Revision of the spider genus Stygopholcus (Araneae, Pholcidae), endemic to the Balkan Peninsula
Abstract
The genus Stygopholcus Kratochvíl, 1932 is endemic to the Balkan Peninsula and includes only four nominal species: the epigean S. photophilus Senglet, 1971 in the south (Greece to Albania) and the ‘northern clade’ consisting of three troglophile species ranging from Croatia to Albania: S. absoloni (Kulczyński, 1914); S. skotophilus Kratochvíl, 1940; and S. montenegrinus Kratochvíl, 1940 (original rank re-established). We present redescriptions of all species, including extensive data on ultrastructure, linear morphometrics of large samples, and numerous new localities. We georeference previously published localities as far as possible, correct several published misidentifications, and clarify nomenclatorial problems regarding the authority of Stygopholcus and the identity of the type species S. absoloni. We suggest that the ‘northern clade’ has a relict distribution, resulting from past and present geologic and climatic factors. Future work on Stygopholcus should focus on the southern Dinarides, combining dense sampling with massive use of molecular data.
References
Absolon K. & Kratochvíl J. 1932. Zur Kenntnis der höhlenbewohnenden Araneae der illyrischen Karstgebiete. Mitteilungen über Höhlen- und Karstforschung 1932: 73–81.
Antić D.Ž., Dražina T., Rađa T., Tomić V.T. & Makarov S.E. 2015. Review of the family Anthogonidae (Diplopoda, Chordeumatida), with descriptions of three new species from the Balkan Peninsula. Zootaxa 3948: 151–181. https://doi.org/10.11646/zootaxa.3948.2.1
Bennett R. 2014. COSEWIC assessment and status report on the northwestern cellar spider Psilochorus hesperus in Canada. Committee on the Status of Endangered Wildlife in Canada, Ottawa. Available from www.registrelep-sararegistry.gc.ca/default_e.cfm [accessed 8 May 2021].
Bilandžija H., Morton B., Podnar M. & Četković H. 2013. Evolutionary history of relict Congeria (Bivalvia: Dreissenidae): unearthing the subterranean biodiversity of the Dinaric Karst. Frontiers in Zoology 10: 5. Available from https://doi.org/10.1186/1742-9994-10-5 [accessed 8 May 2021].
Blackwelder R.E. 1967. Taxonomy. A Text and Reference Book. John Wiley, New York.
Bonnet P. 1958. Bibliographia araneorum. Analyse méthodique de toute la littérature aranéologique jusqu’en 1939. Tome II. Systématique des araignées (Étude par ordre alphabétique) (4me partie: N-S). Douladoure, Toulouse.
Bregović P., Fišer C. & Zagmajster M. 2019. Contribution of rare and common species to subterranean species richness patterns. Ecology and Evolution 9: 11606–11618. https://doi.org/10.1002/ece3.5604
Brignoli P.M. 1971. Beitrag zur Kenntnis der mediterranen Pholcidae (Arachnida, Araneae). Mitteilungen aus dem Zoologischen Museum in Berlin 47: 255–267. https://doi.org/10.1002/mmnz.1971047020
Brignoli P.M. 1976. Ragni di Grecia IX. Specie nuove o interessanti delle famiglie Leptonetidae, Dysderidae, Pholcidae ed Agelenidae (Araneae). Revue suisse de Zoologie 83: 539–578. https://doi.org/10.5962/bhl.part.91452
Brignoli P.M. 1983. A Catalogue of the Araneae described between 1940 and 1981. Manchester University Press, Manchester.
Brown B.V. 1993. A further chemical alternative to critical-point-drying for preparing small (or large) flies. Fly Times 11: 10.
Deeleman-Reinhold C.L. 1978. Revision of the Cave-dwelling and related Spiders of the Genus Troglohyphantes Joseph (Linyphiidae), with special Reference to the Yugoslav Species. Academia Scientiarum et Artium Slovenica, Ljubljana.
Deeleman-Reinhold C.L. 1993. The genus Rhode and the harpacteine genera Stalagtia, Folkia, Minotauria, and Kaemis (Araneae, Dysderidae) of Yugoslavia and Crete, with remarks on the genus Harpactea. Revue arachnologique 10: 105–135.
Deeleman-Reinhold C.L. & Deeleman P.R. 1988. Revision des Dysderinae (Araneae, Dysderidae), les espèces mediterranéennes occidentales exceptées. Tijdschrift voor Entomologie 131: 141–269.
Deltshev C.D. 1985. New data concerning cave spiders (Araneae) in Greece with description of a new Leptonetela (Araneae, Leptonetidae). Acta zoologica Bulgarica 27: 41–45.
Deltshev C.D. 2004. A zoogeographical review of the spiders (Araneae) of the Balkan Peninsula. In: Griffiths H.I., Kryštufek B. & Reed J.M. (eds) Balkan Biodiversity. Pattern and Process in the European Hotspot: 193–200. Springer, Dordrecht. https://doi.org/10.1007/978-1-4020-2854-0
Deltshev C.D., Vrenosi B., Blagoev G. & Lazarov S. 2011. Spiders of Albania – faunistic and zoogeographical review (Arachnida: Araneae). Acta zoologica Bulgarica 63: 125–144.
Džukić G. & Kalezić M.L. 2004. The biodiversity of amphibians and reptiles in the Balkan Peninsula. In: Griffiths H.I., Kryštufek B. & Reed J.M. (eds) Balkan Biodiversity. Pattern and Process in the European Hotspot: 167–192. Springer, Dordrecht. https://doi.org/10.1007/978-1-4020-2854-0
Eastwood W.J. 2004. East Mediterranean vegetation and climate change. In: Griffiths H.I., Kryštufek B. & Reed J.M. (eds) Balkan Biodiversity. Pattern and Process in the European Hotspot: 25–48. Springer, Dordrecht. https://doi.org/10.1007/978-1-4020-2854-0
Eberhard W.G., Huber B.A., Rodriguez R.L., Briceño D., Salas I. & Rodriguez V. 1998. One size fits all? Relationships between the size of genitalia and other body parts in 20 species of insects and spiders. Evolution 52: 415–431. https://doi.org/10.2307/2411078
Eberle J., Dimitrov D., Valdez-Mondragón A. & Huber B.A. 2018. Microhabitat change drives diversifica-tion in pholcid spiders. BMC Evolutionary Biology 18: 141. https://doi.org/10.1186/s12862-018-1244-8
Gottstein Matočec S., Bakran-Petricioli T., Bedek J., Bukovec D., Buzjak S., Franičević M., Jalžić B., Kerovec M., Kletečki E., Kralj J., Kružić P., Kučinić M., Kuhta M., Matočec N., Ozimec R., Rađa T., Štamol V., Ternjej I. & Tvrtković N. 2002. An overview of the cave and interstitial biota of Croatia. Natura Croatica 11: 1–112.
Griffiths H.I., Kryštufek B. & Reed J.M. 2004. Balkan Biodiversity. Pattern and Process in the European Hotspot. Springer, Dordrecht. https://doi.org/10.1007/978-1-4020-2854-0
Hewitt G. 2000. The genetic legacy of the Quaternary ice ages. Nature 405: 907–913. https://doi.org/10.1038/35016000
Hsü K., Ryan W.B.F. & Cita M. 1973. Late Miocene desiccation of the Mediterranean. Nature 242: 240–244. https://doi.org/10.1038/242240a0
Huber B.A. 2000. New World pholcid spiders (Araneae: Pholcidae): a revision at generic level. Bulletin of the American Museum of Natural History 254: 1–348. https://doi.org/10.1206/0003-0090(2000)254<0001:NWPSAP>2.0.CO;2
Huber B.A. 2009. Life on leaves: leaf-dwelling pholcids of Guinea, with emphasis on Crossopriza cylindrogaster, a spider with inverted resting position, pseudo-eyes, lampshade web, and tetrahedral egg-sac (Araneae: Pholcidae). Journal of Natural History 43: 2491–2523. https://doi.org/10.1080/00222930903207876
Huber B.A. 2011a. Revision and cladistic analysis of Pholcus and closely related taxa (Araneae, Pholcidae). Bonner zoologische Monographien 58: 1–509.
Huber B.A. 2011b. Phylogeny and classification of Pholcidae (Araneae): an update. The Journal of Arachnology 39: 211–222. https://doi.org/10.1636/CA10-57.1
Huber B.A. 2012. Revision and cladistic analysis of the Afrotropical endemic genus Smeringopus Simon, 1890 (Araneae: Pholcidae). Zootaxa 3461: 1–138.
Huber B.A. 2013. Revision and cladistic analysis of the Guineo-Congolian spider genus Smeringopina Kraus (Araneae, Pholcidae). Zootaxa 3713: 1–160. https://d.doi.org/10.11646/zootaxa.3713.1.1
Huber B.A. 2014. Pholcidae. In: Roig-Juñent S., Claps L.E. & Morrone J.J. (eds) Biodiversidad de Artrópodos Argentinos, Vol. 3: 131–140. Sociedad Entomológica Argentina.
Huber B.A. 2020. Revision of the spider genus Hoplopholcus Kulczyński (Araneae, Pholcidae). Zootaxa 4726: 1–94. https://doi.org/10.11646/zootaxa.4726.1.1
Huber B.A. & Chao A. 2019. Inferring global species richness from megatransect data and undetected species estimates. Contributions to Zoology 88: 42–53. https://doi.org/10.1163/18759866-20191347
Huber B.A. & Fleckenstein N. 2008. Comb-hairs on the fourth tarsi in pholcid spiders (Araneae, Pholcidae). Journal of Arachnology 36: 232–240. https://doi.org/10.1636/CSh07-71.1
ICZN. 1999. International Code of Zoological Nomenclature. Fourth edition. London, U.K. International Trust for Zoological Nomenclature. Available from https://www.iczn.org/the-code/ [accessed 8 May 2021].
Kissel C., Speranza F. & Milicevic V. 1995. Paleomagnetism of external southern and central Dinarides and northern Albanides: Implications for the Cenozoic activity of the Scutari-Pec transverse zone. Journal of Geophysical Research 100: 14999–15007. https://doi.org/10.1029/95JB01243
Komnenov M. 2009. Checklist of spiders (Araneae) of Bosnia and Herzegovina. Prilozi fauni Bosne i Hercegovine 5: 51–69.
Komnenov M. 2020. Subterranean spiders (Arachnida, Araneae) of Montenegro. The Montenegrin Academy of Sciences and Arts, Proceedings of the Section of Natural Sciences 23 (2019): 119–169.
Kratochvíl J. 1932. Přispĕvek k poznání Araneid střední Slavonie. Sborník Vysoké Školy Zemědělské, Brno (C) 23: 1–16.
Kratochvíl J. 1934. Liste générale des Araignées cavernicoles en Yugoslavie. Pregled pecinskih paukova u Jugoslaviji. Prirodoslovne Razprave 2: 165–226.
Kratochvíl J. 1935. Araignées cavernicoles de Krivošije. Práce Moravské Přírodovědecké Společnosti 9: 1–25.
Kratochvíl J. 1940. Etude sur les araignées cavernicoles du genre Stygopholcus Krat. Acta Societatis Scientiarum Naturalium Moraviae 12: 1–26.
Kratochvíl J. 1978. Araignées cavernicoles des îles Dalmates. Přírodovědné práce ústavů Československé Akademie Věd v Brně (N.S.) 12: 1–59.
Kraus O. 2000. Why no subspecies in spiders? In: Toft S. & Scharff N. (eds) European Arachnology 2000: Proceedings of the 19th European Colloquium of Arachnology, Århus: 303–314. Aarhus University Press, Aarhus.
Kryštufek B. & Reed J.M. 2004. Pattern and process in Balkan biodiversity – an overview. In: Griffiths H.I., Kryštufek B. & Reed J.M. (eds) Balkan Biodiversity. Pattern and Process in the European Hotspot: 1–8. Springer, Dordrecht. https://doi.org/10.1007/978-1-4020-2854-0
Kulczyński W. 1914. Aranearum species novae minusve cognitae, in montibus Kras dictis a Dre C. Absolon aliisque collectae. Bulletin international de l’Academie des Sciences de Cracovie 1914: 353–387, pl. 16.
Lukić M., Delić T., Pavlek M., Deharveng L. & Zagmajster M. 2020. Distribution pattern and radiation of the European subterranean genus Verhoeffiella (Collembola, Entomobryidae). Zoologica Scripta 49: 86–100. https://doi.org/10.1111/zsc.12392
Mammola S., Cardoso P., Angyal D., Balázs G., Blick T., Brustel H., Carter J., Ćurčić S., Danflous S., Dányi L., Déjean S., Deltshev C., Elverici M., Fernández F., Gasparo F., Komnenov M., Komposch C., Kováč L., Kunt K.B., Mock A., Moldovan O.T., Naumova M., Pavlek M., Prieto C.E., Ribera C., Rozwałka R., Růžička V., Vargovitsh R.S., Zaenker S. & Isaia M. 2019. Local- versus broad-scale environmental drivers of continental β–diversity patterns in subterranean spider communities across Europe. Proceedings of the Royal Society B: Biological Sciences 286: 1–9. https://doi.org/10.1098/rspb.2019.1579
Marusik Y.M. & Koponen S. 2002. Diversity of spiders in boreal and arctic zones. The Journal of Arachnology 30: 205–210. https://doi.org/10.1636/0161-8202(2002)030[0205:DOSIBA]2.0.CO;2
Mihevc A., Prelovšek M. & Zupan Hajna N. 2010. Introduction to the Dinaric Karst. Karst Research Institute and Research Centre of the Slovenian Academy of Sciences and Arts, Postojna, Slovenia.
Naumova M., Lazarov S.P., Petrov B.P. & Deltshev C.D. 2016. New faunistic data on the cave-dwelling spiders in the Balkan Peninsula (Araneae). Ecologica Montenegrina 7: 425–438.
Naumova M., Lazarov S. & Deltshev C.D. 2019. Faunistic diversity of the spiders in Montenegro (Arachnida: Araneae). Ecologica Montenegrina 22: 50–89. https://doi.org/10.37828/em.2019.22.5
Nentwig W., Blick T., Bosmans R., Gloor D., Hänggi A. & Kropf C. 2020. Spinnen Europas. Version November 2020. Available from https://www.araneae.nmbe.ch [accessed 5 Nov. 2020]. https://doi.org/10.24436/1
Pavlek M. & Mammola S. 2021. Niche-based processes explaining the distributions of closely related subterranean spiders. Journal of Biogeography 48: 118–133. https://doi.org/10.1111/jbi.13987
Platania L., Pavlek M. & Arnedo M. 2020. Testing the monophyly of the ground-dweller spider genus Harpactea Bristowe, 1939 (Araneae, Dysderidae) with the description of three new species. Systematics and Biodiversity 18: 688–707. https://doi.org/10.1080/14772000.2020.1776786
Platnick N.I. 1993. Advances in Spider Taxonomy 1988–1991, with Synonymies and Transfers 1940–1980. The New York Entomological Society, New York.
Platnick N.I. 2000. The World Spider Catalog, Version 1.0. Available from https://wsc.nmbe.ch/archive/ [accessed 8 May 2021].
Platnick N.I. 2002. The World Spider Catalog, Version 2.5. Available from https://wsc.nmbe.ch/archive/ [accessed 8 May 2021].
Reed J.M., Kryštufek B. & Eastwood W.J. 2004. The physical geography of the Balkans and nomenclature of place names. In: Griffiths H.I., Kryštufek B. & Reed J.M. (eds) Balkan Biodiversity. Pattern and Process in the European Hotspot: 9–22. Springer, Dordrecht. https://doi.org/10.1007/978-1-4020-2854-0
Roewer C.F. 1942. Katalog der Araneae von 1758 bis 1940. 1. Band (Mesothelae, Orthognatha, Labidognatha: Dysderaeformia, Scytodiformia, Pholciformia, Zodariiformia, Hersiliaeformia, Argyopiformia). Natura, Buchhandlung für Naturkunde und exakte Wissenschaften Paul Budy, Bremen.
Růžička V., Kůrka A., Buchar J. & Řezáč M. 2005. Czech Republic – the type material of spiders (Araneae). Časopis Národního muzea, Řada přírodovědná 174: 13–64.
Scarascia-Mugnozza G., Oswald H., Piussi P. & Radoglou K. 2000. Forests in the Mediterranean region: gaps in knowledge and research needs. Forest Ecology and Management 132: 97–109. https://doi.org/10.1016/S0378-1127(00)00383-2
Senglet A. 1971. Note sur les Pholcidae (Arachn.) de Grèce. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 44: 345–359. https://doi.org/10.5169/seals-401666
Senglet A. 2001. Copulatory mechanisms in Hoplopholcus, Stygopholcus (revalidated), Pholcus, Spermophora and Spermophorides (Araneae, Pholcidae), with additional faunistic and taxonomic data. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 74: 43–67. https://doi.org/10.5169/seals-402796
Sinopoli G., Peyron O., Masi A., Holtvoeth J., Francke A., Wagner B. & Sadori L. 2019. Pollen-based temperature and precipitation changes in the Ohrid Basin (western Balkans) between 160 and 70 ka. Climate of the Past 15: 53–71. https://doi.org/10.5194/cp-15-53-2019
Sket B. 1994. Distribution patterns of some subterranean Crustacea in the territory of the former Yugoslavia. Hydrobiologia 287: 65–75. https://doi.org/10.1007/BF00006897
Sket B., Paragamian K. & Trontelj P. 2004. A census of the obligate subterranean fauna of the Balkan Peninsula. In: Griffiths H.I., Kryštufek B. & Reed J.M. (eds) Balkan Biodiversity. Pattern and Process in the European Hotspot: 49–68. Springer, Dordrecht. https://doi.org/10.1007/978-1-4020-2854-0
Tomić V.T., Ćurčić S.B. & Pešić V.M. 2000. Contribution to the study of some cavernicolous spiders (Arachnida, Araneae) in Montenegro. Archives of Biological Sciences, Belgrade 52: 35P–36P.
Tzedakis P.C. 2004. The Balkans as prime glacial refugial territory of European temperate trees. In: Griffiths H.I., Kryštufek B. & Reed J.M. (eds) Balkan Biodiversity. Pattern and Process in the European Hotspot: 49–68. Springer, Dordrecht. https://doi.org/10.1007/978-1-4020-2854-0
Wheeler Q.D. & Meier R. 2000. Species Concepts and Phylogenetic Theory. A Debate. Columbia University Press, New York.
World Spider Catalog. 2020. World Spider Catalog. Version 21.5. Natural History Museum Bern. Available from http://wsc.nmbe.ch [accessed 5 Nov. 2020]. https://doi.org/10.24436/2
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