A CRITICALLY ENDANGERED NEW SPECIES OF CNEMIDOPHORUS (SQUAMATA, TEIIDAE) FROM A CERRADO ENCLAVE IN...

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Herpetologica, 59(1), 2003, 76–88q 2003 by The Herpetologists’ League, Inc.

A CRITICALLY ENDANGERED NEW SPECIES OFCNEMIDOPHORUS (SQUAMATA, TEIIDAE) FROM A CERRADO

ENCLAVE IN SOUTHWESTERN AMAZONIA, BRAZIL

GUARINO R. COLLI1,3, GABRIEL C. COSTA1, ADRIAN A. GARDA1, KATIA A. KOPP2,DANIEL O. MESQUITA1, AYRTON K. PERES, JR.1, PAULA H. VALDUJO1,

GUSTAVO H. C. VIEIRA1, AND HELGA C. WIEDERHECKER1

1Departamento de Zoologia, Universidade de Brasılia, 70910-900 Brasılia, DF Brazil2Departamento de Ciencias Biologicas, 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, Rondonia state, Brazil. This species is apparently endemic to Cerrado enclavesin the vicinity of the city of Vilhena, a region under intensive anthropic pressure due to the expansionof soybean plantations. A discriminant analysis indicated that femoral pores and scales around thetail are the best discriminators among Brazilian species of Cnemidophorus. A naıve Bayesian networkconstructed with categorical (mostly coloration) variables indicated that the new species had highconditional probabilities of dorsolateral fields absent, vertebral field spotted, and paravertebral linesabsent. The analyses revealed clear distinctions between species of Cnemidophorus that range northand 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 latePleniglacial. 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; Rondonia

RANGING from northern United Statesto central Argentina, Cnemidophorus hasthe widest geographic distribution amongall teiid genera (Wright, 1993). Currently,57 species are known in the genus (Uetzet al., 1995). All South American speciesof Cnemidophorus are assigned to the C.lemniscatus species group (Burt, 1931;Wright, 1993), which comprises 17 spe-cies: C. arenivagus Markezich, Cole, andDessauer, 1997; C. arubensis Van Lidth deJeude, 1887; C. cryptus Cole and Des-sauer, 1993; C. gramivagus McCrystal andDixon, 1987; C. lacertoides Dumeril andBibron, 1839; C. leachei Peracca, 1897; C.lemniscatus (Linnaeus, 1758); C. littoralisRocha, Araujo, Vrcibradic, and Costa,2000; C. longicaudus (Bell, 1843); C. mu-rinus (Laurenti, 1768); C. nativo Rocha,Bergallo, and Peccinini-Seale, 1997; C. ni-gricolor Peters, 1873; C. ocellifer (Spix,1825); C. pseudolemniscatus Cole andDessauer, 1993; C. serranus Cei and Mar-tori, 1991; C. vacariensis Feltrim and

3 CORRESPONDENCE: e-mail, [email protected]

Lema, 2000; and C. vanzoi (Baskin andWilliams, 1966).

That about 50% of the species in theCnemidophorus lemniscatus group hasbeen described in the last 15 yr testifies tothe paucity of knowledge of the alpha-leveldiversity in the neotropics and the growingtaxonomic activity in the area. Indeed, un-til recently, all populations of Cnemido-phorus in northern South America wereassigned to a single species, C. lemnisca-tus. Several studies have indicated that C.ocellifer is also a complex of species (e.g.,Rocha et al., 1997, 2000; Rodrigues, 1987).In addition, the availability of tools toprobe molecular markers has only recentlyallowed the detection of morphologicallycryptic species, such as C. cryptus (e.g.,Cole and Dessauer, 1993; Sites et al.,1990).

During the course of a study on theCerrado enclaves in Rondonia, Brazil, wecollected a relatively large series of an un-described species of Cnemidophorus. Afew individuals of this species had beenpreviously collected by Vitt and Caldwell(1993) during a survey sponsored by the

March 2003] 77HERPETOLOGICA

FIG. 1.—Vegetation map of type locality of Cnemi-dophorus parecis.

Polonoroeste program along the BR-364road in Rondonia (Nascimento et al., 1988;Vanzolini, 1986). The individuals were re-ferred by Vanzolini both to Cnemidopho-rus ocellifer (Vanzolini, 1986) and to anunnamed member of the ocellifer speciesgroup (sic) (P. E. Vanzolini, personal com-munication, in Vitt and Caldwell, 1993).Herein, we describe this new species,making detailed comparisons with otherBrazilian congeneric species.

MATERIALS AND METHODS

We collected lizards with a shotgun, pit-fall traps, and drift fences on 4–10 Novem-ber 1998 and 21 August–22 September1999, around the city of Vilhena (128 439S, 608 079 W), Rondonia state, Brazil (Fig.1). The vegetation in the study area con-sists of a mosaic of Cerrado enclaves (Ei-ten, 1972) within a matrix of Terra FirmeAmazonian Forest (Pires and Prance,1985) and human-modified areas consist-ing of soybean plantations and groves ofpine trees. We found lizards exclusively inCerrado enclaves with sandy soils aroundthe city of Vilhena, even though we sam-pled the largest patches of Cerrado en-claves throughout Rondonia, includingthose around the cities of Pimenta Buenoand Guajara-Mirim.

We obtained data on C. cryptus, C. gra-

mivagus, C. lemniscatus, C. littoralis, andC. ocellifer from specimens housed in theColecao Herpetologica da Universidade deBrasılia (CHUNB) (Appendix I). The firstthree species occur predominantly in Am-azonian Savannas north of the AmazonRiver (e.g., Avila-Pires, 1995; Cole andDessauer, 1993), whereas the last two oc-cur exclusively south of the Amazon (e.g.,Cei, 1993; Rocha et al., 2000). It shouldbe noted that C. cryptus is a provisionalname for a complex of parthenogenetic,unisexual populations in the Amazon Basinand that much work is necessary to clarifytheir taxonomic status. We recorded thefollowing meristic variables in each speci-men: supralabials (number of enlargedscales along the upper jaw, total on bothsides), infralabials (number of enlargedscales along the lower jaw, total on bothsides), chinshields (most anterior pair ofchinshields separated from infralabials byrow of small scales), gular folds (numberof folds in gular region), supraoculars(number of supraoculars on right side), pa-rietals (number of parietals plus interpa-rietal scale), scales around midbody(counted midway between fore- and hind-limbs, excluding ventrals), transverse rowsof ventrals (counted along the midline,from gular fold to anterior margin of hind-limbs), ventrals in transverse row (countedmidway between fore- and hindlimbs),femoral pores (total number on bothsides), prefemorals (number of enlargedscales on anterior aspect of thigh, countedmidway between the hip and the knee, ona row from femoral pores to granules ondorsal aspect of thigh), prefemoral rows(counted from hip to knee), infratibials(number of enlarged scales on longitudinalrow from knee to base of first metatarsal),preanals (number of enlarged scales onpreanal plate, from level of medialmostfemoral pores to vent), fourth finger la-mellae (counted under the finger), fourthtoe lamellae (counted under the toe),scales around tail (counted on fifth trans-verse row), and dorsals (counted along themidline, from occiput to first transverserow of scales around tail).

We used a discriminant function analy-sis (Tabachnick and Fidell, 2001) to inves-

78 [Vol. 59, No. 1HERPETOLOGICA

tigate differences among species in meris-tic characters. To identify the most pow-erful meristic discriminators of the speciesof Cnemidophorus, we used a stepwisediscriminant function analysis of meristiccharacters (Tabachnick and Fidell, 2001),using the METHOD 5 STEPWISE op-tion in PROC STEPDISC of SAS (SAS In-stitute Inc., 1988). We evaluated linear dis-criminant functions through posteriorprobability error-rate estimates, based oncrossvalidation (SAS Institute Inc., 1988).

We also recorded the following categor-ical variables: lower lateral fields (LLF,one on each side, area between ventralscales and lower lateral stripes: absent,light, spotted, or dark), upper lateral fields(ULF, one on each side, area between up-per lateral and dorsolateral stripes: absent,light, spotted, or dark), dorsolateral fields(DLF, one on each side, dark area be-tween dorsolateral and paravertebralstripes: absent, light, spotted, or dark), ver-tebral fields (VTF, one on each side, mid-dorsal area between paravertebral stripes:absent, light, spotted, or dark), vertebralline (VTL, light stripe from interparietalscale to base of tail: absent, interrupted,continuous, or double), paravertebralstripes (PVS, one on each side, light stripefrom parietal scale to first third of tail, be-tween vertebral and dorsolateral fields: ab-sent, interrupted, or continuous), dorsolat-eral stripes (DLS, one on each side, lightstripe from superciliaries to first third oftail, between dorsolateral and upper lateralfields: absent, interrupted, or continuous),upper lateral stripes (ULS, one on eachside, light stripe from suborbital region tohindlimb, between upper and lower lateralfields: absent, interrupted, or continuous),lower lateral stripes (LLS, one on eachside, light stripe from axilla to hip, be-tween lower lateral field and ventrals: ab-sent, interrupted, or continuous), lateralspots (LTS, rounded light areas on flanks:absent, present), hindlimb spots (HLS,rounded light areas on hindlimbs: absent,present), chinshield contact (CHS, degreeof contact between most anterior pair ofchinshields: no contact, contact smallerthan half of their lengths, or contact great-er than half of their lengths), semicircles

(SMC, degree of contact between supra-oculars and medial head scales: no contact,no contact with semicircles isolating firstsupraocular, supraoculars contacting fron-tal and parietals, supraoculars contactingfrontal, supraoculars contacting parietals),dorsal caudals (DCA, keels on dorsal, cau-dal scales, from most anterior third of tail:absent, present), ventral caudals (VCA,keels on ventral, caudal scales, from mostanterior third of tail: absent, present), andpreanal spur (PAS: absent, present).

To model the conditional dependenciesof each categorical variable within eachspecies, we constructed a naıve Bayesianmodel using the software UNBARB (avail-able at ftp://ftp.cic.unb.br/pub/cic/wagner/software/Bnets/UnBARB.zip). In thismodel (Fig. 2), the variable ‘‘species’’ in-fluences the chance of the occurrence ofstates of each categorical variable, assum-ing that categorical variables are condition-ally independent for a given species andthat the joint distribution of all variablessatisfies the following:

k

P(S, C , . . . , C ) 5 P(C z S) P(S),P1 k i1 2i51

where S is species, C is a categorical vari-able, P(Ci zS) is the posterior probability ofCi given S, and P(S) is the prior probabilityof S (Cowell et al., 1999). We estimatedP(S) and P(Ci z S) using Bayesian statistics(Bernardo and Smith, 2000; DeGroot,1970) and the software Hugin Lite v. 5.3(Jensen and Nielsen, 1999). Throughoutthe text we report means 61 SD.

SPECIES DESCRIPTION

Cnemidophorus parecis sp. nov.Holotype.—CHUNB 09811 (Figs. 3, 4),

adult male, from Fazenda Cachoeira (128329 070 S, 608 259 360 W), 50 km NW ofVilhena, Estado de Rondonia, Brazil, ele-vation about 612 m, collected on 5 Novem-ber 1998 by Guarino R. Colli and students.

Paratopotypes.—(107) All specimensfrom Cerrado enclaves around Vilhena:CHUNB 09762–810, 09812–14, collectedon November 1998 by same collectors;CHUNB 11651–703, 14229–30 collected onAugust–September 1999 by same collectors.


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

Diagnosis.—A species distinguishedfrom all other members of the lemniscatusgroup by the following combination ofcharacters: (1) large size, maximum SVL90 mm; (2) 34–47 scales around tail; (3)25–33 femoral pores; (4) 190–252 dorsals;(5) dorsolateral fields absent; (6) vertebralfield spotted; (7) paravertebral stripes ab-sent (Fig. 4); and (8) preanal spurs absent.Cnemidophorus parecis differs from C.lemniscatus in having 11–14 supralabials(14–16 in C. lemniscatus), 10–15 infrala-bials (15–19), 25–33 femoral pores (44–54), 34–47 scales around tail (24–30), dor-solateral fields absent (dark), vertebralfield spotted (dark), paravertebral stripesabsent (continuous), vertebral line absent(two continuous stripes) (Fig. 5), hindlimbspots absent (present), and ventral caudalssmooth (keeled). Cnemidophorus parecisdiffers from C. gramivagus in having 25–33 femoral pores (40–51 in C. gramiva-gus), 34–47 scales around tail (20–32),lower lateral fields light (absent), upperlateral fields light (dark), dorsolateral fieldsabsent (dark), vertebral field spotted(dark), paravertebral stripes absent (con-tinuous) (Fig. 5), lower lateral stripes in-terrupted (absent), hindlimb spots absent

(present), and ventral caudals smooth(keeled). Cnemidophorus parecis differsfrom C. cryptus in having 11–14 suprala-bials (14–17 in C. cryptus), 8–10 trans-verse rows of ventrals (7–8), 25–33 femoralpores (38–48), 34–47 scales around tail(22–30), upper lateral fields light (dark),dorsolateral fields absent (dark), paraver-tebral stripes absent (continuous) (Fig. 5),and ventral caudals smooth (keeled).Cnemidophorus parecis differs from C. lit-toralis in having 34–47 scales around tail(25–34 in C. littoralis), upper lateral fieldslight (dark), paravertebral stripes absent(interrupted), vertebral line absent (inter-rupted) (Fig. 5), and dorsal caudals keeled(smooth). Cnemidophorus parecis differsfrom C. ocellifer in having 8–10 transverserows of ventrals (5–8 in C. ocellifer), 25–33 femoral pores (16–28), 34–47 scalesaround tail (21–30), and lateral spots ab-sent (present) (Fig. 5).

Description of holotype.—Rostral pen-tagonal, higher than wide, visible fromabove, bordered posteriorly by nasals. Na-sals in contact along midline, each nasaldivided by an oblique suture. Nostril inlower part of suture, directed posterolat-erally, longer than tall. Frontonasal sub-


FIG. 3.—Cnemidophorus parecis, holotype,CHUNB 09811, adult male, SVL 86 mm.

FIG. 4.—Top: Cerrado enclave in Vilhena, Ron-donia, Brazil. Middle: Adult female of Cnemidopho-rus parecis. Bottom: Adult male of C. parecis (holo-type, CHUNB 09811).

hexagonal, suture with nasals and lorealsforming a semicircle, sutures with prefron-tals forming a wide angle. Prefrontalsquadrilateral, with medial suture twice aslong as that between nasals, in contact lat-erally with loreal, first supraciliary, andfirst supraocular. Frontal approximatelypentagonal, longer than wide, and wideranteriorly. Sutures with prefrontals and su-praoculars roundish, those with frontopar-ietals forming a straight line. Frontal incontact laterally with first, second, andthird supraoculars. Frontoparietals pentag-onal, longer than wide, with long, straightmedial suture; in contact laterally withthird supraocular and small scales border-ing supraocular region posteriorly. Inter-parietal hexagonal, bordered at each sideby hexagonal parietals, divided into two ir-

regular scales by an oblique suture. Occip-ital scales irregular, posterior to interpari-etal and parietals, variable in size, includ-ing two moderately enlarged, medialscales, separated by a smaller scale. Foursupraoculars on each side, second andthird largest, followed posteriorly by groupof small scales. Second, third, and fourthsupraoculars separated from supraciliariesby row of small scales. Supraciliaries sevenon each side, highest anteriorly, third larg-est, others subequal. Loreal single, large,


FIG. 5.—Dorsal coloration patterns in six species of Cnemidophorus from Brazil. (A) C. littoralis, CHUNB08308, (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-tal, first supraciliary, frenocular, first sub-ocular, and second, third, and fourth su-pralabials on the left side, and third andfourth supralabials on the right side. Fren-ocular narrow, higher than wide, in contactwith subocular, loreal, first supraciliary,and small scales in ocular region. Subocu-lars three, large, second largest, all in con-tact with supralabials. A continuous keelruns from frenocular to about midlengthof second subocular. Postoculars small, ar-ranged in two rows with 3–4 larger scales.Lower eyelid with semiopaque disc,formed by transversally enlarged, convexscales. Supralabials six on each side, fifthand sixth below center of eye; followed byseries of small scales extending to com-missure of mouth. Temporal region withirregular scales, granular centrally, mod-erately enlarged peripherally. Ear openinglarge, oblique, with smooth margins, an-terior margin forming a semicircle, poste-rior margin straight. Tympanum recessedin a short auditory meatus. All dorsal andlateral head scales juxtaposed, smooth.

Symphysal anteriorly ellipsoid, in pos-

terior contact with first infralabials andpostsymphysal, forming two wide angles.Postsymphysal single, pentagonal, in con-tact with first and second infralabials, fol-lowed by three pairs of enlarged chin-shields. First pair of chinshields in amplecontact along midline, first and secondpairs in contact with infralabials, third pairseparated from infralabials by row of smallscales, followed posteriorly by enlargedscales. Medial scales on chin small,smooth, elongate, arranged in longitudinal,posteriorly divergent rows; increasing insize and becoming roundish posteriorly.Infralabials six, followed posteriorly by se-ries of small scales extending to commis-sure of mouth; fifth infralabial below cen-ter of eye. Gular region divided in two ar-eas: anterior region with enlarged, round-ed scales, in transverse rows, delimitedposteriorly by line uniting lower margin ofear openings; posterior region coveredwith granules, in transverse rows, bor-dered posteriorly by antegular fold. Gularand antegular folds marked by granules;scales between the two folds larger, irreg-ular, slightly imbricate.


TABLE 1.—Canonical discriminant analysis of meris-tic characters of six species of Cnemidophorus.

Canonicalvariable

Canonical statistics

Eigen-value

Cumula-tive

propor-tion r2 F P

12

42.43821.087

0.5880.879

0.9770.955

165.06101.12

,0.0001,0.0001

Variable

Raw canonical coefficients

Canonicalvariable 1

Canonicalvariable 2

SupralabialsInfralabialsChinshieldsGular foldSupraocularsParietalsScales around midbody

0.0980.3120.120

20.1260.291

20.18320.008

20.3420.2110.464

20.09120.18920.121

0.013Ventral rowsTransverse rows of ventralsFemoral poresPrefemoralsPrefemoral rowsInfratibial rowsPreanalsFourth finger lamellae

20.10320.656

0.51020.090

0.05120.01820.10720.047

0.2841.5410.1200.131

20.0130.1980.161

20.029Fourth toe lamellaeScales around tailDorsals

20.03220.11720.011

20.2330.340

20.021

Species

Means

Canonicalvariable 1

Canonicalvariable 2

C. cryptusC. gramivagusC. lemniscatusC. littoralisC. ocelliferC. parecis

6.2026.6729.383

22.26327.00425.579

21.1230.3020.270

21.63226.188

7.106

Scales on nape and sides of neck similarto dorsals. Dorsals and scales on flanksgranular, rounded, smooth, subimbricate;214 scales from nape to base of tail; 111scales in transverse row around midbody(excluding ventrals). Ventrals large,smooth, rectangular (wider than long), im-bricate, in 31 transverse rows; 10 ventralscales in transverse rows across midbody.Ventral scales separated from scales onflanks by row of moderately large scales.Preanal plate with five rows of enlargedscales, surrounded laterally and posteriorlyby small scales. Preanal spurs absent.Femoral pores in a continuous row alongeach thigh, medially with a short gap; 13pores on each side.

Scales on base of tail rectangular, longerthan wide, smaller than ventrals, keeleddorsally, smooth ventrally, in transverserows. All transverse rows continuousaround tail, except first two rows incom-plete ventrally. Tail scales becoming longerand narrower posteriorly; subcaudal scalesbecoming keeled posteriorly. Tail broken,regenerated from 37th transverse row ofscales. Limbs with large, smooth, imbri-cate scales on dorsal aspect of upper arms,anterodorsal aspect of forearms, antero-ventral aspect of thighs, and ventral aspectof lower legs; elsewhere scales small, gran-ular. Larger scales on upper arms in lon-gitudinal rows. Forearms with one row ofenlarged scales, wider than long. Anteriorscales on thigh decreasing in size proxi-mally. Lower legs with three rows of en-larged, hexagonal scales. Ventral aspect ofhands and feet granular; three enlarged tu-bercles at base of pollex. Subdigital lamel-lae single; lamellae under both fourth fin-gers 14; under left fourth toe 25; underright fourth toe 24.

Color in life.—Head brownish greendorsally, lemon-green laterally; labial re-gions and ventral aspect of head yellowish(Fig. 4). Anterior two-thirds of dorsumbrownish green; remaining dorsal areas,dorsal aspect of tail, hindlimbs, and fore-arms brown. Anterior aspect of thighs,proximal aspect of lower legs, lower flanks,and lateralmost longitudinal rows of ven-trals vivid blue. Remainder of belly andventral aspect of limbs and tail immaculatewhite. Anterior aspect of dorsum with twosubtle paravertebral rows of brown spots.Dorsolateral stripe whitish, continuousfrom supraciliaries to first third of tail; lat-eral stripe whitish, continuous from subor-bital region to hip; lower lateral stripe whit-ish, interrupted, extending from axilla tohip.

Color in fixative (ethanol, after forma-lin).—Head brownish green dorsally, blu-ish laterally; labial regions and ventral as-pect of head whitish. Anterior two-thirdsof dorsum brownish blue; remaining dor-sal areas, dorsal aspect of tail, hindlimbs,and forearms brown. Remaining areas ofthe body as in life.

Variation.—Sexes are dimorphic in


FIG. 6.—Scores on the first two canonical discriminant axes based on meristic variables in six species ofCnemidophorus from Brazil.

adult color pattern (Fig. 4). Dorsal surfaceof head brownish green in males, brownin females. Anteriormost two-thirds ofdorsum greenish in males, whereas fe-males have only a narrow, green blotch,restricted to the first third of the dorsum.Sides of the head lemon-green in males,but brown in females. Labial regions andthroat yellowish in males and whitish infemales. Anterior aspect of thighs, proxi-mal aspect of lower legs, and lower flanksvivid blue in males and whitish in females.Overall, juveniles of both sexes resembleadult females in coloration, but the dor-sum is predominantely brown.

Etymology.—The name parecis refersto the highlands in western Brazil, Cha-pada dos Parecis, that separate the Para-guai and Amazon basins and include theknown range of the species. The Chapadados Parecis was named after the Paresi in-digenous people, a branch of the Aruaklanguage, who inhabit the region.

Distribution and ecology.—Known onlyfrom the type-locality (Fig. 1). Cnemido-phorus parecis is one of the most abun-dant lizard species in the Cerrado enclaves

around Vilhena. It is sympatric with thepolychrotids Anolis meridionalis and Po-lychrus acutirostris, the skinks Mabuyafrenata and M. nigropunctata, the gym-nophthalmids Micrablepharus atticolusand Pantodactylus schreibersii, and the te-iids Ameiva ameiva, Kentropyx vanzoi, andTupinambis merianae. Ecological data onthe species are summarized in Mesquita(2001). The microhabitats used most fre-quently by C. parecis are the open ground(with grasses) and the area under schrubs(Fig. 4). Individuals are active during mid-day, with a mean cloacal temperature of38.2 C. The diet consists mainly of ter-mites, ants, spiders, and beetles. Clutchsize varies from one to two eggs.

RESULTS AND DISCUSSION

The first two canonical discriminant var-iables explained approximately 88% of thetotal variation in the meristic variables (Ta-ble 1). The first canonical variable, repre-senting mainly a contrast between femoralpores and transverse rows of ventrals (Ta-ble 1), indicated a clear separation be-tween species from Amazonian Savannas


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6.50

(98–

126)

28.3

66

0.99

(26–

31)

8.00

60.

00(8

–8)

48.2

36

1.68

(44–

54)

109.

986

5.17

(96–

123)

31.8

76

1.38

(30–

38)

8.28

60.

48(8

–10)

32.5

86

1.76

(28–

36)

98.1

56

8.43

(81–

141)

27.7

06

1.10

(25–

31)

7.97

60.

30(5

–8)

19.9

56

1.98

(16–

28)

110.

406

5.85

(96–

127)

32.0

76

0.97

(29–

35)

9.84

60.

49(8

–10)

28.7

56

1.41

(25–

33)

Pre

fem

oral

s

Pre

fem

oral

row

s

Infr

atib

ial

row

s

Pre

anal

s

6.66

61.

00(5

–9)

11.1

46

0.83

(9–1

3)9.

506

0.78

(7–1

2)3.

616

0.49

(3–4

)

6.58

61.

02(4

–9)

11.1

66

0.74

(10–

13)

10.9

56

0.82

(9–1

4)4.

036

0.53

(3–5

)

6.77

60.

81(5

–8)

11.4

06

0.83

(9–1

3)10

.17

60.

82(9

–12)

4.03

60.

33(3

–5)

7.16

61.

08(4

–11)

11.2

46

1.25

(8–1

4)10

.46

61.

35(6

–13)

3.54

60.

46(3

–5)

6.11

60.

87(4

–8)

9.46

60.

81(8

–11)

8.75

60.

77(7

–10)

3.18

60.

40(3

–5)

8.65

61.

06(5

–12)

11.4

66

0.81

(10–

14)

10.4

46

0.70

(8–1

2)4.

236

0.42

(4–5

)F

ourt

hfin

ger

lam

ella

e

Fou

rth

toe

lam

ella

e

Scal

esar

ound

tail

Dor

sals

16.3

16

0.80

(14–

18)

30.6

06

1.28

(28–

33)

27.0

16

1.82

(22–

30)

211.

886

6.25

(202

–227

)

17.2

66

0.98

(14–

21)

31.6

06

1.06

(27–

34)

28.8

46

1.92

(20–

32)

225.

166

8.55

(210

–244

)

15.8

56

0.96

(14–

19)

28.7

86

1.44

(24–

31)

26.7

76

1.24

(24–

30)

213.

526

7.79

(200

–231

)

18.8

56

1.95

(16–

24)

32.8

16

1.76

(29–

37)

28.6

96

1.93

(25–

34)

174.

866

4.04

(168

–191

)

16.6

06

1.36

(13–

20)

30.9

16

2.42

(24–

38)

25.7

56

1.66

(21–

30)

225.

266

10.5

0(2

03–2

60)

15.5

96

0.90

(13–

19)

26.5

26

1.61

(23–

31)

40.3

06

2.04

(34–

47)

216.

266

10.7

5(1

90–2

52)


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

Step Variable entered F P Error rate

123456789

Femoral poresScales around tailDorsalsVentral rowsFourth toe lamellaeTransverse rows of ventralsInfralabialsParietalsSupralabials

3022.05861.85264.27

72.7950.6338.9632.1628.9721.39

,0.0001,0.0001,0.0001,0.0001,0.0001,0.0001,0.0001,0.0001,0.0001

0.17610.12390.09570.04780.04570.04570.04130.02830.0283

1011121314151617

Infratibial rowsPrefemoralsChinshieldsFourth finger lamellaeGular foldScales around midbodyPreanalsSupraoculars

14.238.238.047.746.975.993.633.05

,0.0001,0.0001,0.0001,0.0001,0.0001,0.0001

0.00310.0101

0.01960.01960.02170.02170.02170.02390.02390.0239

(C. cryptus, C. gramivagus, and C. lem-niscatus) and the remaining species (C. lit-toralis, C. ocellifer, and C. parecis) (Fig.6). Amazonian Savanna species, with pos-itive means on the first canonical variable(Table 1), are characterized by a highnumber of femoral pores and fewer trans-verse rows of ventrals (Table 2).

The second canonical discriminant var-iable, representing a contrast betweentransverse rows of ventrals versus suprala-bials and fourth toe lamellae (Table 1), in-dicated a clear separation among the threespecies that occur south of the AmazonRiver (Fig. 6). Cnemidophorus parecis,with the highest mean on the second ca-nonical variable (Table 1), is characterizedby numerous transverse rows of ventralsand few supralabials and fourth toe lamel-lae (Table 2). Cnemidophorus littoralis,with the lowest mean on the second ca-nonical variable (Table 1), is characterizedby few transverse rows of ventrals and nu-merous supralabials and fourth toe lamel-lae (Table 2). Cnemidophorus ocellifer(and the Amazonian Savanna species) hasintermediate values for these variables.

The stepwise discriminant analysis ofmeristic characters selected 17 variables asthe most powerful discriminators amongthe species of Cnemidophorus (Table 3).Femoral pores was selected first, correctlyclassifying 82.4% of the individuals, fol-

lowed by scales around tail and dorsals,whose addition improved the classificationcriterion to 87.6% and 90.4%, respectively(Table 3). To determine if important vari-ables were excluded from the analysis be-cause of correlations among the originalvariables (Tabachnick and Fidell, 2001),we removed femoral pores and repeatedthe analysis. Scales around the tail werethen entered in the first step, correctlyclassifying 56.3% of the lizards (error rate5 0.437, F 5 895.34, P , 0.001). We thenremoved scales around the tail and re-peated the analysis. This time, infralabialswas the first variable to be selected, cor-rectly classifying about 60% of the lizards(error rate 5 0.3978, F 5 525.49, P ,0.001). These analyses indicated that fem-oral pores and scales around the tail arethe meristic characters that best discrimi-nated the species of Cnemidophorus andthat their discriminatory power was high.

The naıve Bayesian model indicatedthat C. parecis determined high condition-al probabilities (.95%) of 12 categoricalvariables: dorsolateral fields absent, verte-bral field spotted, paravertebral lines ab-sent, lateral line continuous, lower lateralline interrupted, vertebral line absent, lat-eral spots absent, hindlimb spots absent,preanal spur absent, supraoculars contact-ing frontal and parietals, ventral caudalssmooth, and dorsal caudals keeled (Table


TA

BL

E4.

—C

ondi

tion

alpr

obab

iliti

esof

stat

esfo

rqu

alit

ativ

ech

arac

ters

for

six

Bra

zilia

nsp

ecie

sof

Cne

mid

opho

rus,

assu

min

ga

naıv

eB

ayes

ian

mod

el.

Var

iabl

es

Spec

ies

C.

cryp

tus

C.

gram

ivag

usC

.le

mni

scat

usC

.lit

tora

lisC

.oc

ellif

erC

.pa

reci

s

Low

erla

tera

lfie

lds

Upp

erla

tera

lfie

lds

Dor

sola

tera

lfie

lds

Ver

tebr

alfie

ldP

arav

erte

bral

stri

pes

Dor

sola

tera

lst

ripe

s

D(5

5.95

%)

D(9

5.24

%)

D(9

6.43

%)

D(7

1.43

%)

C(9

6.43

%)

C(9

7.59

%)

A95

.59%

D(9

4.12

%)

D(9

5.59

%)

D(8

9.71

%)

C(9

1.18

%)

C(9

5.52

%)

L48

.28%

D(7

0.69

%)

D(9

4.83

%)

D(9

3.10

%)

C(9

4.83

%)

C(8

9.47

%)

L50

.88%

D(9

2.98

%)

D(6

4.91

%)

D(6

4.91

%)

I(9

4.74

%)

C(8

2.14

%)

L71

.43%

D(4

6.67

%)

D(4

3.81

%)

L(8

0.00

%)

I(5

6.19

%)

C(6

4.42

%)

L(9

1.96

%)

L(9

1.07

%)

A(9

6.43

%)

Sp(9

6.43

%)

A(9

7.32

%)

C(8

4.68

%)

Upp

erla

tera

lst

ripe

sL

ower

late

ral

stri

pes

Ver

tebr

allin

eL

ater

alsp

ots

Hin

dlim

bsp

ots

Chi

nshi

eld

cont

act

C(9

6.39

%)

C(6

5.06

%)

A(6

7.86

%)

A(6

4.63

%)

A(9

8.78

%)

.(9

6.39

%)

I(5

3.73

%)

A(9

1.04

%)

A(9

5.59

%)

A(9

6.97

%)

P(9

3.94

%)

.(9

7.01

%)

C(7

3.68

%)

C(6

4.91

%)

C(9

3.10

%)

A(5

7.14

%)

P(9

8.21

%)

.(8

7.72

%)

C(9

6.43

%)

I(6

6.07

%)

I(8

9.47

%)

A(9

8.18

%)

A(9

8.18

%)

.(8

0.36

%)

C(5

2.88

%)

A(5

6.73

%)

A(9

4.29

%)

P(9

0.29

%)

A(9

8.06

%)

.(9

8.08

%)

C(9

6.69

%)

I(9

5.50

%)

A(9

7.32

%)

A(9

9.09

%)

A(9

9.09

%)

.(4

4.14

%)

Pre

anal

spur

Sem

icir

cles

Ven

tral

caud

als

Dor

sal

caud

als

A(9

8.78

%)

IFP

(95.

29%

)K

(98.

78%

)K

(98.

78%

)

A(6

9.70

%)

IFP

(94.

20%

)K

(98.

48%

)K

(98.

48%

)

P(5

1.79

%)

IFP

(93.

22%

)K

(98.

21%

)K

(98.

21%

)

A(9

8.18

%)

IFP

(58.

62%

)S

(98.

18%

)K

(98.

18%

)

A(9

9.03

%)

Co

(51.

89%

)S

(99.

03%

)K

(99.

03%

)

A(9

9.09

%)

IFP

(95.

58%

)S

(99.

09%

)K

(99.

09%

)N

ote:

valu

esin

bold

indi

cate

cond

itio

nal

prob

abili

ties

grea

ter

than

95%

.D

:da

rk;

L:

light

;Sp

:sp

otte

d;A

:ab

sent

;P

:pr

esen

t;C

:co

ntin

uous

;I:

inte

rrup

ted;

K:

keel

ed;

S:sm

ooth

;C

o:co

mpl

ete;

IFP

:in

com

plet

e,su

prao

cula

rsco

ntac

ting

fron

tals

and

pari

etal

s;.

:co

ntac

tbe

twee

nfir

stpa

irof

chin

shie

lds

grea

ter

than

half

ofth

eir

leng

ths.


4). Further, only C. parecis determinedhigh conditional probabilities of dorsolat-eral fields absent, vertebral field spotted,and paravertebral lines absent (Table 4).The analysis also indicated that speciesfrom Amazonian Savannas can be clearlyseparated from species south of the Ama-zon River by the presence of keeled sub-caudals in the former (Table 4).

Our results revealed a greater similaritybetween C. parecis and the other speciesranging south of the Amazon River (C. lit-toralis and C. ocellifer). We also verifiedthat, besides the presence of anal spurs inmales, the species of the C. lemniscatuscomplex can also be distinguished fromtheir congeneric southern relatives by thepresence of keeled subcaudals, a largernumber of femoral pores, and fewer trans-verse rows of ventrals. This distinction sug-gests that C. parecis may be a closer rel-ative of the southern species of Cnemido-phorus. A number of palynological andsedimentary studies indicate that, in Ron-donia, open vegetation replaced the forestin the recent past, approximately 3000 ybp(Absy and Van der Hammen, 1976; deFreitas et al., 2001; Sifeddine et al., 2001;Van der Hammen and Absy, 1994). There-fore, present day enclaves of Cerrado inRondonia may be relicts of a past openvegetation landscape that ranged fromCentral Brazil to Rondonia. Following thebreakup of the connections between Ron-donia and the Central Brazil open land-scapes, C. parecis presumably differenti-ated as a consequence of the geographicalisolation. This view is supported by thepresence of a depauperate Cerrado lizardfauna in the Rondonia enclaves (Vanzolini,1986; Vitt and Caldwell, 1993). However,the affinities of C. parecis can only be as-certained with confidence through rigor-ous phylogenetic analysis.

Apparently, C. parecis is an endemic ofa small group of Cerrado patches withsandy soils around the city of Vilhena.Rondonia is the most deforestated state inAmazonia, having lost about one-third ofits total vegetation cover, whereas the mu-nicipality of Vilhena has less than 10% ofthe original vegetation. The Brazilian leg-islature (Codigo Florestal) stipulates that

80% of each rural property must be pre-served (Reserva Legal), but a state lawpassed recently in Rondonia reduced thispercentage up to 20% in highly productiveregions, favoring the establishment oflarge properties. The Cerrado patchesaround Vilhena are critically threatened bythe expansion of soybean plantations, fa-vored by the gentle topography that aidsmechanization, the low risks of drought,and the recent development of two expor-tation corridors. The first is the FerronorteRailroad, connecting Santa Fe do Sul inSao Paulo to Alto Taquari in Mato Grosso.The other is the Hidrovia do Rio Madeira,connecting Porto Velho in Rondonia stateto Itacoatiara in Amazonas state, along theMadeira and Amazon Rivers. Therefore,C. parecis is apparently at a critical risk ofbecoming extinct within a very short pe-riod of time. Because the presumed rangeof C. parecis is not included in any con-servation unit, being entirely within pri-vately owned land, urgent measures areneeded to protect the species.

Acknowledgments.—This work was developed un-der the project ‘‘Estrutura e dinamica da biota deisolados naturais e antropicos de Cerrado: licoes paraa biologia da conservacao,’’ funded by Programa Na-cional da Diversidade Biologica-PRONABIO, MMA-MCT-CNPq-GEF-BIRD. S. F. Balbino, C. G. Batis-ta, C. A. Freitas, D. Diniz, the staff of IBAMA atVilhena, Joecio Costa, Aparecido Donadoni, Vicentede Paulo Campos Godinho, and M. M. Utumi assist-ed in fieldwork. We thank Dr. Wagner Teixeira daSilva for assistance with Bayesian statistics, ZeniltonJ. G. Miranda for drawing the holotype, and threeanonymous reviewers for their insightful commentson the manuscript. This work was supported by grad-uate student fellowships from Coordenacao de Aper-feicoamento de Pessoal de Nıvel Superior to A. A.Garda (# 146980/99-9), P. H. Valdujo, D. O. Mes-quita, and G. H. C. Vieira; from Conselho Nacionaldo Desenvolvimento Cientıfico e Tecnologico to A.K. Peres, Jr. (# 146980/99-9) and H. C. Wiederhecker(# 141596/00-7); and by a research fellowship fromConselho Nacional de Desenvolvimento Cientıfico eTecnologico–CNPq to G. R. Colli (# 302343/88-1).

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Accepted: 22 April 2002Associate Editor:Joseph Mendelson III

APPENDIX ISpecimens Examined

Cnemidophorus cryptus (80).—BRAZIL: Amapa:Macapa, CHUNB 03436–57, 03459–83, 03487–500;Tartarugalzinho, CHUNB 08489–506.

Cnemidophorus gramivagus (64).—BRAZIL: Ama-zonas: Humaita CHUNB 03501–45, 03547–64; Ilhada Botija CHUNB 07944.

Cnemidophorus lemniscatus (54).—BRAZIL: Para:Santarem, CHUNB 01099–01152.

Cnemidophorus littoralis (53).—BRAZIL: Rio deJaneiro: Barra de Marica CHUNB 03228–264,08300–310, 24110–11, 24117, 24867–68.

Cnemidophorus ocellifer (101).—BRAZIL: Bahia:Ibipeba, CHUNB 03134–39, 03265, 03273–74,03358, 03360, 03369–70, 03372, 03375–76, 03378,03401, 03405, 03414, 03421, 08481, 08484–85; Irece,CHUNB 03266–67, 03269–72, 03359, 03361–68,03373–74, 03377, 03379–80, 03382, 07945–56; Len-cois, CHUNB 05591; Paulo Afonso, CHUNB 08155–57; Salvador, CHUNB 08138–39, 08152; Pernambu-co: Buıque, CHUNB 23529; Exu, CHUNB 12855,13590; Petrolina, CHUNB 03402–04, 03406–13,03415–20, 03422–35, 13614, 24109, 24241.

Cnemidophorus parecis (108).—BRAZIL: Rondonia:Vilhena, CHUNB 09762–814, 11651–703, 14229–30.

A CRITICALLY ENDANGERED NEW SPECIES OF CNEMIDOPHORUS (SQUAMATA, TEIIDAE) FROM A CERRADO ENCLAVE IN...

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