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Novitates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3142, 34 pp., 6 figures June 30, 1995 Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the Taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae) RICHARD ETHERIDGE' ABSTRACT Ctenoblepharys adspersa is a highly derived ar- that have been placed in the genera Abas, Ceio- enicolous lizard that inhabits the sandy coast of laemus, Helocephalus, Pelusaurus, Phrynosaura, central Peru. It differs from other Tropiduridae, Velosaura, and Vilcunia and all ofthe species that and the outgroups Phrynosomatidae and Opluri- have been placed in the subgenera Eulaemus, Rhy- dae, in numerous autapomorphies ofthe skull and tidodeira, Mesolaemus, and Ortholaemus also ex- vertebral column, but shares no derived charac- hibit the diagnostic synapomorphies ofLiolaemus teristics with either Phymaturus or Liolaemus oth- and, although some ofthese may represent mono- er than those that diagnose Liolaeminae. The re- phyletic subsets of Liolaemus, they should not be lationships of Ctenoblepharys, Liolaemus, and used as formal names for taxa until their mono- Phymaturus are unresolved. All species of Liolae- phyletic status has been verified by cladistic anal- minae that have been described in, or referred to ysis. Phymaturus indistinctus, P. nevadoi, P. pa- Ctenoblepharys, other than C. adspersa, exhibit tagonicus, P. payunae, P. somuncurensis, and P. the synapomorphies that diagnose Liolaemus, and zapalensis, all formerly subspecies of P. patagon- should be referred to that genus. All ofthe species icus, are elevated to species status. RESUMEN Ctenoblepharys adspersa es un lagarto arenicolo y de la columna vertebral, pero no comparte nin- altamente derivado que habita en la costa arenosa gun caracter derivado con Phymaturus o Liolae- de Centro Per'u. Se diferencia de otros Tropidur- mus excepto los que se diagnostican Liolaeminae. idae y de los grupos externos Phrynosomatidae y Las relaciones de Ctenoblepharys, Liolaemus, y Opluridae en numerosas autapomorfias del craneo Phymaturus no estan resueltas a(un. Todas las es- I Professor Emeritus, Department of Biology, San Diego State University, San Diego, California, 92182-0057. Copyright © American Museum of Natural History 1995 ISSN 0003-0082 / Price $4.30 2 AMERICAN MUSEUM NOVITATES NO. 3142 pecies de Liolaeminae que han sido describidas diagnosticas de Liolaemus y anque algunas de es- en, o referido a Ctenoblepharys, excepto C. ad- tas sinapomorfias representan un subgrupo de Lio- spera, exhiben sinapomorfilas que diagnostican laemus, no deben ser usados como nombres for- Liolaemus, y deben ser referidas a ese genero. To- males para taxones hasta que su estado mono- das las especies que han sido incluidas en los ge- philetico haya sido comprobado por medio del neros Abas, Ceiolaemus, Helocephalus, Pelusau- analises cladistico. Phymaturus indistinctus, P. rus, Phrynosaura, Velosaura, y Vilcunia, y todas nevadoi, P. patagonicus, P. payunae, P. somun- las especies que han sido incluidas en los subge- curensis, y P. zapalensis, todos antiguamente su- neros Eulaemus, Rhytidodeira, Mesolaemus, y bespecies de P. patagonicus, son elevado al rango Ortholaemus tambien exhiben las sinapomorfias de especies. INTRODUCTION In 1845, Tschudi described the iguanian Queiroz (1988), Frost and Etheridge (1989) lizard Ctenoblepharys adspersa from the Pa- and Nuinez and Jaksic (1992). This disagree- cific coast of central Peru, where it appears ment in the allocation of species to the genus to be restricted to coastal sand dunes and Ctenoblepharys appears to have been due, at beaches (Mertens, 1956; J. Wright, personal least in part, to the lack of an adequate de- commun., 1989). The species is rare in col- scription of the type species, to its rarity and lections and has never been adequately de- thus unavailability to all workers, and to the scribed, but the genus has since had a long fact that Boulenger (1885) based his charac- and complicated taxonomic history, with 12 terization of Ctenoblepharys, and of the spe- species, including 15 specific names, having cies C. adspersa, on a specimen ofLiolaemus. been described in or transferred to Cteno- Furthermore, it is clear that various authors blepharys: multimaculatus Dumeril and Bi- have held different, mutually exclusive views bron, 1837; nigriceps Philippi, 1860; mar- as to what constitutes "generic characters." moratus Burmeister, 1861 (non Gravenhorst, Ctenoblepharys adspersa is one ofover 135 1837); jamesi Boulenger, 1891; stolzmanni species of austral South American lizards Steindachner, 1891; anomalus Koslowsky, which, together with Phymaturus and Lio- 1896; reichei Werner, 1907; werneri Muller, laemus, were referred to the tropidurid sub- 1928 (= anomalus fide Cei, 1979a); schmidti family Liolaeminae by Frost and Etheridge Marx, 1960; erroneus Nuinez and Ya-nez, (1989). Phymaturus includes five species, one 1984a; lentus Gallardo, 1966 (= anomalus with six allopatric subspecies that are treated fide Cei, 1979a); donosobarrosi Cei, 1974; ra- as species in this work (see below). Frost and binoi Cei, 1974; audituvelatus Niu-nez and Etheridge (1989) considered Ctenoblepharys Ya-nez, 1983; and pseudoanomalus Cei, to be monotypic, and all but this species and 1981(substitute name for marmoratus Bur- those ofPhymaturus were referred to Liolae- meister, 1861). Additionally, at various times mus, including all of the species that at one some ofthese same forms have been referred time or another had been referred to Abas, to Abas Niuniez and Yafiez, 1984b; Ceiolae- Ceiolaemus, Ctenoblepharys, Eulaemus, mus Laurent, 1984a; Eulaemus Girard, 1858; Helocephalus, Mesolaemus, Ortholaemus, Helocephalus Philippi, 1860; Liolaemus Pelusaurus, Rhytidodeira, Phrynosaura, Ve- Wiegmann, 1834; Ortholaemus Girard, 1858; losaura, and Vilcunia. In their cladistic anal- Phrynosaura Werner, 1907; and Velosaura ysis, Frost and Etheridge (1989) found two Niu-nez and Ya-nez, 1984b. equally parsimonious topologies of relation- In recent years, Cei (1979b) considered that ships: (Liolaemus (Ctenoblepharys + Phy- Ctenoblepharys contained the species ad- maturus)), and (Phymaturus (Ctenoblepharys spersa, nigriceps, reichei, and stolzmanni, + Liolaemus)). Laurent (1984a, 1984b, N(uinez and Yaiiez (1984a) and Veloso and 1985a), employing morphometric data, and Navarro (1988) included adspersa, stolzman- Nuiinez and Ya-nez (1984b), using different ni, erroneus, and nigriceps, and it was con- combinations of characters, have expressed sidered monotypic, containing only C. ad- very different, mutually exclusive ideas about spersa, by Laurent (1984a), Etheridge and de the content of Ctenoblepharys and its rela- 1995 ETHERIDGE: REDESCRIPTION OF CTENOBLEPHARUS ADSPERSA 3 tionships to other Liolaeminae. Therefore, cording to Boulenger's (1885) description, this the question ofthe content ofCtenoblepharys specimen differs from C. adspersa in a num- and its relationships to other Liolaeminae is ber ofways: the digits have smooth subdigital not a trivial one. If Ctenoblepharys or Cte- lamellae, the tail is a little shorter than the noblepharys + Phymaturus is the sister taxon head and body, the adpressed hind limb of Liolaemus, then the characteristics of C. reaches the neck, the digits are short, and the adspersa are important for the process ofpo- dorsal scales are transversely suboval; in C. larizing character state transformations with- adspersa the subdigital lamellae are keeled, in the extremely speciose genus Liolaemus. the tail is longer than the head and body, the My purpose here is to review the taxonom- adpressed hind limb reaches beyond the ex- ic history of Ctenoblepharys, to describe the ternal auditory meatus, the digits are excep- external and skeletal morphology of C. ad- tionally long, and the dorsal scales are round- spersa, to compare C. adspersa with other ed. Additionally, the locality for this speci- species that have been referred to Ctenoble- men is well to the south of the known range pharys, to comment on the possible relation- of C. adspersa, inland, and at a much higher ships of the genera of Liolaeminae, and to altitude. The identity of this specimen is un- discuss the taxonomic status of various ge- known, but specimens ofa possibly new spe- neric and subgeneric names that have been cies of Liolaemus from Arequipa (3 km SW used within the subfamily. Uchumayo, 2150 m, KU 163589, 1635902- A formal phylogenetic analysis of Liolae- 4, SDSU 1945; 18 km N Matarani, 1000 m, minae is in preparation, but is beyond the KU 163595) possess all of the features de- scope ofthe present study. Nevertheless, it is scribed by Boulenger (1985) for C. adspersa. hoped that the characters described here, and Boulenger (1891) described a second spe- their potential utility as synapomorphies cies of Ctenoblepharys, C. jamesi, (spelled (summarized in an indented classification, jamesii on the figure) from Tarapaca, Chile, Appendix 2) will facilitate future work on this at an altitude of 10,000-12,000 ft (3280-3937 large and important component ofthe austral m). Later, Boulenger (1901), following his de- South American herpetofauna. scription of Liolaemus annectens, said that "this species is very nearly related to L. mul- HISTORICAL REVIEW tiformis, Cope, from which it is to be distin- guished by the larger dorsal scales. The two What follows is a chronological account of species establish a passage from Liolaemus the taxonomic literature relating
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