - Department of Chemistry, Life Sciences & Environmental Sustainability, University of Parma, Parma, Italy 2 - Section of Invertebrate Zoology and Hydrobiology, MUSE - Science Museum of Trento, Trento, Italy 3 - Stelvio National Park, Bormio, Italy Social Parasite Ants in the Alps: a New Site of the Vulnerable Myrmica myrmicoxena and New Uppermost Altitudinal Limit for M. microrubra SHORT NOTE mailto:enrico.schifani@unipr.it Enrico Schifani et al. – m Myrmica microrubra and M. myrmicoxena in the Alps2 one quarter of which is made of obligate or facultative social parasites of other congeneric species (Radchenko & Elmes, 2003; 2010). Most of them are inquilines, which evolved multiple times, independently in the genus (Radchenko & Elmes, 2003; Jansen et al., 2010; Seifert, 2018). Unlike in other ant genera (e.g., Plagiolepis, see Degueldre et al., 2021), most parasite Myrmica spp. do not follow the strict version of Emery’s rule (Emery, 1909), as their host species are not always their closest evolutionary relatives (Jansen et al., 2010). In particular, parasite Myrmica species with a long evolutionary history seemed to have developed the ability to exploit a wider and less closely related set of congeneric species as hosts (Jansen et al., 2010; Seifert, 2018). Some of the free-living species of the M. scabrinodis- complex are the hosts for the overwhelming majority of the West-Palearctic social parasite Myrmica (Radchenko & Elmes, 2003; 2010; Seifert, 2018): these range from the widespread facultative parasite M. vandeli Bondroit, 1920, to more specialized species with narrower geographic ranges (M. bibikoffi Kutter, 1963, M. hirsuta Elmes, 1978, and M. laurae (Emery, 1907)), to forms of extreme morpho-functional adaptation to inquilinism (M. lesmanei Bernard, 1967, M. kabylica Cagniant, 1970, and M. karawajevi Arnol’di, 1930) (Radchenko & Elmes, 2010; Seifert, 2018). Host species range from only one to several, with M. karawajevi exploiting the widest amount of different hosts, probably due to a longer evolutionary history (Jansen et al., 2010; Seifert, 2018). Outside the M. scabrinodis-complex, only two parasite Myrmica species are found in the West-Palearctic. The Alpine endemic M. myrmicoxena is a workerless inquiline collected only in three sites within a narrow altitudinal range for 150 years. M. lobulicornis (from the M. lobicornis-group) is considered to be likely its only host species (Forel, 1895; Glaser et al., 2010). On the other hand, M. microrubra Seifert, 1993, widely distributed in Europe, is either considered as an incipient species or as an intraspecific parasitic form of M. rubra (Linnaeus, 1758) (Pearson, 1980; 1981; Seifert, 1994; 2018; Seppä & Pamilo, 1995; Steiner et al., 2006; Schär & Nasch, 2014; Leppänen et al., 2011; 2015; 2016). Here we treat it as a good species following the latest review of the European ant fauna (Seifert, 2018). From 2018 to 2020, terrestrial arthropods were sampled by 150 pitfall traps in 30 sites across an altitudinal and habitat gradient in the Lombardy Sector of the Stelvio National Park. The traps were built with plastic glasses of c. 6 cm of diameter and c. 7 cm of height, buried in the ground, filled with c. 150 ml cc of an attractive and a preserving mixture of white vinegar, sodium chloride, and a drop of detergent as a surfactant, as described by Gobbi (2020). A total of 1800 samples were taken during six sampling sessions per year (150 x 6 x 2 = 1800). Ants were identified under a Zeiss Stemi 508 stereoscopic microscope and measured with the aid of an Axiocam Erc 5 s and Zeiss Zen Core Software according to the keys provided by Radchenko & Elmes (2010) and Seifert (2018). This effort led to a significant database of over 1700 Alpine ant records and interesting incidental discoveries, such as the first finding of ergatandromoph individuals of M. lobulicornis in 2018 (Schifani et al., 2020). In 2019, two rare social parasite Myrmica species of particular interest were collected, namely M. microrubra and M. myrmicoxena (Fig 1). Both were previously known from only one locality in Italy (Glaser, 2003; Glaser et al., 2011), and for both, the new occurrence sites yielded new important ecological or distributional information. Collecting data of the two species are as follows: - M. microrubra (1♀) alongside M. rubra (103☿☿ 1♀): Valle Messi, Sondrio, 46.296930, 10.503609, 1588 m asl, south- facing slope, peat bog, 24.IX.2019 (plot 6.1.4). - M. myrmicoxena (1♀) alongside Formica lemani Bondroit, 1917 (5 ☿☿), F. lugubris Zetterstedt, 1838 (13☿☿), M. lobulicornis (12☿☿): Sobretta-Valle del Gavia, Sondrio, coordinates hidden in conformity with Glaser et al. (2011), 2175 m asl, north- facing slope, pasture/shrubland, 04.X.2019 (plot 4.5.2). The geographic range of M. microrubra is relatively wide but does not cover the entire distribution of its host M. rubra (Seifert, 2018). Records of M. microrubra around the Alps often come from sites below 700 m asl (e.g., Wagner, 2020). In Italy, M. rubra records are mostly distributed North of the Apennines and especially common in the Po Plain and Alps (Baroni Urbani, 1971; Mei, 1984; Le Moli & Zaccone, 1995; Glaser, 2003; 2004; Sielezniew et al., 2010; Glaser et al., 2012; Scupola, 2018; Castracani et al., 2020). The first discovery of M. microrubra in Italy was published by Glaser (2003) and made in South Tyrol (980 m asl). Since our record also comes from the Alpine region, it is still uncertain where the species’ southernmost distribution limit stands. During our survey, M. rubra was detected in four sites, mostly occurring from 1400 to 1600 m asl, with a single finding at 1900 m asl. A total of 1848 M. rubra workers were found in 113 samples from pastures or peat bogs. Ecological data on M. microrubra are relatively few and scattered; however, to the best of our knowledge, our record significantly extends its altitudinal limit upwards by about 600 m. On the other hand, ecological data over M. myrmicoxena mostly come from only two sites (Glaser et al., 2011), as very little information is contained in its original description (Forel, 1896). Our data are consistent with these previous two findings regarding altitudinal preferences (M. myrmicoxena was previously found at 1700 and 2213 m asl), habitat selection (Alpine pastures), North-facing slopes, and host species. The new site is geographically placed between the two Swiss sites and the Italian site reported by Glaser et al. (2011) (Fig 2). We collected M. lobulicornis, possibly the exclusive host of M. myrmicoxena, on 151 samples distributed in 15 sites from 1600 to 2400 m asl, resulting in 962 workers. Sites above 1900 m mostly represented open habitats, while a few sites at lower altitudes were coniferous and deciduous forests or peat bogs. Sociobiology 68(4): e7276 (December, 2021) 3 In comparison, the related M. lobicornis, which appears to be the only other species that could potentially serve as a host for M. myrmicoxena (Glaser et al., 2011), was collected between 1400 (the lowest sampled altitude) and 2000 m asl in 12 sites. Given the extreme scarcity of data on M. myrmicoxena and the conservation concerns expressed by previous authors, this additional discovery significantly reinforces the ecological and biological hypotheses formulated so far (Glaser et al., 2011). The Italian distribution of M. lobulicornis currently appears poorly documented, and future investigations may contribute to either confirming or dismissing M. myrmicoxena as an Alpine endemic ant (a rare condition only shared with F. paralugubris Seifert, 1996 – see Seifert, 2018). Myrmica microrubra and M. myrmicoxena were caught only in 0.7-0.9% of the samples in which their respective host species were found. During the two-years survey, only a single specimen of another inquiline social parasite (Formicoxenus nitidulus (Nylander, 1846) (also considered Vulnerable by the IUCN, Social Insect Specialists Group, 1996) was found in about 980 samples containing its host species (a queen was found in the same locality of M. myrmicoxena in a trap retrieved on 16.VII.2018). This collection number represents a lower rate compared to the parasite Myrmica spp. despite literature data suggesting this species is frequent in the Italian Alps (Baroni Urbani, 1971). Inquiline social parasites are difficult to detect during standardized field surveys with a generic focus on ants (e.g., Glaser, 2004; Glaser et al., 2012; Fig 1. Myrmica microrubra (left) and M. myrmicoxena (right) from the Stelvio National Park. Scale bars: 0.5 mm. Enrico Schifani et al. – m Myrmica microrubra and M. myrmicoxena in the Alps4 Spotti et al., 2015), and pitfall trapping is not a proper sampling method useful to detect inquiline parasites. Yet, our findings demonstrate that even on these ants, important information can still be recovered thanks to the extensive use of this sampling tool in studies with a broader ecological aim. In both our findings, the parasites were found in the same sample along with several individuals of their respective host species, which suggests they may have been caught during colony relocation. Finally, insect decline and its severe potential outcomes on ecosystems functioning has recently attained much attention showing the need for a deeper commitment in field survey, the development of effective monitoring systems, and biodiversity database implementation in the face of global changes and contexts with different anthropic impact (Campanaro et al., 2011; Gibb et al., 2017; Leather, 2017; Homburg et al., 2019). Acknowledgments This work was supported by the Stelvio National Park as part of the mid-term monitoring of Alpine faunistic biodiversity concerning climate change, carried out in coordination and with the same survey design of the other three Italian Alpine national parks and in cooperation with MUSE- Science Museum of Trento (Italy). The work was granted by the Italian Ministry for Environment, Land and Sea Protection and co-financed by the Autonomous Province of Trento. Alessandro Gugiatti and Paolo Belotti supervised and managed fieldwork for the survey of epigean arthropod fauna. Teresa Boscolo and Michael Bernasconi (MUSE-Science Museum of Trento, Italy) sorted the Formicidae collected by pitfall traps, and Alessandra Franceschini (MUSE -Science Museum of Trento, Italy) managed the ant collection. This work was also supported by grants from the University of Parma (FIL-2020) assigned to A. Mori and benefited from the equipment and framework of the COMP- HUB Initiative, funded by the “Departments of Excellence” program of the Italian Ministry for Education, University and Research (MIUR, 2018–2022). Authors’ Contributions ES: conceptualization, methodology, investigation, data curation, writing-original draft, writing-review & editing, visualization. CC: conceptualization, methodology, validation, resources, data curation, writing-review & editing, supervision, project administration. LP: methodology, validation, resources, data curation, supervision, project administration, funding acquisition. MGo: methodology, validation, resources, data curation, writing-review & editing, supervision, project administration, writing-review & editing. VL: validation, resources, data curation, supervision, project administration, funding acquisition. FAS: validation, resources, writing-review & editing. AM: resources, writing-review & editing, supervision, project administration, funding acquisition. DG: visualization, writing-review & editing. DAG: resources, writing-review & editing, supervision, project administration, funding acquisition. MGh: resources, writing-review & editing. MG: funding acquisition. References Baroni Urbani, C. (1971). Catalogo delle specie di Formicidae d’Italia (Studi sulla mirmecofauna d’Italia X). Memorie della Società Entomologica Italiana, 50: 5-287. Fig 2. The four Alpine sites where Myrmica myrmicoxena has been found in Italy and Switzerland. Sociobiology 68(4): e7276 (December, 2021) 5 Bolton, B. 2021. An online catalog of the ants of the world. https://antcat.org. (accessed 26 November 2021). Buschinger, A. (2009). Social parasitism among ants: a review (Hymenoptera: Formicidae). Myrmecological News, 12: 219-235. Campanaro, A., Toni, I., Handerson, S. & Grasso, D.A. (2011). Monitoring of Lucanus cervus by means of remains of predation (Coleoptera: Lucanidae). Entomologia Generalis, 33: 79-89. doi: 10.1127/entom.gen/33/2011/79 Castracani, C., Spotti, F.A., E. Schifani, Giannetti, D., Ghizzoni, M., Grasso, D.A. & Mori, A. (2020). Public engagement provides first insights on Po Plain ant communities and reveals the ubiquity of the cryptic invader Tetramorium immigrans (Hymenoptera, Formicidae). Insects, 11: 678. doi: 10.3390/insects11100678 Degueldre, F., Mardulyn, P., Kuhn, A., Pinel, A., Karaman, C., Lebas, C., Schifani, E., Bračko, G., Wagner, H.C., Kiran, K., Borowiec, L., Passera, L., Abril, S., Espadaler, X. & Aron, S. (2021). Evolutionary history of inquiline social parasitism in Plagiolepis ants. Molecular Phylogenetics and Evolution, 155: 107016. doi: 10.1016/j.ympev.2020.107016 Emery, C. (1909). Über den Ursprung der dulotischen, parasitischen und myrmekophilen Ameisen. Biologisches Zentralblatt, 29: 352-362. Espadaler, X. & López-Soria, L. (1991). Rareness of certain Mediterranean ant species: fact or artifact? Insectes Sociaux, 38: 365-377. doi: 10.1007/BF01241872 Forel, A. (1895). Ueber den Polymorphismus und Ergatomorphismus der Ameisen. Verhandlungen der Gesellschaft Deutscher Naturforscher und Ärzte, 66: 142-147. Gibb, H., Dunn, R.R., Sanders, N.J., Grossman, B.F., Photakis, M., Abril, S., Agosti, D., Andersen, A.N., Angulo, E., Armbrecht, I., Arnan, X., Baccaro, F.B., Bishop, T.R., Boulay, R., Brühl, C., Castracani, C., Cerda, X., Del Toro, I., Delsinne, I., Diaz, M., Donoso, D.A., Ellison, A.M., Enriquez, M.L., Fayle, T.M., Feener, D.H., Jr., Fisher, B.L., Fisher, R.N., Fitzpatrick, M.C., Gómez, C., Gotelli, N.J., Gove, A., Grasso, D.A., Groc, S., Guenard, B., Gunawardene, N., Heterick, B., Hoffmann, B., Janda, M., Jenkis, C., Kaspari, M., Klimes, P., Lach, L., Laeger, T., Lattke, J., Leponce, M., Lessard, J.- P., Longino, J., Lucky, A., Luke, S.H., Majer, J., McGlynn, T.P., Menke, S., Mezger, D., Mori, A., Moses, J., Munyai, T.C., Pacheco, R., Paknia, O., Pearce-Duvet, J., Pfeiffer, M., Philpott, S.M., Resasco, J., Retana, J., Silva, R.R., Sorger, M.D., Souza, J., Suarez, A., Tista, M., Vasconcelos, H.L., Vonshak, M., Weiser, M.D., Yates, M. & Parr, C.L. (2017). A global database of ant species abundances. Ecology, 98: 883-884. doi: 10.1002/ecy.1682. Glaser F. (2003). Die Ameisenfauna (Hymenoptera, Formicidae) des Vinschgaus (Südtirol, Italien) eine vorläufige Artenliste. Gredleriana, 3: 209-230. Glaser, F. (2004). Verbreitung und Gefährdung von Ameisen (Hymenoptera, Formicidae) in Auen-und Uferlebensräumen der Etsch (Südtirol, Italien). Gredleriana, 4: 209-230. Glaser, F., Michael, J. & Seifert, B. (2011). Rediscovered after 140 years at two localities: Myrmica myrmicoxena Forel, 1895 (Hymenoptera: Formicidae). Myrmecological News, 14: 107-111. Glaser, F., Freitag, A. & Martz, H. (2012). Ants (Hymenoptera: Formicidae) in the Münstertal (Val Müstair) – a hot spot of regional species richness between Italy and Switzerland. Gredleriana, 12: 273 - 284. Gobbi, M. (2020). Global warning: challenges, threats and opportunities for ground beetles (Coleoptera: Carabidae) in high altitude habitats. Acta Zoologica Academiae Scientiarum Hungaricae, 66: 5-20. doi: 10.17109/AZH.66.Suppl.5.2020 Hölldobler, B. & Wilson, E.O. (1990). The ants. Cambridge: Harvard University Press, 732 p Homburg, K., Drees, C., Boutaud, E., Nolte, D., Schuett, W., Zumstein, P., von Ruschkowski, E. & Assmann, T. (2019). Where have all the beetles gone? Long-term study reveals carabid species decline in a nature reserve in Northern Germany. Insect Conservation and Diversity, 12: 268-277. doi: 10.1111/icad.12348 Jansen, G., Savolainen, R. & Vepsäläinen, K. (2010). Phylogeny, divergence-time estimation, biogeography and social parasite- host relationships of the Holarctic ant genus Myrmica (Hymenoptera: Formicidae). Molecular Phylogenetics and Evolution, 56: 294-304. doi: 10.1016/j.ympev.2010.01.029 Latella, L., Pedrotti, L. & Gobbi, M. (2019). Records of Cholevinae (Coleoptera: Leiodidae) sampled by pitfall traps in the Central Italian Alps. Journal of Insect Biodiversity, 13: 36-42. doi: 10.12976/jib/2019.13.2.3. Le Moli, F. & Zaccone, A. (1995). Ricerche sulla mirmecofauna del Cansiglio (Prealpi Carniche). Società Veneziana di Scienza Naturali, 20: 33-52. Leather, S.R. (2017). “Ecological Armageddon”- more evidence for the drastic decline in insect numbers. Annals of Applied Biology, 172: 1-3. doi: 10.1111/aab.12410 Leppänen, J., Vepsäläinen, K. & Savolainen, R. (2011). Phylogeography of the ant Myrmica rubra and its inquiline social parasite. Ecology and Evolution, 1: 46-62. doi: 10.1002/ ece3.6 Leppänen, J., Seppä, P., Vepsäläinen, K. & Savolainen, R. (2015). Genetic divergence between the sympatric queen morphs of the ant Myrmica rubra. Molecular Ecology, 24: 2463-2476. doi: 10.1111/mec.13170 Leppänen, J., Seppä, P., Vepsäläinen, K., & Savolainen, R. (2016). Mating isolation between the ant Myrmica rubra and its microgynous social parasite. Insectes Sociaux, 63: 79-86. doi: 10.1007/s00040-015-0438-y Enrico Schifani et al. – m Myrmica microrubra and M. myrmicoxena in the Alps6 Pearson, B. & Child, A.R. (1980). The distribution of an esterase polymorphism in macrogynes and microgynes of Myrmica rubra Latreille. Evolution, 34: 105-109. doi: 10.2307/2408318 Radchenko, A. & Elmes, G.W. (2003). A taxonomic revision of the socially parasitic Myrmica ants (Hymenoptera: Formicidae) of the Palaearctic region. Annales Zoologici, 53: 217-243. Radchenko, A. & Elmes, G.W. (2010). Myrmica ants (Hymenoptera: Formicidae) of the old world. Warszawa: Natura optima dux Foundation, 789 p Schär, S. & Nash, D.R. (2014). Evidence that microgynes of Myrmica rubra ants are social parasites that attack old host colonies. Journal of Evolutionary Biology, 27: 2396-2407. doi: 10.1111/jeb.12482 Schifani, E. (2017). First record of the vulnerable social parasite ant Plagiolepis grassei in Italy (Hymenoptera: Formicidae). Fragmenta Entomologica, 49: 61-64. doi: 10. 4081/fe.2017.231 Schifani, E., Castracani, C., Spotti, F. A., Giannetti, D., Ghizzoni, M., Gobbi, M., Pedrotti, L., Grasso, D.A. & Mori, A. (2020). Ergatandromorphism in the Ant Myrmica lobulicornis Nylander, 1857 (Formicidae: Myrmicinae). Sociobiology, 67: 330-334. doi: 10.13102/sociobiology.v67i2.5084 Scupola, A. (2018). The ants of Veneto. Verona: WBA Handbooks, 336 p Seifert, B. (1994). Taxonomic description of Myrmica microrubra n. sp.-a social parasitic ant so far known as the microgyne of Myrmica rubra (L.). Abhandlungen und Berichte des Naturkundemuseums Görlitz, 67: 9-12. Seifert B. (2018). Ants of Northern and Central Europe. Tauer: Lutra Verlags- und Vertriebsgesellschaft, 408 p Seppä, P. & Pamilo, P. (1995). Gene flow and population viscosity in Myrmica ants. Heredity, 74: 200-209. Sielezniew, M., Patricelli, D., Dziekanska, I., Barbero, F., Bonelli, S., Casacci, L.P., Witek, M. & Balletto, E. (2010). The first record of Myrmica lonae (Hymenoptera: Formicidae) as a host of the socially parasitic large blue butterfly Phengaris (Maculinea) arion (Lepidoptera: Lycaenidae). Sociobiology 56: 465-475. Mei, M. (1984). Nuovi reperti de formicidi per l’Italia centrale (Hymenoptera, Formicidae). Bollettino dell’Associazione Romana di Entomologia, 37: 49-58. Social Insects Specialist Group. (1996). Formicidae. In: 2006 IUCN Red List of Threatened Species. Available from: http:// www.iucnredlist.org Spotti, F.A., Castracani, C., Grasso, D.A. & Mori, A. (2015). Daily activity patterns and food preferences in an alpine ant community. Ethology, Ecology and Evolution, 27: 306-324. doi: 10.1080/03949370.2014.947634 Steiner, F.M., Schlick-Steiner, B.C., Konrad, H., Moder, K., Christian, E., Seifert, B., Crozier, R., Stauffer, C. & Buschinger, A. (2006). No sympatric speciation here: multiple data sources show that the ant Myrmica microrubra is not a separate species but an alternate reproductive morph of Myrmica rubra. Journal of Evolutionary Biology, 19: 777- 787. doi: 10.1111/j.1420-9101.2005.01053.x Visicchio, R., Mori, A., Grasso, D.A., Castracani, C. & Le Moli, F. (2001). Glandular sources of recruitment, trail, and propaganda semiochemicals in the slave-making ant Polyergus rufescens. Ethology, Ecology and Evolution, 13: 361-372. doi: 10.1080/08927014.2001.9522767 Wagner, H.C. (2020). The geographic distribution of ants (Hymenoptera: Formicidae) in Styria (Austria) with a focus on material housed in the Universalmuseum Joanneum. Joannea Zoologie, 18: 33-152.