Aops oncodactylus, gen. et sp. nov., the first troglobitic urodacid (Urodacidae : Scorpiones), with a re-assessment of cavernicolous, troglobitic and troglomorphic scorpions

Erich S. Volschenk; Lorenzo Prendini

doi:10.1071/IS06054

RESEARCH ARTICLE

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Aops oncodactylus, gen. et sp. nov., the first troglobitic urodacid (Urodacidae : Scorpiones), with a re-assessment of cavernicolous, troglobitic and troglomorphic scorpions

Erich S. Volschenk ^A ^B ^C and Lorenzo Prendini ^A

+ Author Affiliations

- Author Affiliations

^A Division of Invertebrate Zoology, American Museum of Natural of History, Central Park West at 79th Street, New York, NY 10024-5192, USA.

^B Present address: Department of Terrestrial Invertebrates, Western Australian Museum, Locked Bag 49, Welshpool DC, WA 6986, Australia.

^C Corresponding author. Email: evolsche@gmail.com

Invertebrate Systematics 22(2) 235-257 https://doi.org/10.1071/IS06054
Submitted: 7 December 2006 Accepted: 28 December 2007 Published: 12 May 2008

Abstract

We describe a new genus and species of troglomorphic scorpion from Ledge Cave, Barrow Island, off the north-western coast of Western Australia. This troglobite scorpion was initially difficult to place within the superfamily Scorpionoidea Latreille on account of its unusual morphology. Based on cladistic analysis, we demonstrate that this scorpion is phylogenetically most closely related to the endemic Australian genus Urodacus Peters (Urodacidae Pocock). The new taxon, named Aops oncodactylus, gen. et sp. nov., is the first troglobitic urodacid and the first troglobitic scorpion recorded from continental Australia. In light of this discovery, we re-assess the known diversity of troglobitic scorpions and the troglomorphic adaptations (troglomorphies) of cavernicolous scorpions. A distinction is recognised between the exclusive occurrence in cavernicolous habitats and the presence of troglomorphies, which are exhibited by some endogean (humicolous) scorpions. The definition of troglobitic scorpions is limited to species that are both restricted to cavernicolous habitats and exhibit pronounced troglomorphies. Only 20 scorpion species meet both criteria and are considered unequivocally troglobitic according to this definition.

Additional keywords: Australia, cladistics, evolution, morphology, phylogeny, taxonomy.

Acknowledgements

We are grateful to Bill Humphreys for collecting the holotype of A. oncodactylus and to the following for loaning specimens for this study or facilitating access to the collections under their care during our visits: Charles E. Griswold and Darrell Ubick (CAS); Petra Sierwald (FMNH); Matt E. Braunwalder (MEB); Mark Judson, Wilson Lourenço, and Christine Rollard (MNHN); Torbjörn Kronestedt and Julio Ferrer (NHRM); Robert Raven (QM); Oscar Francke and Edmundo González (UNAM); Jonathan Coddington and Dana de Roche (USNM); Mark Harvey and Julianne Waldock (WAM); W. David Sissom (WDS); Jason Dunlop and Shahin Nawai (ZMB); Hieronymus Dastych (ZMH). We thank Craig Eddie, Gonzalo Giribet, Andrew Gluesenkamp, Hamid Goodarzi, Siegfried Huber, the late Adam Locket, Lionel Monod, Peter Sprouse, Valerio Vignoli and John Visser for donating specimens examined during the course of the study. We appreciate the assistance of Randy Mercurio (AMNH) with the habitus photographs. Three anonymous reviewers provided constructive comments on earlier drafts of this paper. While at the AMNH, ESV was supported by a Genomics Postdoctoral Fellowship and a fellowship from the Richard Lounsbery Foundation. This study was partially supported by an Australian Biological Resources Study research grant (ABRS 206-54) to E. S. Volschenk, M. S. Harvey and L. Prendini.

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Appendix 1. Material examined

Specimens deposited in the following collections: American Museum of Natural History (AMNH), New York, including Alexis Harington (AH) Collection and Lorenzo Prendini (LP) Collection; California Academy of Sciences (CAS), San Francisco; Erich S. Volschenk (ESV) personal collection, Brisbane, Australia; Field Museum of Natural History (FMNH), Chicago; Matt. E. Braunwalder (MEB) personal collection, Zurich, Switzerland; Museum National d’Histoire Naturelle (MNHN), Paris; Naturhistoriska Riksmuseet (NHRM), Stockholm; Queensland Museum (QM), Brisbane, Australia; Coleccion Nacional de Acaros y Aracnidos, Instituto de Biología, Universidad Nacional Autónoma de México (UNAM), México City; United States National Museum of Natural History (USNM), Smithsonian Institution, Washington, DC; Western Australian Museum (WAM), Perth; W. David Sissom (WDS) personal collection, Canyon, TX; Zoologisches Museum der Humboldt-Universität, Berlin (ZMB); Zoologisches Institut und Museum der Universität Hamburg (ZMH).

Bothriuridae Simon, 1880

Lisposoma josehermana Lamoral, 1979: Namibia: 16 ♂ 54 ♀ (AMNH), Farm Varianto on Elandshoek 771, 19°22.773′S 17°44.456′E, Tsumeb District, Oshikoto Region, 4.i.2004, L. Prendini, E. Scott, T. & C. Bird, Q. & N. Martins; 5 ♂ 7 ♀ 5 subad. ♂ 5 subad. ♀ (AMNH), Farm Uisib 427, 15 km NW of Otavi, 19°33.132′S 17°14.124′E, Grootfontein District, Otjozondjupa Region, 2.i.2004, L. Prendini, E. Scott, T. & C. Bird, Q. & N. Martins.

Chactidae Pocock, 1893

Belisarius xambeui Simon, 1879: 1 ♂ [dissected by M. Vachon, no hemispermatophore] (MNHN RS 0790); 1 ♀ (MNHN RS 7748), Malhomme; 1 subad. ♀, 2 juv. ♂ (MNHN RS 0797), 18.xii.1949; 1 subad. ♂ (MNHN RS 0793), Amèlie, les Bains, France, iv.1900; 1 ♀ (AMNH LP 2904), 1 juv. (AMNH LP 2851), East Pyrenees, c. 3 km above Prats de Mollo, 42°23.707′N 2°28.546′E, France, 10.vi.2003, S. Huber, 2725 ft, under leaf litter, scree and boulders (drainage); 2 juv. ♂, 1 juv. ♀ (MNHN RS 0789), Fr. De Montbolo, Pyrenees Orientales, France, 17.iv.1946, Coiffat; 1 juv. ♀ (MNHN RS 0794), Grotte pres de la metarie d’en Briscot a La Preste, commun d’Arles-sur-Tech, Pyrenees Orientales, France, 6.iv.1929, J. R. Denis & P. Remy, in cave, 40 m from entrance in total darkness; 1 subad. ♀, 4 juv. (MNHN RS 0792), La Preste, Pyrenees Orientales, France, Nadar; 1 subad. ♀ (MNHN RS 4708), Prats de Mollo, Pyrenees Orientales, France, 23.iv.1935, G. Colas; 2 ♀ (MNHN RS 0795), Pyrenees Orientales, France; 1 juv. (MNHN RS 0796), Pyrenees Orientales, France; 1 juv. (AMNH LP 1891), Pyrenees Orientales, Mts between La Preste and Prats de Mollo, France, 2001, L. Monod; 1 ♀ (MNHN RS 0791), Vernet les Bains, France, Bolivar; 1 ♀ (MNHN), Cav Petit Terrades (Alt Empordà), Spain, 13.iv.1979, X. Bellés & J. Comas; 1 ♀ (MNHN RS 0798), Cav del Ciri, San Lorenzo Muga, Spain, vi.1944, Espanol; 2 ♀, 1 juv. ♀ (MNHN RS 7743), Mataro, Spain, 31.viii.1948, J. Nègre; 1 ♂ (MNHN), Av. Toni (S of Barcelona), Spain, 9.v.1976, O. Escolá; 1 ♀, 1 juv. (MNHN RS 0802), San Bartolo me. Vidra, P. Barcelona, Spain, 12.x.1950, J. Mateu, en foret du Fagus sous les cailloux; 1 juv. (AMNH LP 2325), Montseny, 41°45′45″N 2°21′53″W, Prov. de Barcelona, Spain, 17.vii.2002, G. Giribet, E. Mateos & N. Lopez, 820 m; 1 subad. ♀, 1 juv. ♂ (MNHN), Av. Taleixà, Girona, Spain, 1.xi.1970, E. Vives; 1 juv. ♂ (MNHN RS 0799), Avenc de Taleiczà, Terradas, Girona, Spain, 6.vi.1954, Heurob; 2 ♀ (MNHN RS 0801), Camprodon, Girona, Spain, ix.1954, J. Mateu; 2 ♀ (MNHN RS 0800), Camprodon, Girona, Spain, 17.x.1946, Espanol; 1 ♀ (MNHN RS 3230), Camprodon, Girona, Spain, J. Mateu; 1 ♀, 1 subad. ♀, 1 juv. ♀ (MNHN RS 4582), Camprodon, Forêt de Hêtres, Girona, Spain, viii.1963, J. Mateu; 2 juv. (MNHN RS 0803), Cueva Vera Major, Terrades, Girona, Spain, 23.vi.1944, Espanol; 1 ♀ (AMNH LP 1629), Fajeda d’en Jordà, Girona, Spain, 5.xi.1993, G. Giribet & C. Ribera; 2 ♂ (MNHN), Les Guilleriers, Girona, Spain, 24.v.1981, E. Vives; 1 ♀ (MNHN), Puig Sacalin, Girona, Spain, 30.v.1982, E. Vives; 1 juv. (MNHN RS 4554), San Juan de las Abadesas, Girona, Spain, 3.xi.1962, H. C.; 2 ♀ (MNHN), Santigosa, Girona, Spain, 21.viii.1969, E. Vives; 2 ♀ (MNHN), Santigosa, Girona, Spain, 3.viii.1976, E. Vives; 1 juv. ♀ (MNHN), Vall de Vianya, Fageda, Girona, Spain, 30.iv.1978, Espadaler, 800 m; 2 ♂ (MEB 002), Engracia near Amélie-les-Bains, Pyrenees, France, 16.vi.1983, M. E. Braunwalder; 1 ♀ (MEB 080), Engracia near Amélie-les-Bains, Pyrenees, France, 5.vi.1984, M. E. Braunwalder; 1 ♀ (MEB 081) Banat, near Prat-de-Mollo, Pyrenees, France, M. E. Braunwalder, 6.vi.1984; 2 ♂ (MEB 823), Ravin d’En, Sant Magi near Taulis, Roussillon-Pyrénées, France, 26.v.1995, M. E. Braunwalder; 1 ♂ (MEB 824), Coste de Galy near Marsal, Roussillon, Pyrenees, France, 26.v.1995, M. E. Braunwalder; 1 ♂ (MEB 961), Rivière de Caillan, between Conat and Ria-Sirach Urbanya, Pyrenees, France, 19.v.1997, M. E. Braunwalder; 1 juv. (MEB 964), Roubrio, Riuferrer, Vallée de Bonabosc, Pyrenees, France, 21.v.1997, M. E. Braunwalder; 1 ♀ and 1 juv. (AMNH), Pyrenees, Uaut Vellespir, D54 nach Montferrer, kleine Geröllhalde, France, 1305 ft, 42°27.120′N 2°36.191′E, 5.vi.2003, S. Huber.

Chaerilidae Pocock, 1893

Chaerilus chapmani Vachon & Lourenço, 1985: 1 ♂ (AMNH), Clearwater III Cave, S. Mulu NP, Sarawak, Gunong Api, Kalimantan (Borneo) Island, Malaysia, v.1984.

Diplocentridae Karsch, 1880

Diplocentrus anophthalmus Francke, 1977: Holotype ♀, paratype ♀ (AMNH), Actun Chukum, Yucatán, México, 29.xi.1974, J. Reddell, D. McKenzie, R. Mitchell & S. Wiley.

Diplocentrus cueva Francke, 1978: Holotype ♂ (AMNH), Cueva Desapareciendo, 2 km W Acatlan, Oaxaca, México, 3.i.1976, A. Grubbs.

Diplocentrus mitchelli Francke, 1977: Holotype juv. ♂ (AMNH), Actun Halmensura, Campeche, México, 31.x.1974, J. Reddell, S. Wiley & D. McKenzie.

Nebo hierichonticus (Simon, 1872): 1 ♀ (ZMB 31133), Tabgha, Palestine, 28.ii.1919, P. E. Schmitz; 1 ♀ (ZMB 8229), Palestine, Schluter; 2 subad. ♂, 1 juv. ♂, 1 juv. ♀ (ZMB 31134), Red Sea, Wadi Ghuwer, Palestine, 1912, Dr Brühl; 1 ♀ (NHRM [JF 54]), Nazaret, Israel, iii.1995, J. Ferrer; 5 ♂ 2 ♀ (FMNH), Afule, Palestine, 25.ix.1934, Dr F. R. S. Shaw; 1 juv. ♂ (CAS), Mt Carmel, Israel, 12.xi.1960, Dr M. R. Warburg; 1 subad. ♂ (CAS), Mt Carmel, Israel, 20.vii.1960, Dr M. R. Warburg; 1 ♂ (AMNH [AH 573]), En Gedi, Israel, 2.ii.1978, P. Amitai; 1 ♂ (AMNH [AH 573]), Arad, 7 km E, Judean Desert, Israel, 15.ix.1984, G. Swager.

Euscorpiidae Laurie, 1896

Troglocormus ciego Francke, 1981: Holotype ♂ (AMNH), Cueva de Elias, 13 km N Agua Buena, San Luis Potosí, México, 3.viii.1975, D. Mckenzie.

Troglocormus willis Francke, 1981: México: Tamaulipas: Holotype ♂ (AMNH), Cueva del Brinco (entrance passage), 3.v.1978, P. Sprouse. Paratypes: 1 ♀ (AMNH), Cueva de Esperanza, Yerbabuena, Tamaulipas, México, 20.xi.1979, P. Sprouse; 1 juv. (AMNH), Cueva del Brinco, vi.1978, P. Sprouse; 1 juv. (AMNH), Cueva de la Onza, 300 m N Conrado Castillo, 21.iii.1982, G. Atkins. Additional material: 1 subad. (AMNH LP 2461), Cueva Allarines, Conrado Castillo, 25.xi.2003, A. Gluesenkamp; 1 subad. (AMNH LP 2460), Cueva Cuchilla, Conrado Castillo, 28.xi.2003, A. Gluesenkamp; 1 subad. (AMNH LP 2743), P-2 Cave, Conrado Castillo, 3.iv.2004, B. Shade.

Hemiscorpiidae Pocock, 1893

Hemiscorpius lepturus Peters, 1861: 1 ♂ syntype (ZMH 2321 ex ZMB), 1 ♀ syntype (ZMH 2322 ex ZMB), Baghdad, Iraq, 1893, Peters; 2 ♂, 2 ♀ syntypes (ZMB 43), Mendeli, near Baghdad, Iraq, Petermann. Additional material: 2 ♀ (ZMB 2322), Baghdad, Iraq; 2 ♂ (ZMB 7550); 1 ♂, 2 ♀ (USNM), Gach Saran, 35 km E, Khuzistan Prov., Iran, 6.ii.1964, J. Neal; 1 ♂, 2 ♀ (FMNH), Mandali, Iraq, Y. Lazar, 16.xii.1934; 1 ♀ (AMNH LP 4342), Baghmalek, Khouzestan Prov., Iran, v.2005, H. R. Goodarzi; 1 ♂, 1 ♀ (AMNH LP 4343), Andimeshk, Khouzestan Prov., Iran, v.2005, H. R. Goodarzi.

Heteroscorpionidae Kraepelin, 1905

Heteroscorpion opisthacanthoides (Kraepelin, 1896): Madagascar: Holotype ♀, 1 ♂, 1 juv. ♂ syntypes (ZMH), Nossi Comba, Nossibe, 27.ix.1895, P. Frey; 1 ♀, 4 juv. ♂ syntypes (ZMH), Nossi Comba, 3.x.1896, P. Frey. Additional material: 1 subad. ♂ (ZMB 7809 ex ZMH), Nossi Comba, iii.1896, P. Frey; 2 ♀, 1 subad. ♂, 1 subad. ♀, 1 juv. ♂, 3 (MNHN RS 4813), Nossi Komba, 1945, Millot; 1 ♂, 1 ♀ (MNHN RS 4813), Nossi Bé, Komba, 1945, Millot; 1 juv. ♀ (MNHN RS 3536), Nosy Komba, flanc nord, Prov. Diègo Suarez, A. Robinson; 2 ♂, 3 ♀, 2 subad. ♂, 8 juv. ♂ (ZMH A26/03), Lokobé Reserve, Nosy Be I., 6–9.ii.1983, W. R. Lourenço; 1 ♀ (MNHN RS 5046), Sakavalana, 1901, Alluaud; 1 ♀ (MNHN RS 0557), Ambilobe, Waterlot; 1 juv. ♂ (MNHN RS 4473), Forêt de Mahory, Prov. Diègo Suarez, 16.i.1966, Betsch & J. J. Petter.

Liochelidae Fet & Bechly, 2001

Opisthacanthus validus Thorell, 1876: South Africa: Eastern Cape Province: 1 ♀, 1 juv. ♀ (AMNH LP 2763 [JV 7482, 7483]), Rietvlei, Witmos, Bedford District, 32°32′S 25°44′E, 13.vi.2003, J. Visser; 1 subad. ♀ (AMNH LP 2766 [JV 7726]), Farm Rea’s, c. 7 km E Ramaphosa Village, Hankey District, 19.i.2003, J. Visser; 1 juv. (AMNH LP 2761 [JV 7701]), Swart River, Humansdorp District, 34°04′S 24°54′E, 24.v.2003, J. Visser; 1 subad. ♂, 2 juv. (AMNH LP 2762 [JV 7668, 7669, 7671]), Elands River, at old bridge, Humansdorp District, 33°59′S 24°05′E, 20.i.2003, J. Visser; 1 ♂ (AMNH LP 2764 [JV 7827]), Farm Uitvlugt 194, Humansdorp District, 33°53′S 24°36′E, 7.ix.2003, J. Visser; 3 ♀ (AMNH LP 2765 [JV 7834–7836]), Rietvlei, Humansdorp District, 33°55′S 24°35′E, 7.ix.2003, J. Visser; 1 ♀ (AMNH LP 2767 [JV 7876]), Jeffreys Bay, Humansdorp District, 34°03′S 24°55′E, 13.ix.2003, J. Visser.

Scorpionidae Latreille, 1802

Scorpio maurus mogadorensis Birula, 1910: 1 ♀ (AMNH LP 3497), 8 km after Tagmoute, on road to Tanalt, NW of Tafraoute, Morocco, 29°44′05.5″N 09°09′55.9″W, 14.ix.2004, V. Vignoli; 1 ♂, 2 juv. (AMNH), Tafraoute, 500 m from village (camping), Morocco, 1008 m, 29°43′19.2″N 08°58′46.6″W, 10.ix.2004, V. Vignoli; 1 ♀, 1 juv. (AMNH), Tafraoute, 500 m from village (camping), Morocco, 1008 m, 29°43′19.2″N 08°58′46.6″W, 14.ix.2004, V. Vignoli; 4 juv. (AMNH), Asni, 4 km before, on road Marrakech-Asni, Morocco, 1082 m, 31°16′55.7″N 07°57′35″W, 22.ix.2004, V. Vignoli; 1 ♂, 2 ♀, 2 subad. ♂ (AMNH), Arbalou (Ourika), Marrakech Prov., Morocco, 1000 m, 14–18.v.1975, B. Malkin.

Superstitioniidae Stahnke, 1940

Alacran tartarus Francke, 1982: México: Oaxaca: Holotype ♀ (AMNH), Sótano de San Agustín, 720 m below surface, San Agustín, 5 km SE Huautla de Jiménez, San Agustin Expedition (Huautla Project), Spring 1979. Paratypes: 1 ♀ (MNHN RS 7439), Cueva del Escorpíon, San Miguel Dolina, San Miguel, i.1978, R. Jameson & P. Mothes; 1 ♀ (AMNH), Sótano Li Nita, Huautla de Jiménez, 1.iv.1981, M. Minton, L. Wilk & R. Simmons; 1 juv., Sótano Li Nita, Huautla de Jiménez, 22.iii.1981, M. Minton. Additional material: 2 ♀ (WDS), Sótano de San Agustín Section, Sistema Huautla, 5 km SE Huautla de Jiménez, v.1985, A. G. Grubbs, J. H. Smith & F. Holliday; 1 ♀ (AMNH LP 3499), Cueva del Escorpíon, 18.1065°N 96.7982°W, Municipio San Miguel, 18.ix.2004, A. Gluesenkamp, P. Sprouse & C. Savvas, 1561 m.

Sotanochactas elliotti (Mitchell, 1971): Holotype subad. ♂ (AMNH), Sótano de Yerbaniz, c. 12 km N Ciudad Valles, San Luis Potosí, México, 31.vii.1969, W. R. Elliott; 1 paratype ♂ (WDS), same locality data as holotype, R. W. Mitchell, 4.vii.1970; 1 juv. paratype (MNHN RS 5376), same locality data as holotype, 28.iii.1970, W. Elliott.

Typhlochactas cavicola Francke, 1986: Holotype ♂ (AMNH), Cueva del Vandalismo, 1 km SE Rancho Nuevo, Tamaulipas, México, 15.iii.1982, D. Honae.

Typhlochactas granulosus Sissom & Cokendolpher, 1998: Holotype ♂ (AMNH), Sótano de Poncho, Municipio Tlaquilpa, Veracruz, México, 22.iii.1995, P. Sprouse.

Typhlochactas mitchelli Sissomi, 1988: Holotype ♂, 1 ♂ 1 subad. ♀ paratypes (AMNH), Cerro Ocote, 5 mi S Tenango, Oaxaca, México, iv.1987, A. Grubbs, A. Cressler & P. Smith.

Typhlochactas reddelli Mitchell, 1986: Holotype ♀ (AMNH), Cueva del Ojo de Agua de Tlilapan, Tlilapan, Veracruz, México, 4.viii.1967, J. Reddell, J. Fish & T. R. Evans.

Typhlochactas rhodesi Mitchell, 1968: Holotype ♀ (AMNH), Cueva de la Mina, Rancho del Cielo, Tamaulipas, México, 29.iii.1967, R. Mitchell, R. Mitchell, V. Colvin, D. Falls & K. Pittard. Paratypes: 1 juv. ♀ (MNHN RS 4760), Cueva de la Mina, Rancho El Cielo, Tamaulipas, México, 29.viii.1967, R. Mitchell, F. Abernathy & W. Rhodes; 1 juv. ♀ (AMNH), Cueva de la Mina, 6 mi NW Gómez Farias, Tamaulipas, México, 1.vii.1969, S. J. Peck & R. Norton.

Typhlochactas sylvestris Mitchell & Peck, 1977: Holotype ♀ (AMNH), 15 mi S Valle Nacional, Oaxaca, México, 21.v.1971, S. Peck.

Urodacidae Pocock, 1893

Urodacus macrurus Pocock, 1899: 1 ♀ (ZMH), Australia, 26.iii.1897; 1 ♂ (QM S58508), Carnarvon NP, Mt Moffat Section, Queensland, Australia, 24°58′43″S 147°55′17″E, 31.i.–9.ii.2000, C. Eddie; 1 ♂ (ESV 4424), Carnarvon NP, Mt Moffat Section, Queensland, Australia, 25°01′15″S 147°55′57″E, 13–19.iv.1999, C. Eddie; 1 ♂ (ESV 6020), Carnarvon NP, Mt Moffat Section, Queensland, Australia, 25°04′18″S 147°52′43″E, 20.i. 1999, C. Eddie, R. Johnston, & A. Young; 1 ♂ (AMNH LP 2341), Carnarvon NP, Mt Moffatt section, SE side of road between mailboxes and homestead, Queensland, Australia, 25°02′37″S 147°54′39″E, 14.x.2002, C. Eddie; 1 ♂ (AMNH LP 2879), Carnarvon NP, Mt Moffatt section, SE side of road between mailboxes and homestead, Queensland, Australia, 25°02′37″S 147°54′39″E, 13.x.2002, C. Eddie; 1 ♀ (ESV 6021), Carnarvon NP, Mt Moffat Section, Queensland, Australia, 25°06′51″S 147°51′36″E, 30.iii–8.iv.1998, C. Eddie & A. Dudley.

Urodacus manicatus (Thorell, 1876): 1 ♂, 1 ♀ (ESV 6022), Grampians Ranges, Victoria, Australia, 37°01′02″S 142°25′28″E, 5.xii.1998, E. S. Volschenk & J. Tregear; 1 ♂ (AMNH LP 938, 939), 1 ♂, 1 juv. (AMNH LP 941), 1 ♀ (AMNH LP 940), Canberra, Australian Capital Territory, Australia, xii.1997, H. Wilson; 1 ♀ (AMNH LP 1537), 1 ♀ (AMNH LP 1657), Overland corner, South Australia, Australia, 25.ix.1997, E. S. Volschenk; 1 ♂ (AMNH LP 1538), Rocky River, Kangaroo I., Australia, 1986, N. A. Locket; 3 ♂, 5 ♀, 2 subad. ♀ (CAS), Black Mtn, ACT, Australia, 760 m, 9.xii.1962, E. S. Ross & D. Q. Cavagnaro.

Urodacus mckenziei Volschenk, Smith & Harvey, 2000: Paratype ♀ (WAM 98/1592), Peron Homestead, Shark Bay, Western Australia, Australia, iv.1998, T. Tischler; 2 ♂ (WAM 99/1128, 99/1147), Francois Peron NP, Western Australia, Australia, 25°52′31″S 113°32′59″E, 17.i–25.v.1995, M. S. Harvey, et al. ; 1 ♀ (AMNH LP 2689), Peron Homestead, 100 m W of Visitor’s Centre, Western Australia, Australia, 25°50′S 113°33′E, iv.1996, T. Tischler.

Urodacus megamastigus L. E. Koch, 1977: 1 ♂ (WAM), Roy Hill, Western Australia, Australia, 18.v.1982, A. M. Douglas; 1 ♂ (WAM), Millstream, Western Australia, Australia, 8.v. 1986, D. King, et al.; 1 ♂ (WAM 99/1212), Mt Brockman Station, Western Australia, Australia, 22°18′27″S 117°15′27″E, 11.v. 1999, P. J. West; 1 ♂ (WAM T40692), Mt Brockman (Namuldi mine, Hamersley Iron) Western Australia, Australia, 22°18′38″S 117°47′47″E, Anstee.

Urodacus novaehollandiae Peters, 1861: Australia: 2 ♂, 1 ♀, 1 subad. ♂, 1 juv. ♂ (ZMH), Lion Mill, near Perth, 22.v.1905, Michaelsen; 1 ♂ (ZMH), Karrakatta, near Perth, 28.v.1905, Michaelsen; 1 ♂, 1 subad. ♂, 1 juv. ♂ (ZMH), Wooroloo, near Perth, 29.v.1905, Michaelsen; 1 ♂, 1 subad. ♀, 4 juv. ♂ (ZMH), York, 11.viii.1905, Michaelsen; 1 ♂ (AMNH LP 1540), 1 ♂ (AMNH LP 1658), Jarrahdale, Western Australia, Australia, 25.ii.1998, E. S. Volschenk; 1 ♀ (AMNH LP 2121), Perth Hills (Darling Escarpment), Western Australia, Australia, i.2002.

Urodacus planimanus Pocock, 1893: 1 ♂, 1 ♀, 1 juv. ♀ (ZMH), Jarrahdale, near Perth, Australia, 1905, Michaelsen; 1 ♂ ESV6023, Jarrahdale, Western Australia, 20.iii. 1998, E. S. Volschenk; 1 ♂, 3 ♀ (WAM 69/1/4), Roleystone, Western Australia, Australia, 1968, S. Stevans; 1 ♂, 1 juv. (WAM 97/2662, 97/2663), Mt Cooke (nr Summit), Western Australia, Australia, 32.255°S 116.18°E, 7.viii. 1990, M. Harvey, J. Waldock & M. Peterson; 1 ♀ (WAM 68/298), Mt Cooke, Western Australia, Australia, 26.vii.1967, M. Parking & M. Porter; 1 ♂ (AMNH LP 1539), 1 ♂ (AMNH LP 1661), Jarrahdale, Western Australia, Australia, 25.ii.1998, E. S. Volschenk; 1 ♂, 1 subad. ♀ (AMNH LP 2123), Jarrahdale, Western Australia, Australia, 32°17′35″S 116°10′39″E, 2.vi.2001, E. S. Volschenk; 1 ♂ (WAM 68–297), Mt Cooke, Western Australia, Australia, 26.vii.1967, M. Parkin & M. Porter.

Urodacus varians Glauert, 1963: 3 ♂ (WAM T51161, T41169, T41163), Meridian (Wildcat oilwell), 30 mi. E of Willara Hill, Western Australia, Australia, 19°03′S 122°22′E, 25.v.1982, A.M. Douglas.

Urodacus yaschenkoi (Birula, 1903): 1 ♂ (AMNH LP 1541), 1 subad. ♀ (AMNH LP 1659), Berri, South Australia, Australia, 24.ix.1997, N. A. Locket & E. S. Volschenk; 1 ♀ (AMNH LP 2120), Berri, South Australia, Australia, 34°12′40.5″S 140°12′42.4″E, 10.xii.1998, E. S. Volschenk.

Vaejovidae Thorell, 1876

Uroctonus grahami Gertsch & Soleglad 1972: Holotype ♀ (AMNH), Samwell cave (main entrance), Shasta County, California, USA, 27.vi.1960, R. E. Graham. Additional material: 1 juv. (AMNH), same locality data as holotype, 5.vi.1957, R. de Saussure.

Vaejovis gracilis Gertsch & Soleglad, 1972: México: Veracruz: Holotype juv. (AMNH), Cueva de Atoyac, Atoyac, 6.viii.1969, S. & J. P. Peck. Additional material: 1 ♂ (AMNH), Sótano de las Golondrinas, near Potrero, 8.i.1977, A. Grubbs; 2 ad. (UNAM), Paraje Nuevo, Cueva del Ojo de Agua de Tlilapan, Tlilapan.

Appendix 2. Characters and character states employed in cladistic analysis of nine species of Urodacidae Pocock, 1893 and six outgroup taxa

Character states are scored 0–3, * (polymorphic); ? (unknown); and – (inapplicable). Eight uninformative characters, indicated by †, were excluded from all analyses. Unknown states in Aops oncodactylus, gen. et sp. nov. are attributed to the fact that the adults are presently unknown. Previous character numbers as follows (author/number): P00 = Prendini (2000), P03 = Prendini (2003), PCW03 = Prendini et al. (2003).

(1) Cheliceral movable finger, distal external and distal internal teeth: unequal, with distal external tooth considerably smaller than distal internal tooth, aligned longitudinally and not opposable (0); unequal, with distal external tooth noticeably smaller than distal internal tooth, and opposable, i.e. forming a bicusp (1); subequal, with distal external tooth only slightly smaller than distal internal tooth, and opposable, i.e. forming a bicusp (2). Previous characters: P00/11, P03/10, PCW03/2.

(†2) Cheliceral movable finger, number of subdistal teeth: two (0); one (1). Previous characters: P00/10, P03/9.

(3) Cheliceral fingers, secondary serrations: absent (0); present (1). Previous characters: P00/12.

(4) Carapace, median notch in anterior margin: absent (0); shallow (1); strongly excavated (2). Previous characters: P00/3, P03/3, PCW03/5.

(5) Carapace, lateral ocelli, number of pairs: three (0); two (1); none (2). Previous characters: P00/1, P03/1.

(6) Carapace, median longitudinal furrow: broad, shallow, without suture (0); narrow, suturiform (1). Previous characters: P00/4, P03/4.

(7) Carapace, median ocular tubercle: raised (0); shallow (1); none (2). Previous characters: P00/2, P03/2.

(8) Carapace, posterior sutures: absent (0); present (1). Previous characters: P00/7, P03/6.

(9) Prosoma, mesosoma, metasoma and legs, nongranular surfaces: smooth (0); distinctly punctate (1). Previous characters: P00/8, P03/7.

(10) Pedipalp chela fingers, terminal teeth interlocking when closed: unevenly, with movable finger displaced to exterior, with notch near tip of fixed finger (0); evenly, movable finger not displaced to exterior (1). Previous characters: PCW03/23 [merged states 1 and 2].

(11) Pedipalp chela fingers, dentate margins: entire (0); markedly scalloped (1). Previous characters: P00/34, P03/28.

(12) Pedipalp chela fingers, number of rows of primary denticles: single (0); double along entire length, but often fused basally (1); multiple (more than two), at least basally, sometimes becoming single or double distally (2). Previous characters: P00/33, P03/27.

(13) Pedipalp chela movable finger, lobe (♂): absent or at most weakly developed, close to base of movable finger and lacking an obvious notch in fixed finger (0); well developed, almost midway along movable finger (if dentate margin markedly scalloped, second lobe more strongly developed than other lobes on the movable finger), with a distinct notch in fixed finger, lobe rounded dorsally and lacking a sharp conical tooth (1); unknown (?). Previous characters: P00/35, P03/29, PCW03/25 (part) [corrected state in Nebo].

(14) Pedipalp chela manus, dorsal secondary carina: obsolete (0); distinct (1). Previous characters: P00/22, P03/18.

(†15) Pedipalp chela manus, ventroexternal carina: oblique to longitudinal axis of chela, distal edge disconnected from external movable finger condyle and directed towards (almost connecting) internal movable finger condyle (0); parallel to longitudinal axis of chela, distal edge disconnected from external movable finger condyle and directed towards a point between external and internal movable finger condyles, but closer to external condyle (1). Previous characters: P00/27, P03/21.

(16) Pedipalp chela manus, ventrointernal carina: equally developed as internomedian carina (0); more strongly developed than internomedian carina, which may be obsolete (1). Previous characters: P00/29, P03/23, PCW03/36.

(17) Pedipalp patella, anterior process: absent (0); present (1). Previous characters: P00/18, P03/14.

(18) Pedipalp patella, dorso-external carina: distinct (0); obsolete (1). Previous characters: P00/16, P03/12 [corrected state in U. yaschenkoi].

(19) Pedipalp chela, position trichobothrium it: manus or base of fixed finger (0); midway (Hemiscorpius) to distal (Nebo) along fixed finger (1). Previous characters: P00/47, P03/38.

(†20) Pedipalp chela, position trichobothrium dsb: below db–dst axis (0); in line with db–dst axis (1). Previous characters: P00/60, P03/50.

(21) Pedipalp chela, position trichobothrium db: dorsal surface (0); internal surface (1). Previous characters: P00/59, P03/49.

(22) Pedipalp chela, position trichobothrium esb: manus, behind point of articulation between fixed and movable fingers and below eb–est–et axis (0); midway along fixed finger, in line with eb–est–et axis (1). Previous characters: P00/62, P03/52.

(23) Pedipalp chela, external surface with accessory trichobothria: absent (0); present (1). Previous characters: P00/53, P03/44.

(24) Pedipalp chela, position trichobothrium Dt: manus, at basal end (0); manus, mid-length or slightly less than mid-length (1); manus, distal half, near base of fixed finger (2); proximal end of fixed finger (3). Previous characters: P00/52, P03/43.

(25) Pedipalp chela, position trichobothrium Db: external surface (0); dorsal surface (1). Previous characters: P00/51, P03/42.

(26) Pedipalp chela, number V trichobothria: 4 (0); 6 or more (1). Previous characters: P00/49, P03/40, PCW03/51.

(†27) Pedipalp chela, position trichobothria V₁–V₃: evenly spaced, distance V₁–V₂ similar to distance V₂–V₃ (0); unevenly spaced, distance V₁–V₂ much less than distrance V₂–V₃ (1); inapplicable (–). Previous characters: P00/50, P03/41.

(28) Pedipalp chela, position trichobothria V₂–V₄: evenly spaced, distance V₃–V₄ similar to distance V₂–V₃ (0); unevenly spaced, distance V₃–V₄ much less than distrance V₂–V₃ (1); inapplicable (–). Previous characters: P00/50, P03/41, PCW03/52.

(29) Pedipalp patella, number e trichobothria: 13 (0); 15 or more (1). Previous characters: P00/45, P03/36, PCW03/54 (part).

(30) Pedipalp patella, number v trichobothria: single row of 3 (0); single row of 4–10 (1). Previous characters: P00/43, P03/34, PCW03/55.

(31) Legs I and II, basitarsi, retrolateral margins, setation: no macrosetae (0); row of three or more spiniform macrosetae (1); comb-like row of three or more setiform macrosetae that may or may not be interspersed with slender spiniform macrosetae (2). Previous characters: PCW03/67 (merged P00/72, P00/73 and P03/59).

(32) Legs, telotarsi, ventrosubmedian setae type: stout, spiniform (0); slender, subspiniform (1). Previous characters: P00/70, P03/58.

(†33) Legs IV, telotarsi, prolateral row of ventrosubmedian spiniform setae: present (0); absent (1). Previous characters: P00/71, PCW03/71 (part).

(34) Legs, telotarsi, well developed ventromedian row of setae: setiform (0); spiniform (1); absent (2). Previous characters: P00/68, P03/56.

(35) Legs, telotarsi, laterodistal lobes: truncated, base of median dorsal lobe flush (0); rounded, notches at base of median dorsal lobe (1). Previous characters: P00/65, P03/54.

(†36) Embryonic development: apoikogenic (0); katoikogenic (1); unknown (?). Previous characters: P00/77, P03/62.

(37) Hemispermatophore, sclerotised mating plug: absent (0); present (1); unknown (?). Previous characters: P00/92, P03/75.

(38) Metasomal segments (♂): similar to or slightly longer than ♀ (0); much longer than ♀ (1); unknown (?).

(39) Metasomal segments I–IV, dorsolateral carinae: present and distinct on at least some segments (0); absent or obsolete on all segments (1).

(40) Metasomal segment V, dorsolateral carinae: present and distinct (0); absent or obsolete (1). Previous characters: PCW03/77.

(41) Metasomal segment I, median lateral carinae (♂): present and distinct (0); absent or obsolete (1); unknown (?).

(42) Metasomal segment I, median lateral carinae (♀): present and distinct (0); absent or obsolete (1).

(43) Metasomal segment V, median lateral carinae: present and distinct (0); absent or obsolete (1).

(†44) Metasomal segment V, ventrolateral carinae: present and distinct (0); absent or obsolete (1). [corrected entries in L. josehermana].

(45) Metasomal segments I–IV, ventral carinae: absent on all segments (0); present and distinct on at least some segments (1). Previous characters: P00/95, PCW03/80 (part).

(46) Metasomal segments I–IV, ventral carinae: paired ventrosubmedian carinae (0); single ventromedian carina (1); inapplicable (–). Previous characters: P00/95, P03/77.

(†47) Metasomal segment V, ventromedian carina: present and distinct (0); absent or obsolete (1). [corrected entries in L. josehermana].

(48) Metasomal segment V, ventromedian carina distal portion: straight (0); breaking up into numerous granules (1); bifurcating (2); inapplicable (–). Previous characters: P00/99, P03/81, PCW03/86 [corrected entries in L. josehermana].

(49) Telson vesicle, shape (♂): unmodified (0); elongated, with pair of distal lobes (1); elongated, without pair of distal lobes (2); unknown (?). Previous characters: P00/109, P03/89.

(50) Telson vesicle, shape (♂): unmodified (0); laterally compressed (1); unknown (?). Previous characters: P00/107, P03/87.

(51) Telson vesicle, shape (♀): unmodified (0); laterally compressed (1). Previous characters: P00/108, P03/88.

(52) Telson vesicle, anterodorsal lateral lobes: present (0); absent (1). Previous characters: P00/110, P03/90.

(53) Telson vesicle, surface macrosculpture: entirely smooth (0); longitudinal rows of granules on ventral and lateral surfaces (1); 2 or more longitudinal rows of granules on all or part of ventral surface only (2). Previous characters: P00/111 (modified), P03/91, PCW03/89.

(54) Telson, venom glands: complex (0); simple (1). Previous characters: P00/113, P03/93.

(55) Telson, aculeus: long, shallowly curved (0); very short, sharply curved (1). Previous characters: P00/105, P03/85.

(56) Burrows constructed: absent (0); present (1); unknown (?). Previous characters: PCW03/104, PCW03/105 (modified).