Abstract
As part of the Western Ghats-Sri Lanka biodiversity hotspot, Sri Lanka harbors a rich and diverse fauna, especially in the perhumid southwestern part of the island. However, many invertebrate groups such as spiders continue to be poorly studied. The present paper reviews our knowledge about Pholcidae, a family of spiders that is well represented in Sri Lanka, both by numerous (10) introduced species and by a rich native fauna in five genera (described native Sri Lankan species in parentheses): Belisana Thorell, 1898 (6), Leptopholcus Simon, 1893 (1), Pholcus Walckenaer, 1805 (4), Tissahamia Huber, 2018 (4), and Wanniyala Huber & Benjamin, 2005 (9). Fourteen species are newly described: Belisana minneriya sp. n., B. badulla sp. n., B. gowindahela sp. n.; Pholcus metta sp. n., P. puranappui sp. n., P. uva sp. n.; Tissahamia karuna sp. n.; Wanniyala mudita sp. n., W. orientalis sp. n., W. upekkha sp. n., W. ohiya sp. n., W. viharekele sp. n., W. mapalena sp. n., and W. labugama sp. n.. All new species are described from males and females. New Sri Lankan records are given for 16 previously described species. Distribution data suggest that most or all of the 24 native species might be endemic to Sri Lanka, but the Indian pholcid fauna remains almost entirely unknown.
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
Ali, J.R. & Aitchison, J.C. (2008) Gondwana to Asia: Plate tectonics, paleogeography and the biological connectivity of the Indian subcontinent from the Middle Jurassic through latest Eocene (166–35 ma). Earth-Science Reviews, 88, 145–166.
https://doi.org/10.1016/j.earscirev.2008.01.007
Benjamin, S.P. (2015) Model mimics: antlike jumping spiders of the genus Myrmarachne from Sri Lanka. Journal of Natural History, 49, 2609–2666.
https://doi.org/10.1080/00222933.2015.1034209
Benjamin, S.P. & Bambaradeniya, C.N.B. (2006) Systematics and conservation of spiders in Sri Lanka: current status and future prospects. In: Bambaradeniya, C.N.B (Ed.), The Fauna of Sri Lanka. IUCN—The World Conservation Union, Colombo, pp. 70–76.
Benjamin, S.P. & Kanesharatnam, N. (2016) Description of three new species of the tropical Asian jumping spider genus Onomastus Simon, 1900 from high altitude cloud forests of Sri Lanka (Araneae: Salticidae). Zootaxa, 4205 (5), 431–453.
https://doi.org/10.11646/zootaxa.4205.5.2
Benjamin, S.P., Nanayakkara, R.P. & Sayananda, S.K. (2012) The taxonomy and conservation status of the spiders (Arachnida: Araneae) in Sri Lanka. In: Weerakoon, D.K. & Wijesundara, S. (Eds.), The National Red List of Sri Lanka. Conservation Status of the Fauna and Flora. Ministry of Environment, Colombo, Sri Lanka, pp. 42–44.
Bossuyt, F., Meegaskumbura, M., Beenaerts, N., Gower, D.J., Pethiyagoda, R., Roelants, K., Mannaert, A., Wilkinson, M., Bahir, M.M., Manamendra-Arachchi, K., Ng, P.K.L., Schneider, C.J., Oommen, V.O. & Milinkovitch, M.C. (2004) Local endemism within the Western Ghats-Sri Lanka biodiversity hotspot. Science, 306, 479–481.
https://doi.org/10.1126/science.1100167
Brignoli, P.M. (1972) Ragni di Ceylon I. Missione biospeleologica Aellen-Strinati (1970) (Arachnida, Araneae). Revue suisse de Zoologie, 79, 907–929.
https://doi.org/10.5962/bhl.part.97143
Brignoli, P.M. (1975) Über die Gruppe der Haplogynae (Araneae). Proceedings of the 6th International Arachnological Congress, 1974, 33–38.
Brignoli, P.M. (1980) Sur le genre Leptopholcus Simon, 1893 (Araneae, Pholcidae). Revue de zoologie africaine, 94, 649–655.
Chown, S.L. & Gaston, K.J. (2000) Areas, cradles and museums: the latitudinal gradient in species richness. Trends in Ecology and Evolution, 15, 311–315.
https://doi.org/10.1016/S0169-5347(00)01910-8
Deeleman-Reinhold, C.L. (1986) Leaf-dwelling Pholcidae in Indo-Australian rain forests. In: Eberhard, W.G., Lubin, Y.D. & Robinson, B.C. (Eds.), Proceedings of the Ninth International Congress of Arachnology, Panama 1983. Smithsonian Institution Press, Washington, D.C., pp. 45–48.
Deeleman-Reinhold, C.L. (1995) Redescription of Holocneminus multiguttatus Simon and description of two new species of pholcid spiders from Australia. Beiträge zur Araneologie, 4, 31–41.
Deeleman-Reinhold, C.L. & van Harten, A. (2001) Description of some interesting, new or little known Pholcidae (Araneae) from Yemen. In: Prakash, I. (Ed.), Ecology of Desert Environments. Scientific Publishers, Jodhpur, pp. 193–207.
Dimitrov, D., Astrin, J.J. & Huber, B.A. (2013) Pholcid spider molecular systematics revisited, with new insights into the biogeography and the evolution of the group. Cladistics, 29, 132–146.
https://doi.org/10.1111/j.1096-0031.2012.00419.x
Dong, T., Zheng, G., Yao, Z. & Li, S. (2016) Fifteen new species of the spider genus Pholcus (Araneae: Pholcidae) from Southeast Asia. Zootaxa, 4136 (2), 201–246.
https://doi.org/10.11646/zootaxa.4136.2.1
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
Gunatilleke, N., Gunatilleke, S. & Ashton, P.S. (2017) South-west Sri Lanka: a floristic refugium in South Asia. Ceylon Journal of Science, 46, 65–78.
https://doi.org/10.4038/cjs.v46i5.7454
Huber, B.A. (1997) On the distinction between Modisimus and Hedypsilus (Pholcidae; Araneae), 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., 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 22 Jan. 2019)
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. (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. (2005) High species diversity, male-female coevolution, and metaphyly in Southeast Asian pholcid spiders: the case of Belisana Thorell 1898 (Araneae, Pholcidae). Zoologica, 155, 1–126.
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 (37), 3305–3319.
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.
Huber, B.A., Carvalho, L.S. & Benjamin, S.P. (2014a) 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., Colmenares, P.A. & Ramirez, M.J. (2014b) Fourteen new generic and ten new specific synonymies in Pholcidae (Araneae), and transfer of Mystes Bristowe to Filistatidae. Zootaxa, 3847 (3), 413–422.
https://doi.org/10.11646/zootaxa.3847.3.5
Huber, B.A., Koh, J.K.H., Ghazali, A.R.M., Nuñeza, O., Leh Moi Ung, C. & Petcharad, B. (2016a) 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., Petcharad, B., Leh Moi Ung, C., Koh, J.K.H. & Ghazali, A.R.M. (2016b) 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., 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. (in press) New species reveal unexpected interspecific microhabitat diversity in the genus Uthina Simon, 1893 (Araneae: Pholcidae). Invertebrate Systematics. [in press]
Irie, T. (2001) Notes on the pholcid spider, Crossopriza lyoni (Blackwall 1867). Kishidaia, 80, 7–20.
Kanesharatnam, N. & Benjamin, S.P. (2016) Three new generic records and descriptions of four new species of jumping spiders (Araneae, Salticidae) from Sri Lanka. European Journal of Taxonomy, 228, 1–23.
https://doi.org/10.5852/ejt.2016.228
Karsch, F. (1891) Arachniden von Ceylon und von Minikoy, gesammelt von den Herren Doctoren P. und F. Sarasin. Berliner entomologische Zeitschrift, 36, 267–310.
Kraus, O. (1957) Araneenstudien 1. Pholcidae (Smeringopodinae, Ninetinae). Senckenbergiana biologica, 38 (3/4), 217–243.
Meegaskumbura, M., Bossuyt, F., Pethiyagoda, R., Manamendra-Arachchi, K., Bahir, M., Milinkovitch, M.C. & Schneider, C.J. (2002) Sri Lanka: An amphibian hot spot. Science, 298, 379.
https://doi.org/10.1126/science.298.5592.379
Murphy, F. & Murphy, J. (2000) An Introduction to the Spiders of South East Asia. Malaysian Nature Society, Kuala Lumpur, 624 pp., 32 pls.
Myers, N., Mittermeier, R.A., Mittermeier, C.G., da Fonseca, G.A. & Kent, J. (2000) Biodiversity hotspots for conservation priorities. Nature, 403, 853–858.
https://doi.org/10.1038/35002501
Pickard-Cambridge, O. (1869) Part I of catalogue of a collection of Ceylon Araneidea lately received from Mr. J. Nietner, with descriptions of new species and characters of a new genus. Journal of the Linnean Society London, Zoology, 10, 373–397.
Pocock, R.I. (1900) The Fauna of British India, including Ceylon and Burma. Arachnida. Taylor and Francis, London, pp. 153–279.
Polotow, D. & Griswold, C. (2017) Cleaning old cabinets: revealing the taxonomy of Sri Lankan wolf spiders (Araneae, Udubidae and Zoropsidae). Zootaxa, 4362 (1), 51–74.
https://doi.org/10.11646/zootaxa.4362.1.3
Pyron, R.A., Dushantha Kandambi, H.K., Hendry, C.R., Pushpamal, V., Burbrink, F.T. & Somaweera, R. (2013) Genus-level phylogeny of snakes reveals the origins of species richness in Sri Lanka. Molecular Phylogenetics and Evolution, 66, 969–978.
https://doi.org/10.1016/j.ympev.2012.12.004
Ranasinghe, U.G.S.L. & Benjamin, S.P. (2016a) The goblin spider genus Xestaspis in Sri Lanka (Araneae: Oonopidae). Zootaxa, 4189 (1), 60–80.
https://doi.org/10.11646/zootaxa.4189.1.2
Ranasinghe, U.G.S.L. & Benjamin, S.P. (2016b) A review of Sri Lankan Brignolia including the description of four new species (Araneae: Oonopidae). Zootaxa, 4144 (4), 451–476.
https://doi.org/10.11646/zootaxa.4144.4.1
Ranasinghe, U.G.S.L. & Benjamin, S.P. (2018) Three new species of Aprusia (Araneae: Oonopidae) from Sri Lanka with a phylogenetic analysis of the genus. Journal of Natural History, 52, 713–738.
https://doi.org/10.1080/00222933.2018.1444803
Roelants, K., Jiang, J. & Bossuyt, F. (2004) Endemic ranid (Amphibia: Anura) genera in southern mountain ranges of the Indian subcontinent represent ancient frog lineages: Evidence from molecular data. Molecular Phylogenetics and Evolution, 31, 730–740.
https://doi.org/10.1016/j.ympev.2003.09.011
Saaristo, M.I. (2001) Pholcid spiders of the granitic Seychelles (Araneae, Pholcidae). Phelsuma, 9, 9–28.
Sherriffs, W.R. (1919) A contribution to the study of South Indian arachnology. The Annals and Magazine of Natural History, Series 9, 4 (22), 220–253.
https://doi.org/10.1080/00222931908673888
Simon, E. (1893) Histoire Naturelle des Araignées. 1 (2). 2nd edition. Roret, Paris, 233 pp. [pp. 256–488]
Simon, E. (1903) Histoire Naturelle des Araignées. 2 (4). 2nd edition. Roret Paris, 412 pp. [pp. 669–1080]
Simon, E. (1905) Arachnides de Java. Mitteilungen des naturhistorischen Museums Hamburg, 22, 49–73.
Simon, E. (1909) Étude sur les arachnides du Tonkin (1ère partie). Bulletin Scientifique de la France et de la Belgique, 42, 69–147.
https://doi.org/10.5962/bhl.part.24151
Song, D., Zhu, M. & Chen, J. (1999) The Spiders of China. Hebei Science and Technology Publishing House, Shijiazhuang, 640 pp., 4 pls.
Strand, E. (1907) Süd- und ostasiatische Spinnen. Abhandlungen der naturforschenden Gesellschaft Görlitz, 25, 107–215, pls. 1–21.
Thorell, T. (1898) Viaggio di Leonardo Fea in Birmania e regioni vicine. LXXX. Secondo saggio sui Ragni birmani. II. Retitelariae et Orbitelariae. Annali del Museo Civico di Storia Naturale di Genova, Series 2, 19 [=39], 271–378.
Tong, Y. (2013) Haplogynae Spiders from Hainan, China. Kexue chubanshe, Beijing, vi + 96 pp., 81 pls.
Valdez-Mondragón, A. (2010) Revisión taxonómica de Physocyclus Simon, 1893 (Araneae: Pholcidae), con la descripción de especies nuevas de México. Revista Ibérica de Aracnología, 18, 3–80.
Van Bocxlaer, I., Biju, S.D., Loader, S.P. & Bossuyt, F. (2009) Toad radiation reveals into-India dispersal as a source of endemism in the Western Ghats-Sri Lanka biodiversity hotspot. BMC Evolutionary Biology, 9, 131.
https://doi.org/10.1186/1471-2148-9-131
Williams, E.E. (1972) The origin of faunas. Evolution of lizard congeners in a complex island fauna: a trial analysis. In: Dobzhansky, T., Hecht, M.K. & Steere, W.C. (Eds.), Evolutionary Biology. Meredith Corporation, New York, pp. 47–89.
https://doi.org/10.1007/978-1-4684-9063-3_3
World Spider Catalog (2018) World Spider Catalog. Version 19.0. Natural History Museum Bern. Available from: http://wsc.nmbe.ch (accessed 16 April 2018)
Yao, Z., Dong, T., Zheng, G., Fu, J. & Li, S. (2016) High endemism at cave entrances: a case study of spiders of the genus Uthina. Scientific Reports, 6, 35757.
https://doi.org/10.1038/srep35757
Yao, Z., Pham, D. & Li, S. (2015) Pholcid spiders (Araneae: Pholcidae) from northern Vietnam, with descriptions of nineteen new species. Zootaxa, 3909 (1), 1–82.
https://doi.org/10.11646/zootaxa.3909.1.1
Zhang, F. & Zhu, M.S. (2009) A review of the genus Pholcus (Araneae: Pholcidae) from China. Zootaxa, 2037, 1–114.