Whiptail lizards in South America: a new Ameivula (Squamata, Teiidae) from Planalto dos Gerais,...

Amphibia-Reptilia 35 (2014): 227-242

Whiptail lizards in South America: a new Ameivula (Squamata,Teiidae) from Planalto dos Gerais, Eastern Brazilian Cerrado

Federico José Arias1,2,∗, Mauro Teixeira Junior1, Renato Recoder1, Celso Morato de Carvalho3,

Hussam Zaher4, Miguel Trefaut Rodrigues1

Abstract. We describe a new species of whiptail lizard genus Ameivula from Planalto dos Gerais, a sandstone plateau thatextends along the Cerrado region in the states of Bahia, Tocantins, Piauí, and Minas Gerais, in Brazil. The new species isthe third recognized species of the A. ocellifera group registered for the Cerrado. Quantitative analyses of morphometriccharacters showed that Caatinga species are distinguished from their Cerrado congeners on basis of body size and shape.A discriminant analysis upon meristic characters showed that the new species can be clearly distinguished from the other twoCerrado species A. mumbuca and A. jalapensis, also the morphology of the finger lamellae, and clutch size distinguish thesespecies.

Keywords: Ameivula ocellifera group, Brazil, lizards, new species, Planalto dos Gerais.

Introduction

In a recent phylogenetic analysis based onmorphological traits, the genus Cnemidopho-rus resulted not monophyletic (Harvey et al.,2012), as previously suggested by molecularstudies (Reeder et al., 2002; Guigliano et al.,2006; Guigliano, 2009). To resolve the poly-phyly, Cnemidophorus was divided in four ge-nera: (1) the genus Cnemidophorus, includingthe species of the former C. lemniscatus group;(2) the new genus Aurivela which includes twospecies (A. longicauda and A. tergolaevigata);(3) the new genus Contomastix assembling thespecies previously referred to the Cnemidopho-rus lacertoides group; and (4) the new genusAmeivula, composed by the species of the for-mer Cnemidophorus ocellifer group.

1 - Departamento de Zoologia, Instituto de Biociências,Universidade de São Paulo, Caixa Postal 11.46 1, CEP05422-970, São Paulo, Brazil

2 - Instituto de Bio y Geociencias (IBIGEO) del Noa, Fac-ultad de Ciencias Naturales, Universidad Nacional deSalta, Av. Bolívia 5150, 4400 Salta, Argentina

3 - Instituto Nacional de Pesquisas da Amazônia, Núcleo dePesquisas de Roraima, Rua Coronel Pinto, 327, Centro,CEP 69301-150, Boa Vista, Roraima, Brazil

4 - Museu de Zoologia, Universidade de São Paulo, CaixaPostal 42694, CEP 04299-970, São Paulo, Brazil∗Corresponding author; e-mail: [email protected]

Currently, Ameivula has thirteen recognizedspecies (table 1) distributed throughout the Cis-Andean open-areas south of the Amazon River(Arias et al., 2011b; Harvey et al., 2012; Silvaand Ávila-Pires, 2013). These species generallyoccur in open habitats with sandy soils and hightemperature, in very distinct landscape phys-iognomies, such as the Caatinga (Vanzolini,1974; Vanzolini et al., 1980; Vitt, 1983, 1995;Menezes et al., 2011; Sales et al., 2012), Cer-rado (Vitt, 1991), “campos rupestres” (Menezeset al., 2011; Arias et al., 2014), Restingas (Diasand Rocha, 2004; Dias and Rocha, 2007; San-tana et al., 2010; Menezes and Rocha, 2011),and the Bolivian, Paraguayan and ArgentinianChaco (Williams and Tedesco, 1985; Cei, 1993;Dirksen and De la Riva, 1999; Cabrera, 2012).

The species of Ameivula exhibit morpholo-gical characteristics (both in lepidosis as colorpattern) that distinguish two groups of thespecies (Arias et al., 2011a): (1) the A. littoralisgroup, composed by A. abaetensis, A. littoralis,A. cyanura, and A. venetacauda; and (2) theA. ocellifera group, composed by A. abalosi,A. ocellifera, A. mumbuca, A. jalapensis, A. ni-grigula, A. pyrrhogularis and A. confusioniba.In addition, characters on hemipenis morphol-ogy and statistical analyses on continuous char-

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228 F.J. Arias et al.

Table 1. Summarized distribution of the 13 currently recognized species of Ameivula.

Species Comments and distribution

A. abaetensis (Dias et al., 2002) Endemic to the Restingas of northeastern coast of Brazil, betweennorthern Bahia and northern Sergipe. Occurs sympatrically with A.ocellifera in sand dunes of Abaeté, Salvador (Dias and Rocha, 2011).Member of A. littoralis group.

A. abalosi (Cabrera, 2012) Endemic to Argentinian and Paraguayan Chaco (Cabrera, 2012). Probablyin Bolivian Chaco. Member of A. ocellifera group.

A. cipoensis (Arias et al., 2014) Endemic from the mountain region of Parque Nacional Serra do Cipó,Southern of the Espinhaço chain, Minas Gerais state. This species occursin high elevation between 900 and about 1200 m, were the landscape isthat of “campos rupestres”.

A. confusioniba (Arias et al., 2011a) Occurs in Serra das Confusões, Piauí state, Brazil, sympatrically with A.venetacauda (Arias et al., 2011a), and in Serra Vermelha (Silva andÁvila-Pires, 2013). Member of A. ocellifera group.

A. cyanura (Arias et al., 2011b) Santo Inácio and Morro do Chapéu, northwestern Bahia, Brazil; occurs insympatry with A. nigrigula (Arias et al., 2011b). Member of A. littoralisgroup.

A. jalapensis (Colli et al., 2009) Endemic of Jalapão region, north portion of the Cerrado biome, Tocantinsstate, Brazil (Colli et al., 2009).

A. littoralis (Rocha et al., 2000) Endemic to the coastal Restinga areas of Rio de Janeiro state, southeasternBrazil (Rocha et al., 2000; Menezes et al., 2004). Member of A. littoralisgroup.

A. mumbuca (Colli et al., 2003b) Apparently endemic to the Jalapão region, northern of the Cerrado bioma,Tocantins state, Brazil.

A. nativa (Rocha et al., 1997) Southeastern Brazilian coast, between northern Espírito Santo andsouthern Bahia, Brazil (Rocha et al., 1999). Unisexual, parthenogenetic,hybrid origin.

A. nigrigula (Arias et al., 2011b) Southeastern Caatinga biome, on the right bank of the São Francisco river,Bahia state.

A. ocellifera (Spix, 1825) Type species of the Ameivula genus. Widespread species that ranges fromnortheastern Brazil to Mato Grosso do Sul, state. Currently considered aspecies complex.

A. pyrrhogularis (Silva and Ávila-Pires, 2013) Northern and central part of the Piauí state, Brazil. Transition zonebetween Caatinga and Cerrado (Silva and Ávila-Pires, 2013).

A. venetacauda (Arias et al., 2011a) Occurs in Caatinga Biome, Serra das Confusões (Arias et al., 2011a), Serrada Capivara (this work), Água Mineral Manaíra (Silva and Ávila-Pires,2013), southeastern Piauí, Brazil. Member of A. littoralis group.

acters support these two species groups (Silva

and Ávila-Pires, 2013).

The Cerrado biome is the largest Neotropi-

cal woodland-savanna, covering approximately

2 million km2, bearing high levels of endemism

(Oliveira and Marquis, 2002). Estimates indi-

cate that approximately 80% of its original

cover has been lost (Cavalcanti and Joly, 2002;

Klink and Machado, 2005). For these reasons,

the Brazilian Cerrado is considered a biodi-

versity “hostpot” (Mittermeier et al., 2004).

The Cerrado geomorphology is characterized

by the presence of distinct geomorphic surfaces:

the ancient vast plateaus (locally called cha-

padões) have altitudes of 500-1700 m, and are

dominated by savanna-like vegetation. These

plateaus are separated by a network of periph-

eral depressions (altitudes of 100-500 m) of

younger valleys (Ab’Saber, 1983; Silva, 1997;

Werneck, 2011), where the vegetation is char-

acterized by recently originated plant commu-

nities, including a mosaic of wet grassland,

cerrado vegetation, gallery forest, and patches

of deciduous and semi-deciduous forest (Cole,

1986; Silva, 1997; Werneck, 2011). Recent

studies show that highland areas (chapadões)

New Ameivula from eastern Brazilian Cerrado 229

played an important role in the history andcurrent diversity of reptiles along the Cerrado(Nogueira et al., 2011).

During an extensive field work in the ParqueNacional Cavernas do Peruaçu, on the south-ern portion of Planalto dos Gerais, in the Mi-nas Gerais state of Brazil, we obtained a largeseries of specimens of Ameivula that revealedto be very distinct from their congeners in lepi-dosis, hemipenis, color pattern, and morphome-try. Here, we describe these specimens as a newspecies of Ameivula. The new species is mor-phologically and geographically more similar tothe two recognized species inhabiting the Cer-rado biome: A. mumbuca and A. jalapensis, oc-curring parapatricaly with both species in sandyareas of Jalapão region, at the northern end ofPlanalto dos Gerais.

Materials and methods

Fieldwork

Specimens were collected at Parque Nacional Cavernas doPeruaçu, municipality of Januária, in the northern portion ofthe state of Minas Gerais, Brazil (Teixeira Jr., 2010). Mostspecimens were captured with the aid of pitfall traps thatconsisted in four thirty-liters buckets that were buried atground level and connected to each other with three four-meter long drift fences, forming a Y shaped configuration.Traps were installed in the main vegetation physiognomiesas follows: 15 traps on dry forest habitats; 15 on savanna; 17on “carrascos” and two on riparian evergreen forest. Trapsremained opened during periods of 30 days in January (rainyseason) and July (dry season) 2008 and 2009, totalizing aneffort equivalent to 14 560 buckets × days. Collected in-dividuals were fixed in 10% formalin, preserved in 70%ethanol, and deposited in the Museu de Zoologia da Uni-versidade de São Paulo, Brazil (MZUSP).

Laboratory work

We compared the new species with 13 recognized speciesof Ameivula (listed in the Appendix) and added to ourcomparisons data of the coloration in life taken from theoriginal descriptions of A. littoralis (Rocha et al., 2000),A. mumbuca (Colli et al., 2003b), A. nativa (Rocha et al.,1997), and A. jalapensis (Colli et al., 2009), while for A.pyrrhogularis (Silva and Ávila-Pires, 2013) we take datafrom its lepidosis and coloration. Specimens examined aredeposited at the Museu de Zoologia da Universidade de SãoPaulo, Brazil (MZUSP), Museu Nacional do Rio de Janeiro,Brazil (MNRJ), Fundación Miguel Lillo, Tucumán, Ar-gentina (FML), Colección Herpetológica de la Universidad

Nacional del Nordeste, Corrientes, Argentina (UNNEC),and Colección Herpetológica Museo de Ciencias Naturales,Salta, Argentina (MCN).

We recorded the following characters in each individual:number of supraocular and supraciliar scales on right side;femoral pores on both sides; longitudinal rows of ventralscales; transverse rows of ventral scales, from gular foldto anterior margin of hindlimbs; subdigital lamellae underfourth finger and fourth toe; longitudinal rows of enlargedscales in the dorsal region of the arm; scales around mid-body, counted midway between fore- and hindlimbs ex-cluding ventrals; scales around tail, counted on the fourthtransverse row; number of gular scales, counted along themidline, from postsymphysal scale to mesoptychial scales;dorsal scales, counted along the midline, from occiput tofirst transverse row of scales around tail; presence or ab-sence of enlarged scales in the temporal region, posteriorlyto the third subocular; presence or absence of keel in sub-digital lamellae both in the hand or foot; and presence orabsence of patch of tibiotarsal spurs in the males. In addi-tion, we took the following measurements using a digitalcaliper to the nearest 0.02 mm: snout-vent length (SVL),trunk length (TRL), head length (HL), head width (HW),forearm length (FAL), total hindlimbs length (HLL), andtail length (TAL). Scale observation and measurements weremade under a stereomicroscope (10-40×). Moreover, weanalyzed the color pattern and color in life in both malesand females. We prepared the hemipenis of MZUSP 99677and MZUSP 99623 following the procedures described byManzani and Abe (1988), subsequently modified by Pe-santes (1994) and Zaher (1999; see also Zaher and Prudente,2003). Later, we compared with the hemipenial morphologyof A. confusioniba, A. pyrrhogularis, A. venetacauda (Silvaand Ávila-Pires, 2013), and A. cipoensis (Arias et al., 2014).

Quantitative analysis

Our observations, made both in the field and the collectionson the species of the Ameivula ocellifera group from Cer-rado and from Caatinga biome, let us distinguish two sub-groups. One formed for the Caatinga species with conspicu-ous sexual dimorphism and ontogenetic change in the colorpattern of the males. The other one, formed for the Cer-rado species, being smaller and without ontogenetic changein the color pattern. Here, we hypothesize that both sub-groups can be also distinguished with morphometric charac-ters. Thus, to test for significant morphometric differencesbetween the two species subgroups within the Ameivulaocellifera group, we performed a two-way MANOVA usinggroup and sex as factors and then performed a discriminantfunction analysis to find the best combination of variablesthat distinguish both subgroups. For this purpose, we se-lected six of the seven measurements: SVL, TRL, HL, HW,FAL, HLL. We analyzed TAL separately because of miss-ing data. We used a dataset containing measurements of fivemales and five females of Ameivula sp. nov., A. jalapen-sis and A. mumbuca (Cerrado group), A. aff. ocellifera,A. cipoensis, A. confusioniba and A. nigrigula (Caatingagroup). As sexual dimorphism interacted with group dif-ferences (see results), we performed discriminant analyses


for males and females separately. Then, we tested for dif-ferences among the three species of the Cerrado group (i.e.Ameivula sp. nov., A. jalapensis and A. mumbuca) in size(SVL) with ANOVA and in shape with the remaining fivecharacters using MANOVA after controlling for size. Thesize free database was obtained by regressing each mea-surement on a first principal component (82.3% of varia-tion explained) obtained with a pooled within-group princi-pal component analysis. We analyzed TAL separately usingANCOVA because of missing data. Morphometric measure-ments were log10 transformed prior to analysis and normal-ity was confirmed (Kolmogorov Smirnov Test, p > 0.05)(Zar, 2010).

A discriminant function analysis was conducted to inves-tigate quantitative differences in lepidosis among Ameivulasp. nov., A. jalapensis and A. mumbuca. Analysis wasbased on the following meristic characters: number of gularscales (GUL), transversal rows of ventral scales (VT), scalesaround midbody (SAM), scales around tail (SAT), femoralpores (FP), dorsal scales (D), subdigital lamellae underfourth finger (FFL) and fourth toe (FTL), for which normal-ity was confirmed (Kolmogorov Smirnov Test, p > 0.05).Log10 transformation was also used for selected meristicdata, and we performed all statistical analysis using IBMSPSS Statistics (version 20.0, SPSS Inc., 2011).

Results

Ameivula xacriaba sp. nov. (figs 1, 2)

Holotype. MZUSP 99690, an adult malefrom Parque Nacional Cavernas do Peruaçu(15°9′17.04′′S, 44°18′18.29′′W, 747 m a.s.l.),Januária municipality, state of Minas Gerais,Brazil. Collected by M.T. Rodrigues, R. Re-coder, M. Teixeira Jr., M. Aurelio de Sena andA. Camacho on 17 January 2009.

Paratypes. Adult males: MZUSP 99676,MZUSP 99685, MZUSP 99700, MZUSP99679, MZUSP 99632, 99669, 99622, 99677,99641. Adult females: MZUSP 99642, MZUSP99658, MZUSP 99638, MZUSP 99618,MZUSP 99630, 99678, 99643. Collected byM.T. Rodrigues, R. Recoder, M. Teixeira Jr., M.Aurelio de Sena and A. Camacho on 17 January2009. All paratypes with the same locality dataas for the holotype.

Diagnosis. A slender, medium sized speciesof Ameivula, distinguished from all otherspecies of the genus by the following combina-tion of characters: (1) 6-7 supralabials; (2) 5-6 infralabials; (3) 4-5 supraoculars; (4) five

Figure 1. Dorsal, ventral and lateral view of the head of theholotype of Ameivula xacriaba sp. nov., MZUSP 99690.

supracilaries; (5) 4-6 parietals; (6) 8 longitudi-nal rows of ventral scales; (7) 27-29 transversalventral rows; (8) 14-17 femoral pores; (9) 184-216 dorsal scales; (10) 24-29 scales around tail;(11) 94-102 scales around midbody; (12) 13-15 fourth finger lamellae; (13) 23-28 fourth toelamellae; (14) 41-47 gular scales; (15) absenceof paravertebral stripes; (16) lateral ocelli ex-tending from hindlimbs to forelimbs.

The new species is assigned to the Ameivulaocellifera group (Arias et al., 2011a) by a lownumber of femoral pores (less than 20), en-larged scales in the temporal region (posteriorlyto the third subocular), five supraciliary scales,8 rows of ventral scales, and a brownish tail.

Comparison. Data from species in compari-son are given in parentheses; Ameivula xacriaba


Figure 2. Photo in life of Ameivula xacriaba sp. nov.,showing variation in male (A), female (B) and juvenile (C).Photo: Mauro Teixeira Jr. This figure is published in colorin the online version.

sp. nov. can be distinguished from A. abaeten-sis, A. cyanura, A. littoralis and A. venetacauda(i.e., members of the A. littoralis group) by hav-ing enlarged scales in the lower temporal re-gion, posteriorly to the third subocular (scalessub equal), three rows of enlarged scales in thedorsal part of the arm (one row), 5 supraciliars(6-7), 8 longitudinal rows of ventral scales (8-10), 27-29 transversal ventral rows (29-38), 14-17 femoral pores (21-45), absence of tibiotarsalspurs in the males (with 1-2 rows of 4-6 spurs),and by having a brown tail (bluish green). Fur-thermore, it differs from A. abaetensis, A. cya-nura and A. littoralis by lacking of a mid-dorsal

stripe (present) and a stripe in the tail (presentin A. abaetensis and A. littoralis). Ameivulaxacriaba sp. nov. also differs from A. vene-tacauda by having laterodorsal and lateroven-tral stripes, and light green ocelli (no stripes orocelli). Ameivula xacriaba sp. nov. differs fromA. nativa by being bisexual (parthenogeneticspecies) and having 14-17 femoral pores (22-26); 23-28 fourth toe lamellae (26-33); and bylacking a mid-dorsal stripe (present and with aposteriorly undulating margin). Ameivula xacri-aba sp. nov. differs from A. jalapensis by hav-ing 14-17 femoral pores (10-16); 5 parietals (3-5); 27-29 transverse rows of ventral scales (25-27); 8 longitudinal rows of ventral scales (6-8);41-47 gular scales (46-64); subdigital lamellaesmooth (subdigital lamellae keeled); a mediumbody size, maximum SVL � = 70.2 mm, � =64.7 mm (smaller body size, maximum SVL� = 54.2 mm, � = 58 mm), tail length/SVLmean = 1.9 (mean = 2.6) a white throat (lime-green, Colli et al., 2009), ocelli on lower lat-eral fields (ocelli absent), and conspicuous sex-ual dichromatism (without sexual dichroma-tism, Colli et al., 2009). Ameivula xacriaba sp.nov. differs from A. mumbuca by having 14-17femoral pores (16-20), subdigital lamellae with-out keel (subdigital lamellae keeled), ocelli onlower lateral fields in both males and females(only in males; Colli et al., 2003), lateroven-tral field bright green in males (bright yellowin males), and conspicuous sexual dichroma-tism (without sexual dichromatism). Ameivulaxacriaba sp. nov. differs from A. confusionibaby having 23-28 fourth toe lamellae (29-35),14-17 femoral pores (16-21), 41-47 gular scales(29-39), and bright green ocelli on lower lateralfields extending from forelimbs to hindlimbs(only 2-4 small yellow ocelli). Ameivula xacri-aba sp. nov. differs from A. ocellifera by hav-ing 23-28 fourth toe lamellae (28-31), 191-216dorsal scales (172-188), 41-47 gular scales (32-35), without paravertebral stripes (with markedparavertebral stripes), and sexual dichromatism(without sexual dichromatism). Ameivula xacri-aba sp. nov. differs from A. nigrigula by hav-


ing 13-15 fourth finger lamellae (16-18), 23-28fourth toe lamellae (30-32), 23-29 scales aroundthe tail (27-32), 41-47 gular scales (31-35), ab-sence of a patch of tibiotarsal spurs in males(present), gular region white (dark black), andby lacking ontogenetic color change in males(present). Ameivula xacriaba sp. nov. differsfrom A. abalosi by having 14-17 femoral pores(16-20), 13-15 fourth finger lamellae (15-17),41-47 gular scales (31-35), postnasal and pre-frontal scales contacting each other (no con-tact), and by lacking a distinctive patch of tibio-tarsal spurs in males (present). Ameivula xacri-aba sp. nov. differs from A. pyrrhogularis byhaving 14-17 femoral pores (16-24); maximumSVL � = 70.2 mm, � = 64.7 mm (maximumSVL � = 89 mm, � = 77.5 mm); and gularregion white in males (orange in males) (Silvaand Ávila-Pires, 2013). Ameivula xacriaba sp.nov. differs from A. cipoensis by having 14-17 femoral pores (17-20); 13-15 fourth fingerlamellae (12-14); 23-28 fourth toe lamellae (21-24); 41-47 gular scales (31-38); 191-216 dor-sal scales (202-225); four supraoculars (three);frontoparietal and third supraoculars never con-tact (in contact); the third pair of chinshields thesame size than the others (third pair larger thanthe others).

Description of holotype. Measurements:snout-vent-length (SVL) 60.7 mm, trunk length29.6 mm, head length 14.7 mm, head width9.7 mm, head height 8.1 mm, tail length125 mm (2.1 times longer than SVL), humeruslength 5.6 mm, forearm length 19.7 mm, femurlength 10.7 mm, tibia length 8.4 mm, foot length17.5 mm, and hindlimb length 36.6 mm.

Rostral large, wider than high, visible fromabove, with the posterior edge sharp, sepa-rated from frontonasal by the midline con-tact between nasals. Nasals roughly pentago-nal smaller than prefrontals. Nasal and post-nasal in broad contact by an oblique suture.Nostril subovoid, in lower part of suture. Fron-tonasal hexagonal, wider than long, contactingnasals, postnasals and prefrontals. Prefrontals,longer than wide roughly trapezoidal, pentag-

onal, in broad contact along midline; contact-ing postnasal, loreal, frontal and first supraoc-ular; suture between prefrontals as long asfrontonasal. Frontal approximately pentagonal,longer than wide, wider anteriorly with lateralmargins slightly concave; contacting first leftsupraocular. Two longer than wide, and approx-imately pentagonal frontoparietals; wider pos-teriorly, slightly concave laterally; their mid-line suture straight. Five parietals, external onessmaller; interparietal sub-pentagonal, with lat-eral sides slightly concave, longer and widerthan others, bordered at each side by parietals.Occipital scales forming two transverse rowsregular in size. Four supraoculars on each side,third the longest; first in contact with loreal,prefrontal, frontal (on the left side), and firstsupraciliary; not contacting the second supraoc-ular. Five supraciliaries on each side, secondlargest, the third and fourth sub-equal, the fifthis smallest. Only first supraciliary in contactwith first supraocular, all others separated fromsupraoculars by a row of granules. Loreal sin-gle, large, as wide as third supralabial, aswide as high, in contact with postnasal, pre-frontal, first supraocular, first supraciliary, firstand second suboculars, and third and fourthsupralabial. First subocular narrow, higher thanwide, in contact with second subocular, loreal,and first supraciliary. Four suboculars on eachside, first and second keeled, second in con-tact with the fourth supralabial; third subocu-lar keeled, approximately rectangular, in contactwith fourth, fifth and sixth supralabials; fourthsubocular smooth, as long and wide as the third,with rounded posterior edges. Seven supralabi-als on each side, third the largest, posterior onessmaller. Temporal region with irregular scales,granular centrally, enlarged ventrally. A lon-gitudinal supratemporal row with moderatelylarge scales, decreasing in size posteriorly. Earopening large, round, with smooth margins, el-lipsoid, slightly wider dorsally. All dorsal andlateral head scales juxtaposed, smooth (exceptfor keeled suboculars). Symphysal as long aswide, anteriorly concave, almost straight poste-


riorly where it contacts first infralabial and post-symphysal, forming two wide angles. Postsym-physal single, pentagonal, longer than wide, incontact with first and second infralabials; fol-lowed by five pairs of enlarged chinshields. Firstpair of chinshields in broad contact along mid-line, in contact with third infralabials, slightlylonger that the other. Second to fourth pairs ofchinshields separated from infralabials by a rowof small granules. Six infralabials on each side;followed posteriorly by series of small scalesextending to commissure of mouth; first infral-abial the smallest. Gular region divided in twoareas: anterior one with enlarged, rounded andslightly elongate scales, disposed in irregular30 transverse rows, delimited posteriorly by aintertympanic sulcus; posterior region coveredwith granules, in 17 transverse rows, smaller lat-erally, bordered posteriorly by antegular fold.Gular and antegular folds marked by granules.Mesoptychial region with two irregular rows ofenlarged scales between the two folds. Scaleson nape and sides of neck similar to dorsals.Dorsals and scales on flanks granular, rounded,smooth, sub-imbricate; 192 scales along a mid-dorsal line from nape to base of tail, 99 scalesaround mid-body (excluding ventrals). Ventralscales large, smooth, rectangular (wider thanlong), imbricated, in 28 transverse rows; 8 ven-tral scales in transverse rows across midbody.Ventral scales separated from scales on flanksby row of moderately enlarged scales. Preanalplate with three enlarged scales, a central onesurrounded laterally by smaller ones and twoon posterior region. Femoral pores in a continu-ous row along each thigh, with a short mediallygap; 8 pores on right side, 7 on left. Scales onbase of tail rectangular, smaller than ventrals, intransverse rows; keeled and slightly mucronatedorsal and laterally, smooth ventrally, imbri-cate; 29 scales around tail. Tail scales becominggradually longer and narrower from the base totip; subcaudal scales becoming keeled distally.Limbs with large, smooth, imbricate scales ondorsal face of upper arms, antero-dorsal face offorearms, antero-ventral face of thighs, and ven-

tral face of lower legs; elsewhere scales small,granular. Larger scales in longitudinal rows onupper arms. Forearms with one row of enlargedscales, wider than long. Anterior scales on thighdecreasing in size proximally, with three rowsof enlarged scales. Lower legs with two rowsof enlarged, hexagonal scales. Ventral face ofhands and feet granular; one enlarged tubercleat base of pollex. Sub-digital lamellae single; 14lamellae under fourth fingers; 24 under fourthtoes.

Color in life. Dorsal parts of head, body, limbsand tail are brown. Lateral parts of the head arelight brown, and white ventrally. The laterodor-sal field is black, with irregular green spots ex-tends from neck to base of tail. Below it, a greenlaterodorsal stripe that extends from the supra-ciliar region to the base of the tail. The lateralfield is green and contains an irregular seriesof black spots interspersed with bright greenocelli, between the fore and hindlimbs. Separat-ing the lateral field from the ventrolateral field,a lateroventral stripe that extends from nostril tohindlimbs. The lateroventral field is green withirregular green ocelli. The dorsal and lateral sur-face of the tail is brown, and ventrally is white.The belly is light blushing while ventral parts ofthe limbs, cloacal region and tail are white.

Color in ethanol. Head brownish dorsally,labial regions and ventral aspect of head bluishwhite. Dorsal parts of the body, limbs, andtail are bluish brown. Dorsolateral and lat-eral fields dark black, separated by dorsolat-eral stripe. A longitudinal series of small bluishwhite ocelli, between the fore and hindlimbs,is present in the lateral field. Ventrolateral fieldbluish white with spots white. Belly light blue,ventral aspect of the limbs, cloacal region andtail are white.

Variation. Data based on five adult males andfive adult females, listed as range (mean ±SD): SVL 59.6-70.2 mm (62.8 ± 3.3 mm);maximum male SVL 70.2 mm; maximumfemale SVL 64.8 mm. Head length 13.7-


19.4 mm (15.9 ± 1.7 mm). Head width 8.7-12.9 mm (10.7 ± 1.3 mm). Head height 7.6-11.2 mm (8.9 ± 1.2 mm). Humeral length 4.3-6.4 mm (5.1 ± 0.7 mm). Forelimb length 18.2-23.0 mm (19.8 ± 1.4 mm). Femur length 9.1-11.5 mm (10.5 ± 0.8 mm). Tibia length 8.2-9.9 mm (8.8 ± 0.5 mm). Foot length 17.5-21.5 mm (18.7 ± 1.2 mm). Hindlimb 2.0times longer than foot length. Hindlimb length35.2-42.8 mm (38.0 ± 2.3). Tail length 104-134.6 mm (117.6 ± 12.2 mm), 1.9 times longerthan SVL. Supralabials scales 6-7 (6.3 ± 0.5);infralabials scales 5-6 (5.2 ± 0.4). The speci-men MZUSP 99685 has the frontonasal divided.Variations in other meristic characters are sum-marized in table 2.

An evident sexual dichromatism is present,especially during the reproductive season. Inadult males the flank color (between forelimbsand hindlimbs) is bright green, while in adultfemales the lateroventral areas are bright yellow.In the juveniles of both sexes the lateroventralareas are light green, as for the ocelli (fig. 2).

Hemipenial morphology. Description is basedon the fully everted and expanded right organ ofMZUSP 99677 (fig. 3). Bilobed hemipenis. Thesulcus spermaticus is a broad channel, originat-ing at the base of the organ, reaching the apex.At the distal tip of the body the sulcus is di-vided in two lobes, each lobe has papillae at theouter edges, bellow them there are three sacs ineach lobe. In the distal region there are 4-5 lam-inae, laterally located, separated by two longi-tudinal septa on each side. The central zone ofdistal region and proximal region of the organare completely nude, in both faces (sulcate andasulcate).

The hemipenial morphology of Ameivulaxacriaba sp. nov. show features that are sharedby other members of A. ocellifera group, suchas A. pyrrogularis, A. confusioniba, A. nigrigula(Silva and Ávila-Pires, 2013), and A. cipoensis(Arias et al., 2014). The distal tip of the hemipe-nis of these species are papillate at the outeredges, with two or three sacs at the edge of eachlobe. The hemipenis of the A. littoralis species

group lack these ornamentations, has a protu-berance on each lobule at the distal tip (api-cal region), and the body of the organ is coverby transverse flounces (Silva and Ávila-Pires,2013).

Etymology. The specific name “xacriaba”honors the Xakriabás indians. A remainingcommunity that occupies an Indian Territory inthe vicinities of Peruaçu region and along theleft bank of the river São Francisco, in the mu-nicipalities of São João das Missões, state ofMinas Gerais, Brazil.

Distribution and ecology. The new species isapparently endemic to the Planalto dos Gerais(fig. 4), a large, eroded sandstone plateau thatreaches 900 m in some points and extendsmainly along all the western region of Bahiastate, in the eastern portion of the Brazilian Cer-rado (Teixeira et al., 2013). The type series wasall collected at Parque Nacional Cavernas doPeruaçu, and it is the most common lizard inthis region (Teixeira Jr., 2010). The new specieswas found only in savannas, despite significantsampling efforts involving pitfall traps and ac-tive searches in all vegetation types known inthe area (fig. 5). It is a Cerrado area that waspartially converted into pasturelands in the past,but which is regenerating since 1997, and itis now protected. Except for some areas withbare sandy soil, the savanna vegetation in theregion has an extensive herbaceous cover witha sparse arboreal stratum that is predominantlycomposed by porcada trees (Copaifera martii)and pequi trees (Caryocar brasiliense).

Additional material from MZUSP confirmsthe presence of Ameivula xacriaba sp. nov. inParque Nacional Grande Sertão Veredas, MinasGerais state, and in the municipalities of Jab-orandi and São Desiderio, state of Bahia (seeAppendix). At Parque Nacional Grande SertãoVeredas Ameivula xacriaba sp. nov. is the sec-ond most abundant species, after Vanzosaurarubricauda, being sympatric with at least 25other species of lizards, some of them typical ofthe Serra Geral region (e.g., Stenocercus quinar-


Tabl

e2.

Com

pari

son

ofse

lect

edm

eris

ticch

arac

ters

amon

gth

em

embe

rsof

the

Am

eivu

laoc

elli

fera

grou

pfr

omC

erra

doan

dC

aatin

ga.V

alue

sin

dica

tem

ean

±st

anda

rdde

viat

ion,

with

rang

ein

pare

nthe

ses.

∗ Mer

istic

char

acte

rsta

kefr

omth

eor

igin

alde

scri

ptio

n(S

ilva

and

Ávi

la-P

ires

,201

3).

Cha

ract

ers

A.x

acri

aba

sp.n

ov.

A.m

umbu

caA

.jal

apen

sis

A.c

onfu

sion

iba

A.c

ipoe

nsis

A.n

igri

gula

∗ A.p

yrrh

ogul

aris

(n=

92)

(n=

35)

(n=

38)

(n=

30)

(n=

40)

(n=

25)

(n=

128)

Gul

arsc

ales

45.7

±4

(41-

52)

44.9

±4.

4(3

8-49

)51

.3±

6.6

(41-

64)

34.1

±4.

2(2

9-39

)33

.7±

2.2

(30-

38)

33.3

±2.

1(3

1-35

)?

Fem

oral

pore

s15

±1.

2(1

4-17

)18

.3±

1.3

(16-

20)

13.1

±2.

3(1

0-16

)17

.5±

1.8

(16-

21)

17.7

±1

(17-

20)

17.1

±(1

5-20

)18

.5±

1.4

(16-

22)

Tra

nsve

rsal

row

sof

28±

0.5

(27-

29)

27.1

±0.

6(2

6-28

)25

.7±

0.8

(25-

27)

28.2

±0.

9(2

7-29

)29

.1±

0.8

(28-

30)

27.4

±(2

6-29

)28

.3±

0.8

(26-

31)

vent

rals

cale

sL

ongi

tudi

nalr

ows

of8

±0

8±

07.

33±

1.2

(6-8

)8

±0

8±

0(8

)8

±0

8±

0ve

ntra

lsca

les

%of

cont

actb

etw

een

52.1

7%(n

=92

)62

.86%

(n=

35)

23.6

8%(n

=38

)10

0%(n

=30

)10

0%(n

=20

)72

%(n

=25

)?

1aan

d2a

supr

aocu

lar

Four

finge

rla

mel

lae

14.1

±0.

6(1

3-15

)16

±0.

6(1

5-17

)14

.7±

1.3

(14-

16)

15.6

±1

(15-

17)

12.5

±0.

8(1

2-14

)17

±0.

8(1

6-18

)16

.1±

1.0

(13-

19)

Four

thto

ela

mel

lae

25.4

±1.

7(2

3-28

)26

.8±

1.5

(25-

29)

26±

1.5

(23-

28)

30.4

±2.

1(2

9-35

)22

.5±

1.2

(21-

24)

31.3

±1

(30-

32)

29.8

±2.

0(2

4-34

)Sc

ale

arou

nd99

.3±

4.0

(94-

106)

98.8

±3.

0(9

5-10

2)94

.3±

4.7

(89-

100)

95±

6.8

(87-

105)

92.6

±5

(85-

100)

99.8

±3.

0(9

7-10

5)10

4±

7.1

(87-

124)

mid

body

(SA

M)

Scal

ear

ound

26.3

±2.

3(2

3-29

)25

.1±

2.0

(22-

28)

24±

1.4

(22-

26)

24.9

±2.

0(2

2-28

)26

.7±

1.6

(25-

29)

30±

2.7(

27-3

2)26

.7±

1.6

(23-

33)

tail

(SA

T)

Dor

sal

202

±8.

321

1.5

±9.

221

8±

6.7

201.

6±

8.5

213.

6±

7.3

208.

6±

1322

2.9

±12

.0(1

91-2

16)

(200

-227

)(2

08-2

26)

(188

-211

)(2

02-2

25)

(190

-220

)(1

92-2

55)

SAM

/SA

T×

100

383.

6±

27.5

395.

7±

34.9

406.

1±

30.5

381.

4±

25.5

347.

6±

22.5

328.

4±

3038

9.5

(341

.4-4

17.4

)(3

51.8

-454

.6)

(356

-454

.6)

(332

-408

)(3

07-3

73)

(309

.4-3

63)

SAM

/Dor

sal×

100

49.2

±2

(47.

2-52

.2)

46.8

±2.

4(4

3.2-

50.5

)44

.3±

2.7

(40.

3-49

)47

.1±

2.6

(42-

50.5

)43

.4±

2.8

(39-

49.5

)48

.3±

4.3

(44.

7-55

.3)

46.7


Figure 3. Asulcate (A), and sulcate (B) faces of hemipenis of Ameivula xacriaba sp. nov., MZUSP 99677.

Figure 4. Geographic distribution of the analyzed species of the Ameivula ocellifera group from central-east Brazil. Thetriangle with a dot indicates the type locality of Ameivula xacriaba sp. nov. This figure is published in color in the onlineversion.


Figure 5. Habitat of Ameivula xacriaba sp. nov. at the Parque Nacional do Peruaçu, Minas Gerais state, Brazil. Photo: MauroTeixeira Jr. This figure is published in color in the online version.

Table 3. Results of a discriminant function analysis onmorphometric variables of Ameivula species from Cerradoand Caatinga groups. Scores with highest correlation witheach discriminant function are highlighted in bold.

Males Females

Variables Coeffi- Corre- Coeffi- Corre-cients lation cients lation

SVL −0.175 0.879 −0.565 0.517TRL 0.139 0.677 −0.443 0.372HL 0.434 0.865 0.398 0.558HW −0.291 0.676 0.610 0.496FAL 0.364 0.837 0.516 0.639HLL 0.605 0.954 0.720 0.830

Eigenvalue 2.573 2.351Wilks’ Lambda 0.280 0.298

ius, Bachia geralista; Nogueira and Rodrigues,2006; Recoder and Nogueira, 2007; Teixeira etal., 2013).

Quantitative analyses. Group differenceswere significant (MANOVA, Wilk’s Lambda =0.313, F6,61 = 22.33, P < 0.01). Sexual dimor-phism in morphometry was detected (Wilk’sLambda = 0.288, F6,61 = 25.11, P < 0.01)and interacted significantly with group differ-ences (Wilk’s Lambda = 0.762, F6,61 = 3.17,P < 0.01).

The discriminant function presented high cor-relation with HLL and SVL for males, and HLL

and FAL for females (table 3). The two groupscan be differentiated on basis of the significantlylarger size (ANOVA, males: F1,34 = 65.67,P < 0.01, females: F1,34 = 20.05, P <

0.01) and longer hind limbs of the species fromthe Caatinga group (ANCOVA, males: F1,34 =4.23, P < 0.05, females: F1,34 = 18.96,P < 0.01). Based on the discriminant function,88.6% of the individuals were correctly classi-fied in each group for males and 91.4% for fe-males (table 4).

The three species from Cerrado (A. xacri-aba sp. nov., A. mumbuca and A. jalapensis)showed significant differences in size (ANOVA,F2,29 = 28.88, P < 0.01) but sexes didnot differed (F1,29 = 0.27, P = 0.607)and did not interacted with species differences(F2,29 = 0.49, P = 0.617). In post hoc com-parisons, A. xacriaba sp. nov. is significantlylarger than A. jalapensis (Tukey’s HSD Test,F1,9 = 32.4, P < 0.01) and A. mumbuca(F1,9 = 5.47, P < 0.05). Species did not dif-fered in shape (MANOVA, Wilk’s Lambda =0.477, F10,40 = 1.79, P = 0.09) and sexualdimorphism in shape was not detected (Wilk’sLambda = 0.937, F10,40 = 1.31, P = 0.98).However, A. xacriaba sp. nov. presented shorter


Table 4. Jackknifed classification matrix based on morphometric data of Ameivula xacriaba sp. nov., A. jalapensis, A.mumbuca (Cerrado group), A. confusioniba, A. cipoensis, A. nigrigula and A. aff. ocellifera (Caatinga group).

N Males Females

Cerrado Caatinga % correct Cerrado Caatinga % correct

Cerrado 15 14 1 93.3 14 1 93.3Caatinga 20 3 17 85.0 2 18 90.0

Table 5. Results of a discriminant analysis on meristicvariables with individuals of Ameivula xacriaba sp. nov., A.jalapensis and A. mumbuca. Scores with highest correlationwith each discriminant function are highlighted in bold.

Variables DF1 DF2

Transverse rows of ventral scales (VT) 0.429 −0.257Gular scales (GUL) −0.294 0.056Scales around tail (SAT) 0.179 −0.102Scales around midbody (SAM) 0.149 0.119Femoral pores (FP) 0.285 0.908Dorsals (D) −0.163 0.453Fourth finger lamella (FFL) −0.070 0.220Fourth toe lamella (FTL) −0.052 0.130

Eigenvalue 11.853 1.140% of variance 91.2 8.8Cumulative % 91.2 100

tails than A. jalapensis (ANCOVA, F1,9 = 6.79,P < 0.05).

A discriminant analysis performed on meris-tic characters using individuals of A. xacriabasp. nov., A. jalapensis and A. mumbuca showeda differentiation among the species in the multi-variate space. The first discriminant function ac-counted by 91.2% of variation among the threespecies, and was mainly explained by the num-ber of transversal rows of ventral scales (VT)and number of gular scales (GUL) (table 5). Inthis axis, A. jalapensis present negative loadingsand is differentiated from A. xacriaba sp. nov.and A. mumbuca which present positive load-ings (fig. 6). Thus, A. jalapensis present a sig-nificantly higher number of gulars scales thanA. xacriaba sp. nov. (ANOVA, F1,16 = 20.81,P < 0.01) and A. mumbuca (F1,17 = 11.76,P < 0.01), and a lower number of transverserows of ventrals (F2,25 = 29.22, P < 0.01). Thesecond function accounted by the remaining8.8% of variation among groups and was mainlyexplained by the number of femoral pores (FP)and dorsal scales (D) (table 5). In this axis, A.

Figure 6. Scatter plot of results of a discriminant analysis onmeristic variables using individuals of Ameivula xacriabasp. nov. (triangles), A. jalapensis (circles) and A. mumbuca(squares). The percentage of variation explained by eachdiscriminant function is presented between brackets. Groupcentroids are represented by black crosses. This figure ispublished in color in the online version.

xacriaba sp. nov. is distinguished from A. mum-buca in presenting significantly lower numberof pores (F1,16 = 33.33, P < 0.01) and dor-sals scales (F1,16 = 6.18, P < 0.05). Basedon the discriminant function, 89.3% of the indi-viduals were correctly classified in each species(table 6).

Discussion

In the last 15 years, the taxonomy of lizards cur-rently assigned to the genus Ameivula has un-dergone important changes. For a long time, asingle species (Cnemidophorus ocellifer) wasadmitted, which was widely distributed alongopen habitats of Cis-Andean South Amer-ica with conspicuous geographical variation incolor pattern and lepidosis (Rodrigues, 1987).Subsequent studies led to the description of sev-eral new species: A. nativa, A. littoralis, and A.abaetensis from the coastal Restingas (Rocha


Table 6. Jackknifed classification matrix based on meristic data of Ameivula xacriaba sp. nov., A. jalapensis and A. mumbuca.

N A. xacriaba sp. nov. A. jalapensis A. mumbuca % correct

A. xacriaba sp. nov. 9 7 0 2 77.8A. jalapensis 10 0 10 0 100A. mumbuca 9 1 0 8 88.9

et al., 1997, 2000; Dias et al., 2002); A. vene-tacauda, A. confusioniba, A. nigrigula, A. cya-nura, and A. pyrrhogularis from the Caatingaand transitional zones between Caatinga andCerrado (Arias et al., 2011a, 2011b; Silva andÁvila-Pires, 2013); A. abalosi from the Chaco(Cabrera, 2012); Cnemidophorus parecis, A.mumbuca, and A. jalapensis from the Cer-rado (Colli et al., 2003a, 2003b, 2009). Cur-rently, Cnemidophorus parecis is not consid-ered a member of the genus, because molecularand morphological evidence (Guigliano et al.,2006, 2013; Arias et al., 2011a) indicates thatthis species is a member of Ameiva genus.

Ameivula was recently proposed by Harveyet al. (2012) as a new genus to accommodatethe species formerly attributed to the Cnemi-dophorus ocellifer group that formed a mono-phyletic unit in their phylogenetic analysis. Thegenus was diagnosed as follows: (1) presence ofa long first supraciliary; (2) prefrontal usually incontact with nasal; (3) prefrontal separated fromthe first supraciliary; and (4) papillate catch-ment folds and awns present in the hemipenis(Harvey et al., 2012). However, this diagnosisis somewhat problematical as these charactersshow some degree of variation within the genus.A long first supraciliary is actually absent in allspecies of the A. littoralis group, A. abalosi andA. confusioniba, while the prefrontal does notcontact the nasal in A. abalosi and the prefrontalcontacts the first supraciliary in all species ofthe A. littoralis group (except in the holotypeof A. littoralis) (Arias et al., 2014). Therefore,an improved phylogenetic analysis, includingspecies of both A. littoralis and A. ocelliferagroups, based on additional evidence both tax-onomically and geographically accurate, is stillneeded in order to adequately define and diag-nose the genus.

Our results, based on a multivariate analy-sis of morphometric characters, show that thespecies of the Ameivula ocellifera group clus-ter in two groups, the first one being formedby the Caatinga species (A. nigrigula, A. aff.ocellifera, A. confusioniba, and A. cipoensis)while the second one is composed by the Cer-rado species (A. mumbuca, A. jalapensis and A.xacriaba sp. nov.), were the Caatinga speciesare significantly larger than Cerrado species.This shows that there are more than two mor-phological groups in Ameivula, but their mono-phyly must be tested with phylogenetic analy-sis. Furthermore, the differences between Cer-rado and Caatinga species suggest that environ-ment pressure has an important role in the bodysize of this group of lizards. Futures studies,such as biogeographc, phylogeographic, phylo-genetic (among other), are needed to understandthe evolutionary history of this group of lizards.

Our analysis also separates A. mumbuca, A.jalapensis, and A. xacriaba sp. nov. on thebasis of meristic characters such as numberof transversal rows of ventral scales, numberof gular scales, number of femoral pores, anddorsal scales. Additional characters diagnosingA. xacriaba sp. nov. from A. jalapensis andA. mumbuca are: (1) the morphology of thesubdigital lamellae, keeled in A. jalapensis andA. mumbuca, smooth in A. xacriaba sp. nov.;(2) the clutch size, with reproductive female ofA. mumbuca and A. jalapensis (n = 4 for eachspecies) containing a single egg, confirmingprevious observations (Colli et al., 2003b; Colliet al., 2009) while the reproductive females(n = 4) of A. xacriaba sp. nov. retaining twoeggs; and (3) the color pattern in male andfemale (adult and juveniles) of A. xacriaba sp.nov. with ocelli on lateral fields, while in A.


Figure 7. Photo in life showing variation in the color patternamong male of Ameivula xacriaba sp. nov. (A), A. mumbuca(B) and A. jalapensis (C). This figure is published in colorin the online version.

mumbuca only some males have 2-3 ocelli, andin A. jalapensis lack ocelli (fig. 7).

Nogueira et al. (2011) showed that distribu-tion patterns and endemism of Cerrado squa-mates are affected by geomorphological units,varying between headwater plateaus (between500 and 1700 m) and peripheral depressions(below 500 m). The Planalto dos Gerais (orSerra Geral plateau, including the Jalapão re-gion) was indicated for these authors as one ofthe areas of endemism for Cerrado squamatereptiles. Furthermore, recent studies on Qua-ternary climatic fluctuations indicates that thePlanalto dos Gerais is a large refugial area (Wer-neck et al., 2012), which currently harbors manypotential endemic species, such as: Ameivulamumbuca, A. jalapensis, Ameiva xacriaba sp.nov., Anops acrobeles, Bachia oxyrhina, Bachia

geralista, and other undescribed species. Thefast loss of the original vegetation cover in theregion, which were in the last decades largelyreplaced by soy plantations (Teixeira et al.,2013; Klink and Machado, 2005), threatens se-riously this new and still poorly known species,and the remaining herpetofauna of the Planaltodos Gerais.

Acknowledgements. We thank Consejo de Investigaciónde la Universidad Nacional de Salta (CIUNSa), ConsejoNacional de Ciencia y Tecnología (CONICET-IBIGEO),Fundação de Amparo à Pesquisa do Estado de São Paulo(FAPESP), Conselho Nacional de Desenvolvimento Cien-tífico e Tecnológico (CNPq), IBAMA, and ICMBIO for fi-nancial and legal supports. We also thank Agustin Camacho,Marco Aurélio de Sena, Senhor Norinho and Dona Nita andthe officers from Parque Nacional das Cavernas do Peruaçufor logistical support and help with fieldwork; and PedroNunes for help in the preparation of the hemipenis.

References

Ab’Saber, A.N. (1983): O domínio dos cerrados: introduçãoao conhecimento. Revista do Serviço Público 111: 41-55.

Arias, F., de Carvalho, C.M., Rodrigues, M.T., Zaher, H.(2011a): Two new species of Cnemidophorus (Squa-mata: Teiidae) from the Caatinga, northwest of Brazil.Zootaxa 2787: 37-54.

Arias, F., de Carvalho, C.M., Rodrigues, M.T., Zaher, H.(2011b): Two new species of Cnemidophorus (Squa-mata: Teiidae) of the C. ocellifer group, from Bahia,Brazil. Zootaxa 3022: 1-21.

Arias, F., de Carvalho, C.M., Zaher, H., Rodrigues, M.T.(2014): A new species of Ameivula (Squamata: Tei-idae) from Southern Espinhaço Mountain Range, Brazil.Copeia 2014: 95-105.

Cabrera, M.R. (2012): A new species of Cnemidophorus(Squamata, Teiidae) from South American Chaco. Her-petological Journal 22: 123-131.

Cei, J.M. (1993): Reptiles del Noroeste, Nordeste y Este dela Argentina. Herpetofauna de las Selvas Subtropicales,Puna y Pampas. Museo Regionale di Scienze Naturali,Torino, Monografie XIV.

Cole, M. (1986): The Savannas: Biogeography and Geob-otany. Academic Press, London.

Colli, G.R., Costa, G.C., Garda, A.A., Kopp, K.A.,Mesquita, D.O., Peres, A.K., Valdujo, P.H., Vieira,G.H., Wiederhecker, H.C. (2003a): A critically endan-gered new species of Cnemidophorus (Squamata, Tei-idae) from a Cerrado enclave in southwestern Amazonia,Brazil. Herpetologica 59: 76-88.

Colli, G.R., Caldwell, J.P., Costa, G.C., Gainsbury, A.M.,Garda, A.A., Mesquita, D.O., Filho, C.M.M.R., Soare,A.H.B., Silva, V.N., Valdujo, P.H., Vieira, G.H.C., Vitt,


L.J., Werneck, F.P., Wiederhecker, H.C., Zatz, M.G.(2003b): A new species of Cnemidophorus (Squamata,Teiidae) from the Cerrado biome in central Brazil. Occa-sional Papers Sam Noble Oklahoma Museum of NaturalHistory 14: 1-14.

Colli, G.R., Giugliano, L.G., Mesquita, D.O., França, F.G.R.(2009): A new species of Cnemidophorus from the Jala-pao Region, in the central Brazilian Cerrado. Herpeto-logica 65: 311-327.

Dias, E.J.R., Rocha, C.F.D., Vrcibradic, D. (2002): NewCnemidophorus (Squamata: Teiidae) from Bahia state,northeastern Brazil. Copeia 2002: 1070-1077.

Dias, E.J.R., Rocha, C.F.D. (2004): Thermal ecology, ac-tivity patterns, and microhabitat use by two sympatricwhiptail lizards (Cnemidophorus abaetensis and Cnemi-dophorus ocellifer) from northeastern Brazil. Journal ofHerpetology 38: 586-588.

Dias, E.J.R., Rocha, C.F.D. (2007): Niche differences be-tween two sympatric whiptail lizards (Cnemidophorusabaetensis and C. ocellifer, Teiidae) in the Restingahabitat of northeastern Brazil. Brazilian Journal of Bi-ology 67: 41-46.

Dirksen, L., De la Riva, I. (1999): The lizards and amphis-baenians of Bolivia (Reptilia, Squamata): checklist, lo-calities, and bibliography. Graellsia 55: 199-215.

Giugliano, L.G., Contel, E.P.B., Colli, G.R. (2006): Ge-netic variability and phylogenetic relationship of Cnemi-dophorus parecis (Squamata, Teiidae) from cerrado iso-lates in southwestern Amazonia. Biochemical Systemat-ics and Ecology 34: 383-391.

Giugliano, L.G., Nogueira, C., Valdujo, P.H., Collevatti,R.G. (2013): Cryptic diversity in South America Teiinae(Squamata: Teiidae) lizards. Zoologica Scripta 42: 473-487.

Harvey, M.B., Ugueto, G.N., Gutberlet Jr., R.L. (2012):Review of teiid morphology with a revised taxonomyand phylogeny of the Teiidae (Lepidosauria: Squamata).Zootaxa 3459: 1-156.

Klink, C.A., Machado, R.B. (2005): Conservation of theBrazilian Cerrado. Conservation Biology 19: 707-713.

Manzani, P.R., Abe, A.S. (1988): Sobre dois novos métodosde preparo de hemipenis de serpentes. Memórias doInstituto Butantan 50: 15-20.

Menezes, V.A., Rocha, C.F., Dutra, G.F. (2004): Reproduc-tive ecology of the parthenogenetic whiptail lizard Cne-midophorus nativo in a Brazilian Restinga habitat. Jour-nal of Herpetology 38: 280-282.

Menezes, V.A., Van Sluys, M., Fontes, A.F., Rocha, C.F.(2011): Living in a caatinga-rocky field transitional habi-tat: ecological aspect of the whiptail lizard Cnemidopho-rus ocellifer (Teiidae) in the northeastern Brazil. Zoolo-gica 28: 8-16.

Menezes, V.A., Rocha, C.F. (2011): Thermal ecology offive Cnemidophorus species (Squamata: Teiidae) in eastcoast of Brazil. Journal of Thermal Biology 36: 232-238.

Nogueira, C., Rodrigues, M.T. (2006): The genus Steno-cercus (Squamata: Tropiduridae) in Extra-AmazonianBrazil, with the description of two new species. SouthAmerican Journal of Herpetology 1: 149-165.

Nogueira, C., Ribeiro, S., Costa, G.C., Colli, G.R. (2011):Vicariance and endemism in a Neotropical savannahotspot: distribution patterns of Cerrado squamate rep-tiles. Journal of Biogeography 38: 1907-1922.

Pesantes, O. (1994): A method for preparing the hemipenisof preserved snakes. Journal of Herpetology 28: 93-95.

Recoder, R., Nogueira, C. (2007): Composição e diver-sidade de répteis Squamata na região sul do ParqueNacional Grande Sertão Veredas, Brasil central. BiotaNeotropica 7: 267-278.

Rocha, C.F.D., Bergallo, H.G., Peccinini Seale, D. (1997):Evidence of and unisexual population of the Brazilianwhiptail lizard genus Cnemidophorus (Teiidae), withdescription of a new species. Herpetologica 53: 374-382.

Rocha, C.F.D., Menezes, V.A., Bergallo, H.G., Dutra, G.F.(1999): Cnemidophorus nativo geographic distribution.Herpetological Review 30: 109.

Rocha, C.F.D., Araujo, A.F.B., Vrcibradic, D., Da Costa,E.M.M. (2000): New Cnemidophorus (Squamata; Tei-idae) from coastal Rio de Janeiro state, southeasternBrazil. Copeia 2000: 501-509.

Rodrigues, M.T. (1987): Sistemática, ecologia e zoo-geografía dos Tropidurus do grupo torquatus ao sul doRio Amazonas (Sauria, Iguanidae). Arquivos de Zoolo-gia. São Paulo 31: 205-230.

Sales, R.F.D., Ribeiro, L.B., Jorge, J.S., Freire, E.M.X.(2012): Feeding habits and predator-prey size relation-ships in the whiptail lizard Cnemidophorus ocellifer(Teiidae) in the semiarid region of Brazil. South Ameri-can Journal of Herpetology 7: 149-156.

Santana, G.G., Vasconcellos, A., Gadelha, Y.E.A., Vieira,W.L.S., Almeida, W.O., Nóbrega, R.P., Alves, R.R.N.(2010): Feeding habits, sexual dimorphism and size atmaturity of the lizard Cnemidophorus ocellifera (Spix,1825) (Teiidae) in a reforested Restingas habitat inNortheastern Brazil. Brazilian Journal of Biology 70:409-416.

Silva, J.M.C. (1997): Endemic bird species and conserva-tion in the Cerrado region, South America. Biodiversityand Conservation 6: 435-450.

Silva, M.B., Ávila-Pires, M.T. (2013): The genus Cne-midophorus (Squamata: Teiidae) in States of Piauí,northeastern Brazil, with description of a new species.Zootaxa 3681: 455-477.

Teixeira Jr., M. (2010): Os lagartos do vale do Rio Peruaçu,MG, Brasil: Aspectos biogeográficos, história natural eimplicações para a conservação. Universidade de SãoPaulo, São Paulo.

Teixeira Jr., M., Recoder, R.S., Camacho, A., De Sena,M.A., Navas, C.A., Rodrigues, M.T. (2013): A newspecies of Bachia Gray, 1845 (Squamata: Gymnoph-thalmidae) from the Eastern Brazilian Cerrado, and dataon its ecology, physiology and behavior. Zootaxa 3616:173-189.

Vanzolini, P.E. (1974): Ecological and geographical dis-tribution of lizards in Pernambuco, northeastern Brasil(Sauria). Papéis Avulsos de Zoologia, São Paulo 28: 61-90.


Vanzolini, P.E. (1976): On the lizards of a Cerrado-Caatingacontact, evolutionary and zoogeographical implications(Sauria). Papéis Avulsos de Zoologia, São Paulo 29:111-119.

Vanzolini, P.E., Ramos, A., Vitt, L.J. (1980): Répteis dasCaatingas. Academia Brasileira de Ciências, Rio deJaneiro, Brazil.

Vitt, L.J. (1991): An introduction to the ecology of Cerradolizards. Journal of Herpetology 25: 79-90.

Werneck, F.P. (2011): The diversification of eastern SouthAmerican open vegetation biomes: historical biogeogra-phy and perspectives. Quaternary Science Reviews 30:1630-1648.

Werneck, F.P., Nogueira, C., Colli, G.R., Sites Jr., J., Costa,G.C. (2012): Climatic stability in the Brazilian Cerrado:implications for biogeographical connections of SouthAmerican savannas, species richness and conservationin a biodiversity hotspot. Journal of Biogeography 3: 1-12.

Zaher, H. (1999): Hemipenial morphology of the SouthAmerican Xenodontine snakes, with a proposal for amonophyletic Xenodontinae and a reappraisal of Colu-broid hemipenes. Bulletin of the American Museum ofNatural History 240: 1-168.

Zaher, H., Prudente, A.L.C. (2003): Hemipenes ofSiphlophis (Serpentes, Xenodontinae) and techniques ofhemipenial preparation in snakes: a response to Dowl-ing. Herpetological Review 34: 302-307.

Zar, J.H. (2010): Biostatistical Analysis, 5th Edition. Pren-tice Hall, New Jersey, 960 pp.

Submitted: August 26, 2013. Final revision received: March19, 2014. Accepted: April 21, 2014.Associate Editor: Sylvain Ursenbacher.

Appendix. Examined specimens

Ameivula abaetensis. Brazil, Sergipe: MZUSP 79610,80982-80983, Barra dos Coqueiros. MZUSP 88104,88106, 88108, Estação Ecológica Serra de Itabaiana.MZUSP 49385-49387, Santo Amaro das Brotas. Bahia:MZUSP 56559-56567, Guarajuba. MZUSP 96847-50,Jandaíra. MNRJ 8618, 8623, 8631, 8640, 8642, 8644,8645, 8647, 8650, 8658, 8687, 8681, 8685, 9300-9301,Salvador (paratypes).

Ameivula abalosi. Argentina, Salta: MCN 2807, Irigoyen,Oran. FML 8278-8279, La Quena, Dpto. Gral. SanMartín; Santiago del Estero: MCN 1105-1119. San-tiago del Estero Parque Nacional Copo, Monte Que-mado. FML 1346-1, 1346-2, Caspi Corral, Dpto.Figueroa. Chaco UNNE 1096, 10274, 9131, 1, 3, 5, 6,1095, 10464, 4586. Comandante Frias, Dpto. GeneralGuemes. UNNEC 5676, 5681, 5692, 6826, La Libertad,Dpto. Bermejo, Formosa.

Ameivula cipoensis. Brazil, Minas Gerais: MZUSP 103264-103670, MZUSP 103272-103280, MZUSP 54899-54900; MZUSP 54941-54943; MZUSP 55476; MZUSP

55534-55539; MZUSP 68628; MZUSP 72416-72418;MZUSP 78126-78127; MZUSP 78224; MZUSP 78586-78589; MZUSP 95176. Serra do Cipó.

Ameivula confusioniba. Brazil, Piauí: MZUSP 100187-1000205. Parque Nacional da Serra das Confusões,Caracol.

Ameivula cyanura. Brazil, Bahia: MZUSP 65779-65784,65801, 62801-62810, 56115-56127, 74215-74222,Morro do Chapéu, MZUSP 56286-56287, 72420-72422,Santo Inácio. MZUSP 56286.

Ameivula jalapensis. Brazil, Tocantins: MZUSP 100168and 100169, São Félix do Tocantins. MTR 14574,14503, 14616, 14576, 14502, 14573, 14714, 14596,14627, 14399. Almas (Estação Ecológica Serra Geral doTocantins).

Ameivula littoralis. Brazil, Rio de Janeiro: MNRJ 6555-6556, 6559-6560, 6564-6566, 6578-6579, 6585, 6587,6592, 6599, 6601, 6603-6604, 6607, 6621, 6665, Barrade Maricá (paratypes).

Ameivula mumbuca. Brazil, Tocantins: MZUSP 92547-92548, Mateiros (paratypes). MZUSP 94102-94109,93415, Mateiros (Estação Ecológica Serra Geral do To-cantins).

Ameivula aff. mumbuca. Brazil, Tocantins: MZUSP 96473-96518, Luis Eduardo Magalhães. MZUSP 98313-98331; São Salvador. Goiás: MZUSP 97054-97064,Serra da Mesa. MZUSP 97248-97264, Cana Brava.MZUSP 69367, Monte Alegre.

Ameivula nativa. Brazil, Espírito Santo: MNRJ 39545,4713, 4715, 4718-4719, 4722, 4724-4725, 4730, 4732,4734-4738, Linhares (paratypes). MZUSP 95100-95101, Conceição da Barra. Bahia: MZUSP 96833-96834, Itacaré.

Ameivula nigrigula. Brazil, Bahia: MZUSP 93706-93772,93813-93828, Santo Inácio. MZUSP 55290-55310,Catinga do Moura. Minas Gerais: MZUSP 78577-78589, Mocambinhos. MZUSP 56919-56932, GrãoMogol.

Ameivula ocellifera. Brazil, Bahia: MZUSP 191-196, 201-206, 6149-6152, 6700, 8468, 26827-26859, 65762, Sal-vador.

Ameivula aff. ocellifera. Brazil, Bahia: MZUSP 30308-30320, Cocorobó. MZUSP 12202-12210, 60875,65812-65816, Raso da Catarina. MZUSP 57243-57248,Mucugê.

Ameivula venetacauda. Brazil, Piauí: MZUSP 100115-100119, 100120-100127. Parque Nacional da Serra dasConfusões. Caracol.

Ameivula xacriaba sp. nov. Brazil, Bahia: MTR 15618 Jab-orandi, MTR 17857-17859, 17890-17895, São Deside-rio. Minas Gerais: MZUSP 94077-94080, Formoso.MZUSP 67085-67088, Arinos. MZUSP 99690; 99676,99685, Januária (Parque Nacional Caverna do Peruaçu).

Whiptail lizards in South America: a new Ameivula (Squamata, Teiidae) from Planalto dos Gerais,...

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