On Venezuelan pholcid spiders (Araneae, Pholcidae)

Keywords: Synspermiata, Venezuela, taxonomy, microhabitat shifts, color polymorphism

Abstract

We present a comprehensive revision of the pholcid spider collection of M.A. González-Sponga, who between 1998 and 2011 described 22 new genera and 51 new species of Pholcidae from Venezuela. In addition, we treat the pholcid material collected during three expeditions to Venezuela conducted between 2002 and 2020. Of González-Sponga’s pholcid taxa we recognize three genera and 24 species as valid. We describe 43 new species (all from males and females) in one new and 13 previously described genera; four genera are newly recorded for Venezuela. We describe the previously unknown females of 15 species, present new records for 46 previously described species, synonymize one genus and one species, and correct numerous minor errors in previous publications on Venezuelan pholcids. At the generic level, the Venezuelan pholcid fauna now appears fairly well known, but available data on distribution and endemism suggest that many species remain undiscovered and undescribed. Despite the obvious gaps, our data are congruent with previous studies on other taxa that have the highest levels of endemism in the Venezuelan Andes, the Coastal Ranges, and the Guyana Highlands. The Falcón Region in particular shows a complex mosaic of biogeographic relationships with other regions. We provide new biological data on numerous species. We document the first cases of evolutionary microhabitat shifts in the genera Mecolaesthus Simon, 1893 and Priscula Simon, 1893. We document several cases of close congeners sharing localities, usually in slightly to conspicuously different microhabitats, sometimes apparently in identical microhabitats. We document several cases of color polymorphism, mostly intersexual, in Metagonia conica (Simon, 1893) both intersexual and among males. We document further cases of two rare phenomena in Pholcidae: use of specific non-silken structures for retreats (in Pisaboa Huber, 2000) and egg parasitism (in Priscula).

References

Aharon S., Huber B.A. & Gavish-Regev E. 2017. Daddy-long-leg giants: revision of the spider genus Artema Walckenaer, 1837 (Araneae, Pholcidae). European Journal of Taxonomy 376: 1–57. https://doi.org/10.5852/ejt.2017.376

Astrin J.J., Huber B.A., Misof B. & Klütsch C.F.C. 2006. Molecular taxonomy in pholcid spiders (Pholcidae, Araneae): evaluation of species identification methods using CO1 and 16S rRNA. Zoologica Scripta 35: 441–457. https://doi.org/10.1111/j.1463-6409.2006.00239.x

Astrin J.J., Misof B. & Huber B.A. 2007. The pitfalls of exaggeration: molecular and morphological evidence suggests Kaliana is a synonym of Mesabolivar (Araneae: Pholcidae). Zootaxa 1646: 17–30. https://doi.org/10.11646/zootaxa.1646.1.2

Austin A.D. 1985. The function of spider egg sacs in relation to parasitoids and predators, with special reference to the Australian fauna. Journal of Natural History 19: 359–376. https://doi.org/10.1080/00222938500770261

Baert L. 2014. New spider species (Araneae) from the Galápagos Islands (Ecuador). Bulletin de la Société royale belge d’Entomologie 149: 263–271.

Barrio-Amorós C.L. 1999. Sistematica y biogeografía de los anfibios (Amphibia) de Venezuela. Acta Biológica Venezuelica, Caracas 18: 1–93.

Beatty J.A., Berry J.W. & Huber B.A. 2008. The pholcid spiders of Micronesia and Polynesia (Araneae, Pholcidae). Journal of Arachnology 36: 1–25. https://doi.org/10.1636/H05-66.1

Bonnet P. 1957. Bibliographia araneorum. Analyse méthodique de toute la littérature aranéologique jusqu’en 1939. Tome II. Systématique des araignées (Étude par ordre alphabétique) (3me partie: G-M). Douladoure, Toulouse.

Brignoli P.M. 1975. Über die Gruppe der Haplogynae (Araneae). In: Vlijm L. (ed.) Proceedings of the 6th International Arachnological Congress (1974): 33–38. Nederlandse Entomologische Vereniging, Amsterdam.

Brignoli P.M. 1981. Studies on the Pholcidae, I. Notes on the genera Artema and Physocyclus (Araneae). Bulletin of the American Museum of Natural History 170: 90–100.

Brown K.S. & Fernández Y.F. 1984. Los Heliconiini (Lepidoptera, Nymphalidae) de Venezuela. Boletín de Entomología Venezolana 3: 29–73.

Bruvo-Mađarić B., Huber B.A., Steinacher A. & Pass G. 2005. Phylogeny of pholcid spiders (Araneae: Pholcidae): combined analysis using morphology and molecules. Molecular Phylogenetics and Evolution 37: 661–673. https://doi.org/10.1016/j.ympev.2005.08.016

Caporiacco L. di 1955. Estudios sobre los aracnidos de Venezuela. 2a parte: Araneae. Acta Biologica Venezuelica 1: 265–448.

Carico J.E. 2008. Revision of the Neotropical arboreal spider genus Syntrechalea (Araneae, Lycosoidea, Trechaleidae). Journal of Arachnology 36: 118–130. https://doi.org/10.1636/H07-23.1

Carico J.E. & da Silva E.L.C. 2010. Taxonomic review of the Neotropical spider genus Paradossenus (Araneae: Lycosoidea: Trechaleidae: Trechaleinae) with a new erection of the subfamily Trechaleinae and a key to included genera. Journal of Arachnology 38: 212–236. https://doi.org/10.1636/A09-22.1

Carvalho L.S., Dias S.C., Candiani D.F. & Bonaldo A.B. 2010. On the female of Metagonia taruma (Araneae: Pholcidae), ecology of the pholcid spiders in the Urucu River Basin, Amazonas, Brazil and new records from Brazilian Amazonia. Zoologia (Curitiba) 27: 431–439. https://doi.org/10.1590/S1984-46702010000300016

Carvalho L.S., Martins P.H., Schneider M.C. & Cabra-García J.J. 2017. New records of spiders (Arachnida, Araneae) from the state of Roraima, northern Brazil. Check List 13 (1): 1–23. https://doi.org/10.15560/13.1.2040

Cloudsley-Thompson J.L. 1995. A review of the anti-predator devices of spiders. Bulletin of the British Arachnological Society 10: 81–96.

Colmenares G.P.A. 2008. Tres nuevos registros para la araneofauna Venezolana (Arachnida, Araneae, Pholcidae). Boletín del Centro de Investigaciones Biológicas, Universidad del Zulia 42: 85–92.

Cotoras D.D., Brewer M.S., Croucher P.J.P., Oxford G.S., Lindberg D.R. & Gillespie R.G. 2017. Convergent evolution in the colour polymorphism of Selkirkiella spiders (Theridiidae) from the South American temperate rainforest. Biological Journal of the Linnean Society 120: 649–663. https://doi.org/10.1111/bij.12908

Díaz de Gamero M.L. 1996. The changing course of the Orinoco River during the Neogene: a review. Palaeogeography, Palaeoclimatology, Palaeoecology 123: 385–402. https://doi.org/10.1016/0031-0182(96)00115-0

Eberhard W.G., Agnarsson I. & Levi H.W. 2008. Web forms and the phylogeny of theridiid spiders (Araneae: Theridiidae): chaos from order. Systematics and Biodiversity 6: 415–474. https://doi.org/10.1017/S1477200008002855

Eberle J., Dimitrov D., Valdez-Mondragón A. & Huber B.A. 2018. Microhabitat change drives diversification in pholcid spiders. BMC Evolutionary Biology 18: 141. https://doi.org/10.1186/s12862-018-1244-8

Gertsch W.J. 1982. The spider genera Pholcophora and Anopsicus (Araneae, Pholcidae) in North America, Central America and the West Indies. Association for Mexican Cave Studies Bulletin 8: 95–144.

Gertsch W.J. 1986. The spider genus Metagonia (Araneae: Pholcidae) in North America, Central America, and the West Indies. Texas Memorial Museum Speleological Monographs 1: 39–62.

Gertsch W.J. & Peck S.B. 1992. The pholcid spiders of the Galápagos Islands, Ecuador (Araneae: Pholcidae). Canadian Journal of Zoology 70: 1185–1199. https://doi.org/10.1139/z92-166

González-Sponga M.A. 1998. Arácnidos de Venezuela. Cuatro nuevos géneros y cuatro nuevas especies de la familia Pholcidae Koch, 1850 (Araneae). Memorias de la Sociedad de Ciencias Naturales La Salle 57: 17–31.

González-Sponga M.A. 1999. Arácnidos de Venezuela. Ocho especies nuevas del género Priscula y descripción de Priscula venezuelana Simon, 1893 (Arachnida: Araneae: Pholcidae). Boletin de la Academia de Ciencias Fisicas, Matematicas y Naturales, Caracas 54: 123–168.

González-Sponga M.A. 2003. Arácnidos de Venezuela. Cuatro géneros y cuatro especies nuevas de la familia Pholcidae. Memoria de la Fundación La Salle de Ciencias Naturales 155: 91–104.

González-Sponga M.A. 2004. Arácnidos de Venezuela. Un nuevo género y nuevas especies de la familia Pholcidae (Araneae). Aula y Ambiente 8: 63–76.

González-Sponga M.A. 2005. Arácnidos de Venezuela. Tres nuevos géneros y cuatro nuevas especies de la familia Pholcidae (Araneae). Saber 17: 99–109.

González-Sponga M.A. 2006. Arácnidos de Venezuela. Un nuevo género y cinco nuevas especies de la familia Pholcidae (Araneae). Sapiens. Revista Universitaria de Investigación 7: 9–27.

González-Sponga M.A. 2007. Biodiversidad. Arácnidos de Venezuela. Descripción de cinco nuevas especies del género Physocyclus (Araneae, Pholcidae). Sapiens. Revista Universitaria de Investigación 8: 53–69.

González-Sponga M.A. 2009. Arácnidos de Venezuela. Cuatro nuevos géneros y cinco especies nuevas de la familia Pholcidae (Araneae). Anartia 21: 1–16.

González-Sponga M.A. 2010. Biodiversidad. Arácnidos de Venezuela. Redescripción de seis géneros, cinco especies y descripción de una nueva especie de la familie Pholcidae Koch, 1850 (Araneae). Boletin de da Academia de Ciencias Físicas, Matemáticas y Naturales de Venezuela 68: 9–26.

González-Sponga M.A. 2011a. Descripción de seis nuevas especies del género Blechroscelis (Araneae: Pholcidae). Acta Biologica Venezuelica 28: 23–38.

González-Sponga M.A. 2011b. Biodiversidad de Venezuela. Aracnidos. Descripción de cinco nuevos géneros y cinco nuevas especies de la familia Pholcidae Koch, 1850. Acta Biologica Venezuelica 28: 39–51.

Hieber C.S. 1992. Spider cocoons and their suspension systems as barriers to generalist and specialist predators. Oecologia 91: 530–535. https://doi.org/10.1007/BF00650327

Huber B.A. 1994. Genital morphology, copulatory mechanism and reproductive biology in Psilochorus simoni (Berland, 1911) (Pholcidae; Araneae). Netherlands Journal of Zoology 44: 85–99. https://doi.org/10.1163/156854294X00079

Huber B.A. 1997a. Evidence for gustatorial courtship in a haplogyne spider (Hedypsilus culicinus: Pholcidae: Araneae). Netherlands Journal of Zoology 47: 95–98. https://doi.org/10.1163/156854297X00265

Huber B.A. 1997b. On the distinction between Modisimus and Hedypsilus (Araneae, Pholcidae), with notes on behavior and natural history. Zoologica Scripta 25: 233–240. https://doi.org/10.1111/j.1463-6409.1996.tb00164.x

Huber B.A. 1997c. On American ‘Micromerys’ and Metagonia (Araneae, Pholcidae), with notes on natural history and genital mechanics. Zoologica Scripta 25: 341–363. https://doi.org/10.1111/j.1463-6409.1996.tb00170.x

Huber B.A. 1997d. Redescriptions of Eugène Simon’s neotropical pholcids (Araneae, Pholcidae). Zoosystema 19: 573–612.

Huber B.A. 1998a. Genital mechanics in some Neotropical pholcid spiders (Araneae: Pholcidae), with implications for systematics. Journal of Zoology, London 244: 587–599. https://doi.org/10.1111/j.1469-7998.1998.tb00063.x

Huber B.A. 1998b. The pholcid spiders of Costa Rica (Araneae: Pholcidae). Revista de Biologia Tropical 45: 1583–1634.

Huber B.A. 1998c. Report of some pholcid spiders collected in Guatemala and Honduras (Araneae, Pholcidae). Revue suisse de Zoologie 105: 49-80. https://doi.org/10.5962/bhl.part.80031

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%3C0001:NWPSAP%3E2.0.CO;2

Huber B.A. 2001. The pholcids of Australia (Araneae; Pholcidae): taxonomy, biogeography, and relationships. Bulletin of the American Museum of Natural History 260: 1–144. https://doi.org/10.1206/0003-0090(2001)260%3C0001:TPOAAP%3E2.0.CO;2

Huber B.A. 2004. Evidence for functional segregation in the directionally asymmetric male genitalia of the spider Metagonia mariguitarensis (González-Sponga) (Pholcidae: Araneae). Journal of Zoology, London 262: 317–326. https://doi.org/10.1017/S0952836903004709

Huber B.A. 2005a. Evidence for directional selection on male abdomen size in Mecolaesthus longissimus (Araneae, Pholcidae). Journal of Arachnology 33: 573–581. https://doi.org/10.1636/04-69.1

Huber B.A. 2005b. Revision and cladistic analysis of the spider genus Carapoia González-Sponga (Araneae: Pholcidae), with descriptions of new species from Brazil’s Atlantic forest. Invertebrate Systematics 19: 541–556. https://doi.org/10.1071/IS05038

Huber B.A. 2009. Four new generic and 14 new specific synonymies in Pholcidae, and transfer of Pholcoides Roewer to Filistatidae (Araneae). Zootaxa 1970: 64–68. https://doi.org/10.11646/zootaxa.1970.1.3

Huber B.A. 2011. Revision and cladistic analysis of Pholcus and closely related taxa (Araneae, Pholcidae). Bonner zoologische Monographien 58: 1–509.

Huber B.A. 2012. Revision and cladistic analysis of the Afrotropical endemic genus Smeringopus Simon, 1890 (Araneae: Pholcidae). Zootaxa 3461: 1–138. https://doi.org/10.11646/zootaxa.3461.1.1

Huber B.A. 2014. Pholcidae. In: Roig-Juñent S., Claps L.E. & Morrone J.J. (eds) Biodiversidad de Artrópodos Argentinos 3: 131–140. Sociedad Entomológica Argentina, Tucumán, Argentina.

Huber B.A. 2015. Small scale endemism in Brazil’s Atlantic Forest: 14 new species of Mesabolivar (Araneae, Pholcidae), each known from a single locality. Zootaxa 3942: 1–60. https://doi.org/10.11646/zootaxa.3942.1.1

Huber B.A. 2016. Spider diversity and endemism in a South American hotspot: 20 new species of Carapoia (Araneae: Pholcidae) from Brazil’s Atlantic Forest. Zootaxa 4177: 1–69. https://doi.org/10.11646/zootaxa.4177.1.1

Huber B.A. 2018. The South American spider genera Mesabolivar and Carapoia (Araneae, Pholcidae): new species and a framework for redrawing generic limits. Zootaxa 4395: 1–178. https://doi.org/10.11646/zootaxa.4395.1.1

Huber B.A. 2019. The pholcid spiders of Sri Lanka (Araneae: Pholcidae). Zootaxa 4550: 1–57. https://doi.org/10.11646/zootaxa.4550.1.1

Huber B.A. & Arias C.Q. 2017. Revision of the Venezuelan spider genus Stenosfemuraia González-Sponga, with new generic and specific synonymies (Araneae, Pholcidae). Zootaxa 4341: 495–508. https://doi.org/10.11646/zootaxa.4341.4.3

Huber B.A. & Benjamin S. 2005. The pholcid spiders from Sri Lanka: redescription of Pholcus ceylonicus and description of Wanniyala new genus (Araneae: Pholcidae). Journal of Natural History 39: 3305–3319. https://doi.org/10.1080/00222930500145123

Huber B.A. & Brescovit A.D. 2003. Ibotyporanga Mello-Leitão: tropical spiders in Brazilian semi-arid habitats (Araneae: Pholcidae). Insect Systematics and Evolution 34: 15–20. https://doi.org/10.1163/187631203788964926

Huber B.A. & Carvalho L.S. 2019. Filling the gaps: descriptions of unnamed species included in the latest molecular phylogeny of Pholcidae (Araneae). Zootaxa 4546: 1–96. https://doi.org/10.11646/zootaxa.4546.1.1

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. & Eberhard W.G. 1997. Courtship, copulation, and genital mechanics in Physocyclus globosus (Araneae, Pholcidae). Canadian Journal of Zoology 74: 905–918. https://doi.org/10.1139/z97-109

Huber B.A. & Kwapong P. 2013. West African pholcid spiders: an overview, with descriptions of five new species (Araneae, Pholcidae). European Journal of Taxonomy 59: 1–44. https://doi.org/10.5852/ejt.2013.59

Huber B.A. & Rheims C.A. 2011. Diversity and endemism of pholcid spiders in Brazil’s Atlantic Forest, with descriptions of four new species of the Atlantic Forest endemic genus Tupigea (Araneae: Pholcidae). Journal of Natural History 45: 275–301. https://doi.org/10.1080/00222933.2010.524319

Huber B.A. & Schütte A. 2009. Preliminary notes on leaf-dwelling Metagonia spiders (Araneae: Pholcidae) in the Esquinas Rainforest near La Gamba, Costa Rica: leaf preference, mimesis, and web structure. Contributions to Natural History (Bern) 12: 681–697.

Huber B.A. & Warui C.M. 2012. East African pholcid spiders: an overview, with descriptions of eight new species (Araneae, Pholcidae). European Journal of Taxonomy 29: 1–44. https://doi.org/10.5852/ejt.2012.29

Huber B.A. & Wunderlich J. 2006. Fossil and extant species of the genus Leptopholcus in the Dominican Republic, with the first case of egg-parasitism in pholcid spiders (Araneae: Pholcidae). Journal of Natural History 40: 2341–2360. https://doi.org/10.1080/00222930601051196

Huber B.A., Deeleman-Reinhold C.L. & Pérez-González A. 1999. The spider genus Crossopriza (Araneae, Pholcidae) in the New World. American Museum Novitates 3262: 1–10. Available from http://hdl.handle.net/2246/3105 [accessed 27 Aug. 2020].

Huber B.A., Pérez-González A. & Baptista R.L.C. 2005. Leptopholcus (Araneae: Pholcidae) in continental America: rare relicts in low precipitation areas. Bonner zoologische Beiträge 53: 99–107.

Huber B.A., Fischer N. & Astrin J.J. 2010. High level of endemism in Haiti’s last remaining forests: a revision of Modisimus (Araneae: Pholcidae) on Hispaniola, using morphology and molecules. Zoological Journal of the Linnean Society 158: 244–299. https://doi.org/10.1111/j.1096-3642.2009.00559.x

Huber B.A., Pérez-González A., Astrin J.J., Blume C. & Baptista R. 2013. Litoporus iguassuensis Mello-Leitão, 1918 (Araneae, Pholcidae): camouflaged retreat, sexual dimorphism, female color polymorphism, intra-specific genital variation, and description of the male. Zoologischer Anzeiger 252: 511–521. https://doi.org/10.1016/j.jcz.2012.12.001

Huber B.A., Colmenares P.A. & Ramirez M.J. 2014a. Fourteen new generic and ten new specific synonymies in Pholcidae (Araneae), and transfer of Mystes Bristowe to Filistatidae. Zootaxa 3847: 413–422. https://doi.org/10.11646/zootaxa.3847.3.5

Huber B.A., Carvalho L.S. & Benjamin S.P. 2014b. On the New World spiders previously misplaced in Leptopholcus: molecular and morphological analyses and descriptions of four new species (Araneae, Pholcidae). Invertebrate Systematics 28: 432–450. https://doi.org/10.1071/IS13050

Huber B.A., Le Gall P. & Mavoungou J.F. 2014c. Pholcid spiders from the Lower Guinean region of Central Africa: an overview, with descriptions of seven new species (Araneae, Pholcidae). European Journal of Taxonomy 81: 1–46. https://doi.org/10.5852/ejt.2014.81

Huber B.A., Nuñeza O.M. & Leh Moi Ung C. 2015. Revision, phylogeny, and microhabitat shifts in the Southeast Asian spider genus Aetana (Araneae, Pholcidae). European Journal of Taxonomy 162: 1–78. https://doi.org/10.5852/ejt.2015.162

Huber B.A., Petcharad B., Leh Moi Ung C., Koh J.K.H. & Ghazali A.R.M. 2016a. The Southeast Asian Pholcus halabala species group (Araneae, Pholcidae), new data from field observations and ultrastructure. European Journal of Taxonomy 190: 1–55. https://doi.org/10.5852/ejt.2016.190

Huber B.A., Koh J.K.H., Ghazali A.R.M., Nuñeza O., Leh Moi Ung C. & Petcharad B. 2016b. New leaf- and litter-dwelling species of the genus Pholcus from Southeast Asia (Araneae, Pholcidae). European Journal of Taxonomy 200: 1–45. https://doi.org/10.5852/ejt.2016.200

Huber B.A., Neumann J., Grabolle A. & Hula V. 2017. Aliens in Europe: updates on the distributions of Modisimus culicinus and Micropholcus fauroti (Araneae, Pholcidae). Arachnologische Mitteilungen 53: 12–18. https://doi.org/10.5431/aramit5303

Huber B.A., Eberle J. & Dimitrov D. 2018. The phylogeny of pholcid spiders (Araneae, Pholcidae): a critical evaluation of relationships suggested by molecular data. ZooKeys 789: 51–101. https://doi.org/10.3897/zookeys.789.22781

Huber B.A., Caspar K. & Eberle J. 2019. New species reveal unexpected interspecific microhabitat diversity in the genus Uthina Simon, 1893 (Araneae: Pholcidae). Invertebrate Systematics 33: 181–207. https://doi.org/10.1071/IS18002

Johnson N.F., Chen H. & Huber B.A. 2018. New species of Idris Förster (Hymenoptera, Platygastroidea) from southeast Asia, parasitoids of the eggs of pholcid spiders (Araneae, Pholcidae). ZooKeys 811: 65–80. https://doi.org/10.3897/zookeys.811.29725

Koçak A.Ö. & Kemal M. 2008. New synonyms and replacement names in the genus group taxa of Araneida. Centre for Entomological Studies Ankara, Miscellaneous Papers 139–140: 1–4.

Lessert R. de. 1938. Araignées du Congo Belge. Revue de Zoologie et de Botanique africaines 30: 424–457.

Machado É.O. 2007. Análise cladística do gênero neotropical Mesabolivar González-Sponga, 1998 (Araneae: Pholcidae). Master thesis, University of São Paulo, Brazil.

Machado É.O. 2011. Estratificação de habitat, diversidade e evolução do gênero Mesabolivar González-Sponga, 1998 (Araneae: Pholcidae). PhD thesis, University of São Paulo, Brazil.

Manicom C., Schwarzkopf L., Alford R.A. & Schoener T.W. 2008. Self-made shelters protect spiders from predation. Proceedings of the National Academy of Sciences 105: 14903–14907. https://doi.org/10.1073/pnas.0807107105

Marcuzzi G. 1954. Studies on the fauna of Curaçao and other Caribbean Islands: no 22. Tenebrionid beetles of Curaçao, Aruba, Bonaire, and the Venezuelan Islands. Natuurwetenschappelijke Studiekring voor Suriname en de Nederlandse Antillen 10: 1–36.

Mayr E. & Ashlock P.D. 1991. Principles of Systematic Zoology. McGraw-Hill, New York.

Mello-Leitão C. de. 1947. Some new pholcids of the British Museum. Anais da Academia Brasileira de Ciências 19: 159–164.

Mijares-Urrutia A., Manzanilla-Puppo J. & La Marca E. 1999. Una nueva especie de Tepuihyla (Anura: Hylidae) del noroeste de Venezuela, con comentarios sobre su biogeografía. Revista de Biologia Tropical 47: 1099–1110.

Nuessly G.S. & Goeden R.D. 1984. Aspects of the biology and ecology of Diguetia mojavea Gertsch (Araneae, Diguetidae). Journal of Arachnology 12: 75–85.

Peck S.B. 1982. A contribution to the knowledge of the invertebrate cave faunas of Venezuela: Invertebrate faunas of tropical American caves, part 4. International Journal of Speleology 12: 75–81. https://doi.org/10.5038/1827-806X.12.1.8

Petrunkevitch A. 1929. The spiders of Porto Rico. Transactions of the Connecticut Academy of Arts and Sciences 30: 1–158.

Rivero J.A. 1961. Salientia of Venezuela. Bulletin of the Museum of Comparative Zoology 126: 1–207.

Saaristo M.I. 1999. An arachnological excursion to the granitic Seychelles, 1–26th January 1999. Arachnid species lists for Silhouette, Cousine & Mahé. Phelsuma 7 (suppl.): 1–12.

Saaristo M.I. 2001. Pholcid spiders of the granitic Seychelles (Araneae, Pholcidae). Phelsuma 9: 9–28.

Saaristo M.I. 2010. Araneae. In: Gerlach J. & Marusik Y. (eds) Arachnida and Myriapoda of the Seychelles islands. Siri Scientific Press, Manchester, UK.

Scharff N. & Coddington J.A. 1997. A phylogenetic analysis of the orb-weaving spider family Araneidae (Arachnida, Araneae). Zoological Journal of the Linnean Society 120: 355–434. https://doi.org/10.1111/j.1096-3642.1997.tb01281.x

Simon E. 1893a. Descriptions d’espèces et de genres nouveaux de l’ordre des Araneae. Annales de la Société entomologique de France 62: 299–330.

Simon E. 1893b. Histoire naturelle des araignées. Deuxième édition, tome premier. Roret, Paris.

Tong Y. 2013. Haplogynae Spiders from Hainan, China. Ke xue chu ban she, Beijing.

Tong Y.F. & Li S.Q. 2009. Six new cave-dwelling pholcid spiders (Araneae: Pholcidae) from Hainan Island, with two newly recorded genera from China. Zootaxa 1988: 17–32.

Uhl G. 1993. Mating behaviour and female sperm storage in Pholcus phalangioides (Fuesslin) (Araneae). Memoirs of the Queensland Museum 33: 667–674.

Uhl G. 1994. Genital morphology and sperm storage in Pholcus phalangioides (Fuesslin) (Pholcidae; Araneae). Acta Zoologica, Stockholm 75: 13–25. https://doi.org/10.1111/j.1463-6395.1994.tb00957.x

Uhl G., Nessler S.H. & Schneider J.M. 2010. Securing paternity in spiders? A review on occurrence and effects of mating plugs and male genital mutilation. Genetica 138: 75–104. https://doi.org/10.1007/s10709-009-9388-5

Valdez-Mondragón A. 2010. Revisión taxonómica de Physocyclus Simon, 1893 (Araneae: Pholcidae), con la descripción de especies nuevas de México. Revista Iberica de Aracnología 18: 3–80.

Published
2020-10-01
How to Cite
Huber, B. A., & Villarreal, O. (2020). On Venezuelan pholcid spiders (Araneae, Pholcidae). European Journal of Taxonomy, 718(1), 1-317. https://doi.org/10.5852/ejt.2020.718.1101
Section
Monograph