Zootaxa 1849: 35–58 (2008) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ ZOOTAXA Copyright © 2008 · Magnolia Press ISSN 1175-5334 (online edition) Review and cladistic analysis of the Neotropical tarantula genus Ephebopus Simon 1892 (Araneae: Theraphosidae) with notes on the Aviculariinae RICK C. WEST1, SAMUEL D. MARSHALL2, CAROLINE SAYURI FUKUSHIMA3,4 & ROGÉRIO BERTANI4 1 3436 Blue Sky Place, Victoria, B.C., CA. E-mail: [email protected] 2 Department of Biological Sciences, 317 Bienvenu Hall, Northwestern State University Natchitoches, LA. E-mail: [email protected] 3 Instituto de Biociências, Departamento de Zoologia, Universidade de São Paulo. E-mail: [email protected] 4 Instituto Butantan, Av. Vital Brazil 1500, 05503–900, São Paulo SP. E-mail: [email protected] Abstract The tarantula genus Ephebopus Simon 1892 is reviewed and includes the type species, E. murinus (Walckenaer 1837), and E. uatuman Lucas, Silva & Bertani 1992, E. cyanognathus West & Marshall 2000, E. rufescens West & Marshall 2000 and Ephebopus foliatus, sp. nov., from Guyana. Ephebopus violaceus Mello-Leitão 1930 is transferred to Tap- inauchenius Ausserer, where it is a senior synonym of Tapinauchenius purpureus Schmidt 1995 new synonymy. Ephebo- pus fossor Pocock 1903 is considered a nomen dubium. Ephebopus occurs in northeastern South America where it is known only from Brazil, Guyana, Suriname, and French Guiana. Spiders of the genus are generally fossorial; however, Ephebopus murinus has a developmental stage that is arboreal. A cladistic analysis of the Theraphosidae retrieves the Aviculariinae as monophyletic, including Avicularia Lamarck, Iridopelma Pocock 1901, Pachistopelma Pocock 1901, Tapinauchenius, Psalmopoeus Pocock, Ephebopus, Stromatopelma Karsch and Heteroscodra Pocock, having as a syna- pomorphy the well-developed scopulae on tarsi and metatarsi I–II that is very laterally extended. Key words: Aviculariinae, systematics, Ephebopus, Guyana, cladistic analysis, theraphosid behavior Introduction The Theraphosidae are found on all continents, except Antarctica. More than 900 species are described, con- sisting roughly of a third of all described mygalomorph species (Platnick 2008). Most theraphosids are found in tropical areas but some representatives live in subtropical and temperate regions. They are mainly terrestrial in habit, living in burrows and other natural cavities or under rocks and fallen logs. Arboreal forms are found mainly in the tropical New World but also in Africa and Asia. The taxonomy of this family is very confused and taxonomic revisions are rare (Raven 1990). Cladistic analyses of relationship of species and genera are still rare and there was only one attempt to recover the cladistic relationships of all theraphosid subfamilies (Raven 1985). Therefore, the taxonomic position of several genera in theraphosid subfamilies is controversial, as is the composition of the Aviculariinae. Simon (1892) created the Avicularieae, including only Avicularia and Tapinauchenius, then characterized by the absence of a line of setae dividing the tarsal scopulae (i.e., entire or undivided scopula) of leg IV, legs without spines, and posterior legs longer than anterior ones. Later, Pocock (1901) included Pachistopelma and Iridopelma as well as Ephebopus and Psalmopoeus in the Avicularieae. He also suggested the inclusion of the West African genera Scodra Becker 1879 (= Stromatopelma) and Heteroscodra. Simon (1903) included the Accepted by R. Raven: 10 Jul. 2008; published: 13 Aug. 2008 35 two African genera in Avicularieae, but transferred Ephebopus to the Phoneyuseae and synonymized Iri- dopelma with Avicularia. Mello-Leitão (1923) considered only American genera belonged in the Avicularii- nae, including his monotypic genus Ancylochiros Mello-Leitão 1920. He also considered Typhochlaena C. L. Koch to be valid and Iridopelma its junior synonym. Roewer (1942) transferred Heteroscodra and Scodra back to the Aviculariinae. He also included the Middle Eastern genus, Avicuscodra Strand (= Chaetopelma Ausserer, Ischnocolinae), in the Aviculariinae but considered Typhochlaena a junior synonym of Avicularia and placed Ancylochiros (= Avicularia) in the Ischnocolinae and Ephebopus in the Eumenophoriinae. Raven (1985) transferred Ephebopus to the Theraphosinae, Stromatopelma and Heteroscodra to the Eumenophorinae, Psalmopoeus to the Selenocosmiinae, revalidated Iridopelma and considered Ancylochiros a junior synonym of Avicularia. Raven (1985) also included only the genera Avicularia, Iridopelma, Pachis- topelma, and Tapinauchenius in the Aviculariinae based on the following shared traits: spinose process between the lobes of the male palpal tarsi, tarsi as broad as or broader than the metatarsi, and the legs being weakly spined or lacking spines. Raven (1985: 119) noted, based on a curator’s drawing of the type, that the type species of Tapinauchenius, T. plumipes (C. L. Koch 1842), appeared to lack a well-developed spinose process on the male palpal tarsi. Lucas et al. (1991) described the male of E. murinus and based on the posses- sion of a sinuous embolus lacking keels (in addition to characters used by Raven 1985), and transferred Ephe- bopus to the Aviculariinae. Based on their observations, they noted that males of both Ephebopus and Tapinauchenius lacked the spinose process on the palpal tarsi, but considered the wide tarsal pads, elongated embolus, and absence of spines on the legs as synapomorphies uniting the Aviculariinae As for Ephebopus itself, no other theraphosid genus has had its taxonomic position so uncertain. First included in the Selenocosmieae by Simon (1892), the genus was later transferred variously to the Avicularieae (Pocock 1901), Phoneyuseae (Simon 1903), back to Aviculariinae (Mello-Leitão 1923), Eumenophorinae (Roewer 1942), Theraphosinae (Raven 1985), and again to Aviculariinae (Lucas et al. 1991). Thus, it was placed at least once in four of the eight theraphosid subfamilies recognized by Raven (1985). The type species of Ephebopus, Mygale murina Walckenaer 1837, was based on a brief but illustration- free description but was fairly accurate. After some decades, Simon (1892) transferred M. murina to Ephebo- pus. Simon (1903) examined the holotype of Santaremia pococki F. O. P.-Cambridge 1896 and concluded it was a junior synonym of Ephebopus murinus. Pocock (1903) described a second species, Ephebopus fossor Pocock 1903 from Ecuador, and Mello-Leitão (1930) added the third, E. violaceus Mello-Leitão 1930 from Cuminá River region, Brazil. Only females of Ephebopus species were known until Lucas et al. (1991) described the male of E. muri- nus, and males and females of a new species E. uatuman from Presidente Figueredo, Amazonas, Brazil. These authors also transferred E. violaceus to Avicularia. Two more species of Ephebopus were described recently, Ephebopus cyanognathus and Ephebopus rufescens, both from French Guiana (West & Marshall 2000). We describe a new species of Ephebopus from Guyana, and present a cladistic analysis of Ephebopus spe- cies, including representatives of seven theraphosid subfamilies and all aviculariine genera, in order to test the cladistic relationships of Ephebopus and the Aviculariinae. Material and methods All measurements are given in mm and were made to the nearest 0.01 mm with the aid of a Wild-Heerbrugg dissecting microscope using an ocular micrometer or with dial vernier calipers. Leg and pedipalp measure- ments were taken from the dorsal aspect on the left side (unless appendages were lost or obviously regener- ated) of all specimens whereas coxae and trochanters are measured from their ventral aspect, on the same side. Leg span is taken on left side in a straight line from the tip of leg I to that of leg IV. The extent of metatarsal scopulae is presented as a fraction (maximum 1.0) measured from the base ventrally. Claws are not included 36 · Zootaxa 1849 © 2008 Magnolia Press WEST ET AL. in measurements of tarsi. Images were made with a Nikon Coolpix 5000 digital camera and a WILD stereomi- croscope. Scanning electron micrographs were obtained with a LEO 440I from the Instituto de Geociências da Universidade de São Paulo. Standard abbreviations and measurements are used for ocular and spination descriptions and follow those of Prentice (1992) and Reichling and West (1996). Male palpal bulb keel termi- nology follows Bertani (2000); urticating hair type terminology follows Cooke et al. (1972) and Marshall & Uetz (1990). Material of the following institutions were examined: AMNH – American Museum of Natural History, New York; BMNH – The Natural History Museum, London; IBSP – Instituto Butantan, São Paulo; MIZA – Instituto de Zoologia, Universidad Central de Venezuela, Maracay; MNHN – Muséum National d'Histoire Naturelle, Paris; MZSP – Museu de Zoologia, Universidade de São Paulo, São Paulo; NMNH – National Museum of Natural History, Washington, DC; MNRJ – Museu Nacional, Rio de Janeiro; SMF – Senckenberg Museum, Frankfurt. Geographical coordinates: primary sources are between round brackets, secondary sources between square brackets. Cladistics: A data matrix with 21 taxa and 48 characters (Table 3) was used with three programs using equal (Hennig86 1.5, Farris 1988; Nona 2.0 for Windows ,Goloboff 1993a) successive (Hennig86 1.5) and implied character weighting (X-Pee-Wee 1.3 for Windows Goloboff 1997). For Hennig86, the exact algorithm "ie". For X-Pee-Wee 1.3 and Nona 2.0, the commands h100, h/20, amb- and mult*50 were used. For X-Pee- Wee, concavities