Blackwell Science, LtdOxford, UKZOJZoological Journal of the Linnean Society0024-4082The Lin- nean Society of London, 2005? 2005 1433 405416 Original Article PHYLOGENETIC TAXONOMY OF THE CERCOSAURINIT. M. DOAN and T. A. CASTOE Zoological Journal of the Linnean Society, 2005, 143, 405–416. With 1 figure Phylogenetic taxonomy of the Cercosaurini (Squamata: Gymnophthalmidae), with new genera for species of Neusticurus and Proctoporus TIFFANY M. DOAN1* and TODD A. CASTOE2 1Biology Department, Vassar College, Poughkeepsie, NY 12604–0555, USA 2Department of Biology, University of Central Florida, 4000 Central Florida Blvd, Orlando, FL 32816– 2368, USA Received December 2003; accepted for publication June 2004 The tribe Cercosaurini is one of the most poorly studied groups of the lizard family Gymnophthalmidae. Recent stud- ies have suggested that two cercosauriine genera, Neusticurus and Proctoporus, are polyphyletic. The aim of the cur- rent study was to rectify the polyphyletic relationships and construct a phylogenetic taxonomy of the Cercosaurini that is congruent with evolutionary history. Neusticurus is divided into two genera, one of them new (Potamites), based on the clades recovered by molecular studies and previously discussed morphological data. Proctoporus is divided into three genera, one of which is new (Petracola), while an older name (Riama) is resurrected for another. All five genera are described and defined and taxonomic keys are presented. This study represents an important advance in rectifying the taxonomy of the Cercosaurini. Many other para- and polyphyletic genera remain in the Gymnophthalmidae and much future work on this group is warranted. © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 143, 405–416. ADDITIONAL KEYWORDS: lizards – phylogenetic systematics – Reptilia – taxonomy. INTRODUCTION those genera. Several speciose genera, including Anadia, Euspondylus, and Proctoporus, were omitted The family Gymnophthalmidae has endured a turbu- completely from the study and taxon sampling was lent taxonomic history plagued by poorly defined gen- poor for other speciose genera such as Bachia, Pholi- era and the apparent presence of many homoplasious dobolus, and Ptychoglossus. Using morphological char- morphological characters. In addition, until recently acters, Doan (2003b) reconstructed a phylogeny of 11 no attempt has been made to classify the genera species formerly belonging to the genera Cercosaura, within a phylogenetic context. Presch (1980) and Pantodactylus, and Prionodactylus, which were all Hoyos (1998) constructed phylogenetic hypotheses transferred to Cercosaura in that publication. based on osteological and myological characters, but The most recent phylogenetic hypotheses advanced their conclusions provided little phylogenetic for the Gymnophthalmidae by Castoe, Doan & Parkin- resolution. Pellegrino et al. (2001) reconstructed the son (2004) included 12 additional species and one phylogenetic relationships of the family using gene additional genus (Proctoporus). Moreover, that study sequences, including 49 species in 24 genera. Their filled in some of the gene fragments that had been species sampling was excellent for taxa in the sub- missing from Pellegrino et al.’s (2001) study and fur- family Gymnophthalminae but relatively poor in the ther clarified some of the relationships. One major Cercosaurinae; although they placed 22 genera in the change involved re-allocating Ptychoglossus from the latter, they had not examined any members of nine of Cercosaurinae to a more basal position sister to Alopoglossus in subfamily Alopoglossinae. Another taxonomic alteration involved raising the tribe Ecple- *Corresponding author. Present address: Department of opini to subfamily status, the Ecpleopinae. A third Biological Sciences, Central Connecticut State University, 1615 Stanley Street, New Britain, CT 06050, USA. change raised the genus Bachia to tribal status, the E-mail: [email protected] Bachini. Other relationships that were suggested by © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 143, 405–416 405 406 T. M. DOAN and T. A. CASTOE Castoe et al. (2004) were that Neusticurus and Procto- P. pachyurus. Since that time, three additional genera porus were polyphyletic. The current study attempts have been described for species currently considered to rectify those polyphyletic relationships by examin- to be Proctoporus. Gray (1858) described Riama uni- ing the phylogeny of tribe Cercosaurini and altering color and O’Shaughnessy (1879) described Emphras- the generic taxonomy to recognize monophyletic lin- sotis simoterus as monotypic genera. The third genus, eages. Thus, we alter the classification to make it more Oreosaurus, was described by Peters (1862) as congruent with evolutionary history. Ecpleopus (Oreosaurus) striatus, a subgenus of Ecple- opus, which is a genus now considered to be distantly related (in a separate subfamily; Castoe et al., 2004). TAXONOMIC HISTORY OF NEUSTICURUS DUMÉRIL Boulenger (1885, 1902, 1908) considered six species & BIBRON to be members of Oreosaurus (O. laevis Boulenger, The genus Neusticurus consists of 11 species ranging O. luctuosus Peters, O. ocellifer Boulenger, O. oculatus from central Bolivia and south-western Brazil to O’Shaughnessy, O. petersii Boettger, and O. striatus). Colombia on the eastern versant of the Andes and in Boettger (1891) additionally described O. guentheri. the Guianan Shield region of Venezuela, Guyana, Andersson (1914) made Oreosaurus a junior synonym Suriname, French Guiana, and northern Brazil. One of Proctoporus because he could not find sufficient dif- additional species is from central Costa Rica (Uzzell, ferences between the two genera. This synonymy was 1966; Avila-Pires, 1995). The species occur at low ele- followed by all subsequent authors. vations from sea level to 1800 m and most are semi- Uzzell (1958, 1970) suggested preliminary group- aquatic, a unique trait for the family (Uzzell, 1966; ings within Proctoporus. The P. luctuosus group, Hoogmoed, 1973; Avila-Pires & Vitt, 1998; Vitt et al., unified by a divided palpebral eye disc, no median 1998). Neusticurus appears to be most closely related occipital, four supraoculars, and legs overlapping to Echinosaura (Burt & Burt, 1931; Uzzell, 1966) and when adpressed, included P. achlyens Uzzell, P. laevis, Teuchocercus (Fritts & Smith, 1969) based on external P. luctuosus, P. oculatus, and P. shrevei Parker (Uzzell, morphology. 1958). The P. pachyurus group, unified by the presence Duméril & Bibron (1839) created the genus Neusti- of a single palpebral scale over the eye, a median curus to contain Lacerta bicarinata Linnaeus. Uzzell occipital, and squarish pregular scales that do not (1966) divided the genus into two separate groups: form chevrons (Uzzell, 1970; Doan & Castoe, 2003), the N. bicarinatus group, containing N. bicarinatus, included P. bolivianus Werner, P. guentheri, and N. medemi Dixon & Lamar, N. racenisi Roze, N. rudis P. pachyurus. Doan & Castoe (2003) added two addi- Boulenger, and N. tatei Burt & Burt (Dixon & Lamar, tional new species, P. sucullucu and P. unsaacae. The 1981); and the N. strangulatus group, containing N. monotypic P. ventrimaculatus Boulenger group was strangulatus Cope, N. apodemus Uzzell, N. cochranae defined by a divided palpebral disc, two supraoculars Burt & Burt, N. ecpleopus Cope, N. juruazensis Avila- with a greatly expanded first superciliary, and dark Pires & Vitt, and N. ocellatus Sinitsin (Van Devender, ventral scales (Uzzell, 1970). 1969; Vanzolini, 1995; Avila-Pires & Vitt, 1998). These three taxonomic groups are allopatric in Uzzell’s divisions were based primarily on distinctive distribution, with the P. luctuosus group occurring in hemipenial morphology and he suggested that these Ecuador, Colombia, Venezuela, and Trinidad, the two groups may actually be linked by convergence P. pachyurus group in central and southern Peru and and not common evolutionary history (Uzzell, 1966; Bolivia, and the P. ventrimaculatus group in northern Pellegrino et al., 2001). Pellegrino et al. (2001) pointed Peru (Uzzell, 1958, 1970). out the nonmonophyly of Neusticurus, with the four Kizirian (1996) performed a much more extensive species that they included appearing in two or three study of the 16 species of Proctoporus from Ecuador, different positions on their trees (depending on the describing nine previously unrecognized species. data sets employed). Castoe et al. (2004) also found Neither that study, nor others (Kizirian & Coloma, the genus Neusticurus to be polyphyletic based on the 1991; Kizirian, 1995, 1996), have attempted to place five species that they included. the Ecuadorian species in any groupings. Kizirian (1996) provided a full nomenclatural history of Proc- toporus, but remarked that monophyly of the genus TAXONOMIC HISTORY OF PROCTOPORUS TSCHUDI had never been ascertained unambiguously. Tradi- The genus Proctoporus currently consists of 31 species tionally, the key diagnostic character for inclusion in that occur in several mountainous habitats in South the genus was the absence of prefrontal scales (Peters America and the Caribbean (Duellman, 1979; Kizir- & Donosos-Barros, 1970) but because three species of ian, 1996; Doan & Castoe, 2003; Doan & Schargel, the gymnophthalmid genus Pholidobolus also lack 2003; Doan, 2003a; Köhler & Lehr, 2004). In 1845, prefrontals (Montanucci, 1973) the character cannot Tschudi erected the genus for the new species be considered