Evolution of genital asymmetry, exaggerated eye stalks, and extreme palpal elongation in Panjange spiders (Araneae: Pholcidae)

Bernhard Huber, Olga M. Nuñeza


We describe several new species of the previously monotypic Panjange lanthana species group from the Philippines and document their extraordinary morphology. Some species show strong male genital asymmetry, a phenomenon that seems to be exceedingly rare in spiders. Males of most species have eye stalks, and in two species these eye stalks are among the longest ever recorded in spiders. Some species show a tendency for male genital (pedipalp) elongation, and one species has the longest and thinnest palps ever recorded in Pholcidae. A cladistic analysis is performed including all described and several undescribed species of Panjange (except for one “problem species”), supporting the lanthana group and its close relationship with members of the possibly paraphyletic cavicola group. The following eight new species are described: Panjange malagos Huber sp. nov.; Pa. casaroro Huber sp. nov.; Pa. camiguin Huber sp. nov.; Pa. hamiguitan Huber sp. nov.; Pa. isarog Huber sp. nov.; Pa. dinagat Huber sp. nov.; Pa. marilog Huber sp. nov.; Pa. bukidnon Huber sp. nov.


Genital asymmetry; eye stalks; palpal elongation; sexual selection; Panjange; Philippines

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DOI: https://doi.org/10.5852/ejt.2015.169


Agnarsson I. 2006. Asymmetric female genitalia and other remarkable morphology in a new genus of cobweb spiders (Theridiidae, Araneae) from Madagascar. Biological Journal of the Linnean Society 87: 211–232. http://dx.doi.org/10.1111/j.1095-8312.2006.00569.x

Baker R.H. & Wilkinson G.S. 2001. Phylogenetic analysis of sexual dimorphism and eye-span allometry in stalk-eyed flies (Diopsidae). Evolution 55 (7): 1373–1385. http://dx.doi.org/10.1111/j.0014-3820.2001.tb00659.x

Barnes R.S.K. 1968. On the evolution of elongated ocular peduncles in the Brachyura. Systematic Zoology 17: 182–187.

Bowman T.E. 1984. Stalking the wild crustacean: The significance of sessile and stalked eyes in phylogeny. Journal of Crustacean Biology 4: 7–11. http://dx.doi.org/10.2307/1547891

Brady A.R. 1964. The lynx spiders of North America, north of Mexico (Araneae: Oxyopidae). Bulletin of the Museum of comparative Zoology 131 (3): 429–518.

Brady A.R. 1975. The lynx spider genus Oxyopes in Mexico and Central America (Araneae: Oxyopidae). Psyche 82 (2): 189–243. http://dx.doi.org/10.1155/1975/24938

Brown B.V. 1993. A further chemical alternative to critical-point-drying for preparing small (or large) flies. Fly Times 11: 10.

Burger M. 2010. Complex female genitalia indicate sperm dumping in armored goblin spiders (Arachnida, Araneae, Oonopidae). Zoology 113: 19–32. http://dx.doi.org/10.1016/j.zool.2009.04.002

Burkhardt D. & de la Motte I.1988. Big ‘antlers’ are favoured: female choice in stalk-eyed flies (Diptera, Insecta), field collected harems and laboratory experiments. Journal of Comparative Physiology A 162 (5): 649–652. http://dx.doi.org/10.1007/BF01342640

Deeleman-Reinhold C.L. 1986a. 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: 45–48. Smithsonian Institution Press, Washington.

Deeleman-Reinhold C.L. 1986b. Studies on tropical Pholcidae II. Redescription of Micromerys gracilis Bradley and Calapnita vermiformis Simon (Araneae, Pholcidae) and description of some related new species. Memoirs of the Queensland Museum 22 (2): 205–224.

Deeleman-Reinhold C.L. & Deeleman P.R. 1983. Studies on tropical Pholcidae I. Panjange, a new genus of Indo-Australian leaf- and rock-dwelling pholcid spiders (Araneae). Zoologische Medelelingen 57 (14): 121–130.

Deeleman-Reinhold C.L. & Platnick N.I. 1986. A new Panjange from northern Borneo (Araneae, Pholcidae). Journal of the New York Entomological Society 94 (4): 559–561.

Eberhard W.G. & Huber B.A. 2010. Spider genitalia: precise maneuvers with a numb structure in a complex lock. In: Leonard J.L. & Córdoba-Aguilar A. (eds) Evolution of Primary Sexual Characters in Animals: 249–284. Oxford University Press, Oxford, U.K.

Gerhardt U. 1928. Biologische Studien an griechischen, corsischen und deutschen Spinnen. Zeitschrift für Morphologie und Ökologie der Tiere 10 (4): 576–675. http://dx.doi.org/10.1007/BF00419324

Gerhardt U. 1930. Biologische Untersuchungen an südfranzösischen Spinnen. Zeitschrift für Morphologie und Ökologie der Tiere 19 (1): 184–227. http://dx.doi.org/10.1007/BF00412295

Goloboff P.A. 1993. NONA (NO NAME) ver. 2. Published by the author, Tucumán, Argentina. Available from http://www.lillo.org.ar/phylogeny/ [accessed 21 Nov. 2015]

Goloboff P.A. 1997. Pee-Wee, ver. 2.8. Published by the author, New York. Available from http://www.lillo.org.ar/phylogeny/ [accessed 21 Nov. 2015]

Goloboff P.A., Farris J.S. & Nixon K. 2004. TNT, tree analysis using new technology, version 1.1. sponsored by the Willi Hennig Society. Available from http://www.lillo.org.ar/phylogeny/ [accessed 21 Nov. 2015]

Goloboff P.A., Farris J.S. & Nixon K. 2008. TNT, a free program for phylogenetic analysis. Cladistics 24 (5): 774–786. http://dx.doi.org/10.1111/j.1096-0031.2008.00217.x

Huber B.A. 1997a. Evidence for gustatorial courtship in a haplogyne spider (Hedypsilus culicinus: Pholcidae: Araneae). Netherlands Journal of Zoology 47 (1): 95–98. http://dx.doi.org/10.1163/156854297X00265

Huber B.A. 1997b. On American ‘Micromerys’ and Metagonia (Araneae, Pholcidae), with notes on natural history and genital mechanics. Zoologica Scripta 25 (4): 341–363. http://dx.doi.org/10.1111/j.1463-6409.1996.tb00170.x

Huber B.A. 1999. Sexual selection in pholcid spiders (Araneae, Pholcidae): artful chelicerae and forceful genitalia. Journal of Arachnology 27 (1): 135–141.

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. http://dx.doi.org/10.1206/0003-0090(2001)260<0001:TPOAAP>2.0.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. http://dx.doi.org/10.1017/S0952836903004709

Huber B.A. 2006. Cryptic female exaggeration: the asymmetric female internal genitalia of Kaliana yuruani (Araneae: Pholcidae). Journal of Morphology 276 (6): 705–712. http://dx.doi.org/10.1002/jmor.10431

Huber B.A. 2010. Mating positions and the evolution of asymmetric insect genitalia. Genetica 138: 19–25. http://dx.doi.org/10.1007/s10709-008-9339-6

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. 2013. Revision and cladistic analysis of the Guineo-Congolian spider genus Smeringopina Kraus (Araneae, Pholcidae). Zootaxa 3713: 1–160. http://dx.doi.org/10.11646/zootaxa.3713.1.1

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

Huber B.A. & Fleckenstein N. 2008. Comb-hairs on the fourth tarsi in pholcid spiders (Araneae, Pholcidae). Journal of Arachnology 36: 232–240. http://dx.doi.org/10.1636/CSh07-71.1

Huber B.A., Sinclair B. & Schmitt M. 2007. The evolution of asymmetric genitalia in spiders and insects. Biological Reviews 82: 647–698. http://dx.doi.org/10.1111/j.1469-185X.2007.00029.x

Jocqué R., Bosselaers J. & Henrard A. 2012. A new species of Hortipes (Araneae, Corinnidae), the first spider with an insertable retrolateral tibial apophysis on the male palp. European Journal of Taxonomy 26: 1–11. http://dx.doi.org/10.5852/ejt.2012.26

Kuntner M., Coddington J.A. & Schneider J.M. 2009. Intersexual arms race? Genital coevolution in nephilid spiders (Araneae, Nephilidae). Evolution 63 (6): 1451–1463. http://dx.doi.org/10.1111/j.1558-5646.2009.00634.x

Kunz K., Garbe S. & Uhl G. 2012. The function of the secretory cephalic hump in males of the dwarf spider Oedothorax retusus (Linyphiidae: Erigoninae). Animal Behaviour 83 (2): 511–517. http://dx.doi.org/10.1016/j.anbehav.2011.11.028

Levi H.W. 1964. American spiders of the genus Phoroncidia (Araneae: Theridiidae). Bulletin of the Museum of Comparative Zoology at Harvard College 131: 65–86.

Marusik Y.M. & Zamani A. 2015. The spider family Filistatidae (Araneae) in Iran. ZooKeys 516: 123–135. http://dx.doi.org/10.3897/zookeys.516.10146

Nixon K.C. 2002. Winclada, version 1.00.08. Program and documentation. Available from http://www.cladistics.com/wincDownload.htm [accessed 21 Nov. 2015]

Panhuis T.M. & Wilkinson G.S. 1999. Exaggerated male eye span influences contest outcome in stalkeyed flies (Diopsidae). Behavioral Ecology and Sociobiology 46 (4): 221–227. http://dx.doi.org/10.1007/s002650050613

Platnick N.I. & Dupérré N. 2009. The American goblin spiders of the new genus Escaphiella (Araneae, Oonopidae). Bulletin of the American Museum of Natural History 328: 1–151. http://dx.doi.org/10.1206/679.1

Platnick N.I. & Dupérré N. 2011. The Andean goblin spiders of the new genera Paradysderina and Semidysderina (Araneae, Oonopidae). Bulletin of the American Museum of Natural History 364: 1–121. http://dx.doi.org/10.1206/771.1

Platnick N.I., Dupérré N., Berniker L. & Bonaldo A.B. 2013. The goblin spider genera Prodysderina, Aschnaoonops, and Bidysderina (Araneae, Oonopidae). Bulletin of the American Museum of Natural History 373: 1–102. http://dx.doi.org/10.1206/822.1

Schaible U., Gack C. & Paulus H.F. 1986. Zur Morphologie, Histologie und biologischen Bedeutung der Kopfstrukturen männlicher Zwergspinnen (Linyphiidae, Erigoninae). Zoologische Jahrbücher. Abteilung für Systematik, Ökologie und Geographie der Tiere 113: 389–408.

Vanacker D., Maes L., Pardo S., Hendrickx F. & Maelfait J.-P. 2003. Is the hairy groove in the gibbosus male morph of Oedothorax gibbosus (Blackwall 1841) a nuptial feeding device? Journal of Arachnology 31: 309–315. http://dx.doi.org/10.1636/s02-06

Wilkinson G.S. & Dodson G.N. 1997. Function and evolution of antlers and eye stalks in flies. In: Choe J.C. & Crespi B.J. (eds) The Evolution of Mating Systems in Insects and Arachnids: 310–328. Cambridge University Press, Cambridge.

World Spider Catalog. 2015. World Spider Catalog. version 16.5. Natural History Museum Bern. Available from http://wsc.nmbe.ch [accessed 17 Aug. 2015]

Zeil J., Nalbach G. & Nalbach H.-O. 1986. Eyes, eye stalks and the visual world of semi-terrestrial crabs. Journal of Comparative Physiology A 159 (6): 801–811. http://dx.doi.org/10.1007/BF00603733


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