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 distinguished by the larger dorsal scales. The two What follows is a chronological account of species establish a passage from Liolaemus the taxonomic literature relating to Cteno- to Ctenoblepharis, especially through C. Ja- blepharys. mesii, Blg." Subsequently L. annectens was Ctenoblepharys adspersa was described by synonymized with L. multiformis Cope, 1876, Tschudi (1845), from Hacienda Acaray, 2 by Burt and Burt (1933), which was in turn leagues from Huacho (11°07'S,77°37'W) on synonymized with L. signifer, Dumeril and the Pacific coast of Peru. A moderately de- Bibron, 1837, by Laurent (1992), who con- tailed description of the genus, and a very sidered L. annectens to be a subspecies ofL. brief description of the color pattern of the signifer. species was provided in Latin. The generic Steindachner (1891) described Ctenoble- description was repeated and the species de- pharys stolzmanni from "Hoch Peru," based scription amplified the following year on three syntypes in Vienna (NMW 1 3580[3]), (Tschudi, 1846). and provided a brief description of a speci- Boulenger (1885) included Ctenoblepharys men he referred to as the "typische Exem- adspersa in his Catalogue of Lizards in the plar" of Ctenoblepharys adspersa in the Vi- British Museum, changing the spelling to enna Museum. This specimen appears to have Ctenoblepharis adspersus, an emendation been one ofthree upon which Tschudi based followed by most subsequent authors until it his description (Ortiz-Zapata, 1989a). was found to be unjustified by Frost and Eth- Werner (1907), in his description of the eridge (1989). Boulenger's (1885) description new genus and species Phrynosaura reichei, was based on a specimen from Arequipa, suggested that the genus closest to Phryno- Peru, at an altitude of 7500 ft (2286 m). Ac- saura was Ctenoblepharys. 4 AMERICAN MUSEUM NOVITATES NO. 3142

Burt and Burt (1933) listed Ctenoblepharys tipunctatus is currently considered a syn- adspersa, C. jamesi, and C. stolzmanni as onym of Pristidactylus scapulatus (Etheridge members of the genus. and Williams, 1985). Burmeister's (1861) Donoso-Barros (1958a) did not have a Liolaemus marmoratus was transferred to specimen of Ctenoblepharys adspersa avail- Phrynosaura by Muller (1928), to Ctenoble- able, but using the illustration ofthis species pharys by Cei (1974), and back to Liolaemus in Tschudi (1846) for comparison, he con- by Cei (1980a); then Cei (1981) provided it cluded that although it showed certain sim- with the substitute name L. pseudoanomalus ilarities with Phrynosaura reichei, the char- because of the preoccupation ofL. marmor- acteristics of C. adspersa listed by Boulenger atus Burmeister, 1861, by L. marmoratus (1885) permitted separation of the two gen- Gravenhorst, 1837 (= Tropidurus nitidus era. Additionally, he (Donoso-Barros, 1958a) Wiegmann, 1834). Laurent (1984a) placed stated that the differences between C. jamesi this species in his newly described genus and Phrynosaura reichei were so great that Ceiolaemus, thus reverting the name to Ceio- there was no point in discussing them. Short- laemus marmoratus. Lataste (1892) consid- ly thereafter, Donoso-Barros (1958b) recog- ered Helocephalus nigriceps to be a synonym nized two species of Ctenoblepharys in Chile, of Ctenoblepharys adspersa, Koslowsky C. adspersa and C. jamesi, citing Hellmich's (1898) considered it to be a variety of Lio- (1934) statement that C. adspersa was a prob- laemus signifer Dumeril and Bibron, 1837, lematic member ofthe herpetofauna ofChile. and Burt and Burt (1933) referred to it as a In the same work (Donoso-Barros, 1958b) he subspecies of L. signifer. Hellmich (1934) provided measurements and descriptions of recognized Helocephalus as a subgenus of an adult male, female, andjuvenile, and pho- Liolaemus. tographs of an adult female of C. jamesi. He Donoso-Barros (1969) synonymized Lio- pointed out that Codoceo (1950) had listed laemus lentus Gallardo, 1966, with Phryno- this species under the name Liolaemus mul- saura werneri Miller, 1928, and transferred tiformis multiformis, and believed the error Liolaemus anomalus Koslowsky, 1896, to was due to their possession of convergent Ctenoblepharys, based on its presumed mor- structures. Donoso-Barros (1958b) then list- phological and ecological similarities with C. ed the differences that distinguish Ctenoble- nigriceps and C. schmidti. This action was pharys from Liolaemus multiformis. followed by Peters and Donoso-Barros (1970), Marx (1960) described Ctenoblepharys who stated that theirjustification had not yet schmidti from 40 miles east of San Pedro de been published, but was in a manuscript pre- Atacama, Antofagasta Province, Chile. Fol- pared by Donoso-Barros; the latter presum- lowing Donoso-Barros (1958b), he recog- ably referred to the work, cited above, that nized C. adspersa, C. stolzmanni, and C. ja- had actually appeared in the previous year. mesi as belonging to the genus, and provided Donoso-Barros (1971, 1972) examined a a key to these four species. specimen said to be the type of Ctenoble- Donoso-Barros (1966) provided a briefde- pharys adspersa in the Musem of Natural scription of Ctenoblepharys; however, al- History ofNeuchatel, and the type specimen though C. adspersa was mentioned as the ofPhrynosaura reichei in the Zoological Mu- nominal form ofthe genus, no description of seum ofthe University ofConcepcion, Chile. the species was given because only the lizards He concluded that both species were valid, ofChile were included. In the same work, he but that they were congeneric, and so placed transferred Helocephalus nigriceps Philippi, Phrynosaura in the synonymy of Ctenoble- 1860, ofthe Atacama desert ofChile, to Cte- pharys. noblepharys. This species has had a long and Cei (1974) reviewed the taxonomic history complex taxonomic history'tangled with that of Ctenoblepharys and recognized 11 species of Ctenoblepharys. Boulenger (1885) recog- in the genus: C. adspersa, C. anomalus, C. nized Helocephalus nigriceps but considered marmoratus, C. nigriceps, C. schmidti, C. Leiosaurus multipunctatus Burmeister, 1861, werneri, C. reichei, C.jamesi, C. stolzmanni, and Liolaemus marmoratus Burmeister, and two described as new from central Ar- 1861, to be its synonyms. Leiosaurus mul- gentina, C. donosobarrosi and C. rabinoi. In 1995 ETHERIDGE: REDESCRIPTION OF CTENOBLEPHARUS ADSPERSA 5 the following year, Cei et al. (1975) recog- laemus pseudoanomalus and L. anomalus, nized the similarity of Liolaemus multima- thus removing these species from Ctenoble- culatus and Ctenoblepharys rabinoi, and pharys and revalidating the name Liolaemus transfered the former to Ctenoblepharys. Cei marmoratus Burmeister, 1861, for Liolae- (1979a) returned C. anomalus to Liolaemus, mus pseudoanomalus Cei, 1981. He (Lau- and placed Phrynosaura werneri in its syn- rent, 1 984a) also agreed with Cei (1 979b) that onymy. Subsequently Cei (1979b) reexam- Ctenoblepharys rabinoi, Liolaemus multi- ined the basis for placing the remaining spe- maculatus (Dumeril and Bibron, 1837), and cies in Ctenoblepharys, and concluded that L. riojanus Cei, 1979b, are not related to these forms "exhibit a spectrum of common Ctenoblepharys, and considered these, to- structural characteristics that are apparently gether with L. occipitalis Boulenger, 1885, L. adaptive to live in an arid, sandy environ- lutzae Mertens, 1938, L. scapularis Laurent, ment" and also that "several unrelated, geo- 1982, L. wiegmannii (Dumeril and Bibron, graphically noncontiguous groups of species 1837), and L. cranwelli Donoso-Barros, 1973, are involved." He limited Ctenoblepharys to to form "a natural group, for which the ge- include only C. adspersa, C. stolzmanni, C. neric name Ortholaemus Girard is available, reichei, and C. nigriceps and returned the re- should it deserve generic or subgeneric rec- maining forms to Liolaemus. ognition." Laurent (1984a) and Nunez and N(uinez and Ya-nez (1983) described Cte- Yainez (1984b) apparently were not aware of noblepharys audituvelatus from San Pedro de each others' works. Atacama on the Atacama Plateau, Segunda Ortiz-Zapata and N(uinez (1986) followed Region, northern Chile, referring it to this N(uinez and Yaiiez (1984b) in transferring C. genus on the basis ofthe characteristics cited audituvelatus to Phrynosaura. Ortiz-Zapata by Cei (1979b). The same authors (Nuinez (1989a), after studying the lizards in the col- and Yanez, 1984b) then described two new lection of the Museum of Natural History of genera from northern Chile, Abas and Velo- Neuchatel, stated that the description of saura, resurrected the genus Phrynosaura, and Ctenoblepharys adspersa (Tschudi, 1845) was redefined Ctenoblepharys. In the genus Abas based on three specimens. He designated one were placed A. anomalus, A. pseudoanoma- of these, a male (MHNN 229 1-1), as the lec- lus, A. insolitus Cei and Pefaur, 1982, and A. totype, and the other two, both females, as fabiani Yanez and Ntuniez, 1983. In Velosaura paralectotypes, one of which (MHNN 2291 - were placed V. aymararum Veloso et al., 2) remains in the museum at Neuchatel, the 1982, and V. jamesi. In Phrynosaura were other (NHMW 18905) having been trans- placed P. reichei and P. audituvelatus, in ferred to the Natural History Museum in Vi- Ctenoblepharys were placed C. adspersa, C. enna. nigriceps, and C. stolzmanni, and all others In their cladistic analyses of iguanian liz- were referred to Liolaemus. A fourth species ards, Etheridge and de Queiroz (1988), and ofCtenoblepharys, C. erroneus, probably from Frost and Etheridge (1989) followed Laurent near San Pedro de Atacama in northern Chile, (1984a) in recognizing Ctenoblepharys as was then added by Nuiinez and Yanez (1984a). monotypic. Etheridge and de Queiroz (1988) Laurent (1984a), primarily based on mor- considered Ctenoblepharys to be the sister phometric comparisons with other Liolae- taxon of Vilcunia + Liolaemus. Frost and minae, considered Ctenoblepharys to be Etheridge (1989) included Vilcunia in the ge- monotypic, containing only C. adspersa. In nus Liolaemus, and considered the relation- Liolaemus he placed L. nigriceps, L. jamesi, ships of Ctenoblepharys, Liolaemus, and and L. schmidti. Ctenoblepharys and Phry- Phymaturus to be unresolved. nosaura were said to be "closely related," but Laurent (1992) summarized his earlier he stated that there is a wide gap between C. morphometric studies of the genus Liolae- adspersa on the one hand and P. reichei, C. mus (Laurent, 1984a, 1984b, 1985a), and stolzmanni, and C. audituvelatus on the oth- recognized two large species groups that he er, and thus the latter two were transferred earlier (1983a) had referred to as the Chileno to Phrynosaura. He (Laurent, 1984a) also and Argentino groups, as subgenera, L. (Lio- proposed a new genus, Ceiolaemus, for Lio- laemus) for the former, and L. (Eulaemus) 6 AMERICAN MUSEUM NOVITATES NO. 3142 for the latter. Within Eulaemus he recognized italis, M. peruvianus, Tropidurus etheridgei, a fitzingerii group and a signifer group, the and T. hygomi were used as exemplars. latter containing L. jamesi, L. schmidti and Of the 12 unrooted networks discovered L. nigriceps. by Frost and Etheridge (1989), Opluridae was the first and Phrynosomatidae the second outgroup of Tropiduridae in nine; in one (Opluridae + Polychrotidae) was the first, and MATERIALS AND METHODS Phrynosomatidae the second outgroup of This study was based on an examination Tropiduridae; in one, Phrynosomatidae was of 15 preserved specimens ofCtenoblepharys the first and (Opluridae + Polychrotidae) the adspersa and two complete skeletons pre- second; and in one, Opluridae was the first pared by hand from preserved specimens. and Polychrotidae the second outgroup. For Preserved specimens of 107 species of Lio- the purpose of this study, Phrynosomatidae laemus and eight species ofPhymaturus, and and Opluridae were considered potential sec- skeletons of 86 species of Liolaemus and six ond outgroups for Liolaeminae. Relation- species of Phymaturus were also examined. ships within Polychrotidae have not been re- These specimens, together with examplars of solved (Frost and Etheridge, 1989), and this the putative outgroups Leiocephalinae + family was not considered due to time con- Tropidurinae, Phrynosomatidae and Oplur- straints. In the absence of an explicit phylog- idae, are listed in Appendix 1. Some data eny for Opluridae, the exemplars chosen were were obtained from literature accounts of Chalarodon madagascariensis, Oplurus cu- species not examined; these also are listed in vieri, 0. cyclurus, 0. quadrimaculatus, and Appendix 1. 0. saxicola. The studies of Montanucci Squamation terminology follows Smith (1987), de Queiroz (1992), andWiens (1993a, (1946), and osteological terminology follows 1993b) formed the basis for selection ofphry- Oelrich (1956) for the skull, and Etheridge nosomatids examined: Petrosaurus mearnsi, (1964, 1965, 1966) for the postcranial skel- P. thalassinus, Uta stansburiana, U. palmeri, eton. Terminology of lateral neck folds fol- Urosaurus graciosus, U. nigricaudus, U. /ah- lows Frost (1992). teli, U. ornatus, Phrynosoma asio, P. doug- No formal analysis of the internal rela- lassii, P. orbiculare, P. coronatum, Uma no- tionships of Liolaeminae was undertaken. tata, U. exsul, and U. scoparia. However, for the purpose of discussing pos- Illustrations ofthe skulls and vertebrae were sible synapomorphies for groups within Lio- prepared by Callie Mack using a camera lu- laeminae, the outgroup method was used for cida. the polarization of character-state transformations (Watrous and Wheeler, 1981; Mad- GENUS CTENOBLEPHARYS dison et al., 1984). All remaining Tropidur- TSCHUDI, 1845 idae (= Leiocephalinae + Tropidurinae) con- Ctenoblepharys Tschudi, 1845: 150. Type species: stitute the first outgroup for Liolaeminae Ctenoblepharys adspersa Tschudi, 1845, by (Frost and Etheridge, 1989). According to monotypy. Pregill (1992), the best candidate for the least Ctenoblepharis: Boulenger 1885: 165. Unjustified apomorphic extant species ofLeiocephalus is emendation of Ctenoblepharys Tschudi, 1845 L. carinatus, and this species, plus L. schrei- (but see text). bersi and L. cubensis were used as exemplars for Leiocephalinae. No phylogeny for the Ctenoblepharys adspersa Tschudi Stenocercus group is available, so S. crassi- Figures 1, 2 caudatus, S. praeornatus, S. chlorostictus, S. Ct.[enoblepharys] adspersa Tschudi, 1845: 150 chrysopygus, S. empetrus, S. guentheri, S. (type locality: not given; lectotype [Ortiz, 1989] imitator, S. modestus, S. percultus, and S. MHNN 229-1). roseiventris were used as exemplars of the Ct.[enoblepharys] adspersa: Tschudi, 1846: 36 (re- group. Following the phylogeny of the Tro- stricted type locality: Hacienda Acaray (1 1o07'S- pidurus group presented by Frost (1992), Ur- 77°37'W), 2 leagues from Huacho, in the coastal anoscodon superciliosus, Microlophus occip- region). 1995 ETHERIDGE: REDESCRIPTION OF CTENOBLEPHARUS ADSPERSA 7

Fig. 1. Adult male Ctenoblepharys adspersa (MVZ 85415) from 5.5 km NE San Bartolo, Peru.

Ctenoblepharis adspersus: Boulenger, 1885: 165. from 6 km N Tacna at 1000 m elevation, was Unjustified emendation of Ctenoblepharys ad- identified by Pearson and Ralph (1978) as spersa Tschudi, 1845 (but see text). Ctenoblepharys sp. This locality is about 470 ETYMoLoGY: The generic name is formed km SW of Paracas, and 46 km inland from from the Greek words ktenos, meaning comb, the coast, and would represent a considerable and blepharis, meaning eyelash, presumably range extension if the specimen was one of in reference to the pronounced, serrate comb C. adspersa. formed by the pointed and projecting outer DiAGNosIs: Ctenoblepharys adspersa is a ciliaries. The specific name is from the Latin lizard of the family Tropiduridae, subfamily word adspersus, meaning a sprinkling, prob- Liolaeminae (Frost and Etheridge, 1989) ably in reference to the dorsal color pattern which differs from other members ofthe sub- that is dominated by numerous, scattered family (i.e., Phymaturus and Liolaemus) in whitish dots. having a wider skull, larger orbits, wider tem- DISTIUBuTIoN: Sandy beaches and dunes of poral fenestrae, prefrontals wider than long, the Pacific coast of Peru, from Hacienda lacrimal foramen large, a wide maxillary pro- Acaray ( 11°05'S, 77°32'W) southward to cess of the palatines, a short ectopterygoid, Paracas (' 13°50'S, 76°14'W). A specimen the maxillary process of the ectopterygoid 8 AMERICAN MUSEUM NOVITATES NO. 3142

panded crowns with large secondary cusps, three rather than four sternal ribs, and a slender body with a long, slender tail rather than a depressed body with a short, spinose tail. DESCRIPTION: Squamation and Propor- tions (figs. 1, 2): Head large, distinct from neck, widest across temporal region, 0.82- 0.92 times wider (as measured across widest part of temporal region) than long (as measured from inferior apex ofexternal auditory meatus to anterior surface of rostral). Snout short (as measured from tip of snout to anterior corner or orbit), 0.29-0.35 times head length, projecting slightly beyond lowerjaws; orbit (as measured along its greatest horizontal length) large, 0.43-0.47 times head length. Nasal region swollen, convex in profile; frontonasal region slightly concave in profile. Rostral narrow, 2.60-3.50 times wider than high, bordered by 5-7 postrostrals. Nasal scales large, projecting, separated from rostral and anterior supralabials by two scale rows; nostril oriented anterolaterally, occu- pying most ofscale. Dorsal head scales small, poorly differentiated, somewhat irregularly convex, especially in frontonasal region; in some specimens slightly concave (perhaps due to preservation). Supraorbital semicircles apparent only in prefrontal region; 2 or 3 irregular scale rows, or 1-3 azygous frontals between orbits; supraoculars all very small, subequal, mostly hexagonal, 10-13 in a horizon- Fig. 2. (A) Dorsal view; (B) lateral view of tal line across widest part of supraocular head of an adult male Ctenoblepharys adspersa region between superciliaries and frontals; in- (MVZ 85415). Scale = 0.5 cm. terparietal a little larger than adjacent parietal scales, bordered by 9 or 10 scales, with a distinct "eye"; no pair of enlarged parietals posterior to interparietal. Superciliaries short, truncate, the retroarticular process of the not strongly keeled, about 2-3 times longer mandible much shorter than the angular pro- than wide, anterior 6-7 with oblique sutures, cess, the parapophyses of the atlas flat and followed by a row of 6-8 small, nonoverlap- oriented posterolaterally, and the neural ping scales, preceded by one large anterior arches of the body vertebrae very wide. It and one slightly smaller posterior canthal. further differs from Liolaemus in having the Palpebrals small, convex, juxtaposed; inner lateral borders of the orbitonasal fenestra ciliaries rectangular, about twice as high as formed by the prefrontals rather than the wide; outer ciliaries of lower lid 11-14, tri- frontals, the supratemporal exposed on the angular, sharply pointed distally, altogether lateral surface of the supratemporal process forming a strongly projecting serrate comb; of the parietal, a shorter dentary, a longer outer ciliaries ofupper lid 13-18, the anterior angular, and no posterior coracoid fenestra. and posterior ones triangular, but not as It further differs from Phymaturus in having sharply pointed or as strongly projecting as tricuspid marginal teeth with tapered crowns those oflower lid, those in middle oflid more and small secondary cusps rather than ex- nearly rectangular, scarcely projecting, with 1995 ETHERIDGE: REDESCRIPTION OF CTENOBLEPHAR US ADSPERSA 9 a convex margin. Scales of preocular-sub- continuing posteriorly as longitudinal neck ocular-postocular arc 7-10, the third, fourth, fold, crossed by oblique neck fold and ante- or fifth distinctly elongate, about 2-3 times humeral fold; no supra-auricular, supernu- longer than high and with a blunt keel, or merary antegular, or antegular folds; gular none distinctly elongate; suboculars and pos- fold represented laterally by short folds sep- toculars swollen, forming an interrupted, arate medially by 10-12 scales. Lateral nu- strongly projecting shelf. A single row of 9- chal pouches absent. 11 lorilabials, slightly smaller than suprala- Limbs moderately slender; hind limbs bials, separating suboculars from supralabi- 0.64-0.78 times snout-vent length. Brachial als. Anterior loreals about equal in size to and antebrachial scales, except preantebra- lorilabials, followed by row ofmuch smaller, chials, convex, nonoverlapping, with inter- often elongate scales between anterior subo- stitial granules, about equal in size to lateral culars and lorilabials. Supralabials 7-1 1, sub- body scales; several longitudinal rows of equal. Temporals small, convex, juxtaposed, smooth, flat, rhomboidal, imbricate prean- with minute interstitial granules, about 14- tebrachials, about equal in size to ventral body 16 between postocular and anterior margin scales. Suprafemorals, postfemorals, and of ear. External auditory meatus large, obo- posterior infrafemorals similar in form to vate, 1.54-1.87 times higher than greatest dorsal body scales but smaller; prefemorals width, vertical diameter 0.41-0.50 times lon- similar to ventral body scales, grading into gitudinal diameter oforbit, bordered by small, smooth, rounded posterior infrafemorals, convex scales, some slightly enlarged ante- with interstitial granules. Supratibials and riorly, or not differentiated from posterior pretibials small, convex, similar to dorsal temporals. Mental small, 0.51-0.76 times as body scales; posttibials and infratibials wide as rostral, bordered by two infralabials smooth, flat, subimbricate, about equal in size and two postmentals, not in contact with an- to ventral body scales. Supracarpals and su- terior sublabials. Infralabials 9-10. Gulars pratarsals smooth, rhomboidal, imbricate; small, smooth, convex, oval or rounded, with infracarpals and infratarsals imbricate, minute interstitial granules. somewhat projecting, mucronate, some with Dorsal scales of neck and body small, flat a tridentate margin. Supradigitals smooth, or in some specimens slightly concave (per- imbricate, with a slightly concave distal mar- haps due to state ofpreservation), aligned in gin on manus, horizontal or slightly convex more-or-less transverse rows, nonoverlap- on pes; distal margin of terminal supradigi- ping, with tiny interstitial granules, becoming tals distinctly notched. Lateral digitals tri- smaller and strongly convex on the sides, angular, forming a serrate comb, more pro- grading gradually into smooth, flat, subim- nounced on pes. Subdigital lamellae with three bricate ventral scales. Ventrals rhomboidal or four low, blunt keels, each terminating in with rounded corners, about three times larg- a blunt mucron; subdigital lamellae offourth er than largest dorsals, becoming scarcely im- toe 33-38. Claws long and slender, variable bricate with interstitial granules on posterior in length perhaps due to wear between shed- abdomen. Scales of precloacal region (fig. 3 ding cycles; fourth toe claw about as long as A, B) similar to posterior ventrals in both 3-5 distal supradigitals. sexes. Precloacal pores 6-8 (x = 7.1, sd = Tail rather thick and somewhat depressed, 0.8), present in males (N = 8) only. Scales 0.53-0.60 percent total length in males, 0.53- around midbody 95-119 (x = 103.9, sd = 0.57 in females; proximal 86-90 percent of 6.6); middorsal scales from occiput to point tail wider than high, rounded distally. Dorsal even with anterior margin of thigh 113-135 and lateral caudal scales like those of dorsal (X = 120.8, sd = 7.0). Lateral nuchal skin body, becoming subimbricate on middle third folds well-developed and complex: two short of tail, bluntly keeled and weakly mucronate folds, one (possibly postauricular) originating on distal third; ventral caudals, like ventral at superior, the other at inferior apex of ex- body scales, becoming more distinctly im- ternal auditory meatus, converging posteri- bricate, bluntly keeled and mucronate on dis- orly to form, with posteriorborder ofexternal tal third of tail. Autotomic part of tail with auditory meatus, a shallow depression, and five dorsal and four ventral transverse rows 10 AMERICAN MUSEUM NOVITATES NO. 3142

Fig. 3. Ventral view ofpygal area of(A) male (MVZ 85415), (B) female (FML 0464) ofCtenoblepharys adspersa; and (C) male (SDSU 1532), and (D) female (SDSU 1319) of Liolaemus multicolor. Scale = 1.0 cm. in each segment, the first two dorsal rows in pale tan with small brown spots; lower sur- each segment above the first ventral row. faces of lower ciliaries white. Outer ciliaries Color Pattern (figs. 1, 2): In preservative, each with a single light brown scale organ, dorsal and lateral surfaces of head medium most ofthem subterminal in position. Dorsal brown with scattered whitish dots. Palpebrals surfaces ofneck, body and tail speckled, with whitish, those of upper lid with small brown indistinct crossbands of brown or dark gray, spots; inner ciliaries dark brown proximally, alternating with light tan, two on neck and fading to pale tan distally; upper and lower six on body, becoming progressively more surfaces of outer ciliaries of upper lid, and obscure distally on tail; banded pattern ren- dorsal surfaces of outer ciliaries of lower lid dered indistinct by numerous small, whitish 1995 ETHERIDGE: REDESCRIPTION OF CTENOBLEPHARUS ADSPERSA I1I dots. Upper surfaces oflimbs similarly band- suture with one another extending posteriorly ed, also with small, whitish dots. In males, well beyond level ofanterior corner oforbits, throat and chest light grayish-brown with their sutures with the frontal forming a shal- scattered, small whitish spots, especially low W. Descending prefrontal processes of prominent on sides; an ill-defined unpig- frontals short, widely separated from pala- mented zone on anterior chest, between fore- tines, prefrontals contributing to lateral walls limb insertions; remainder of abdomen and of orbitonasal fenestra. Prefrontal short and ventral surfaces of limbs and tail whitish. wide, much wider than long, flat above, with Throat of females as in males, other ventral a slight medial concavity; antorbital process surfaces unpigmented. Color photograph of of prefrontal flattened, posterolaterally ori- living individual from Paracas Beach with ented, strongly projecting. Parietal short and pattern as above; dark crossbands brownish wide, width ofanterior margin 1.1-1.4 times gray, light background color yellowish tan. length as measured from anterior margin to Measurements: Snout-vent length (SVL) of posterior extremity of supratemporal pro- largest male (LACM 49145) 75 mm, tail in- cesses. Parietal foramen formed by a median complete; largest female (SMF 75966) 67 mm, notch in anterior margin of parietal and an tail 80 mm. Measurements of an adult male apposing notch in posterior margin offrontal. (MVZ 85415) in millimeters: SVL 72, tail Postfrontals very small, not or scarcely vis- length 110, forelimb 31, hind limb 49, head ible in dorsal view. Supratemporal exposed length (inferior apex ofexternal auditory me- on lateral face of supratemporal process of atus to rostral) 18.0, maximum head width parietal, not hidden within groove on ventral (across temporal region) 16.2, snout length face of latter; supratemporal fenestra wide, (anterior corner of orbit to rostral) 5.8, di- 1.5-1.6 times longer than wide. Supratem- ameter of orbit (between posterior and an- poral process of squamosal without a dorsal terior corner of orbit) 7.5, vertical diameter hooklike process, separated from contact with of external auditory meatus 3.7. parietal by supratemporal. Skeleton: The following descriptions are Lacrimal very small, scarcely visible in lat- based on two complete skeletons prepared eral view; lacrimal foramen very large, its from preserved specimens. The larger, LACM inferior margin formed by maxilla so that 49147, is a male 68 mm SVL, with a skull prefrontal fails to contact lacrimal below. 15.8 mm long. The occipital sutures are part- Maxillary process of jugal slopes distinctly ly fused at the condyle, the remaining brain- outward, its lateral aspect tapered anteriorly case sutures unfused; scapulocoracoid su- almost to a point where it contacts the lac- tures fused, acetabular sutures partly fused, rimal. epiphyses of long bones not fused to diaph- Interpterygoid vacuity miter-shaped, wide yses. Based on these observations, this in- posteriorly, abruptly narrowing at the pala- dividual had not attained maximum size. The topterygoid suture, continuing forward to smaller specimen, REE 2513, is a female, 57 separate palatines and posterior portion of mm SVL, with a skull 13.8 mm long. All vomers from medial contact. Infraorbital fe- sutures remain unfused. nestra obovate, its anterior margin nearly Skull (fig. 4A-C) short and broad, 1.27- horizontal due to wide maxillary process of 1.29 times longer than wide; orbit large, its palatine, its posterior margin formed by rel- greatest longitudinal diameter 0.49-0.50 atively short, anterolaterally oriented ectop- times skull length; snout short, 0.29 times terygoid. Maxillary process of ectopterygoid skull length; postorbital part of skull short, anteriorly truncate, not tapered to a point. 0.20-0.22 times skull length. Nasal process Basisphenoid short and wide, its pterygoid of premaxilla not extremely wide or narrow, processes short and widely divergent, its posteriorly clamped between nasals; arch spheno-occipital processes short, not extend- formed by premaxillary processes of nasals ing to spheno-occipital tubercles. and nasal process of premaxilla separated Meckel's groove unfused, its upper and from underlying septomaxilla by a wide gap. lower borders in contact between teeth 10- Nasals wide, not tapering to a point poste- 14, open from tooth 9 to mandibular sym- riorly between frontal and prefrontals, their physis, occupied by splenial under teeth 15- 12 AMERICAN MUSEUM NOVITATES NO. 3142

A pmxD t_e~~~~~~~~c

Fig. 4. Skulls of(A-C) Ctenoblepharyvs adspersa (LACM 49147) and (D-F) Liolaemus nigriceps(REE 2537); A, D dorsal view; B, E ventral view; C, F view of anterior wall of left orbit. Abbreviations: bso- basioccipital, bsp-basisphenoid, ect-ectopterygoid, fr-frontal, ju-jugal, lac-lacrimal, max-maxilla, na-nasal, onf-orbitonasal fenestra, par-parietal, pmx-premaxilla, p0o-postorbital, prf-prefrontal, ptr-pterygoid, qu-quadrate, sq-squamosal, st-supratemporal, vom-vomer. Scales for A, B, D and E = 1 cm; for Cand F = 0.5 cm. 1995 ETHERIDGE: REDESCRIPTION OF CTENOBLEPHARUS ADSPERSA 13

20. Angular large, labial process wide, lingual process narrow, extending forward to position of last tooth of dentary; posterior my- lohyoid foramen not apparent. Splenial large, extending forward between inferior apex of posterior lingual process of coronoid to level of dentary tooth number 15; anterior mylo- I hyoid foramen represented by one or two openings between inferior margin ofanterior A splenial and dentary; anterior inferior alve- olar foramen represented by a notch in dentary, with superior margin ofanterior splenial forming inferior border. Posterior limit of dentary on labial face ofmandible about even with posterior margin of posterior lingual process of coronoid. Labial process of coronoid constricted proximally, well separated from anterior supra-angular foramen. Re- troarticular process short, shorter than angular process. All marginal teeth slightly tapered, with crowns slightly compressed linguolabially, I and slightly curved inward. Both upper and lower jaws with 20 teeth on each side. Pre- maxilla with six simply pointed teeth; first four maxillary teeth similar to premaxillary teeth, followed by 13 teeth tricuspid with Fig. 5. Dorsal view of the 11th body vertebra small secondary cusps. Anterior 6-8 dentary of (A), Ctenoblepharys adspersa (LACM 49247); teeth simply pointed, followed by 12-14 tri- and (B), Liolaemus nigriceps (REE 2537). Scale = cuspid teeth with small secondary cusps. 1.0 mm. Pterygoids with 1-3 small, pointed teeth; palatine teeth absent. Parapophyses of atlas somewhat flattened, posteriorly oriented and tapering distally. processes gradually reduced in length poste- Presacral vertebrae 23, all with an unusually riorly, absent on vertebrae 33 through 43. wide neural arch and slight constriction be- Proximal (interclavicle) process ofclavicle tween zygapophyses (fig. 5A). Ratios based expanded posteriorly to form a thin plate, on measurements of 11th vertebra are: (1) fenestrate or not on one or both sides, with maximum length of neural arch, including posterior margin irregular or forming a zygapophyses, divided by narrowest width of smooth curve; proximal process about equal arch between zygapophyses 0.98-1.00; (2) in length to distal (suprascapular) process, the maximum length of neural arch divided by latter narrow and rounded. Interclavicle ar- maximum width across prezygapophyses row-shaped, proximal 65 percent of lateral 0.78-0.88; (3) ventral length of centrum, ex- processes contacting posterior margins of cluding condyle, divided by narrowest width clavicles, medial process widest just anterior of neural arch between zygapophyses 0.69. to sternum, extending posteriorly into sternal Caudal vertebrae 43, the first eight nonau- fontanelle to about level of articulations of totomic, with posterolaterally oriented pro- second pair of sternal ribs. Scapulocoracoid cesses, the anterior ones projecting not quite and primary coracoid fenestrae large; scap- as far as sacral diapophyses; remaining au- ular fenestra absent, with no trace ofthinning totomic caudal numbers 9-23 with slender, in the scapula; secondary coracoid fenestra laterally oriented processes, numbers 24-32 absent, but a thin region present between sec- with anterolaterally oriented processes; all ondary coracoid ray and coracoid plate. Su- 14 AMERICAN MUSEUM NOVITATES NO. 3142 prascapula curves gradually dorsomedially also point out here that the status of Cte- above scapula-suprascapular suture. Ster- noblepharys erroneus remains uncertain. The num pentagonal, articulating laterally with unique type, which I have not examined, is three pair ofsternal ribs and posteriorly with said to lack projecting outer ciliaries and the xiphisternal rods; fontanelle a narrow oval, tail is shorter than the snout-vent length, but about three times longer than wide. Two pair it has poorly differentiated cephalic scales and ofxiphisternal ribs, lacking free posterior ex- a truncate snout (Nu(niez and Ya-nez, 1984b). tensions of xiphisternal rod. Pubic tubercle In view of its lack of a ciliary comb and its of pelvis flattened vertically. short tail, and its locality in the Atacama des- Long bones of appendicular skeleton grac- ert of northern Chile, it seem unlikely that ile; metacarpal of fourth digit about as long this species will be referred to Ctenoblepha- as combined length of metacarpal and prox- rys. imal phalanx offifth digit; proximal and dis- Cei (1986) recognized four species ofPhy- tal phalanges of fifth digit about equal in maturus under the generic name Centrura: P. length; femur with a distinct sigmoid curve; flagellifer (= P. palluma), P. mallimaccii, P. tibia without a posterior distal bladelike pro- punae and P. patagonicus, the latter with six cess; distal extremity of distal penultimate subspecies: P. p. patagonicus, P. p. indistinc- phalanx of fifth toe extends slightly beyond ta, P. p. nevadoi, P. p. payunae, P. p. so- distal extremity of metatarsal of fourth toe. muncurensis, and P. p. zapalensis. Pereyra Plantar tubercles of fifth metatarsal widely (1985, 1991) recently described a fifth spe- separated. Claws slightly over 1.5 times length cies, P. antofagastensis. The subspecies ofP. of penultimate phalanx. patagonicus are diagnosable on the basis of Muscles (fig. 6A): Medial head ofM. flexor squamation, proportions, color pattern, and tibialis internus exposed, not covered by hy- the presence or absence of sexual dichro- pertrophied M. puboischiotibialis. Insertion matism; and, they are all allopatric, with no ofM. tibialis anterior not hypertrophied. Me- evidence of intergradation (Cei and Castro, lanic pigment not present within median por- 1973; Cei and Roig, 1975; Cei, 1986). They tion of epimysium of M. pterygomandibu- therefore appear to meet the criteria for evo- laris. lutionary species sensu Frost and Hillis (1990) and Frost et al. (1992), and for this reason are here elevated to specific rank. Neverthe- RELATIONSHIPS OF LIOLAEMUS, less, the species of Phymaturus fall into two CTENOBLEPHARYS, AND groups that are phenetically similar in squa- PHYMATUR US mation and skeletal morphology, the P. pal- For the purpose of this discussion, and in luma group (P. antofagastensis, P. palluma, the remainder of this work, all of the species P. punae, and P. mallimaccii) characterized ofLiolaeminae except for Ctenoblepharys ad- by a larger adult body size (maximum snout- spersa and those referred to Phymaturus, are vent length 95-110 mm), more strongly spi- included in the genus Liolaemus (sensu lato). nose caudal scales, and more fragmented head Evidence for monophyly ofthis taxon will be scales, including several rows of lorilabials given below. Thus, the generalizations as to and the absence of an elongate subocular, an character states of Liolaemus described be- open Meckel's groove, and a large splenial low also apply to the species that have, at one that extends at least as far forward as the time or another, been referred to Ctenoble- midpoint of the dentary tooth row. The P. pharys (except the type), as well as to those patagonicus group (P. patagonicus, P. indis- referred to Abas, Ceiolaemus, Eulaemus, tinctus, P. nevadoi, P. payunae, P. somun- Mesolaemus, Ortholaemus, Pelusaurus, curensis, and P. zapalensis) is characterized Phrynosaura, Rhytidodeira, Velosaura, and by the alternatives to the above character Vilcunia. This action does not necessarily im- states. ply that any one of these taxa is not mono- Comparisons of the skeleton and integu- phyletic, but rather that all ofthem are mem- ment of Ctenoblepharys adspersa with Phy- bers of a single clade that does not include maturus and Liolaemus indicate that C. ad- Phymaturus or Ctenoblepharys adspersa. I spersa possesses a large number of charac- 1995 ETHERIDGE: REDESCRIPTION OF CTENOBLEPHARUS ADSPERSA 15

teristics, chiefly of the skeleton, not found AZ publsht,bpublahtib amb iiitlbilltib femtib elsewhere in Liolaeminae. These are (1) skull wide, length/width ratio 0.77-0.79 (0.63-0.72 in Phymaturus, 0.63-0.74 in Liolaemus); (2) orbit large, skull/orbit ratio 1.99-2.04 .N- - (2.18-2.92 in Phymaturus, 2.12-2.72 in Lio- laemus); (3) temporal fenestra wide, fixtib cm f CxtSbsp A7jflength/width ratio 1.49-1.59 (1.78-3.10 in Phymaturus, 1.72-3.57 in Liolaemus); (4) median suture between nasal bones extends gast /////,SE p1111 posteriorly far beyond level ofanterior corner of orbits (rather than to about level of ante- 1E1bFtibant rior corner of orbits), (5) prefrontal much wider than long, the antorbital process prominent and strongly projecting (approached in some species of Liolaemus, e.g., L. anomalus, L. pseudoanomalus); (6) lacrimal bone B pubIshtib amb reduced, not forming inferior margin of lac- B , iiitb femtib rimal foramen (rather than large and forming inferior margin oflacrimal foramen; lacrimal absent in L. nigriceps); (7) lacrimal foramen very large; (8) maxillary process ofjugal ex- ternally tapering almost to a point (rather than truncate); (9) orbital surface of jugal slopes fixtAbcm /////t1///2 strongly outward (rather than vertical or slopes slightly outward; (10) palatines and gast tibant posterior vomers not in medial contact (rather than vomers always in full contact and usually also palatines in contact anteriorly); (11) anterior margin of infraorbital fenestra nearly horizontal due to wide maxillary process of palatine (rather than rounded, with C5publshtlb *mb WtI,b f,emtib narrow maxillary process of palatine); (12) ectopterygoid short, that part forming posterior margin of infraorbital fenestra shorter than that part of palatine forming medial margin offoramen (rather than long, that part forming posterior margin of infraorbital foramen longer than that part ofpalatine form- ga8s,SlIIling medial!{/D margin of foramen); (13) ectopterygoid orientation more nearly lateral than anterolateral; (14) dorsal aspect of maxillary tibant process ofectopterygoid truncate (ratherthan pointed); (15) retroarticular process of mandible much shorter than angular process NVW(rather than equal to or longer); (16) par- Fig. 6. Dorsal view of the musculature of the right hind limb of (A) male Ctenoblepharys adspersa (LACM 49147), (B) female Liolaemus fitzingerii, (SDSU 1191) and (C) male Liolaemus cp-M. flexor tibialis internus capitis posterior, elongatus (SDSU 1857). Abbreviations: amb-M. gast-M. gastrocnemius, ilitib-M. iliotibialis, ambiens, femtib-M. femerotibialis, ffixtib cm- pubishtib-M. puboischiotibialis, tibant-M. ti- M. flexor tibialis internus capitis medialis, flxtib bialis anterior. 16 AMERICAN MUSEUM NOVITATES NO. 3142 apophyses of atlas flat and oriented poster- the ventral surfaces of the paroccipital pro- olaterally (rather than round and laterally ori- cess of parietal (Frost and Etheridge, 1989; ented); and (17) neural arch ofbody vertebrae fig. 2C), (3) dentary long, extending well be- very wide, arch of 1 th vertebra length/width yond posterior lingual process of coronoid ratio 0.98-1.08 (1.26-1.84 in Phymaturus, (except in a single specimen of L. sylvanae 1.12-1.81 in Liolaemus). [MCZ 156906]), (4) anterior process of an- There appear to be no characteristics of gular reduced or absent, not extending for- squamation in Ctenoblepharys adspersa that ward beyond level of posterior lingual pro- are unique within Liolaeminae, although sev- cess of coronoid, and (5) posterior coracoid eral are found in only a few species. The small, fenestra present (rarely absent as an individ- undifferentiated head scales, including nu- ual variant). merous subequal supraoculars and parietals In Ctenoblepharys adspersa and Phyma- of C. adspersa, appear to occur in only Lio- turus, the cloacal region (the triangular region laemus erroneus. As in Ctenoblepharys ad- between the hind limb insertions and vent) spersa, the mental is narrower than the rostral ofmales is about the same size as in females, in most individuals of Phymaturus, and in but in Liolaemus it is much larger in males, those species of Liolaemus assigned to Or- due to the anterior location and hypertrophy tholaemus by Laurent (1984a), but in the lat- of the anterior retractor muscles of the hem- ter the mental has a different shape due to its ipenes (Arnold, 1984) (fig. 3C, D). In most contact with the anterior sublabials. A sub- Liolaemus males the scales in the central part ocular fragmented into several short scales ofthis region are approximately the same size also occurs in the Phymaturuspalluma group as the ventral body scales, whereas they are and in Liolaemus stolzmanni and L. erro- conspicuously reduced in females. In C. ad- neus. The pattern of dorsal body scales in spersa and Phymaturus the scales of this re- Ctenoblepharys adspersa, i.e., small, flat, oval gion are the same size in both sexes. Although or rounded, nonoverlapping scales arranged Arnold (1984) stated that the anterior re- in more-or-less transverse rows with con- tractor muscle was larger in Ctenoblepharys spicuous interstitial granules, is not exactly and Liolaemus than in Phymaturus, he has matched in any other Liolaeminae, although informed me (Arnold, personal commun., similar patterns of subimbricate to nonov- 1989) that his specimens of Ctenoblepharys erlapping scales with interstitial granules oc- adspersa were, in fact, specimens of Liolae- cur in a number ofspecies ofLiolaemus, e.g., mus monticola chillensis (BMNH 1910.- L. anomalus, L. audituvelatus, L. pseudoan- 9.20.7-9). omalus, L. stolzmanni, L. schmidti, L. in- Although absent in Phymaturus and Cte- solitus, L. nigriceps, L. reichei, and L. an- noblepharys adspersa, in a large number of dinus. Liolaemus, melanic pigment is present with- Osteological character states shared by in the median portion of the epimysium of Ctenoblepharys adspersa and Phymaturus, but the M. ptergomandibularis, the medial head not found in Liolaemus, are: (1) lateral bor- of the M. flexor tibialis internus is covered ders of orbitonasal fenestra formed by pre- by a hypertrophied M. puboischiotibialis, and frontals (fig. 4A), (2) supratemporal exposed the insertion of the M. tibialis anterior is hy- on lateral face ofparaoccipital process of pa- pertrophied in association with the presence rietal, (3) dentary short, not extending pos- ofa sharp, bladelike process ofthe tibia (Cei, teriorly beyond posterior lingual process of 1993: fig 36) (fig. 6C). The tibial blade char- coronoid, (4) anterior process ofangular long, acter was first described and illustrated for extending forward to level with posterior Liolaemus occipitalis by Keller and Krause dentary tooth, and (5) posterior coracoid fe- (1986). These character states have been con- nestra absent. Alternative states of these firmed in 58 species ofLiolaemus (Appendix characters found in Liolaemus are: (1) lateral 2), including L. multimaculatus, L. nigriceps, borders of orbitonasal fenestra formed en- L. jamesi, L. schmidti, L. donosobarrosi, L. tirely or almost entirely by frontal down- rabinoi, and L. audituvelatus, all ofwhich, at growths (fig. 4F), (2) supratemporal mostly one time or another, have been referred to or entirely enclosed within a deep groove on Ctenoblepharys. A pigmented epimysium of 1995 ETHERIDGE: REDESCRIPTION OF CTENOBLEPHARUS ADSPERSA 17 the M. pterygomandibularis and a hypertro- mains. In their cladistic analysis, Frost and phied M. puboischiotibials also occur in L. Etheridge (1989) found two topologies that anomalus and L. pseudoanomalus, but a tib- are independent of network. In Topology 1, ial blade and a hypertrophied M. tibialis an- Phymaturus was the sister taxon of Cteno- terior are lacking in these species. All other blepharys + Liolaemus, and in Topology 2 Liolaemus examined (57 species), as well as Liolaemus was the sister taxon of Phyma- Ctenoblepharys, Phymaturus, other Tropi- turus + Ctenoblepharys. In Topology 1 the duride, Phrynosomatidae, and Opluridae, linkage ofCtenoblepharys with Liolaemus was lack melanic pigment in the epimysium of supported by the supratemporal fitting in a M. pterygomandibularis, a tibial blade is groove of the supratemporal process of the lacking, and the M. puboischiotibialis and M. parietal. Unfortunately, this character was in- tibialis anterior are not hypertrophied (Hoy- correctly coded for Ctenoblepharys. In Cte- os, 1990; personal obs.). noblepharys, as in Phymaturus, the supra- Of the character states found in Cteno- temporal occupies its primitive position on blepharys adspersa that are unique within the lateral face of the supratemporal process Liolaeminae, the following likewise do not of the parietal. In Topology 2, the linkage of occur in other Tropiduridae, nor in Opluri- Ctenoblepharys with Phymaturus is support- dae or Phrynosomatidae, and are thus pre- ed by the presence of a divided subocular, sumed to be autapomorphies for the species: present in C. adspersa and in the P. palluma (1) skull wide, length/width ratio 0.77-0.79 group, but not the P. patagonicus group. Frost (0.57-0.71 in other Tropiduridae, 0.58-0.73 and Etheridge (1989) also pointed out that in Phrynosomatidae, 0.62-0.70 in Opluri- Arnold (1984) noted the presence of a well- dae), (2) orbit large, skulVorbit ratio 1.99- defined fleshy insertion of the M. retractor 2.04 (2.08-2.48 in other Tropiduridae, 2.13- lateralis posterior of the hemipenis as a pos- 2.64 in Phrynosomatidae, 2.13-2.61 in sible synapomorphy for the Liolaemus group Opluridae); (3) median suture between nasals (i.e., Liolaeminae), and that it is better de- extends posteriorly far beyond anterior cor- veloped in Ctenoblepharys and Liolaemus ner of orbits, (4) lacrimal bone very small, than in Phymaturus. However, as pointed out not forming part of inferior margin of lacri- above, Arnold's specimen representative of mal foramen (lacrimal absent in some phry- Ctenoblepharys was actually a specimen of nosomatids), (5) lacrimal foramen large, (6) Liolaemus monticola. Thus, there are no anterior margin ofinfraorbital fenestra nearly known derived features shared by Phyma- horizontal due to wide maxillary process of turus and Ctenoblepharys to the exclusion of palatine, (7) maxillary process of ectoptery- Liolaemus, nor are there any derived features goid truncate, (8) parapophysis of atlas flat shared by Ctenoblepharys and Liolaemus to and oriented posterolaterally, and (9) neural the exclusion of Phymaturus, or by Phyma- arch of body vertebrae very wide, length/ turus and Liolaemus to the exclusion of Cte- width ratio of 11th vertebra 0.98-1.08 (1.53- noblepharys. Relationships of the three Lio- 2.04 in other Tropiduridae, 1.46-1.62 in laeminae genera therefore remain unre- Phrynosomatidae, 1.57-1.71 in Opluridae). solved. All of the character states listed earlier as shared by Ctenoblepharys and Phymaturus to the exclusion of Liolaemus are judged to be COMMENTS ON THE STATUS OF THE plesiomorphic based on comparisons with GENERA ABAS, CEIOLAEMUS, other Tropiduridae, Phrynosomatidae, and PEL USA URUS, PHR YNOSA URA, Opluridae, and the alternative states listed for RHYTIDODEIRA, VELOSAURA, AND Liolaemus are therefore interpreted as evi- VILCUNIA, AND THE SUBGENERA dence for the monophyly of those Liolae- EULAEMUS, LIOLAEMUS SENSU minae exclusive of Phymaturus and Cte- STRICTO, MESOLAEMUS, AND noblepharys adspersa, i.e., Liolaemus sensu ORTHOLAEMUS lato. Monophyly ofLiolaemus appears to be well The question of relationships of Phyma- supported by the possession offrontal down- turus, Ctenoblepharys, and Liolaemus re- growths that-exclude (or nearly exclude) the 18 AMERICAN MUSEUM NOVITATES NO. 3142 prefrontals from the orbitonasal fenestra, the trophied M. tibialis anterior, while L. an- supratemporal mostly or entirely enclosed omalus and L. pseudoanomalus do not. Thus, within a deep groove on the ventral surface monophyly ofAbas is unsupported. of the paroccipital process of the parietal, a long dentary that extends posteriorly well be- CEIOLAEMUS yond the posterior lingual process ofthe coronoid, the anterior process of the angular re- Laurent (1984a) described Ceiolaemus, in- duced or absent, the presence of a posterior cluding in it C. anomalus and C. marmoratus coracoid fenestra, and the anterior location (= pseudoanomalus), and designated the lat- and hypertrophy of the anterior retractor ter as its type species. It was distinguished muscles of the hemipenes. However, generic from Phymaturus by its larger and less nu- or subgeneric status has been proposed for merous scales and from other liolaemine gen- several groups of species within Liolaemus. era by a number ofmorphometric characters, In this section, their status as monophyletic plus palatine teeth and smooth dorsal scales. Liolaemus will be discussed. Palatine teeth do not occur in any tropidurid subsets of lizards, and it seems likely that this actually was a reference to the presence of pterygoid ABAS teeth, which are present in most Liolaemus, N(uinez and Ya-nez (1 984b) described Abas, including both Liolaemus anomalus and L. and included in it L. anomalus, L. pseu- pseudoanomalus. Also, smooth dorsal scales doanomalus, L. insolitus Cei and Pefaur, occur in a number of other Liolaemus. The 1982, and L.fabiani Ya-nez and Niu-nez, 1983, two species are, however, phenetically ex- the latter designated as the type. The genus tremely similar in squamation, skeletal mor- was characterized as having (1) eyelids with phology, and dorsal color pattern, and may a short comb, (2) diameter of eye less than well be sister taxa. length from anterior border of eye to rostral scale, (3) tail equal to or longer than snout- PELUSA UR US vent length, (4) head scales differentiated, (5) profile isognathus, (6) loreal region slightly Donoso-Barros (1973) described Pelusau- depressed, and (7) dorsal scales imbricate and rus, and P. cranwelli as its only species, based smooth, without companion scales (my on a single female from Macho, Nueva Moka translation). My examination ofthese species (17019'S, 63033'W), Santa Cruz Prov., Boliv- indicates that not all of them have a short ia. Laurent (1 983b) pointed out the similarity comb on the eyelid; the outer lower ciliaries ofthis species to Liolaemus wiegmannii (Du- of L. anomalus and L. pseudoanomalus are meril and Bibron, 1837), and assigned it to more projecting and more nearly pointed than the subgenus Ortholaemus. I have compared in L.fabiani and L. insolitus. In most species the unique type (MACN 3632) with speci- ofLiolaemus the diameter ofthe orbit is less mens of L. wiegmannii and find the squa- than the length of the snout, the head scales mation and color pattern of the type to be are differentiated, the profile is isognathus, well within the limits ofvariation ofL. wieg- and the loreal region is slightly depressed. mannii, although Laurent (1983b) indicated The tail is shorter than the snout-vent length that there may be proportional differences. in L. insolitus (also shown by the measure- The locality for L. wiegmannii most proxi- ments in Cei and Pefaur [1982: table 2]) and mate to that ofL. cranwelli is Yuto (230381S, in female L. pseudoanomalus, and the dorsal 64028'W), Depto Ledesma, Prov. Jujuy, Ar- body scales are nonoverlapping, with con- gentina (FML 256[2], 258[5]), approximately spicuous interstitial granules in all four spe- 680 km to the south. Pelusaurus cranwelli cies. The characteristics said to be shared by may be a synonym ofLiolaemus wiegmannii, the species allocated to Abas (Nutn-ez and or if valid, likely its sister taxon. Ya'nez, 1984b) either do not apply to all of them, or are shared with most other species PHR YNOSA URA of Liolaemus. Furthermore, L. fabiani and Werner (1907) described Phrynosaura, and L. insolitus possess a tibial blade and hyper- its only species, P. reichei. Muller (1928) sub- 1995 ETHERIDGE: REDESCRIPTION OF CTENOBLEPHARUS ADSPERSA sequently designated P. reichei as its type spe- that it may be used as a species-group name cies. He also included in the genus P. mar- for L. kingii, L. archeforus Donoso-Barros moratus and his newly described P. werneri, and Cei, 1971, and L. ruizleali Donoso-Bar- a synonym ofLiolaemus anomalus according ros and Cei, 1971 (= L. kingii fide Cei and to Cei (1979a). N(uinez and Y'a-nez (1984b) Scolaro, 1987). Liolaemus kingii and L. ar- restricted the genus to P. reichei and P. au- cheforus have all ofthe synapomorphies that dituvelatus, the latter transferred from Cte- diagnose Liolaemus, and, according to Lau- noblepharys. They cited as characteristics of rent (1985a), differ from other Liolaemus in the genus (1) eyelid with a conspicuous comb, being "primitive." No derived characters are (2) diameter of eye greater than length be- known to unite L. kingii and L. archeforus, tween anterior border of eye and rostral, (3) the species assigned to Rhytidodeira by Lau- tail shorter than snout-vent length, (4) head rent (1985a). scales poorly differentiated, (5) jaws isognathus, (6) loreal region depressed, and (7) dor- VELOSA URA sal scales imbricate, smooth, and with ac- companying scales (my translation). My ob- N(uinez and Yanez (1984b) described Ve- servations of these two species are in accord losaura, including in it L. aymararum Veloso with this list of characters, except that the et al., 1982, and L. jamesi, designating the head scales of both species are as well differ- former as its type species. Earlier, Veloso et entiated as in many species of Liolaemus, al., (1982), proposed the generic name Jar- e.g., supraorbital semicircles are distinct, sep- arancus for these two species, but failed to arated from transversely expanded supraocu- provide a description. Thus, Jararancus is a lar by a distinct arc of circumorbitals, inter- nomen nudum. N(uinez and Ya-nez (1984b) parietal prominent, flanked by a pair oflarger characterized Velosaura as having (1) eyelid parietal scales, etc. Compared with most oth- with a short comb, (2) diameter of eye larger er Liolaemus, the snout is distinctly truncate, than the length from anterior border of eye and the ciliary fringe is very prominent. Lau- to rostral, (3) tail equal to or longer than snout- rent (1984a) included L. reichei, L. auditu- vent length, (4) head scales differentiated, (5) velatus, and L. stolzmanni in Phrynosaura, profile isognathus, (6) loreal region slightly and distinguished the genus from Ctenoble- depressed, and (7) dorsal scales separated, pharys, Liolaemus (sensu stricto), and Ceio- leaving spaces between them, the borders laemus by its differentiated supraoculars and rounded, with slight keels only on some scales. a number of morphometric differences. Lio- My notes on the holotype of L. jamesi laemus reichi and L. audituvelatus possess a (BMNH 90.6.9.2 [RR1946.8.12.39]; see also tibial blade and hypertrophied M. tibialis an- Boulenger [1891, pl. 1]) are in accord with terior, but the condition in L. stolzmanni is these observations except that the doral body unknown. However, the prominent ciliary scales are subimbricate with faint, blunt keels. fringe and truncated snout may prove to be Additionally, both species possess a tibial synapomorphies for the three species placed blade and associated hypertrophied M. tibi- in Phrynosaura by Laurent (1984a). alis anterior, and although data are not available for L. aymararum, L. jamesi also has a hypertrophied M. puboischiotibialis and pig- RHYTIDODEIRA mented epimysium of the M. pterygoman- Girard (1858) proposed the genus Rhyti- dibularis. Laurent (1992) referred both spe- dodeira for six previously described species cies to the signifer group ofthe subgenus Eu- of Liolaemus: Proctotretus kingii Bell, 1842, laemus (see below), but Nuiinez and Jaksic P. magellanicus Dumeril and Dumeril, 1851, (1992) and N'uniez (1992) continued to rec- P. bibronii Bell, 1842, P. wiegmannii, Tro- ognize Velosaura as a valid genus. None of pidurus nigromaculatus Wiegmann, 1834, and the characters listed for L. aymararum and T. oxycephalus Wiegmann, 1834 (= L. ni- L. jamesi are unique to them, although the gromaculatus). Subsequently, Laurent combination itself may be. Evidence for (1 985a) resurrected Rhytidodeira, designated monophyly of Velosaura appears to be weak L. kingii as its type species, and suggested or lacking. Liolaemus aymararum and L. ja- 20 AMERICAN MUSEUM NOVITATES NO. 3142 mesi are extremely similar phenetically, and femoral scales ofLiolaeminae are small, con- may be synonymous, or, if valid, sister spe- vex, and nonoverlapping in most species, but cies. in those with large, strongly imbricate and lanceolate dorsal body scales, e.g., Liolaemus VILCUNIA chiliensis, L. gravenhorstii, L. lemniscatus, and L. nitidus, the postfemorals are flat and Donoso-Barros and Cei (1971) described subimbricate, as in Vilcunia. The distinctly Vilcunia, with a single species, V. sylvanae, tridentate dorsal scales do indeed appear to and indicated that it differed from Liolaemus be unique within Liolaeminae (Cei and Sco- by its tail shorter than snout-vent length, short laro, 1982), and apparently also within Tro- hind limbs, presence of hemigular fold, ab- piduridae, and may well represent a syna- sence of precloacal pores in both sexes and pomorphy for L. sylvanae, L. periglacialis, posterior border of the thigh not granular. A and L. lineomaculatus. However, precloacal second species, V. periglacialis, was added to pores are also lacking in both sexes of Lio- the genus by Cei and Scolaro (1982), who laemus coeruleus (Cei and Ortiz-Zapata, 1983) indicated that the primary character separat- and L. cristiani (Navarro and Ntuniez, 1992), ing Vilcunia from Liolaemus was the pres- and in some males of several other species ence of lateral mucrons on the dorsal scales, (Laurent, 1984a) that do not otherwise bear giving them a "trifid" appearance, but also a close resemblance to Vilcunia. pointed out that Liolaemus lineomaculatus Boulenger, 1885, lacks precloacal pores as well, and sometimes exhibits dorsal scales EULAEMUS AND LIOLAEMUS, slightly notched on the edges (see Cei and SENSU STRICTO Scolaro, 1982: fig. 4; Cei, 1986: fig. 56k-m). Within Liolaemus, Laurent (1983b) rec- On the basis of morphometric studies, Lau- ognized two large species groups that includ- rent (1985a) recognized Vilcunia and trans- ed the majority of species in the genus. He ferred Liolaemus lineomaculatus to it. Vil- referred to them as the Argentino group and cunia was recognized by Etheridge and de the Chileno group. The Chileno group con- Queiroz (1988) based on its lack ofprecloacal tained 37 species, most of them occurring in pores and tridentate dorsal scales, both pre- Chile, with some extending into Argentina, sumed to be synapomorphies, and its pos- Bolivia, and southern Peru. The Argentino session of a shorter dentary, thought to be a group contained 28 species, most of them plesiomorphic state shared with Phymaturus from Argentina, but with some species in (a longer dentary thought to be a synapo- Chile, Bolivia, Paraguay, and Peru. He stated morphy for Liolaemus); however, they in- that ifthese groups eventually were to be rec- cluded in the genus only V. sylvanae and V. ognized at the generic or subgeneric level, the periglacialis. Frost and Etheridge (1989) con- name Eulaemus Girard, 1858 (type species sidered Vilcunia to be a synonym of Liolae- Proctotretusfitzingerii Dumeril and Bibron, mus because, while not doubting the mono- 1837) was available for the Argentino group, phyly of Vilcunia, a short dentary, which and Liolaemus sensu stricto (type species formed the basis for its exclusion from Lio- Calotes chiliensis Lesson, 1830) should be laemus, was found to be variable within the used for the Chileno group. These groupings genus, and because all preliminary analyses were based primarily on his morphometric of liolaemine relationships (Etheridge, un- studies, which he summarized two years later publ.) had found Vilcunia nested well within (Laurent, 1985a), and proposed the formal Liolaemus. Of the characters listed for Vil- recognition ofEulaemus and Liolaemus sen- cunia by Donoso-Barros and Cei (1971), all su stricto as subgenera. but the tridentate dorsal scales and lack of In 1992, Laurent characterized the two precloacal pores occur in a number of other subgenera and listed the species assigned to species of Liolaemus. The "hemigular fold" each. Liolaemus (48 species) was character- presumably refers to the lateral vestiges of ized as having fewer precloacal pores (x = the medially interrupted transverse gular fold, 2.19 and < 5 in 91% ofspecimens Liolaemus found in almost all Liolaeminae. The post- versus x = 6.40 and > 4 in 92% ofspecimens 1995 ETHERIDGE: REDESCRIPTION OF CTENOBLEPHAR US ADSPERSA ofEulaemus), lateral rather than dorsolateral Liolaemus sensu stricto but in Eulaemus it nostrils, generally flat and long supralabials, usually is anteroposteriorly expanded, often the fourth below the eye with an oblique bor- with an irregular posterior margin, and oc- der, rather than short, high supralabials, fifth- casionally fenestrate. The latter condition oc- seventh below the eye, with posterior border curs in Ctenoblepharys but not Phymaturus vertical. The supralabials of Liolaemus usu- or other Liolaemus sensu lato. Meckel's ally number only four, all of them slender groove is fused in Liolaemus sensu stricto, and at least twice as long as wide, the fourth unfused in Eulaemus except in about 60% of usually the most elongate, more than three L. darwinii Bell, 1843 (sensu Etheridge, 1993). times longer than wide, with its posterior The polarity ofthis character is uncertain be- margin oblique, whereas in Eulaemus the su- cause of interspecific variation in other Lio- pralabials characteristically number five or laemus (sensu lato) and Phymaturus. In some more, the anterior ones square or not much species of both groups the body scales are longer than wide, none of them more than moderately small, rhomboidal, and imbri- three times longer than wide, and the pos- cate, with granular lateral nuchal scales and terior one with its posterior margin nearly complex lateral nuchal skin folds; however, vertical. Laurent (1992) also listed the dis- within Liolaemus sensu stricto, most species tance between the upper border of the sub- exhibit varying degrees of more sharply ocular and lip as distinct: less than the dis- keeled, strongly imbricate, lanceolate dorsal tance between the nasal plates in Liolaemus, body scales, and larger, keeled, imbricate lat- greater in Eulaemus. However, this differ- eral nuchal scales accompanied by the re- ence results from the combined effects oftwo duction or disappearance of lateral nuchal previously mentioned characters, i.e., the na- skin folds. In contrast, within Eulaemus many sals being lateral, and therefore closer to- species exhibit, to varying degrees, dorsal gether, and the supralabials narrower in Lio- body scales that are rounded or oval, sub- laemus than in Eulaemus. imbricate to nonoverlapping, often with con- As indicated earlier, a large number ofLio- spicuous interstitial granules, and the lateral laemus (sensu lato) possess melanic pigment nuchal scales are granular with prominent skin within the median portion of the epimysium folds. Other species of Liolaemus sensu lato of the M. pterygomandibularis, the medial exhibit interspecific variation in these char- head of the M. flexor tibialis internus is cov- acteristics of the dorsal body and lateral nu- ered by an hypertrophied M. puboischioti- chal scales. bialis, and the insertion ofthe M. tibialis an- In the same work, Laurent (1992) recog- terior is hypertrophied in association with the nized within Eulaemus a fitzingerii group, presence of a sharp, bladelike process of the characterized by the presence of a patch of tibia. These characters, all unique within enlarged scales on the posterior thigh, and a Tropiduridae, Phrynosomatidae, and Oplur- signifer group characterized by its absence. idae, and apparently also unique within Ig- The femoral patch referred to as character- uania, are present in the 35 species listed for istic of the fitzingerii group is a patch of Eulaemus by Laurent (1992), save for L. cha- abruptly enlarged, often spiny scales on the coensis Shreve, 1948, which lacks them all. posteriorproximal surface ofthe thigh, which They are also present in all species of Ortho- is otherwise beset with small, nonoverlap- laemus (see below). The alternative, and pre- ping scales. It is present in both sexes, and sumably plesiomorphic states, of these mus- in adult males the patch forms a hemispher- culoskeletal characters occur in the 24 species ical bulge due to even greater hypertrophy of listed for Liolaemus sensu stricto, except L. the proximal part ofthe M. puboischiotibialis duellmani Cei, 1978, which has them. The (Etheridge, 1993: pl. 1.3). As Laurent (1992) plesiomorphic state is present in all other pointed out, the patch of enlarged scales is Liolaeminae as well. unique within Tropiduridae, and it occurs Other differences between the subgenera elsewhere within Iguania only in the phry- appear as trends or tendencies, the polarities nosomatid genus Uma (in which the thigh of which are uncertain. The proximal (inter- muscles are normal) and is almost certainly clavicle) process of the clavicle is slender in derived within Liolaeminae. 22 AMERICAN MUSEUM NOVITATES NO. 3142

A femoral patch, underlain by a hypertro- er iguanians except for a few Agaminae, and phied M. puboischiotibialis, that character- in other Tropiduridae Meckel's groove is izes the fitzingerii group, as well as the mus- fused and the labials are narrow. culoskeletal characteristics of other Eulae- mus, also occur in the species assigned by MESOLAEMUS Laurent (1983a) to the subgenus Ortholae- mus (see below). As indicated above, the In a brief note, Laurent (1985b) proposed musculoskeletal characters that distinguish the new subgeneric name Mesolaemus for Eulaemus from Liolaemus sensu stricto, i.e., Liolaemus cuyanus Cei and Scolaro, 1980. pigmented epimysium of the M. pterygo- The reason stated for this action was that L. mandibularis, hypertrophy of M. pubois- cuyanus "is similar to Ortholaemus in one chiotibialis, and presence of a tibial blade important character, but also is similar to associated with a hypertrophied M. tibialis Eulaemus in another, no less important char- anterior, appear to be unique within Iguania, acter" (my translation). He further stated that and provide evidence for the monophyly of "the data considered does not permit one to a clade composed of Eulaemus + Ortholae- decide if Mesolaemus is the sister group of mus. Furthermore, the femoral patch with an Ortholaemus, of Eulaemus, or of a part of underlying hypertrophied thigh muscle also Eulaemus .. ." (my translation). The char- appears to be unique within Iguania, and may acters referred to, and the data considered, be considered a potential synapomorphy of were not specified. a clade formed by Ortholaemus + the fitzin- Liolaemus cuyanus was initially described gerii group of Eulaemus. However, the rec- as a subspecies ofL. fitzingerii (Cei and Sco- ognition of Eulaemus and Ortholaemus as laro, 1980), and raised to full species status subgenera would render Eulaemus paraphy- by Laurent (1983b). Presumably, the char- letic. Indeed, Laurent (1 983a) considered Or- acter referred to by Laurent (1 985b) as a sim- tholaemus obviously to have been derived ilarity between L. cuyanus and Orytholae- from the Argentino group, e.g., Eulaemus. mus, is the contact of the mental with six Thus, without Ortholaemus as a subset of scales, i.e., the sublabials in addition to the Eulaemus there are no known synapomor- anterior infralabials and postmentals. L. cuy- phies that would unite the species assigned anus is otherwise a typical member of the to Eulaemus by Laurent (1992). fitzingerii group of Laurent (1992), and is Monophyly of the subgenus Liolaemus is phenetically very similar to L. fitzingerii, L. more problematic. Ofthe characters that dis- canqueli, L. melanops, and L. xanthoviridis, tinguish Liolaemus from Eulaemus, the ab- but it is unlike any species of Ortholaemus sence of a hypertrophied M. puboischioti- in its squamation, large body size (maximum bialis, tibial blade and hypertrophied M. ti- SVL 98 mm [Etheridge, 1992]), and expand- bialis anterior, and pigmented M. pterygo- ed, deeply tricuspid posterior marginal teeth. mandibularis epimysium are clearly It is uncertain, as Laurent (1985b) pointed plesiomorphic. Several characteristics of the out, whether the sublabial-mental contact is group probably are derived within Liolae- a synapomorphy for L. cuyanus + Ortholae- minae: the presence of a fused Meckel's mus, or is homoplastic. However, recogni- groove; supralabials narrow, the fourth the tion ofMesolaemus as a monotypic subgenus most posterior and greatly elongated; and of Liolaemus seems unwarranted. precloacal pores usually four or fewer (or absent). However, outgroup comparisons in- ORTHOLAEMUS dicate that the polarity of these states is equivocal. Precloacal pore numbers are high Cei (1979b) pointed out that Liolaemus ra- (5-15) in Eulaemus, Ortholaemus, Cteno- binoi, L. multimaculatus, and L. riojanus (as blepharys, and Phymaturus. Meckel's groove L. multimaculatus riojanus; see Etheridge, is open in Eulaemus (except in some Liolae- 1993) "appear to represent a very specialized mus darwinii), Ortholaemus (except Liolae- group ofpsammophilous lizards," and stated mus occipitalus [Keller and Krause, 1986]), that should the group "be accorded special, Ctenoblepharys, and the Phymaturuspaluma formal recognition, the subgeneric name Or- group, but precloacal pores are lacking in oth- tholaemus (Girard 1858; type species Ortho- 1995 ETHERIDGE: REDESCRIPTION OF CTENOBLEPHARUS ADSPERSA 23 laemus beaglii Girard 1858 = Liolaemus referred to Abas, Ceiolaemus, Eulaemus, multimaculatus) is available." This appears Mesolaemus, Ortholaemus, Pelusaurus, to be the first formal designation of a type Phrynosaura, Rhytidodeira, Velosaura, and species for Ortholaemus. Laurent (1983a) Vilcunia exhibit all of the synapomorphies added L. scapularis, L. lutzae, L. occipitalis, that distinguish Liolaemus from other Lio- L. wiegmannii and L. cranwelli to Ortholae- laeminae. Furthermore, there is some evi- mus, and characterized the group as having dence that Vilcunia, Ceiolaemus, and Phry- (1) more than one row of scales between the nosaura may represent monophyletic groups subocular and supralabials (although in L. within Liolaemus. There also is evidence that scapularis the subocular is often separated Ortholaemus is a monophyletic group, which, from the labials by a single scale), (2) mental together with the the other species that pos- in contact with six (at times eight) scales in- sess a femoral patch (i.e., fitzingerii group), stead offour, (3) mental much narrower than forms a more inclusive clade. This clade, to- rostral, (4) mental wider posteriorly than at gether with Eulaemus, forms a still more in- the border with the lip, and (5) claws longer clusive clade (see Appendix 2). and yellowish in the arenicolous species. He also pointed out that in L. cuyanus Cei and Scolaro, 1980, the rostral is bordered by six DISCUSSION AND SUMMARY scales, but that it is not narrower posteriorly, Ctenoblepharys adspersa is a moderate-size and there is a single row of scales between (maximum SVL 74 mm) iguanian lizard en- the subocular and supralabials. Laurent demic to the sandy beaches and sand dunes (1984a) formally proposed the recognition of of the Pacific coast of central Peru between Ortholaemus as a subgenus ofLiolaemus, and about 11°05' and 13°50'S. Little is known of added to it L. rabinoi and L. riojanus. He its habits except that it runs swiftly over the also designated Ortholaemusfitzroyii Girard sand and that its color pattern is highly cryp- 1858 = (Proctotretus wiegmannii Dumeril and tic on this substrate. It possesses a number Bibron 1837), as the type species, apparently of derived characteristics that are unique having overlooked Cei's (1979b) earlier des- within Tropiduridae and its putative out- ignation ofLiolaemus multimaculatus as the groups Phrynosomatidae and Opluridae; six type species. Later, Laurent (1986) described of these autapomorphies involve the skull, Liolaemus salinicola as a new species in the and two the vertebral column. Additionally, subgenus Ortholaemus. C. adspersa differs from most other Liolae- As indicated above, the species of Ortho- minae in having small, nonoverlapping dor- laemus have all ofthe apomorphic states that sal body scales arranged in more-or-less diagnose the fitzingerdi group of Eulaemus, transverse rows, and numerous, poorly dif- i.e., melanic pigment within the epimysium ferentiated head scales, including small, sub- of the M. pterygomandibularis, the medial equal supraoculars and several suboculars, head ofthe M. flexor tibialis internus covered but with greatly elongate, triangular outer cil- by an hypertrophied M. puboischiotibialis, iaries that form a prominent comb. Cte- and the insertion of the M. tibialis anterior noblepharys adspersa, along with Phymatu- hypertrophied in association with the pres- rus, lacks the synapomorphies that specify a ence ofa sharp, bladelike process ofthe tibia. clade consisting ofall other Liolaeminae, the Additionally, they differ from all other Lio- species of which are here referred to Liolae- laemus in having smaller lorilabial scales, mus. Furthermore, C. adspersa and Phy- usually in two or more rows between the sub- maturus are not known to share any derived ocular and supralabials, flat or concave in- characteristics to the exclusion ofLiolaemus, fralabials (personal obs.), and the mental scale nor does Liolaemus share any derived char- narrower anteriorly than posteriorly, and from acteristics with either Ctenonlepharys or Phy- all other Liolaemus except L. cuyanus in hav- maturus to the exclusion of the other, and ing six, rather than four scales in contact with thus relationships of Ctenoblepharys, Phy- the mental, resulting from contact between maturus, and Liolaemus are unresolved. the mental and sublabials. Thus, Ortholae- All ofthe species ofLiolaeminae that have mus may well be a monophyletic group. been described in or referred to Ctenoble- In summary, the species that have been pharys, with the possible exception of L. er- 24 AMERICAN MUSEUM NOVITATES NO. 3142 roneus, lack the autapomorphies of C. ad- (KU); P. Alberch and J. Rosado, Museum of spersa, and furthermore all of them possess Comparative Zoology, Harvard University, the synapomorphies that diagnose Liolae- Cambridge (MCZ); H. Greene, Museum of mus. This is also true for those species that Vertebrate Zoology, University ofCalifornia, have been described in, or referred to, other Berkeley (MVZ); J. Eiselt, Naturhistorisches genera of Liolaeminae, or as subgenera and Museum, Wien (NMW); K. Klemmer and G. species groups of Liolaemus. Evidence for Kohler, Natur-Museum Senckenberg, Frank- monophyly of some of these groups is lack- furt-am-Main (SMF), G. Zug and R. Heyer, ing, while others may well represent clades United States National Museum of Natural within Liolaemus. However, until such time History, Washington D.C. (USNM), J. Pe- as the historical relationships of the species faur, Universidad de Los Andes (CV-ULA), of Liolaemus have been determined by cla- and H. N'uinez, Museo Nacional de Historia distic analysis, it would seem to be the wisest Natural de Chile (MNHNC) and Departa- course to avoid using these names as formally mento de Biologia Celular y Genetica, Univ- recognized taxa. To do so could lead to the ersidad de Chile (DEGUCH). Thanks also go formation ofa paraphyletic group formed by to John Wright for permission to prepare a the remaining species that then would com- skeleton of Ctenoblepharys adspersa in the pose the genus Liolaemus. LACM collection, to James R. Dixon for providing me with a color photograph and notes on a living individual of C. adspersa, and to ACKNOWLEDGMENTS Herman N(uinez for permission to dissect the For permission to examine specimens in hind limbs and jaw muscles of Liolaemus their care, I am grateful to C. Myers and D. audituvelatus and L. fabiani, and for infor- Frost, American Museum of Natural Histo- mation concerning the presence of a tibial ry, New York (AMNH); R. Laurent, Fun- blade in L. reichei. Thanks also go to Brad- dacion Miguel Lillo, San Miguel de Tucuman ford Hollingsworth for preparation of figures (FML); A. Leviton and J. Vindum, California 1 and 2, to Roland Sosa and Fernando Lobo Academy of Sciences, San Francisco (CAS); for providing the Spanish translation of the H. Marx, Field Museum of Natural History, abstract, and to Darrel Frost, Jimmy Mc- Chicago (FMNH); G. Macola and L. P. Cas- Guire, Bradford Hollingsworth, Lee Gris- tro, Instituto de Biologia, Universidad Na- mer, and John Wiens, who provided helpful cional de Cuyo, Mendoza (IBUNC); W. E. suggestions for improvement of the manu- Duellman, University of Kansas, Lawrence script.

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Lamborot, M., and J. C. Ortiz-Zapata Maddison, W. P., M. J. Donoghue, and D. R. 1990. Liolaemus pseudoanomalus, una nueva Maddison especie de lagarto del Norte Chico de 1984. Outgroup analysis and parsimony. Syst. Chile. Gayana Zool. 54(3-4): 135-142. Zool. 33(1): 83-103. Lataste. F. Marx, H. 1892. Etudes sur la faune chilienne. I. Note 1960. A new iguanid lizard ofthe genus Cten- sur les l¢zards. Actas Soc. Cien. Chili 1: oblepharis. FieldianaZool. 39(37): 407- 7-40. 409. Laurent, R. F. Mertens, R. 1982. Description de trois especes nouvellae 1938. Bemerkungen iuber die brasilianischen du genere Liolaemus (Sauria, Iguani- Arten der Gattung Liolaemus. Zool. dae). Spixiana 5(2): 139-147. Anz. 123(7/9): 220-222. 1983a. Contribucion al conocimiento de la es- 1956. Biologische Reiseskizzen aus Peru. II. tructura taxonomica del genero Liolae- In den Wusten Peru. Natur und Folk, mus Wiegmann (Iguanidae). Bol. Asoc. Senckenb. Natur. Ges. Frankfurt 86: Herpetol. Argentinas 1(3): 16-18. 318-327. 1983b. Sinonimia del genero Pelusaurus Don- Montanucci, R. R. oso-Barros con Liolaemus (Sauria, 1987. A phylogenetic study of the horned liz- Iguanidae). Ibid. 1(1): 9-10. ards, genus Phrynosoma, based on skel- 1984a. On some iguanid genera related to or etal and external morphology. Contrib. previously confused with Liolaemus Sci. Nat. Hist. Mus. Los Angeles 390: Wiegmann. J. Herpetol. 18(4): 357-373. 1-36. 1984b. Fenogramas de algunas representivas del Muller, L. genero Liolaemus y generos vecino (Iguanidae, Reptilia). Acta Zool. Lil- 1928. Herpetologische Mitteilungen. Zool. loana 38(1): 5-17. Anz. 77: 61-84. 1985a. Segunda contribucion al conocimiento Muller, L., and W. Hellmich de la estructura taxonomica del genero 1932a. Beitriige zur Kenntnis der Herpetofauna Liolaemus Wiegmann (Iguanidae). Chiles. III. Liolaemus altissimus altis- Cuad. Herpetol., Asoc. Herpetol. Ar- simus, Liolaemus altissimus araucan- gentina 1(6): 1-37. iensis. Zool. Anz. 98(7-8): 197-208. 1985b. Relaciones entre algunos taxa tradicion- 1932b. Beitriige zur Kenntnis der Herpetofauna almente incluidos en el genero Liolae- Chiles. II. Neue Liolaemus -Arten und mus Wiegmann (Iguanidae). Bol. Asoc. Rassen aus den Hoch-Anden Chiles. Herpetol. Argentina 2(4): 3. Ibid. 97(11-12): 307-329. 1986. Descripciones de nuevos Iguanidae del 1939a. Liolaemus -Arten aus den Westlichen genero Liolaemus. Acta Zool. Lilloana Argentinien. II. Ueber eine neue Lio- 38(2): 87-105. laemus altissimus Rasse von Volcan 1992. On some overlooked species of the ge- Copahue. Ibid. 125(5-6): 113-119. nus Liolaemus Wiegmann (Reptilia 1939b. Liolaemus -Arten aus den westlichen Tropiduriudae) from Peru. Breviora Argentinien. IV. Ueber Liolaemus -Ar- 494: 1-33. ten aus dem Territorien Rio Negro und Lesson, R. P. Neuquen. Ibid. 128(1-2): 1-17. 1830. Observations generales sur les reptile re- Navarro, J., and H. NCuinez cueillis dan la voyage de la corvett La 1992. Acera de la ausencia de poros precloa- Coquille. In M. L. I. Duperry (ed.), Voy- cales en Liolaemus cristiani, nomina- age autour du monde execute par ordre cion del alotipo y cariotypo de la es- du Roi sur la corvette de Sa Majeste La pecie. Not. Mensual Mus. Nac. Hist. Coquille pendant les annees 1822, 1823, Nat. Chile 323: 35-38 1824 et 1845. Zoologie, Reptiles 1-66. 1993. Liolaemus patriciaiturrae y Liolaemus Paris: Arthus Berrand. isabelae, dos nuevos especies de lagar- Leviton, A. E., R. H. Gibbs, Jr., E. Heal, and C. tijas para el norte de Chile. Aspectos E. Dawson biogeograficos y citotaxonomicos 1985. Standards in herpetology and ichthy- (Squamata, Tropiduridae). Bol. Mus. ology: Part I. Standard symbolic codes Nac. Hist. Nat. Chile 44: 99-113. for institutional resource collections in Nfunez, H. herpetology and ichthyology. Copeia 1992. Geographical data ofChilean lizards and 1985(3): 802-832. snakes in the Museo Nacional de His- 28 AMERICAN MUSEUM NOVITATES NO. 3142

toria Natural Santiago, Chile. Smith- Peru. Mem. Mus. Hist. Nat. "Javier son. Herpetol. Inf. Serv. 91: 1-29. Prado" 18: 1-97 Niuniez, H., and F. Jaksic Pereyra, E. A. 1992. Lista comentada de los reptiles ter- 1985. Nuevo iguanido del genero Phymaturus restres de Chile continental. Bol. Mus. del noroeste Argentino. Bol. Asoc. Her- Nac. Hist. Nat. Chile 43: 63-91. petol. Argentina 2(4): 4. Nufnez, H., and J. Navarro 1991. Phymturus antofagastensis (Pereyra 1992. Liolaemus rosenmanni, una nueva es- 1985) (Tropiduridae, Liolaeminae): pecie Chilena de lagartija relacionada al amplicacion descriptiva. Ibid. 7(2): 24- groupo "ruibali." Bol. Mus. Nac. Hist. 27. Nat. Chile 43: 55-62. 1992. Nueva especie de lagarto andino: Lio- N(uinez, H., J. Navarro, and J. Loyola laemus vallecurensis (Tropiduridae, 1991. Liolaemus maldonadae y Liolaemus Liolaeminae). Not. Mensual Mus. Nac. cristiani, dos especies nuevas de lagar- Hist. Nat. Chile 321: 10-14. tijas para Chile (Reptilia, Squamata). Peters, J. A., and R. Donoso-Barros Bol. Mus. Nac. Hist. Nat. Chile 42: 79- 1970. Catalogue ofthe Neotropical Squamata: 88. Part II. Lizards and amphisbaenians. U.S. Natl. Mus. Bull. 297: viii + 293 and N(uinez, H., J. Ya-nez PP. 1983. Ctenoblepharis audituvelatus new spe- Philippi, R. A. cies, a lizard from northern Chile (Rep- 1860. Reise durch die Wiiste Atacama, aufBe- tilia: Iguanidae). Copeia 1983(2): 454- fehl der chilenischen Regierung in Som- 457. mer 1853-1854. Halle: Eduard Anton 1984a. Ctenoblepharis erroneus nov. sp. de ix + 192 pp. Iguanidae para la Zona Norte de Chile. Pregill, G. K. Bol. Mus. Nac. Hist. Nat. Chile 40: 91- 1992. Systematics of the West Indian lizard 95. genus Leiocephalus (Squamata: Iguania: 1984b. Abas y Velosaura, nuevos generos de Tropiduridae). Misc. Publ. Mus. Nat. lagartos iguanidae y proposiciones sis- Hist. Univ. Kansas 84: 1-69. tematicas respecto de los generos alia- Shreve, B. dos (Reptilia: Squamata). Ibid. 40: 97- 1948. A new Liolaemus from Paraguay. Co- 105. peia 1948(2): 111-112. Oelrich, T. M. Steindachner, F. 1956. The anatomy ofthe head of Ctenosaura 1891. Uber einige neue und seltene Reptilien- pectinata (Iguanidae). Univ. Michigan und Amphibien-Arten. Sitzber. Akad. Mus. Zool. Misc. Publ. 94: 1-122. Wiss. Wien 100(1): 291-316. Ortiz-Zapata, J. C. Smith, H. M. 1989a. Catalogue des types du Musee d'His- 1946. Handbook ofLizards. Ithaca, New York: toire Naturelle de Neuchatel. III. Sau- Comstock xxi + 557 pp. riens. Bull. Soc. Neuchateloise Sci. Nat. Tschudi, J. J. von 112: 47-64. 1845. Reptilium conspectum quae in Repub- 1989b. Descripton de Liolaemus silvai sp. nov. lica Peruana reperiuntur et pleraque ob- (Sauria, Iguanidae) du "Nord Chico" du servata vel collecta sunt in itinere a Dr. Chile. Bull. Mus. Nac. Hist. Nat. Paris, J. D de Tschudi. Arch. Naturgeschichte 4e ser, sec. A(l): 247-252 11(1): 150-168. Ortiz-Zapata, J. C., and P. Marquet 1846. Untersuchungen iuber die Fauna Per- 1987. Una nueva especie de lagarto altoan- uana. Herpetologie. St. Gallen: Sceitlin dino: Liolaemus isulgensis (Reptilia- und Zollikofer, 80 pp. Iguanidae). Gayana Zool. 51(1-4): 59- Urbina, M., and 0. Zunigia 63. 1977. Liolaemuspictus talcanensis nov. subsp. Ortiz-Zapata, J. C., and H. Nuiiez (Squamata, Iguanidae), nuevo reptil para 1986. Catalogo critico de los tipos de reptiles el Archipi6lago de Chiloe. An. Mus. Hist. conservados en el Museo Nacional de Nat. Valparaiso 10: 69-74. Historia Natural Santiago. Publ. Ocas. Veloso, A., and J. Navarro Mus. Nac. Hist. Nat. Chile 43: 1-23. 1988. Lista sistematica y distribucion geo- Pearson, 0. P., and C. P. Ralph grafica de anfibios y reptiles de Chile. 1978. The diversity and abundance of verte- Boll. Mus. Reg. Sci. Natur. Torino 6(2): brates along an altitudinal gradient in 481-539. 1995 ETHERIDGE: REDESCRIPTION OF CTENOBLEPHARUS ADSPERSA 29

Veloso, A., M. Sallaberry, J. Navarro, P. Iturra, Wiegmann, A. F. A. J. Valencia, M. Penna, and N. Diaz 1834. Beitriige zur Zoologie gesammelt aufei- 1982. Contribucion sistematica al conoci- ner Reise um die Erde, von Dr. F. J. F. miento de la herpetofauna del extremo Meyen. Siebente Abhandlung. Amphi- norte de Chile. In La vegetacion y los bien. Nova Acta Acad. Caesar. Leop.- vertebrados inferiores y los pisos alti- Carol. 17(1): 184-268d. tudinales entre Arica y Lago Chungara, Wiens, J. J. pp. 135-268. Vol. sintesis Proy. MAB- 1993a. Phylogenetic relationships of phryno- 6-UNEP-UNESCO-1 105-77-0 1. Mon- somatid lizards and monophyly of the tevideo. Sceloporus group. Copeia 1993(2): 287- Watrous, L. E., and Q. D. Wheeler 299. 1981. The outgroup comparison method of 1993b. Phylogenetic systematics ofthe tree liz- character analysis. Syst. Zool. 30(1): 1- ards (genus Urosaurus). Herpetologica 11. 49(4): 399-420. Werner, F. Wiley, E. 0. 1904. Ergebinsse der Hamburger Magalhaen- 1979. An annotated Linnean hiearchy, with ischen Sammelreise. I. Band. Allgeen- comments on natural taxa and compet- ines, Chordonier, Echinodermen und ing systems. Syst. Zool. 28(3): 309-337. Coleleneraten. IV. Reptilien und Batra- Ya-nez, J. L., and H. Niuinez chier. Hamburg: L. Friedrichsen 21 pp. 1983. Liolaemusfabiani, a new species ofliz- 1907. In 0. Burger (ed.), Estudios sobre rep- ard from northern Chile (Reptilia: Igua- tiles chilenos. An. Univ. Chile 121(2): nidae). Copeia 1983(3): 788-790. 147-155.

APPENDIX 1 SPECIMENS EXAMINED Preserved specimens and skeletons of Cteno- gentinus (Muller and Hellmich, 1939b), L. p. ma- blepharys adspersa examined are listed below, fol- jor (Boulenger, 1885), L. p. talcanensis (Urbina lowed by their localities. Preserved specimens and and Zunigia, 1977), L. pseudolemniscatus (Lam- skeletons ofPhymaturus and Liolaemus examined borot and Ortiz-Zapata, 1990), L. rosenmanni for comparison with Ctenoblepharys adspersa, and (Nu(nfez and Navarro 1992), L. silvai (Ortiz-Za- specimens ofoutgroup species examined for com- pata, 1989b), L. vallecurensis (Pereyra, 1992), L. parison with Liolaeminae, are also listed below. zapallarensis sieversi (Donoso-Barros, 1954), Phy- Museum acronyms follow Leviton et al. (1985). maturus mallimaccii (Cei, 1980b), P. indistinctus Numbers preceded by REE are all skeletons; skel- (Cei and Castro, 1973), P. nevadoi (Cei and Roig, etons preceded by other acronyms are indicated 1975). by "sk." Species unavailable for examination, for which Liolaeminae some data were obtained from the literature, include: Liolaemus anomalus ditadai (Cei, 1993), L. Ctenoblepharys adspersa: Peru: Ventanilla, near archeforus gallardoi (Cei, 1986), L. a. sarmientoi Lima, SMF 75966-69; between Ventanilla and (Cei, 1986), L. aymararum (Veloso et al., 1982), Puenta Piedras, SMF 64373-74; Playa Ventanilla L. belli araucaniensis (Muller & Hellmich, 1939a), near Lima, REE 2513; near Ancon north ofLima, L. b. neuquensis (Muller & Hellmich, 1932a), L. SMF 64369-70; 5.5 km NE San Bartolo, 100 m, ceii (Cei, 1986), L. cristiani (Nu(niez et al., 1991), MVZ 85415-16; Ciudad de Dios, FML 0368,0464; L. cyanogaster brattstroemi (Donoso-Barros, Ica: beach south of Paracas, within 100-200 m of 1961), L. erroneus (Nuniez and Ya-nez, 1984b), L. ocean, MCZ 145039-40; Museo Paracas, 30.2 km hellmichi (Donoso-Barros, 1975), L. isabelae (Na- S Pisco, 7.2 km SW Paracas, LACM 49145-6, varro and NuOnez, 1993), L. lativittatus (Werner, 49147(sk); Peru (no additional data), NMW 13578, 1904), L. islugensis (Ortiz-Zapata and Marquet, 18905. 1987), L. maldonadae (Nuinhez et al., 1991), L. Liolaemus abaucan: SDSU 1446-64, REE 2845. melanopleurus (Philippi, 1860), L. modestus (Phi- L. alticolor: SDSU 1697-707, REE 2520, 2641- lippi, 1860), L. nigroviridis minor (Miiller and 50. L. andinus andinus: FML 1764(10). L. a. poe- Hellmich, 1932b), L. n. nigroroseus (Donoso-Bar- cilochromus: SDSU 1597-99, REE 2548, 2670- ros, 1966), L. ortizi (Laurent, 1982), L. patriciai- 78. L. anomalus anomalus: SDSU 1674-75. L. turrae (Navarro and NCuinez, 1993), L. pictus ar- archeforus archeforus: MCZ 162002-03, 30 AMERICAN MUSEUM NOVITATES NO. 3142

164001(sk). L. audituvelatus: MNHNC 980-81. lemniscatus: SDSU 1882-86, 2079-80, REE 2530, L. austromendocinus: SDSU 1799-80, REE 2340- 2889-91. L. leopardinus: MVZ 187765, 187767. 47, 2358-59. L. belli belli: SDSU 1802, REE 1559. L. lineomaculatus: SDSU 1666-68, REE 2463- L. b. moradoensis: SDSU 1803-04. L. bibronii: 69. L. lutzae: SDSU 1187,2047-51, MCZ 46241- SDSU 1805-13, REE 2305,2351-55,2380,2406- 42,46963, 79136-40,92772-74, 119880, AMNH 09, 2461. L. bisignatus: SDSU 1814, REE 2535. 70922-24, 92870-72, FML 1287(7), REE 2524, L. bitaeneatus: SDSU 1815-17, REE 2597-600. 2860, CAS 15802(sk). L. magellanicus: SDSU L. boulengeri: SDSU 1212-16, MVZ 125722-26, 1673, MVZ 180134-35,180136(sk), 180118, REE 125728, 125730-48, 125750-57. AMNH 17022, 2517. L. melanops: SDSU 1178, KU 182045-48, 46431, 95960, REE 2348-50, 2396-97, 2404-05, FML 1609(2), REE 2411. L. montanus: SDSU 2458-60. L. buergeri: MVZ 188691, 188716. L. 1621, FML 1723(9), REE 2540. L. monticola canqueli: KU 182038, 187513, FML 795, 1607, monticola: SDSU 1888-96, REE 2527. L. m. chil- 2115, 2786(3), 2874(7+ 1 sk), 2915, IBAUNC lanensis: SDSU 1898, REE 2596. L. m.villaricensis: 861-9. L. capillitas: SDSU 1818-22. L. chacoen- SDSU 1897, REE 2566. L. multicolor: SDSU sis: SDSU 1823-25, FMNH 44162-70, REE 2549. 1314-19, 1520-32, REE 2547, 2634-40. L. mul- L. chiliensis: SDSU 1837-42, REE 2515. L. coe- timaculatus: SDSU 1181-1185, 1312, 1756-79, ruleus: SDSU 1826-32, REE 2545. L. constanzae: FML 1596(20), 1826(18), REE 2550, 2846. L. ni- SDSU 1834-45, REE 2516. L. copiapensis: SDSU griceps: SDSU 1620, FML 1635(10), AMNH 1846-47, REE 2560, 2765, 2833. L. cranwelli: 131845-46, REE 2537. L. nigromaculatus: SDSU MACN 15.233. L. curicensis: SDSU 1848. L. cur- 1903-05, REE 2551, 2832. L. nigroviridis nigro- is: SDSU 1849-51, REE 2564. L. cuyanus: SDSU viridis: SDSU 1906-08, REE 2558. L. n. cam- 1010, 1051-55, 1089-95, 1229-31, 1465-508, panae: SDSU 1908, REE 2595. L. nitidus: SDSU FML 2097(24), REE 2316-20, 2723-2727. L. 1899-1902, REE 2519, 2834, 2841. L. occipitalis: cyanogaster: SDSU 1833-36, REE 2525. L. dar- SDSU 1186, 2052-57, MCZ 96034, 154189-90, winii: SDSU 1249-50, 1443-45, REE 2321-22, KU 176526-31, CAS 87093-94, REE 2521, 2859. 2493-95. L. disjunctus: FML 1201(5). L. dono- L. olongasta: SDSU 1099-104, 1424-30, 1708- sobarrosi: SDSU 1188, FML 2687(2), 2770(3), 17, REE 2844. L. orientalis orientalis: AMNH 2871(5 + lsk). L. donosoi: FML 1340. L. dorbig- 80076. L. o. chlorostictus: SDSU 1613-1619, REE nyi: SDSU 1625-29, FML 1757(17), REE 2541, 2653-55, 2685-96. L. ornatus: SDSU 1232-48, 2668-69, 2679-84. L. duellmani: KU 161126. L. 1320-38, AMNH 131849, KU 160034-35, eleodori: SDSU 1609-12, REE 2368-79. L. elon- 160210-15, 161182, 183459-68, REE 2522, 2623- gatus elongatus: SDSU 1857-61, REE 2291-96, 33. L. paulinae: SDSU 1910-11, REE 2561. L. 2366-67,2386-89. L. e. petrophilus: SDSU 1862- periglacialis: SDSU 1677, MCZ 162007,162009- 64, REE 2423-32. L. exploratorum: MLP 571. L. 10, 162008(sk). L. pictus pictus: SDSU 1912-15, fabiani: DBGUCH 0350, 0336. L. famatinae: REE 1874, 1890, 1894-99, 2704, 2842. L. p. chil- SDSU 1624, FML 1720(14), REE 2542. L. fitt- oeensis: SDSU 1916-18, REE 2592. L. platei: kaui: FML 1612. L.fitzgeraldi: SDSU 1865, MVZ SDSU 1919-23, REE 2523. L. polystictus: FML 186557-58. L. fitzingerii: SDSU 1191-93, MVZ 1683(2). L. pseudoanomalus: SDSU 1040, 1310- 181615-18, 188785-94, KU 182043-44, FML 11, 1676, REE 2728-29. L pulcherrimus: FML 2128(13), 2130(4), REE 2457. L. forsteri: FML 2184(7), 1961. L. quilmes: SDSU 1013-16, 1021, 2211. L. fuscus: SDSU 1866-68, REE 2529. L. 1058-59, 1251-73, 1340-408, 1533, REE 2578- gracilis: SDSU 1869. L. gravenhorstii: SDSU 1870- 86, 2707-08. L. rabinoi: IBAUNC 818.1-2, 1296. 72, 2081-83, REE 2528, 2831, 2892-95. L. gri- L. ramonensis: SDSU 1887. L. reichei: LACM seus: FrML 1354(4), 1502, 1586(4). L. hernani: 9312. L. riojanus: SDSU 1 105-13, 1146-58, 1169- SDSU 1874-76, REE 2565. L. huacahuasicus: 74, KU 182057, MLP 2730, 2734, 2636, 2752, SDSU 1623, FML 2303(4),2246(3),2297(3), REE REE 2532, 2730-37. L. robertmertensi: SDSU 2543. L. insolitus: CV-ULA IV-0641-49, 2434- 1313, 1518-19, 2587-88. L. robustus: FML 36. L. irregularis: SDSU 1194-210, MVZ 126523- 1682(2). L. rothi: SDSU 1175-77, KU 161166, 47, REE 2656-2667. L.jamesi: SDSU 2623, FML 187517-18, MVZ 180041,188087-94,18824-39, 1193, KU 161741, LACM 134120. L. kingii kin- REE 2398-403. L. ruibali: SDSU 1604-08, REE gii: SDSU 1671-72, REE 2479-86. L. k. baguali: 2297-2304. L. salinicola: SDSU 1011-12, 1189- FML 1247. L. k. somuncurae: SDSU 1669-70, 1190, FML 1909(10), 1912(9), 1807(13), 2020(21), REE 2390-95. L. koslowskyi: SDSU 1005-09, REE 2568, 2738-39. L. sanjuanensis: FML 1016. 1041-50, 1067-69, 1096-97, 1409-23, 1431-42, L. saxatilis: SDSU 1736-39. L. scapularis: SDSU REE 2705-06. L. kriegi: SDSU 1877, REE 2412- 1017-19, 1022-23, 1057, 1060-62, 1220-28, 2422. L. kuhlmanni: SDSU 1878-8 1, REE 2526. 1274-303, REE 2569-77, 2709-22. L. schmidti: L. laurenti: SDSU 1013, 1024-1038, 1070-88, SDSU 1602-03, FMNH 5759-60, AMNH 1114-17, 1121-44, 1159-67, REE 2741-50. L. 131850-61, REE 2518. L. schroederi: MVZ 1995 ETHERIDGE: REDESCRIPTION OF CTENOBLEPHARUS ADSPERSA 31

187760-61. L. signifer signifer: SDSU 1600, S. roseiventris: SDSU 1686, REE 2284. Microlo- AMNH 90457-60,90464-68, REE 1825-29,2562. phus occipitalis: SDSU 2028-29, 3060-61, REE L. s. annectens: FML 1543. L. stolzmanni: NMW 649, 658, 1859, 1861-62. M. peruvianus: SDSU 13580(3). L. sylvanae: MCZ 162004-06, 2031-32. Tropidurus etheridgei: SDSU 2014-17, 156906(sk), KU 190418(sk). L. tacnae: SDSU REE 1954. T. hygomi: SDSU 2022, REE 275. 1924, REE 2533, 2697. L. tenuis tenuis: SDSU Uranoscodon superciliosus: SDSU 2110-14, REE 1925-35, 2084-89, REE 1817, 2591, 2857. L. t. 2589, 2883-84. punctatissimus: SDSU 1936, REE 2536. L. uspallatensis: FML 1541(10), MVZ 126869,188858, 188860-63, 188859(sk). L. variegatus: KU Opluridae 133862-71. L. velosoi: IZUC uncataloged. L. Chalarodon madagascariensis: SDSU 2123-29, walkeri: SDSU 1937, REE 1818, 2534. L. wieg- USNM 149311, REE 455, 457, 547. Oplurus cu- mannii: SDSU 1168,1217-19, CAS 174160-200, vieri: REE 620, 558, 1835. 0. cyclurus: USNM REE 2567, 2601-04. L. williamsi: FML 1701. L. 149330. 0. quadrimaculatus: SDSU 2120-22, REE xanthoviridis: SDSU 1179, KU 182049, 187514. 658. 0. saxicola: SDSU 2119. L. zapallarensis zapallarensis: SDSU 1938,2040- 46, REE 2769, 2843. L. z. ater: SDSU 1939-42, REE 2850-56. Phrynosomatidae Phymaturus palluma: SDSU 1946-79, REE 1950, 2306-15, 2323-28, 2361-62. P. antofagas- Petrosaurus mearnsi mearnsi: SDSU 2253, REE tensis: SDSU 1991. P. patagonicus: SDSU 1980, 287-88, 351, 761, 1557-58. P. m. sleveni: REE REE 2471-72. P. payune: SDSU 1981-84, REE 655. P. thalassinus: REE 575, 765-66, 797, 1381. 2330-39, 2360. P. punae: SDSU 1978-79, REE Uta stansburiana: SDSU 2522-30, REE 274-75, 2356-57,2383-85. P. somuncurensis: SDSU 1780- 1877-78, 1892. Utapalmeri: SDSU 2520-2 1, REE 84, REE 2433-40. P. zapalensis: SDSU 1985-90, 1990. Urosaurus graciosus: SDSU 2543-46, REE REE 2451-53. 271, 1544, 1547, 1625. U. nigricaudus: SDSU 2536-37, REE 492, 494. U. lahteli: SDSU 2854- Leiocephalinae + Tropidurinae 55. U. bicarinatus: SDSU 2533. U. ornatus: SDSU 2534-35,2538-42,2547-53, REE 403, 757, 1553, Leiocephalus carinatus: SDSU 1996-97, REE 1556, 1559. Phrynosoma asio: SDSU 2308-09, 1469, 1805, 1816. L. cubensis: SDSU 1999-2001. REE 1489,1580,1676. P. douglassii: SDSU 2283- L. schreibersi: SDSU 1998, REE 1808. Stenocer- 84. P. orbiculare: SDSU 1109, 1181, 1725, 1931. cus chlorostictus: SDSU 1535. S. crassicaudatus: P. coronatum: SDSU 2305-07, REE 28, 188, 609, SDSU 1680-83, REE 2286, 2593. S. empetris: 1108,1439,1501,1786,1999. Umanotata:SDSU SDSU 3059. S. guentheri: SDSU 1685. S. imita- 2554-72, REE 263-64, 316, 396, 1535-38. U. tor: SDSU 1534. S. modestus: SDSU 1686. S. exsul: SDSU 2274-77, REE 2880-81. U. scoparia: percultus: SDSU 1596. S. praeornatus: REE 2544. REE 509, 551.

APPENDIX 2 AN INDENTED CLASSIFICATION OF LIOLAEMINAE Part A. A tentative classification of Liolae- the oldest names. Formal generic or subgeneric minae, intended to represent historical internest- names have been proposed for some ofthese (e.g., ing, is presented here in indented form. The con- Vilcunia Donoso-Barros and Cei = the sylvanae ventions ofWiley (1979) that are applicable (con- group), but their use is discouraged at this time. ventions 1-4) are followed. An abbreviated in- The term "group" is employed for collations of dented classification is presented first to avoid taxa that are thought to be monophyletic. Species confusion that may result from the large number names separated by an arrow (- ) refer to the spe- ofspecies listed in Part B. Supraspecific groups are cies pairs or polytomies listed alphabetically in followed by characters that are likely synapomor- Part B. phies based on comparisons with the putative outgroups Leiocephalinae + Tropidurinae, Phryno- Phymaturus sedis mutabilis: head and body flat- somatidae and Opluridae. On the same basis, tened; body wide; lateral nuchal skin folds ob- characters not listed are considered plesiomorph- scured by fat-filled pouches; tail with regular whorls ic, or their status is equivocal. Species-group names ofspinose scales; interclavicle short; suprascapula are chosen arbitrarily, in most cases using one of medially inflected; four stemal ribs; marginal teeth, 32 AMERICAN MUSEUM NOVITATES NO. 3142 including at least some premaxillary teeth, ex- hypertrophied M. tibialis anterior. andinus panded, with three (sometimes four) large cusps. williamsi palluma group: superciliaries short; five or more boulengeri group: a patch of abruptly en- subequal suboculars; three or four rows of lor- larged, spinose scales on the posterior me- ilabials; mental narrower than rostral, usually dial surface of thigh, bulged out in adult in contact with sublabials, sometimes frag- males due to hypertrophy ofunderlying M. mented; caudal spines very well developed, two puboischiotibialis. abaucan - xanthovir- annuli per segment. palluma -- punae idis patagonicus group: splenial short; Meckel's wiegmannii group: lorilabials distinctly groove fused. indistinctus -- zapalensis smaller than supralabials, usually in two Ctenoblepharys sedis mutabilis: head short and rows between subocular and supralabials; broad; superciliaries short; outer ciliaries strongly supralabials narrow, but posteriormost projecting, triangular on lower lid; skull wide (about not elongate; sublabials contact mental 1.3 x longer than wide); orbit large (about 0.43- scale, mental widest posteriorly; infrala- 0.47 x skull length); snout short (about 0.29 x bials flat to concave. cranwelli -- wieg- skull length); temporal fenestrae wide (1.5-1.6 x mannii longer than wide); prefrontals wider than long; lacrimal foramen large; maxillary process ofpalatine Part B. It is intended that the following wide; ectopterygoid short; maxillary process ofec- list include all specific and subspecific names topterygoid truncate distally; retroarticular pro- in current usage, except Liolaemus erroneus, cess of mandible much shorter than angular pro- within Liolaeminae. Subspecific names are cess; parapophyses of atlas flat and oriented pos- included for the sake of completeness, with- terolaterally; neural arches ofbody vertebrae very out reference to their status as unitary evolv- wide. adspera ing entities. Inclusion and placement of spe- Liolaemus sedis mutabilis: frontal downgrowths cies and subspecies not examined are based reach, or almost reach, palatines excluding prefrontals from orbitonasal fenestra; supratemporals on data available from the literature. These mostly enclosed in groove along inferior margin species and the bibliographic sources oftheir ofsupratemporal processes ofparietal; dentary ex- data are given in the introduction to Appen- tends posterior to superior apex of coronoid; lin- dix 1. gual process ofangular short or absent; secondary coracoid fenestra present; pygal region of males Phymaturus Gravenhorst, 1838, sedis mu- much larger than in females, scales ofpygal region tabilis usually smaller in females than in males; hemi- palluma group penial retractor muscles located anteriorly and hy- antofagastensis Pereyra, 1985 pertrophied. archeforus -- kingii mallimaccii Cei, 1980 nitidus group: supralabials narrow, width equal palluma (Molina, 1782) to or less than that of lorilabials, usually four, punae Cei, Etheridge & Videla, 1983 the posterior one elongate and usually upturned posteriorly. magellanicus patagonicus group lineomaculatus group: precloacal pores lost; indistinctus Cei & Castro, 1973 at least some dorsal scales triden- nevadoi Cei & Castro, 1975 tate. lineomaculatus patagonicus Koslowsky, 1898 sylvanae group: lateral nuchal scales keeled payunae Cei & Castro, 1973 and imbricate; postfemoral scales subim- somuncurensis Cei & Castro, 1973 bricate. periglacialis -+ sylvanae zapalensis Cei & Castro, 1973 chiliensis group precloacal pores usually four Ctenoblepharys Tschudi, 1845, sedis muta- or fewer, rarely absent; Meckel's groove bilis fused. alticolor -- zapallarensis adspersa Tschudi, 1845 signifer group: epimysium of M. pterygomandibularis pigmented; medial head of M. flexor Liolaemus Wiegmann, 1834, sedis mutabilis tibialis internus covered by hypetrophied M. archeforus puboischiotibialis. anomalus -- pseudoan- a. archeforus Donoso-Barros & Cei, 1975 omalus a. gallardoi Cei & Scolaro, 1982 montanus group: a sharp, bladelike process on a. sarmientoi Donoso-Barros, 1973 posterior distal tibia, associated with greatly kingil 1995 ETHERIDGE: REDESCRIPTION OF CTENOBLEPHARUS ADSPERSA 33

k. kingii (Bell, 1843) kriegi Miller & Hellmich, 1939 k. baguali Cei & Scolaro, 1983 kuhlmanni Miller & Hellmich, 1932 k. somuncurae Cei & Scolaro, 1981 lativittatus Werner, 1904 nitidus group lemniscatus Gravenhorst, 1837 magellanicus (Hombron & Jacquinot, leopardinus Miller and Hellmich, 1847) 1932 lineomaculatus group lorenzmulleri Hellmich, 1950 lineomaculatus Boulenger, 1885 maldonadae Nunez et al., 1991 sylvanae group modestus (Philippi, 1860) periglacialis (Cei & Scolaro, 1982) monticola sylvanae (Donoso-Barros & Cei, m. monticola Muller & Hellmich, 1971) 1932 chiliensis group m. chillanensis Miller & Hllmich, alticolor Barbour, 1909 1932 atacamensis Muller & Hellmich, 1933 m. villaricensis Miiller and Hell- austromendocinus Cei, 1974 mich, 1932 bellii nigromaculatus (Wiegmann, 1834) bellii bell/i Gray, 1845 nigroviridis b. araucaniensis Muller & Hell- n. nigroviridis Miller & Hellmich, mich, 1932 1932 b. moradoensis Hellmich, 1950 n. campanae Hellmich, 1950 b. neuquensis Muller & Hellmich, n. minor Miller & Hellmich, 1932 1939 n. nigroroseus Donoso-Barros, 1966 bibronii (Bell, 1843) nitidus (Wiegmann, 1834) bisignatus (Philippi, 1860) paulinae Donoso-Barros, 1961 bitaeniatus Laurent, 1984 pictus buergeri Werner, 1907 p. pictus (Dumeril & Bibron, 1837) capillitas Hulse, 1979 p. argentinus Muller & Heilmich, ceii Donoso-Barros, 1971 1939 chacoensis Shreeve, 1948 p. chiloeensis Muller & Hellmich, chiliensis (Lesson, 1831) 1939 coeruleus Cei & Ortiz-Zapata, 1983 p. major Boulenger, 1885 constanzae Donoso-Barros, 1961 p. talcanensis Urbina & Zufngia, copiapensis Miller & Hellmich, 1933 1977 cristiani N'unez et al., 1991 platei Werner, 1898 curicensis Miller & Hellmich, 1938 pseudolemniscatus Lamborot & Ortiz- curis NuOnez & Labra, 1985 Zapata, 1990 cyanogaster ramonensis Muller & Hellmich, 1932 c. cyanogaster (Dumeril & Bibron, robertmertensi Hellmich, 1964 1837) sanjuanensis Cei, 1982 c. brattstroemi Donoso-Barros, 1961 saxitilis Avila et al, 1992 donosoi Ortiz-Zapata, 1975 schroederi Miller & Hellmich, 1938 elongatus silvai Ortiz-Zapata, 1989 e. elongatus Koslowsky, 1896 tacnae (Shreve, 1941) e. petrophilus Donoso-Barros & Cei, tenuis 1971 t. tenuis (Dumeril & Bibron, 1837) exploratorum Cei & Williams, 1984 t. punctatissimus Muller & Hell- fitzgeraldi Boulenger, 1899 mich, 1933 fuscus Boulenger, 1885 valdesianus Hellmich, 1950 gracilis (Bell, 1843) variegatus Laurent, 1984 gravenhorstii (Gray, 1845) velosoi Ortiz-Zapata, 1987 hellmichi Donoso-Barros, 1975 walkeri Shreve, 1938 hernani Sallaberry et al, 1982 zapallarensis 34 AMERICAN MUSEUM NOVITATES NO. 3142

z. zapallarensis Muller & Hellmich, reichei (Werner, 1907) 1933 robustus Laurent, 1991 z. ater Muller & Hellmich, 1933 rosenmanni N'uinez & Navarro, 1992 z. sieversi Donoso-Barros, 1954 ruibali Donoso-Barros, 1961 signifer group schmidti (Marx, 1960) anomalus signifer a. anomalus Koslowsky, 1896 s. signifer (Dumeril & Bibron, 1837) a. ditadai Cei, 1986 s. annectens Boulenger, 1901 pseudoanomalus Cei, 1981 stolzmanni (Steindachner, 1891) montanus group vallecurensis Pereyra, 1992 andinus williamsi Laurent, 1992 a. andinus Koslowsky, 1895 boulengeri group a. poecilochromus Laurent, 1986 abaucan Etheridge, 1993 audituvelatus N(uniez & Yaniez, 1983 boulengeri Koslowsky, 1898 aymararum Veloso et al., 1982 canqueli Cei, 1975 disjunctus Laurent, 1990 cuyanus Cei & Scolaro, 1989 dorbignyi Koslowsky, 1898 darwinii (Bell, 1843) duellmani Cei, 1978 donosobarrosi (Cei, 1974) eleodori Cei et al., 1983 fitzingerii (Dumeril & Bibron, 1837) fabiani Ya-nez & Niiunez, 1983 irregularis Laurent, 1986 famatinae Cei, 1980 koslowskyi Etheridge, 1993 fittkaui Laurent, 1986 laurenti Etheridge, 1992 forsteri Laurent, 1982 melanops Burmeister, 1888 griseus Laurent, 1984 olongasta Etheridge, 1993 huacahuasicus Laurent, 1985 ornatus Koslowsky, 1898 insolitus Cei & Pefaur, 1982 quilmes Etheridge, 1993 isabelae Navarro & Ntuniez, 1993 rothi Koslowsky, 1898 islugensis Ortiz-Zapata & Marquet, uspallatensis Macola & Castro, 1982 1987 xanthoviridis Cei & Scolaro, 1980 jamesi (Boulenger, 1891) wiegmannii group melanopleurus (Philippi, 1860) cranwelli (Donoso-Barros, 1973) montanus Koslowsky, 1898 lutzae Mertens, 1938 multicolor Koslowsky, 1898 multimaculatus (Dumeril & Bi- nigriceps (Philippi, 1860) bron, 1837) orientalis occipitalis Boulenger, 1885 o. orientalis Muller, 1923 rabinoi (Cei, 1974) o. chlorostictus Laurent, 1993 riojanus Cei, 1979 ortizi Laurent, 1982 salinicola Laurent, 1986 patriciaiturrae Navarro and N(uinez, scapularis Laurent, 1982 1993 wiegmannii (Dumeril & Bibron, polystictus Laurent, 1991 1837) pulcherrimus Laurent, 1993

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