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

Herpetologica, 59(1), 2003, 76–88 ᭧ 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,KA´ TIA A. KOPP2, DANIEL O. MESQUITA1,AYRTON K. PE´ RES,JR.1,PAULA H. VALDUJO1, GUSTAVO H. C. VIEIRA1, AND HELGA C. WIEDERHECKER1 1Departamento de Zoologia, Universidade de Brası´lia, 70910-900 Brası´lia, DF Brazil 2Departamento de Cieˆncias Biolo´gicas, Universidade Federal de Santa Maria, 97105-900 Santa Maria, RS Brazil

ABSTRACT: We describe a new species of Cnemidophorus from a Cerrado enclave in southwestern Amazonia, Rondoˆnia 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 naı¨ve Bayesian network constructed with categorical (mostly coloration) variables indicated that the new species had high conditional probabilities of dorsolateral ﬁelds absent, vertebral ﬁeld 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; Rondoˆnia

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 testiﬁes 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 Dume´ril 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, Arau´jo, 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 Rondoˆnia, 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 Coleçaõ Herpetolo´gica da Universidade de Brası´lia (CHUNB) (Appendix I). The first three species occur predominantly in Am- azonian Savannas north of the Amazon River (e.g., A´ vila-Pires, 1995; Cole and Dessauer, 1993), whereas the last two occur 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.—Vegetation 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 Rondoˆnia (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 (12Њ 43Ј a row from femoral pores to granules on S, 60Њ 07Ј W), Rondoˆnia 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 first metatarsal), Amazonian Forest (Pires and Prance, preanals (number of enlarged scales on 1985) and human-modified areas consist- preanal plate, from level of medialmost ing of soybean plantations and groves of femoral pores to vent), fourth finger la- pine trees. We found lizards exclusively in mellae (counted under the finger), 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 fifth trans- pled the largest patches of Cerrado en- verse row), and dorsals (counted along the claves throughout Rondoˆnia, including midline, from occiput to first transverse those around the cities of Pimenta Bueno row of scales around tail). and Guajara´-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 first 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 ϭ 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 fields (LLF, of each categorical variable within each one on each side, area between ventral species, we constructed a naı¨ve Bayesian scales and lower lateral stripes: absent, model using the software UNBARB (avail- light, spotted, or dark), upper lateral fields 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 fields fluences 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 fields (VTF, one on each side, mid- that the joint distribution of all variables dorsal area between paravertebral stripes: satisfies the following: absent, light, spotted, or dark), vertebral line (VTL, light stripe from interparietal k P(S, C , ...,C)ϭ P(C ͦ S) P(S), 1k͹ i scale to base of tail: absent, interrupted, ΂΃iϭ1 continuous, or double), paravertebral stripes (PVS, one on each side, light stripe where S is species, C is a categorical vari- from parietal scale to first third of tail, be- able, P(Ci ͦS) is the posterior probability of tween vertebral and dorsolateral fields: ab- Ci given S, and P(S) is the prior probability sent, interrupted, or continuous), dorsolat- of S (Cowell et al., 1999). We estimated eral stripes (DLS, one on each side, light P(S) and P(Ci ͦ S) using Bayesian statistics stripe from superciliaries to first third of (Bernardo and Smith, 2000; DeGroot, tail, between dorsolateral and upper lateral 1970) and the software Hugin Lite v. 5.3 fields: absent, interrupted, or continuous), (Jensen and Nielsen, 1999). Throughout upper lateral stripes (ULS, one on each the text we report means Ϯ1 SD. side, light stripe from suborbital region to SPECIES DESCRIPTION hindlimb, between upper and lower lateral fields: absent, interrupted, or continuous), Cnemidophorus parecis sp. nov. lower lateral stripes (LLS, one on each Holotype.—CHUNB 09811 (Figs. 3, 4), side, light stripe from axilla to hip, be- adult male, from Fazenda Cachoeira (12Њ tween lower lateral field and ventrals: ab- 32Ј 07Љ S, 60Њ 25Ј 36Љ W), 50 km NW of sent, interrupted, or continuous), lateral Vilhena, Estado de Rondoˆnia, Brazil, ele- spots (LTS, rounded light areas on flanks: vation about 612 m, collected on 5 Novem- absent, present), hindlimb spots (HLS, ber 1998 by Guarino R. Colli and students. rounded light areas on hindlimbs: absent, Paratopotypes.—(107) All specimens present), chinshield contact (CHS, degree from Cerrado enclaves around Vilhena: of contact between most anterior pair of CHUNB 09762–810, 09812–14, collected chinshields: no contact, contact smaller on November 1998 by same collectors; than half of their lengths, or contact great- CHUNB 11651–703, 14229–30 collected on er than half of their lengths), semicircles August–September 1999 by same collectors. March 2003] HERPETOLOGICA 79

FIG. 2.—Diagram depicting naı¨ve Bayesian network constructed to evaluate conditional probabilities of 16 categorical variables from species, assuming independence among categorical variables. See text for abbreviations.

Diagnosis.—A species distinguished (present), and ventral caudals smooth from all other members of the lemniscatus (keeled). Cnemidophorus parecis differs group by the following combination of from C. cryptus in having 11–14 suprala- characters: (1) large size, maximum SVL bials (14–17 in C. cryptus), 8–10 trans- 90 mm; (2) 34–47 scales around tail; (3) verse rows of ventrals (7–8), 25–33 femoral 25–33 femoral pores; (4) 190–252 dorsals; pores (38–48), 34–47 scales around tail (5) dorsolateral fields absent; (6) vertebral (22–30), upper lateral fields light (dark), field spotted; (7) paravertebral stripes ab- dorsolateral fields absent (dark), paraver- sent (Fig. 4); and (8) preanal spurs absent. tebral stripes absent (continuous) (Fig. 5), Cnemidophorus parecis differs from C. and ventral caudals smooth (keeled). lemniscatus in having 11–14 supralabials Cnemidophorus parecis differs from C. lit- (14–16 in C. lemniscatus), 10–15 infrala- toralis in having 34–47 scales around tail bials (15–19), 25–33 femoral pores (44– (25–34 in C. littoralis), upper lateral fields 54), 34–47 scales around tail (24–30), dor- light (dark), paravertebral stripes absent solateral fields absent (dark), vertebral (interrupted), vertebral line absent (inter- field spotted (dark), paravertebral stripes rupted) (Fig. 5), and dorsal caudals keeled absent (continuous), vertebral line absent (smooth). Cnemidophorus parecis differs (two continuous stripes) (Fig. 5), hindlimb from C. ocellifer in having 8–10 transverse spots absent (present), and ventral caudals rows of ventrals (5–8 in C. ocellifer), 25– smooth (keeled). Cnemidophorus parecis 33 femoral pores (16–28), 34–47 scales differs from C. gramivagus in having 25– around tail (21–30), and lateral spots ab- 33 femoral pores (40–51 in C. gramiva- sent (present) (Fig. 5). gus), 34–47 scales around tail (20–32), Description of holotype.—Rostral pen- lower lateral fields light (absent), upper tagonal, higher than wide, visible from lateral fields light (dark), dorsolateral fields above, bordered posteriorly by nasals. Na- absent (dark), vertebral field spotted sals in contact along midline, each nasal (dark), paravertebral stripes absent (con- divided by an oblique suture. Nostril in tinuous) (Fig. 5), lower lateral stripes in- lower part of suture, directed posterolat- terrupted (absent), hindlimb spots absent erally, longer than tall. Frontonasal sub- 80 HERPETOLOGICA [Vol. 59, No. 1

FIG. 3.—Cnemidophorus parecis, holotype, CHUNB 09811, adult male, SVL 86 mm. hexagonal, suture with nasals and loreals forming a semicircle, sutures with prefrontals forming a wide angle. Prefrontals FIG. 4.—Top: Cerrado enclave in Vilhena, Ron- quadrilateral, with medial suture twice as doˆnia, Brazil. Middle: Adult female of Cnemidopho- rus parecis. Bottom: Adult male of C. parecis (holo- long as that between nasals, in contact lat- type, CHUNB 09811). erally with loreal, first supraciliary, and first supraocular. Frontal approximately pentagonal, longer than wide, and wider regular scales by an oblique suture. Occip- anteriorly. Sutures with prefrontals and su- ital scales irregular, posterior to interpari- praoculars roundish, those with frontopar- etal and parietals, variable in size, includ- ietals forming a straight line. Frontal in ing two moderately enlarged, medial contact laterally with first, second, and scales, separated by a smaller scale. Four third supraoculars. Frontoparietals pentag- supraoculars on each side, second and onal, longer than wide, with long, straight third largest, followed posteriorly by group medial suture; in contact laterally with of small scales. Second, third, and fourth third supraocular and small scales border- supraoculars separated from supraciliaries ing supraocular region posteriorly. Inter- by row of small scales. Supraciliaries seven parietal hexagonal, bordered at each side on each side, highest anteriorly, third larg- by hexagonal parietals, divided into two ir- est, others subequal. Loreal single, large, March 2003] HERPETOLOGICA 81

FIG. 5.—Dorsal coloration patterns in six species of Cnemidophorus from Brazil. (A) C. littoralis, CHUNB 08308, (B) C. ocellifer, CHUNB 12964, (C) C. parecis, CHUNB 11652, (D) C. lemniscatus, CHUNB 01101, (E) C. cryptus, CHUNB 08491, (F) C. gramivagus, CHUNB 03527. in contact with nasal, frontonasal, prefron- terior contact with first infralabials and tal, first supraciliary, frenocular, first sub- postsymphysal, forming two wide angles. ocular, and second, third, and fourth su- Postsymphysal single, pentagonal, in con- pralabials on the left side, and third and tact with first and second infralabials, fol- fourth supralabials on the right side. Fren- lowed by three pairs of enlarged chin- ocular narrow, higher than wide, in contact shields. First pair of chinshields in ample with subocular, loreal, first supraciliary, contact along midline, first and second and small scales in ocular region. Subocu- pairs in contact with infralabials, third pair lars three, large, second largest, all in con- separated from infralabials by row of small tact with supralabials. A continuous keel scales, followed posteriorly by enlarged runs from frenocular to about midlength scales. Medial scales on chin small, of second subocular. Postoculars small, ar- smooth, elongate, arranged in longitudinal, ranged in two rows with 3–4 larger scales. posteriorly divergent rows; increasing in Lower eyelid with semiopaque disc, size and becoming roundish posteriorly. formed by transversally enlarged, convex Infralabials six, followed posteriorly by se- scales. Supralabials six on each side, fifth ries of small scales extending to commis- and sixth below center of eye; followed by sure of mouth; fifth infralabial below cen- series of small scales extending to com- ter of eye. Gular region divided in two ar- missure of mouth. Temporal region with eas: anterior region with enlarged, round- irregular scales, granular centrally, mod- ed scales, in transverse rows, delimited erately enlarged peripherally. Ear opening posteriorly by line uniting lower margin of large, oblique, with smooth margins, an- ear openings; posterior region covered terior margin forming a semicircle, poste- with granules, in transverse rows, bor- rior margin straight. Tympanum recessed dered posteriorly by antegular fold. Gular in a short auditory meatus. All dorsal and and antegular folds marked by granules; lateral head scales juxtaposed, smooth. scales between the two folds larger, irreg- Symphysal anteriorly ellipsoid, in pos- ular, slightly imbricate. 82 HERPETOLOGICA [Vol. 59, No. 1

TABLE 1.—Canonical discriminant analysis of meris- Scales on base of tail rectangular, longer tic characters of six species of Cnemidophorus. than wide, smaller than ventrals, keeled Canonical statistics dorsally, smooth ventrally, in transverse Cumula- rows. All transverse rows continuous tive Canonical Eigen- propor- around tail, except first two rows incom- variable value tion r2 FPplete ventrally. Tail scales becoming longer 1 42.438 0.588 0.977 165.06 Ͻ0.0001 and narrower posteriorly; subcaudal scales 2 21.087 0.879 0.955 101.12 Ͻ0.0001 becoming keeled posteriorly. Tail broken, Raw canonical coefficients regenerated from 37th transverse row of Canonical Canonical scales. Limbs with large, smooth, imbri- Variable variable 1 variable 2 cate scales on dorsal aspect of upper arms, Supralabials 0.098 Ϫ0.342 anterodorsal aspect of forearms, antero- Infralabials 0.312 0.211 Chinshields 0.120 0.464 ventral aspect of thighs, and ventral aspect Gular fold Ϫ0.126 Ϫ0.091 of lower legs; elsewhere scales small, gran- Supraoculars 0.291 Ϫ0.189 ular. Larger scales on upper arms in lon- Parietals Ϫ0.183 Ϫ0.121 gitudinal rows. Forearms with one row of Scales around midbody Ϫ0.008 0.013 enlarged scales, wider than long. Anterior Ventral rows Ϫ0.103 0.284 Transverse rows of ventrals Ϫ0.656 1.541 scales on thigh decreasing in size proxi- Femoral pores 0.510 0.120 mally. Lower legs with three rows of en- Prefemorals Ϫ0.090 0.131 larged, hexagonal scales. Ventral aspect of Prefemoral rows 0.051 Ϫ0.013 hands and feet granular; three enlarged tu- Infratibial rows Ϫ0.018 0.198 Preanals Ϫ0.107 0.161 bercles at base of pollex. Subdigital lamel- Fourth finger lamellae Ϫ0.047 Ϫ0.029 lae single; lamellae under both fourth fin- Fourth toe lamellae Ϫ0.032 Ϫ0.233 gers 14; under left fourth toe 25; under Scales around tail Ϫ0.117 0.340 right fourth toe 24. Dorsals Ϫ0.011 Ϫ0.021 Color in life.—Head brownish green Means dorsally, lemon-green laterally; labial re- Canonical Canonical Species variable 1 variable 2 gions and ventral aspect of head yellowish (Fig. 4). Anterior two-thirds of dorsum C. cryptus 6.202 Ϫ1.123 C. gramivagus 6.672 0.302 brownish green; remaining dorsal areas, C. lemniscatus 9.383 0.270 dorsal aspect of tail, hindlimbs, and fore- C. littoralis Ϫ2.263 Ϫ1.632 arms brown. Anterior aspect of thighs, C. ocellifer Ϫ7.004 Ϫ6.188 proximal aspect of lower legs, lower flanks, C. parecis Ϫ5.579 7.106 and lateralmost longitudinal rows of ventrals vivid blue. Remainder of belly and ventral aspect of limbs and tail immaculate Scales on nape and sides of neck similar white. Anterior aspect of dorsum with two to dorsals. Dorsals and scales on flanks subtle paravertebral rows of brown spots. granular, rounded, smooth, subimbricate; Dorsolateral stripe whitish, continuous 214 scales from nape to base of tail; 111 from supraciliaries to first third of tail; lat- scales in transverse row around midbody eral stripe whitish, continuous from subor- (excluding ventrals). Ventrals large, bital region to hip; lower lateral stripe whit- smooth, rectangular (wider than long), im- ish, interrupted, extending from axilla to bricate, in 31 transverse rows; 10 ventral hip. scales in transverse rows across midbody. Color in fixative (ethanol, after forma- Ventral scales separated from scales on lin).—Head brownish green dorsally, blu- flanks by row of moderately large scales. ish laterally; labial regions and ventral as- Preanal plate with five rows of enlarged pect of head whitish. Anterior two-thirds scales, surrounded laterally and posteriorly of dorsum brownish blue; remaining dor- by small scales. Preanal spurs absent. sal areas, dorsal aspect of tail, hindlimbs, Femoral pores in a continuous row along and forearms brown. Remaining areas of each thigh, medially with a short gap; 13 the body as in life. pores on each side. Variation.—Sexes are dimorphic in March 2003] HERPETOLOGICA 83

FIG. 6.—Scores on the first two canonical discriminant axes based on meristic variables in six species of Cnemidophorus from Brazil. adult color pattern (Fig. 4). Dorsal surface around Vilhena. It is sympatric with the of head brownish green in males, brown polychrotids Anolis meridionalis and Po- in females. Anteriormost two-thirds of lychrus acutirostris, the skinks Mabuya dorsum greenish in males, whereas fe- frenata and M. nigropunctata, the gym- males have only a narrow, green blotch, nophthalmids Micrablepharus atticolus restricted to the first third of the dorsum. and Pantodactylus schreibersii, and the te- Sides of the head lemon-green in males, iids Ameiva ameiva, Kentropyx vanzoi, and but brown in females. Labial regions and Tupinambis merianae. Ecological data on throat yellowish in males and whitish in the species are summarized in Mesquita females. Anterior aspect of thighs, proxi- (2001). The microhabitats used most fre- mal aspect of lower legs, and lower flanks quently by C. parecis are the open ground vivid blue in males and whitish in females. (with grasses) and the area under schrubs Overall, juveniles of both sexes resemble (Fig. 4). Individuals are active during mid- adult females in coloration, but the dor- day, with a mean cloacal temperature of sum is predominantely brown. 38.2 C. The diet consists mainly of ter- Etymology.—The name parecis refers mites, ants, spiders, and beetles. Clutch to the highlands in western Brazil, Cha- size varies from one to two eggs. pada dos Parecis, that separate the Para- guai and Amazon basins and include the RESULTS AND DISCUSSION known range of the species. The Chapada The first two canonical discriminant var- dos Parecis was named after the Paresi in- iables explained approximately 88% of the digenous people, a branch of the Aruak total variation in the meristic variables (Ta- language, who inhabit the region. ble 1). The first canonical variable, repre- Distribution and ecology.—Known only senting mainly a contrast between femoral from the type-locality (Fig. 1). Cnemido- pores and transverse rows of ventrals (Ta- phorus parecis is one of the most abun- ble 1), indicated a clear separation be- dant lizard species in the Cerrado enclaves tween species from Amazonian Savannas 84 HERPETOLOGICA [Vol. 59, No. 1 0.35 1.01 0.30 0.40 0.10 0.82 5.85 0.97 0.49 1.41 1.06 0.81 0.70 0.42 0.90 1.61 2.04 10.75 108) Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ ϭ (1–3) (1–2) (4–5) (2–5) (8–10) (5–12) (8–12) (4–5) n ( C. parecis (11–14) (10–15) (96–127) (29–35) (25–33) (10–14) (13–19) (23–31) (34–47) 8.65 2.04 1.79 4.01 4.19 9.84 4.23 (190–252) 12.10 15.59 12.50 32.07 28.75 11.46 10.44 26.52 40.30 110.40 216.26 0.99 1.08 0.39 0.47 0.33 0.86 8.43 1.10 0.30 1.98 0.87 0.81 0.77 0.40 1.36 2.42 1.66 10.50 101) Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ ϭ (4–8) (3–5) (0–2) (1–2) (3–4) (3–5) (5–8) (8–11) (7–10) n ( (11–16) (10–14) (81–141) (25–31) (16–28) (13–20) (24–38) (21–30) C. ocellifer 9.46 6.11 1.14 1.64 3.88 4.34 7.97 8.75 3.18 (203–260) 10.89 27.70 19.95 30.91 25.75 12.70 98.15 16.60 225.26 0.99 0.80 0.14 0.44 0.20 0.83 5.17 1.38 0.48 1.76 1.08 1.25 1.35 0.46 1.95 1.76 1.93 4.04 53) Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ ϭ (3–5) 1 SD, with range in parentheses. (1–2) (1–2) (3–5) (3–5) (8–10) (4–11) (8–14) (6–13) n ( (13–17) (10–13) (96–123) (30–38) (28–36) (16–24) (29–37) (25–34) Ϯ C. littoralis 1.02 8.28 3.54 7.16 1.64 4.00 4.20 (168–191) ¯ x 10.59 31.87 32.81 28.69 32.58 11.24 10.46 14.79 18.85 174.86 109.98 Species 0.50 1.02 0.49 0.39 0.19 0.88 6.50 0.99 0.00 1.68 0.81 0.83 0.82 0.33 0.96 1.44 1.24 7.79 54) Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Values indicate ϭ (1–2) (1–2) (3–5) (3–6) (8–8) (5–8) (3–5) (9–13) (9–12) n ( (14–16) (15–19) (98–126) (26–31) (44–54) (14–19) (24–31) (24–30) 1.63 8.00 4.03 6.77 1.19 4.00 3.68 (200–231) C. lemniscatus 17.09 28.36 28.78 26.77 10.17 14.28 15.85 48.23 11.40 213.52 107.80 Cnemidophorus. 0.42 0.87 0.29 0.40 0.00 0.25 5.86 0.98 0.00 2.42 1.02 0.74 0.82 0.53 0.98 1.06 1.92 8.55 64) Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ ϭ (1–2) (1–2) (4–4) (3–5) (8–8) (4–9) (3–5) (9–14) n ( (13–16) (13–18) (95–120) (29–33) (40–51) (10–13) (14–21) (27–34) (20–32) 1.90 1.79 4.00 4.97 8.00 4.03 6.58 (210–244) C. gramivagus 15.93 30.95 45.30 11.16 10.95 31.60 28.84 14.10 17.26 225.16 109.50 0.58 0.95 0.57 0.46 0.61 0.57 4.16 1.05 0.11 1.55 1.00 0.83 0.78 0.49 0.80 1.28 1.82 6.25 80) Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ ϭ (1–3) (1–2) (3–5) (2–5) (7–8) (5–9) (3–4) (9–13) (7–12) n ( (14–17) (13–17) (98–118) (26–32) (38–48) (14–18) (28–33) (22–30) C. cryptus 1.56 7.99 9.50 3.61 6.66 1.70 4.14 3.29 (202–227) 15.08 29.20 30.60 27.01 14.27 16.31 43.42 11.14 211.88 106.38 2.—Meristic characters of six species of ABLE T Variables Supralabials Infralabials Chinshields Gular folds Supraoculars Parietals Scales around midbody Ventral rows Transverse rows of ventrals Femoral pores Fourth finger lamellae Fourth toe lamellae Scales around tail Dorsals Prefemorals Prefemoral rows Infratibial rows Preanals March 2003] HERPETOLOGICA 85

TABLE 3.—Stepwise discriminant analysis of meristic characters from six species of Cnemidophorus, with error rate estimates based on cross validation.

Step Variable entered FPError rate 1 Femoral pores 3022.05 Ͻ0.0001 0.1761 2 Scales around tail 861.85 Ͻ0.0001 0.1239 3 Dorsals 264.27 Ͻ0.0001 0.0957 4 Ventral rows 72.79 Ͻ0.0001 0.0478 5 Fourth toe lamellae 50.63 Ͻ0.0001 0.0457 6 Transverse rows of ventrals 38.96 Ͻ0.0001 0.0457 7 Infralabials 32.16 Ͻ0.0001 0.0413 8 Parietals 28.97 Ͻ0.0001 0.0283 9 Supralabials 21.39 Ͻ0.0001 0.0283 10 Infratibial rows 14.23 Ͻ0.0001 0.0196 11 Prefemorals 8.23 Ͻ0.0001 0.0196 12 Chinshields 8.04 Ͻ0.0001 0.0217 13 Fourth ﬁnger lamellae 7.74 Ͻ0.0001 0.0217 14 Gular fold 6.97 Ͻ0.0001 0.0217 15 Scales around midbody 5.99 Ͻ0.0001 0.0239 16 Preanals 3.63 0.0031 0.0239 17 Supraoculars 3.05 0.0101 0.0239

(C. cryptus, C. gramivagus, and C. lem- lowed by scales around tail and dorsals, niscatus) and the remaining species (C. lit- whose addition improved the classification toralis, C. ocellifer, and C. parecis) (Fig. criterion to 87.6% and 90.4%, respectively 6). Amazonian Savanna species, with pos- (Table 3). To determine if important vari- itive means on the first canonical variable ables were excluded from the analysis be- (Table 1), are characterized by a high cause of correlations among the original number of femoral pores and fewer trans- variables (Tabachnick and Fidell, 2001), verse rows of ventrals (Table 2). we removed femoral pores and repeated The second canonical discriminant var- the analysis. Scales around the tail were iable, representing a contrast between then entered in the first step, correctly transverse rows of ventrals versus suprala- classifying 56.3% of the lizards (error rate bials and fourth toe lamellae (Table 1), in- ϭ 0.437, F ϭ 895.34, P Ͻ 0.001). We then dicated a clear separation among the three removed scales around the tail and re- species that occur south of the Amazon peated the analysis. This time, infralabials River (Fig. 6). Cnemidophorus parecis, was the first variable to be selected, cor- with the highest mean on the second ca- rectly classifying about 60% of the lizards nonical variable (Table 1), is characterized (error rate ϭ 0.3978, F ϭ 525.49, P Ͻ by numerous transverse rows of ventrals 0.001). These analyses indicated that fem- and few supralabials and fourth toe lamel- oral pores and scales around the tail are lae (Table 2). Cnemidophorus littoralis, the meristic characters that best discrimi- with the lowest mean on the second ca- nated the species of Cnemidophorus and nonical variable (Table 1), is characterized that their discriminatory power was high. by few transverse rows of ventrals and nu- The naı¨ve Bayesian model indicated merous supralabials and fourth toe lamel- that C. parecis determined high condition- lae (Table 2). Cnemidophorus ocellifer al probabilities (Ͼ95%) of 12 categorical (and the Amazonian Savanna species) has variables: dorsolateral fields absent, verte- intermediate values for these variables. bral field spotted, paravertebral lines ab- The stepwise discriminant analysis of sent, lateral line continuous, lower lateral meristic characters selected 17 variables as line interrupted, vertebral line absent, lat- the most powerful discriminators among eral spots absent, hindlimb spots absent, the species of Cnemidophorus (Table 3). preanal spur absent, supraoculars contact- Femoral pores was selected first, correctly ing frontal and parietals, ventral caudals classifying 82.4% of the individuals, fol- smooth, and dorsal caudals keeled (Table 86 HERPETOLOGICA [Vol. 59, No. 1 (44.14%) L (91.07%) C (84.68%) I (95.50%) Ͼ S (99.09%) L (91.96%) A (96.43%) A (97.32%) A (97.32%) A (99.09%) A (99.09%) A (99.09%) C (96.69%) K (99.09%) Sp (96.43%) IFP (95.58%) ¨ve Bayesian model. (98.08%) L 71.43% I (56.19%) P (90.29%) L (80.00%) A (56.73%) A (94.29%) C (64.42%) C (52.88%) D (46.67%) D (43.81%) S (99.03%) A (98.06%) A (99.03%) K (99.03%) Ͼ Co (51.89%) assuming a naı errupted; K: keeled; S: smooth; Co: complete; IFP: incomplete, Cnemidophorus, (80.36%) L 50.88% I (94.74%) I (66.07%) I (89.47%) C (82.14%) Ͼ D (92.98%) D (64.91%) D (64.91%) S (98.18%) A (98.18%) A (98.18%) A (98.18%) C (96.43%) K (98.18%) IFP (58.62%) Species (87.72%) L 48.28% P (51.79%) A (57.14%) C (94.83%) C (89.47%) C (73.68%) C (64.91%) C (93.10%) Ͼ D (70.69%) D (94.83%) D (93.10%) P (98.21%) K (98.21%) K (98.21%) IFP (93.22%) (97.01%) I (53.73%) P (93.94%) A (69.70%) A (91.04%) A 95.59% C (91.18%) D (94.12%) D (89.71%) A (95.59%) A (96.97%) C (95.52%) K (98.48%) K (98.48%) Ͼ D (95.59%) IFP (94.20%) : contact between first pair of chinshields greater than half of their lengths. (96.39%) Ͼ A (67.86%) A (64.63%) C. cryptus C. gramivagus C. lemniscatus C. littoralis C. ocellifer C. parecis C (65.06%) D (55.95%) D (71.43%) A (98.78%) A (98.78%) C (96.39%) C (96.43%) C (97.59%) K (98.78%) K (98.78%) Ͼ D (95.24%) D (96.43%) IFP (95.29%) 4.—Conditional probabilities of states for qualitative characters for six Brazilian species of Variables ABLE T Upper lateral stripes Lower lateral stripes Vertebral line Lateral spots Hindlimb spots Chinshield contact Lower lateral fields Upper lateral fields Dorsolateral fields Vertebral field Paravertebral stripes Dorsolateral stripes Note: values in bold indicate conditional probabilities greater than 95%. D: dark; L: light; Sp: spotted; A: absent; P: present; C: continuous; I: int supraoculars contacting frontals and parietals; Preanal spur Semicircles Ventral caudals Dorsal caudals March 2003] HERPETOLOGICA 87

4). Further, only C. parecis determined 80% of each rural property must be pre- high conditional probabilities of dorsolat- served (Reserva Legal), but a state law eral fields absent, vertebral field spotted, passed recently in Rondoˆnia reduced this and paravertebral lines absent (Table 4). percentage up to 20% in highly productive The analysis also indicated that species regions, favoring the establishment of from Amazonian Savannas can be clearly large properties. The Cerrado patches separated from species south of the Ama- around Vilhena are critically threatened by zon River by the presence of keeled sub- the expansion of soybean plantations, fa- caudals in the former (Table 4). vored by the gentle topography that aids Our results revealed a greater similarity mechanization, the low risks of drought, between C. parecis and the other species and the recent development of two expor- ranging south of the Amazon River (C. lit- tation corridors. The first is the Ferronorte toralis and C. ocellifer). We also verified Railroad, connecting Santa Fe´ do Sul in that, besides the presence of anal spurs in Saõ Paulo to Alto Taquari in Mato Grosso. males, the species of the C. lemniscatus The other is the Hidrovia do Rio Madeira, complex can also be distinguished from connecting Porto Velho in Rondoˆnia state their congeneric southern relatives by the to Itacoatiara in Amazonas state, along the presence of keeled subcaudals, a larger Madeira and Amazon Rivers. Therefore, number of femoral pores, and fewer trans- C. parecis is apparently at a critical risk of verse rows of ventrals. This distinction sug- becoming extinct within a very short pe- gests that C. parecis may be a closer rel- riod of time. Because the presumed range ative of the southern species of Cnemido- of C. parecis is not included in any con- phorus. A number of palynological and servation unit, being entirely within pri- sedimentary studies indicate that, in Ron- vately owned land, urgent measures are doˆnia, open vegetation replaced the forest needed to protect the species. in the recent past, approximately 3000 ybp Acknowledgments.—This work was developed un- (Absy and Van der Hammen, 1976; de der the project ‘‘Estrutura e dinaˆmica da biota de Freitas et al., 2001; Sifeddine et al., 2001; isolados naturais e antro´picos de Cerrado: liço˜es para Van der Hammen and Absy, 1994). There- a biologia da conservaçaõ,’’ funded by Programa Na- fore, present day enclaves of Cerrado in cional da Diversidade Biolo´gica-PRONABIO, MMA- MCT-CNPq-GEF-BIRD. S. F. Balbino, C. G. Batis- Rondoˆnia may be relicts of a past open ta, C. A. Freitas, D. Diniz, the staff of IBAMA at vegetation landscape that ranged from Vilhena, Joećio Costa, Aparecido Donadoni, Vicente Central Brazil to Rondoˆnia. Following the de Paulo Campos Godinho, and M. M. Utumi assist- breakup of the connections between Ron- ed in fieldwork. We thank Dr. Wagner Teixeira da Silva for assistance with Bayesian statistics, Zenilton doˆnia and the Central Brazil open land- J. G. Miranda for drawing the holotype, and three scapes, C. parecis presumably differenti- anonymous reviewers for their insightful comments ated as a consequence of the geographical on the manuscript. This work was supported by grad- isolation. This view is supported by the uate student fellowships from Coordenaçaõ de Aper- presence of a depauperate Cerrado lizard feiçoamento de Pessoal de Nı´vel Superior to A. A. Garda (# 146980/99-9), P. H. Valdujo, D. O. Mes- fauna in the Rondoˆnia enclaves (Vanzolini, quita, and G. H. C. Vieira; from Conselho Nacional 1986; Vitt and Caldwell, 1993). However, do Desenvolvimento Cientı´fico e Tecnolo´gico to A. the affinities of C. parecis can only be as- K. Pe´res, Jr. (# 146980/99-9) and H. C. Wiederhecker certained with confidence through rigor- (# 141596/00-7); and by a research fellowship from Conselho Nacional de Desenvolvimento Cientı´fico e ous phylogenetic analysis. Tecnolo´gico–CNPq to G. R. Colli (# 302343/88-1). Apparently, C. parecis is an endemic of a small group of Cerrado patches with LITERATURE CITED sandy soils around the city of Vilhena. ABSY, M. L., AND T. VAN DER HAMMEN. 1976. Some Rondoˆnia is the most deforestated state in paleoecological data from Rondoˆnia, southern part Amazonia, having lost about one-third of of the Amazon basin. Acta Amazonica 6:293–299. its total vegetation cover, whereas the mu- A´ VILA-PIRES, T. C. S. 1995. Lizards of Brazilian Ama- zonia (Reptilia: Squamata). Zoologische Verhande- nicipality of Vilhena has less than 10% of lingen, Leiden 1995:3–706. the original vegetation. The Brazilian leg- BERNARDO,J.M.,AND A. F. M. SMITH. 2000. Bayes- islature (Co´digo Florestal) stipulates that ian Theory. Wiley, New York, New York, U.S.A. 88 HERPETOLOGICA [Vol. 59, No. 1

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