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A Critically Endangered New Species of Cnemidophorus (Squamata, Teiidae) from a Cerrado Enclave in Southwestern Amazonia, Brazil

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A Critically Endangered New Species of Cnemidophorus (Squamata, Teiidae) from a Cerrado Enclave in Southwestern Amazonia, Brazil Herpetologica, 59(1), 2003, 76±88 q 2003 by The Herpetologists' League, Inc. A CRITICALLY ENDANGERED NEW SPECIES OF CNEMIDOPHORUS (SQUAMATA, TEIIDAE) FROM A CERRADO ENCLAVE IN SOUTHWESTERN AMAZONIA, BRAZIL GUARINO R. COLLI1,3,GABRIEL C. COSTA1,ADRIAN A. GARDA1,KAÂ TIA A. KOPP2, DANIEL O. MESQUITA1,AYRTON K. PEÂ RES,JR.1,PAULA H. VALDUJO1, GUSTAVO H. C. VIEIRA1, AND HELGA C. WIEDERHECKER1 1Departamento de Zoologia, Universidade de BrasõÂlia, 70910-900 BrasõÂlia, DF Brazil 2Departamento de CieÃncias BioloÂgicas, Universidade Federal de Santa Maria, 97105-900 Santa Maria, RS Brazil ABSTRACT: We describe a new species of Cnemidophorus from a Cerrado enclave in south- western Amazonia, RondoÃnia state, Brazil. This species is apparently endemic to Cerrado enclaves in the vicinity of the city of Vilhena, a region under intensive anthropic pressure due to the expansion of soybean plantations. A discriminant analysis indicated that femoral pores and scales around the tail are the best discriminators among Brazilian species of Cnemidophorus. A naõÈve Bayesian network constructed with categorical (mostly coloration) variables indicated that the new species had high conditional probabilities of dorsolateral ®elds absent, vertebral ®eld spotted, and paravertebral lines absent. The analyses revealed clear distinctions between species of Cnemidophorus that range north and south of the the Amazon River. The new species may have evolved as a result of vicariance, following the isolation of peripheral enclaves of Cerrado in southwestern Amazonia after the late Pleniglacial. The restricted range in small areas, under extreme human pressure around Vilhena, makes this species one of the most critically endangered elements of the Brazilian herpetofauna. Key words: Amazonia; Brazil; Cerrado; Cnemidophorus; Extinction; Lizard; RondoÃnia RANGING from northern United States Lema, 2000; and C. vanzoi (Baskin and to central Argentina, Cnemidophorus has Williams, 1966). the widest geographic distribution among That about 50% of the species in the all teiid genera (Wright, 1993). Currently, Cnemidophorus lemniscatus group has 57 species are known in the genus (Uetz been described in the last 15 yr testi®es to et al., 1995). All South American species the paucity of knowledge of the alpha-level of Cnemidophorus are assigned to the C. diversity in the neotropics and the growing lemniscatus species group (Burt, 1931; taxonomic activity in the area. Indeed, un- Wright, 1993), which comprises 17 spe- til recently, all populations of Cnemido- cies: C. arenivagus Markezich, Cole, and phorus in northern South America were Dessauer, 1997; C. arubensis Van Lidth de assigned to a single species, C. lemnisca- Jeude, 1887; C. cryptus Cole and Des- tus. Several studies have indicated that C. sauer, 1993; C. gramivagus McCrystal and ocellifer is also a complex of species (e.g., Dixon, 1987; C. lacertoides DumeÂril and Rocha et al., 1997, 2000; Rodrigues, 1987). Bibron, 1839; C. leachei Peracca, 1897; C. In addition, the availability of tools to lemniscatus (Linnaeus, 1758); C. littoralis probe molecular markers has only recently Rocha, ArauÂjo, Vrcibradic, and Costa, allowed the detection of morphologically 2000; C. longicaudus (Bell, 1843); C. mu- cryptic species, such as C. cryptus (e.g., rinus (Laurenti, 1768); C. nativo Rocha, Cole and Dessauer, 1993; Sites et al., Bergallo, and Peccinini-Seale, 1997; C. ni- 1990). gricolor Peters, 1873; C. ocellifer (Spix, During the course of a study on the 1825); C. pseudolemniscatus Cole and Cerrado enclaves in RondoÃnia, Brazil, we Dessauer, 1993; C. serranus Cei and Mar- collected a relatively large series of an un- tori, 1991; C. vacariensis Feltrim and described species of Cnemidophorus. A few individuals of this species had been previously collected by Vitt and Caldwell 3 CORRESPONDENCE: e-mail, [email protected] (1993) during a survey sponsored by the 76 March 2003] HERPETOLOGICA 77 mivagus, C. lemniscatus, C. littoralis, and C. ocellifer from specimens housed in the ColecËaÄo HerpetoloÂgica da Universidade de BrasõÂlia (CHUNB) (Appendix I). The ®rst three species occur predominantly in Am- azonian Savannas north of the Amazon River (e.g., AÂ vila-Pires, 1995; Cole and Dessauer, 1993), whereas the last two oc- cur exclusively south of the Amazon (e.g., Cei, 1993; Rocha et al., 2000). It should be noted that C. cryptus is a provisional name for a complex of parthenogenetic, unisexual populations in the Amazon Basin and that much work is necessary to clarify their taxonomic status. We recorded the following meristic variables in each speci- men: supralabials (number of enlarged scales along the upper jaw, total on both FIG. 1.ÐVegetation map of type locality of Cnemi- dophorus parecis. sides), infralabials (number of enlarged scales along the lower jaw, total on both sides), chinshields (most anterior pair of Polonoroeste program along the BR-364 chinshields separated from infralabials by road in RondoÃnia (Nascimento et al., 1988; row of small scales), gular folds (number Vanzolini, 1986). The individuals were re- of folds in gular region), supraoculars ferred by Vanzolini both to Cnemidopho- (number of supraoculars on right side), pa- rus ocellifer (Vanzolini, 1986) and to an rietals (number of parietals plus interpa- unnamed member of the ocellifer species rietal scale), scales around midbody group (sic) (P. E. Vanzolini, personal com- (counted midway between fore- and hind- munication, in Vitt and Caldwell, 1993). limbs, excluding ventrals), transverse rows Herein, we describe this new species, of ventrals (counted along the midline, making detailed comparisons with other from gular fold to anterior margin of hind- Brazilian congeneric species. limbs), ventrals in transverse row (counted midway between fore- and hindlimbs), MATERIALS AND METHODS femoral pores (total number on both We collected lizards with a shotgun, pit- sides), prefemorals (number of enlarged fall traps, and drift fences on 4±10 Novem- scales on anterior aspect of thigh, counted ber 1998 and 21 August±22 September midway between the hip and the knee, on 1999, around the city of Vilhena (128 439 a row from femoral pores to granules on S, 608 079 W), RondoÃnia state, Brazil (Fig. dorsal aspect of thigh), prefemoral rows 1). The vegetation in the study area con- (counted from hip to knee), infratibials sists of a mosaic of Cerrado enclaves (Ei- (number of enlarged scales on longitudinal ten, 1972) within a matrix of Terra Firme row from knee to base of ®rst metatarsal), Amazonian Forest (Pires and Prance, preanals (number of enlarged scales on 1985) and human-modi®ed areas consist- preanal plate, from level of medialmost ing of soybean plantations and groves of femoral pores to vent), fourth ®nger la- pine trees. We found lizards exclusively in mellae (counted under the ®nger), fourth Cerrado enclaves with sandy soils around toe lamellae (counted under the toe), the city of Vilhena, even though we sam- scales around tail (counted on ®fth trans- pled the largest patches of Cerrado en- verse row), and dorsals (counted along the claves throughout RondoÃnia, including midline, from occiput to ®rst transverse those around the cities of Pimenta Bueno row of scales around tail). and GuajaraÂ-Mirim. We used a discriminant function analy- We obtained data on C. cryptus, C. gra- sis (Tabachnick and Fidell, 2001) to inves- 78 HERPETOLOGICA [Vol. 59, No. 1 tigate differences among species in meris- (SMC, degree of contact between supra- tic characters. To identify the most pow- oculars and medial head scales: no contact, erful meristic discriminators of the species no contact with semicircles isolating ®rst of Cnemidophorus, we used a stepwise supraocular, supraoculars contacting fron- discriminant function analysis of meristic tal and parietals, supraoculars contacting characters (Tabachnick and Fidell, 2001), frontal, supraoculars contacting parietals), using the METHOD 5 STEPWISE op- dorsal caudals (DCA, keels on dorsal, cau- tion in PROC STEPDISC of SAS (SAS In- dal scales, from most anterior third of tail: stitute Inc., 1988). We evaluated linear dis- absent, present), ventral caudals (VCA, criminant functions through posterior keels on ventral, caudal scales, from most probability error-rate estimates, based on anterior third of tail: absent, present), and crossvalidation (SAS Institute Inc., 1988). preanal spur (PAS: absent, present). We also recorded the following categor- To model the conditional dependencies ical variables: lower lateral ®elds (LLF, of each categorical variable within each one on each side, area between ventral species, we constructed a naõÈve Bayesian scales and lower lateral stripes: absent, model using the software UNBARB (avail- light, spotted, or dark), upper lateral ®elds able at ftp://ftp.cic.unb.br/pub/cic/wagner/ (ULF, one on each side, area between up- software/Bnets/UnBARB.zip). In this per lateral and dorsolateral stripes: absent, model (Fig. 2), the variable ``species'' in- light, spotted, or dark), dorsolateral ®elds ¯uences the chance of the occurrence of (DLF, one on each side, dark area be- states of each categorical variable, assum- tween dorsolateral and paravertebral ing that categorical variables are condition- stripes: absent, light, spotted, or dark), ver- ally independent for a given species and tebral ®elds (VTF, one on each side, mid- that the joint distribution of all variables dorsal area between paravertebral stripes: satis®es the following: absent, light, spotted, or dark), vertebral line (VTL, light stripe
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