Whiptail Lizards in South America: a New Ameivula (Squamata, Teiidae) from Planalto Dos Gerais, Eastern Brazilian Cerrado

Home , Ameivula, Cnemidophorus, Contomastix, Teiidae

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 that extends along the Cerrado region in the states of Bahia, Tocantins, Piauí, and Minas Gerais, in Brazil. The new species is the third recognized species of the A. ocellifera group registered for the Cerrado. Quantitative analyses of morphometric characters 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 two Cerrado species A. mumbuca and A. jalapensis, also the morphology of the ﬁnger lamellae, and clutch size distinguish these species.

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

Introduction Currently, Ameivula has thirteen recognized species (table 1) distributed throughout the Cis- In a recent phylogenetic analysis based on Andean open-areas south of the Amazon River morphological traits, the genus Cnemidopho- (Arias et al., 2011b; Harvey et al., 2012; Silva rus resulted not monophyletic (Harvey et al., and Ávila-Pires, 2013). These species generally 2012), as previously suggested by molecular occur in open habitats with sandy soils and high studies (Reeder et al., 2002; Guigliano et al., temperature, in very distinct landscape phys- 2006; Guigliano, 2009). To resolve the poly- iognomies, such as the Caatinga (Vanzolini, phyly, Cnemidophorus was divided in four ge- nera: (1) the genus Cnemidophorus, including 1974; Vanzolini et al., 1980; Vitt, 1983, 1995; the species of the former C. lemniscatus group; Menezes et al., 2011; Sales et al., 2012), Cer- (2) the new genus Aurivela which includes two rado (Vitt, 1991), “campos rupestres” (Menezes species (A. longicauda and A. tergolaevigata); et al., 2011; Arias et al., 2014), Restingas (Dias (3) the new genus Contomastix assembling the and Rocha, 2004; Dias and Rocha, 2007; San- species previously referred to the Cnemidopho- tana et al., 2010; Menezes and Rocha, 2011), rus lacertoides group; and (4) the new genus and the Bolivian, Paraguayan and Argentinian Ameivula, composed by the species of the for- Chaco (Williams and Tedesco, 1985; Cei, 1993; mer Cnemidophorus ocellifer group. Dirksen and De la Riva, 1999; Cabrera, 2012). The species of Ameivula exhibit morphological characteristics (both in lepidosis as color 1 - Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Caixa Postal 11.46 1, CEP pattern) that distinguish two groups of the 05422-970, São Paulo, Brazil species (Arias et al., 2011a): (1) the A. littoralis 2 - Instituto de Bio y Geociencias (IBIGEO) del Noa, Fac- group, composed by A. abaetensis, A. littoralis, ultad de Ciencias Naturales, Universidad Nacional de Salta, Av. Bolívia 5150, 4400 Salta, Argentina A. cyanura, and A. venetacauda; and (2) the 3 - Instituto Nacional de Pesquisas da Amazônia, Núcleo de A. ocellifera group, composed by A. abalosi, Pesquisas de Roraima, Rua Coronel Pinto, 327, Centro, A. ocellifera, A. mumbuca, A. jalapensis, A. ni- CEP 69301-150, Boa Vista, Roraima, Brazil grigula, A. pyrrhogularis and A. confusioniba. 4 - Museu de Zoologia, Universidade de São Paulo, Caixa Postal 42694, CEP 04299-970, São Paulo, Brazil In addition, characters on hemipenis morphol- ∗Corresponding author; e-mail: [email protected] ogy and statistical analyses on continuous char-

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, between northern 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). Probably in 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 occurs in high elevation between 900 and about 1200 m, were the landscape is that 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 and Ávila-Pires, 2013). Member of A. ocellifera group. A. cyanura (Arias et al., 2011b) Santo Inácio and Morro do Chapéu, northwestern Bahia, Brazil; occurs in sympatry with A. nigrigula (Arias et al., 2011b). Member of A. littoralis group. A. jalapensis (Colli et al., 2009) Endemic of Jalapão region, north portion of the Cerrado biome, Tocantins state, Brazil (Colli et al., 2009). A. littoralis (Rocha et al., 2000) Endemic to the coastal Restinga areas of Rio de Janeiro state, southeastern Brazil (Rocha et al., 2000; Menezes et al., 2004). Member of A. littoralis group. 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 and southern 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 from northeastern Brazil to Mato Grosso do Sul, state. Currently considered a species complex. A. pyrrhogularis (Silva and Ávila-Pires, 2013) Northern and central part of the Piauí state, Brazil. Transition zone between 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), Serra da 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 the ancient vast plateaus (locally called cha- and Ávila-Pires, 2013). padões) have altitudes of 500-1700 m, and are The Cerrado biome is the largest Neotropi- dominated by savanna-like vegetation. These cal woodland-savanna, covering approximately plateaus are separated by a network of periph- 2 million km2, bearing high levels of endemism eral depressions (altitudes of 100-500 m) of (Oliveira and Marquis, 2002). Estimates indi- younger valleys (Ab’Saber, 1983; Silva, 1997; cate that approximately 80% of its original Werneck, 2011), where the vegetation is char- cover has been lost (Cavalcanti and Joly, 2002; acterized by recently originated plant commu- Klink and Machado, 2005). For these reasons, nities, including a mosaic of wet grassland, the Brazilian Cerrado is considered a biodi- cerrado vegetation, gallery forest, and patches versity “hostpot” (Mittermeier et al., 2004). of deciduous and semi-deciduous forest (Cole, The Cerrado geomorphology is characterized 1986; Silva, 1997; Werneck, 2011). Recent by the presence of distinct geomorphic surfaces: studies show that highland areas (chapadões) New Ameivula from eastern Brazilian Cerrado 229 played an important role in the history and Nacional del Nordeste, Corrientes, Argentina (UNNEC), current diversity of reptiles along the Cerrado and Colección Herpetológica Museo de Ciencias Naturales, Salta, Argentina (MCN). (Nogueira et al., 2011). We recorded the following characters in each individual: During an extensive field work in the Parque number of supraocular and supraciliar scales on right side; Nacional Cavernas do Peruaçu, on the south- femoral pores on both sides; longitudinal rows of ventral ern portion of Planalto dos Gerais, in the Mi- scales; transverse rows of ventral scales, from gular fold to anterior margin of hindlimbs; subdigital lamellae under nas Gerais state of Brazil, we obtained a large fourth finger and fourth toe; longitudinal rows of enlarged series of specimens of Ameivula that revealed scales in the dorsal region of the arm; scales around mid- to be very distinct from their congeners in lepi- body, counted midway between fore- and hindlimbs excluding ventrals; scales around tail, counted on the fourth dosis, hemipenis, color pattern, and morphome- transverse row; number of gular scales, counted along the try. Here, we describe these specimens as a new midline, from postsymphysal scale to mesoptychial scales; species of Ameivula. The new species is mor- dorsal scales, counted along the midline, from occiput to first transverse row of scales around tail; presence or ab- phologically and geographically more similar to sence of enlarged scales in the temporal region, posteriorly the two recognized species inhabiting the Cer- to the third subocular; presence or absence of keel in sub- rado biome: A. mumbuca and A. jalapensis, oc- digital lamellae both in the hand or foot; and presence or curring parapatricaly with both species in sandy absence of patch of tibiotarsal spurs in the males. In addition, we took the following measurements using a digital areas of Jalapão region, at the northern end of caliper to the nearest 0.02 mm: snout-vent length (SVL), Planalto dos Gerais. trunk length (TRL), head length (HL), head width (HW), forearm length (FAL), total hindlimbs length (HLL), and tail length (TAL). Scale observation and measurements were made under a stereomicroscope (10-40×). Moreover, we Materials and methods analyzed the color pattern and color in life in both males and females. We prepared the hemipenis of MZUSP 99677 Fieldwork and MZUSP 99623 following the procedures described by Specimens were collected at Parque Nacional Cavernas do Manzani and Abe (1988), subsequently modified by Pe- Peruaçu, municipality of Januária, in the northern portion of santes (1994) and Zaher (1999; see also Zaher and Prudente, the state of Minas Gerais, Brazil (Teixeira Jr., 2010). Most 2003). Later, we compared with the hemipenial morphology specimens were captured with the aid of pitfall traps that of A. confusioniba, A. pyrrhogularis, A. venetacauda (Silva consisted in four thirty-liters buckets that were buried at and Ávila-Pires, 2013), and A. cipoensis (Arias et al., 2014). ground level and connected to each other with three four- meter long drift fences, forming a Y shaped configuration. Quantitative analysis Traps were installed in the main vegetation physiognomies as follows: 15 traps on dry forest habitats; 15 on savanna; 17 Our observations, made both in the field and the collections on “carrascos” and two on riparian evergreen forest. Traps on the species of the Ameivula ocellifera group from Cer- remained opened during periods of 30 days in January (rainy rado and from Caatinga biome, let us distinguish two sub- season) and July (dry season) 2008 and 2009, totalizing an groups. One formed for the Caatinga species with conspicu- effort equivalent to 14 560 buckets × days. Collected in- ous sexual dimorphism and ontogenetic change in the color dividuals were fixed in 10% formalin, preserved in 70% pattern of the males. The other one, formed for the Cer- ethanol, and deposited in the Museu de Zoologia da Uni- rado species, being smaller and without ontogenetic change versidade de São Paulo, Brazil (MZUSP). in the color pattern. Here, we hypothesize that both subgroups can be also distinguished with morphometric characters. Thus, to test for significant morphometric differences Laboratory work between the two species subgroups within the Ameivula We compared the new species with 13 recognized species ocellifera group, we performed a two-way MANOVA using of Ameivula (listed in the Appendix) and added to our group and sex as factors and then performed a discriminant comparisons data of the coloration in life taken from the function analysis to find the best combination of variables original descriptions of A. littoralis (Rocha et al., 2000), that distinguish both subgroups. For this purpose, we se- A. mumbuca (Colli et al., 2003b), A. nativa (Rocha et al., lected six of the seven measurements: SVL, TRL, HL, HW, 1997), and A. jalapensis (Colli et al., 2009), while for A. FAL, HLL. We analyzed TAL separately because of miss- pyrrhogularis (Silva and Ávila-Pires, 2013) we take data ing data. We used a dataset containing measurements of five from its lepidosis and coloration. Specimens examined are males and five females of Ameivula sp. nov., A. jalapen- deposited at the Museu de Zoologia da Universidade de São sis and A. mumbuca (Cerrado group), A. aff. ocellifera, Paulo, Brazil (MZUSP), Museu Nacional do Rio de Janeiro, A. cipoensis, A. confusioniba and A. nigrigula (Caatinga Brazil (MNRJ), Fundación Miguel Lillo, Tucumán, Ar- group). As sexual dimorphism interacted with group dif- gentina (FML), Colección Herpetológica de la Universidad ferences (see results), we performed discriminant analyses 230 F.J. Arias et al. for males and females separately. Then, we tested for differences among the three species of the Cerrado group (i.e. Ameivula sp. nov., A. jalapensis and A. mumbuca)insize (SVL) with ANOVA and in shape with the remaining five characters using MANOVA after controlling for size. The size free database was obtained by regressing each mea- surement on a first principal component (82.3% of variation explained) obtained with a pooled within-group principal component analysis. We analyzed TAL separately using ANCOVA because of missing data. Morphometric measurements 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 Ameivula sp. nov., A. jalapensis and A. mumbuca.Analysiswas based on the following meristic characters: number of gular scales (GUL), transversal rows of ventral scales (VT), scales around midbody (SAM), scales around tail (SAT), femoral pores (FP), dorsal scales (D), subdigital lamellae under fourth 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 meristic data, and we performed all statistical analysis using IBM SPSS Statistics (version 20.0, SPSS Inc., 2011).

Results Ameivula xacriaba sp. nov. (figs 1, 2) Holotype. MZUSP 99690, an adult male from Parque Nacional Cavernas do Peruaçu (15°9 17.04 S, 44°18 18.29 W, 747 m a.s.l.), Figure 1. Dorsal, ventral and lateral view of the head of the Januária municipality, state of Minas Gerais, holotype of Ameivula xacriaba sp. nov., MZUSP 99690. Brazil. Collected by M.T. Rodrigues, R. Re- coder, M. Teixeira Jr., M. Aurelio de Sena and supracilaries; (5) 4-6 parietals; (6) 8 longitudi- A. Camacho on 17 January 2009. nal rows of ventral scales; (7) 27-29 transversal Paratypes. Adult males: MZUSP 99676, ventral rows; (8) 14-17 femoral pores; (9) 184- MZUSP 99685, MZUSP 99700, MZUSP 216 dorsal scales; (10) 24-29 scales around tail; 99679, MZUSP 99632, 99669, 99622, 99677, (11) 94-102 scales around midbody; (12) 13- 99641. Adult females: MZUSP 99642, MZUSP 15 fourth finger lamellae; (13) 23-28 fourth toe 99658, MZUSP 99638, MZUSP 99618, lamellae; (14) 41-47 gular scales; (15) absence MZUSP 99630, 99678, 99643. Collected by of paravertebral stripes; (16) lateral ocelli ex- M.T. Rodrigues, R. Recoder, M. Teixeira Jr., M. tending from hindlimbs to forelimbs. Aurelio de Sena and A. Camacho on 17 January The new species is assigned to the Ameivula 2009. All paratypes with the same locality data ocellifera group (Arias et al., 2011a) by a low as for the holotype. number of femoral pores (less than 20), enlarged scales in the temporal region (posteriorly Diagnosis. A slender, medium sized species to the third subocular), five supraciliary scales, of Ameivula, distinguished from all other 8 rows of ventral scales, and a brownish tail. species of the genus by the following combination of characters: (1) 6-7 supralabials; (2) 5- Comparison. Data from species in compari- 6 infralabials; (3) 4-5 supraoculars; (4) five son are given in parentheses; Ameivula xacriaba New Ameivula from eastern Brazilian Cerrado 231

stripe (present) and a stripe in the tail (present in A. abaetensis and A. littoralis). Ameivula xacriaba sp. nov. also differs from A. venetacauda by having laterodorsal and lateroventral stripes, and light green ocelli (no stripes or ocelli). Ameivula xacriaba sp.nov.differsfrom A. nativa by being bisexual (parthenogenetic species) and having 14-17 femoral pores (22- 26); 23-28 fourth toe lamellae (26-33); and by lacking a mid-dorsal stripe (present and with a posteriorly undulating margin). Ameivula xacriaba sp. nov. differs from A. jalapensis by having 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 lamellae smooth (subdigital lamellae keeled); a medium body size, maximum SVL = 70.2 mm, = 64.7 mm (smaller body size, maximum SVL = 54.2 mm, = 58 mm), tail length/SVL mean = 1.9 (mean = 2.6) a white throat (lime- green, Colli et al., 2009), ocelli on lower lateral fields (ocelli absent), and conspicuous sexual dichromatism (without sexual dichromatism, Colli et al., 2009). Ameivula xacriaba sp. nov. differs from A. mumbuca by having 14-17 femoral pores (16-20), subdigital lamellae without keel (subdigital lamellae keeled), ocelli on Figure 2. Photo in life of Ameivula xacriaba sp. nov., lower lateral fields in both males and females showing variation in male (A), female (B) and juvenile (C). Photo: Mauro Teixeira Jr. This figure is published in color (only in males; Colli et al., 2003), lateroven- in the online version. tral field bright green in males (bright yellow in males), and conspicuous sexual dichroma- sp. nov. can be distinguished from A. abaeten- tism (without sexual dichromatism). Ameivula sis, A. cyanura, A. littoralis and A. venetacauda xacriaba sp. nov. differs from A. confusioniba (i.e., members of the A. littoralis group) by hav- by having 23-28 fourth toe lamellae (29-35), ing enlarged scales in the lower temporal re- 14-17 femoral pores (16-21), 41-47 gular scales gion, posteriorly to the third subocular (scales (29-39), and bright green ocelli on lower lateral sub equal), three rows of enlarged scales in the fields extending from forelimbs to hindlimbs dorsal part of the arm (one row), 5 supraciliars (only 2-4 small yellow ocelli). Ameivula xacri- (6-7), 8 longitudinal rows of ventral scales (8- aba sp. nov. differs from A. ocellifera by hav- 10), 27-29 transversal ventral rows (29-38), 14- ing 23-28 fourth toe lamellae (28-31), 191-216 17 femoral pores (21-45), absence of tibiotarsal dorsal scales (172-188), 41-47 gular scales (32- spurs in the males (with 1-2 rows of 4-6 spurs), 35), without paravertebral stripes (with marked and by having a brown tail (bluish green). Fur- paravertebral stripes), and sexual dichromatism thermore, it differs from A. abaetensis, A. cya- (without sexual dichromatism). Ameivula xacri- nura and A. littoralis by lacking of a mid-dorsal aba sp. nov. differs from A. nigrigula by hav- 232 F.J. Arias et al. ing 13-15 fourth finger lamellae (16-18), 23-28 onal, in broad contact along midline; contact- fourth toe lamellae (30-32), 23-29 scales around ing postnasal, loreal, frontal and first supraoc- the tail (27-32), 41-47 gular scales (31-35), ab- ular; suture between prefrontals as long as sence of a patch of tibiotarsal spurs in males frontonasal. Frontal approximately pentagonal, (present), gular region white (dark black), and longer than wide, wider anteriorly with lateral by lacking ontogenetic color change in males margins slightly concave; contacting first left (present). Ameivula xacriaba sp. nov. differs supraocular. Two longer than wide, and approx- from A. abalosi by having 14-17 femoral pores imately pentagonal frontoparietals; wider pos- (16-20), 13-15 fourth finger lamellae (15-17), teriorly, slightly concave laterally; their mid- 41-47 gular scales (31-35), postnasal and pre- line suture straight. Five parietals, external ones frontal scales contacting each other (no con- smaller; interparietal sub-pentagonal, with lat- tact), and by lacking a distinctive patch of tibio- eral sides slightly concave, longer and wider tarsal spurs in males (present). Ameivula xacri- than others, bordered at each side by parietals. aba sp. nov. differs from A. pyrrhogularis by Occipital scales forming two transverse rows having 14-17 femoral pores (16-24); maximum regular in size. Four supraoculars on each side, SVL = 70.2 mm, = 64.7 mm (maximum third the longest; first in contact with loreal, SVL = 89 mm, = 77.5 mm); and gular prefrontal, frontal (on the left side), and first region white in males (orange in males) (Silva supraciliary; not contacting the second supraoc- and Ávila-Pires, 2013). Ameivula xacriaba sp. ular. Five supraciliaries on each side, second nov. differs from A. cipoensis by having 14- largest, the third and fourth sub-equal, the fifth 17 femoral pores (17-20); 13-15 fourth finger is smallest. Only first supraciliary in contact lamellae (12-14); 23-28 fourth toe lamellae (21- with first supraocular, all others separated from 24); 41-47 gular scales (31-38); 191-216 dor- supraoculars by a row of granules. Loreal sin- sal scales (202-225); four supraoculars (three); gle, large, as wide as third supralabial, as frontoparietal and third supraoculars never con- wide as high, in contact with postnasal, pre- tact (in contact); the third pair of chinshields the frontal, first supraocular, first supraciliary, first same size than the others (third pair larger than and second suboculars, and third and fourth the others). supralabial. First subocular narrow, higher than Description of holotype. Measurements: wide, in contact with second subocular, loreal, snout-vent-length (SVL) 60.7 mm, trunk length and first supraciliary. Four suboculars on each 29.6 mm, head length 14.7 mm, head width side, first and second keeled, second in con- 9.7 mm, head height 8.1 mm, tail length tact with the fourth supralabial; third subocu- 125 mm (2.1 times longer than SVL), humerus lar keeled, approximately rectangular, in contact length 5.6 mm, forearm length 19.7 mm, femur with fourth, fifth and sixth supralabials; fourth length 10.7 mm, tibia length 8.4 mm, foot length subocular smooth, as long and wide as the third, 17.5 mm, and hindlimb length 36.6 mm. with rounded posterior edges. Seven supralabi- Rostral large, wider than high, visible from als on each side, third the largest, posterior ones above, with the posterior edge sharp, sepa- smaller. Temporal region with irregular scales, rated from frontonasal by the midline con- granular centrally, enlarged ventrally. A lon- tact between nasals. Nasals roughly pentago- gitudinal supratemporal row with moderately nal smaller than prefrontals. Nasal and post- large scales, decreasing in size posteriorly. Ear nasal in broad contact by an oblique suture. opening large, round, with smooth margins, el- Nostril subovoid, in lower part of suture. Fron- lipsoid, slightly wider dorsally. All dorsal and tonasal hexagonal, wider than long, contacting lateral head scales juxtaposed, smooth (except nasals, postnasals and prefrontals. Prefrontals, for keeled suboculars). Symphysal as long as longer than wide roughly trapezoidal, pentag- wide, anteriorly concave, almost straight poste- New Ameivula from eastern Brazilian Cerrado 233 riorly where it contacts first infralabial and post- tral face of lower legs; elsewhere scales small, symphysal, forming two wide angles. Postsym- granular. Larger scales in longitudinal rows on physal single, pentagonal, longer than wide, in upper arms. Forearms with one row of enlarged contact with first and second infralabials; fol- scales, wider than long. Anterior scales on thigh lowed by five pairs of enlarged chinshields. First decreasing in size proximally, with three rows pair of chinshields in broad contact along mid- of enlarged scales. Lower legs with two rows line, in contact with third infralabials, slightly of enlarged, hexagonal scales. Ventral face of longer that the other. Second to fourth pairs of hands and feet granular; one enlarged tubercle chinshields separated from infralabials by a row at base of pollex. Sub-digital lamellae single; 14 of small granules. Six infralabials on each side; lamellae under fourth fingers; 24 under fourth followed posteriorly by series of small scales toes. extending to commissure of mouth; first infralabial the smallest. Gular region divided in two Color in life. Dorsal parts of head, body, limbs areas: anterior one with enlarged, rounded and and tail are brown. Lateral parts of the head are slightly elongate scales, disposed in irregular light brown, and white ventrally. The laterodor- 30 transverse rows, delimited posteriorly by a sal field is black, with irregular green spots ex- intertympanic sulcus; posterior region covered tends from neck to base of tail. Below it, a green with granules, in 17 transverse rows, smaller lat- laterodorsal stripe that extends from the supra- erally, bordered posteriorly by antegular fold. ciliar region to the base of the tail. The lateral Gular and antegular folds marked by granules. field is green and contains an irregular series Mesoptychial region with two irregular rows of of black spots interspersed with bright green enlarged scales between the two folds. Scales ocelli, between the fore and hindlimbs. Separat- on nape and sides of neck similar to dorsals. ing the lateral field from the ventrolateral field, Dorsals and scales on flanks granular, rounded, a lateroventral stripe that extends from nostril to smooth, sub-imbricate; 192 scales along a mid- hindlimbs. The lateroventral field is green with dorsal line from nape to base of tail, 99 scales irregular green ocelli. The dorsal and lateral sur- around mid-body (excluding ventrals). Ventral face of the tail is brown, and ventrally is white. scales large, smooth, rectangular (wider than The belly is light blushing while ventral parts of long), imbricated, in 28 transverse rows; 8 ven- the limbs, cloacal region and tail are white. tral scales in transverse rows across midbody. Color in ethanol. Head brownish dorsally, Ventral scales separated from scales on flanks labial regions and ventral aspect of head bluish by row of moderately enlarged scales. Preanal white. Dorsal parts of the body, limbs, and plate with three enlarged scales, a central one tail are bluish brown. Dorsolateral and lat- surrounded laterally by smaller ones and two eral fields dark black, separated by dorsolat- on posterior region. Femoral pores in a continu- eral stripe. A longitudinal series of small bluish ous row along each thigh, with a short medially white ocelli, between the fore and hindlimbs, gap; 8 pores on right side, 7 on left. Scales on is present in the lateral field. Ventrolateral field base of tail rectangular, smaller than ventrals, in bluish white with spots white. Belly light blue, transverse rows; keeled and slightly mucronate ventral aspect of the limbs, cloacal region and dorsal and laterally, smooth ventrally, imbri- tail are white. cate; 29 scales around tail. Tail scales becoming gradually longer and narrower from the base to Variation. Data based on five adult males and tip; subcaudal scales becoming keeled distally. five adult females, listed as range (mean ± Limbs with large, smooth, imbricate scales on SD): SVL 59.6-70.2 mm (62.8 ± 3.3 mm); dorsal face of upper arms, antero-dorsal face of maximum male SVL 70.2 mm; maximum forearms, antero-ventral face of thighs, and ven- female SVL 64.8 mm. Head length 13.7- 234 F.J. Arias et al.

19.4 mm (15.9 ± 1.7 mm). Head width 8.7- group lack these ornamentations, has a protu- 12.9 mm (10.7 ± 1.3 mm). Head height 7.6- berance on each lobule at the distal tip (api- 11.2 mm (8.9 ± 1.2 mm). Humeral length 4.3- cal region), and the body of the organ is cover 6.4 mm (5.1 ± 0.7 mm). Forelimb length 18.2- by transverse flounces (Silva and Ávila-Pires, 23.0 mm (19.8 ± 1.4 mm). Femur length 9.1- 2013). 11.5 mm (10.5 ± 0.8 mm). Tibia length 8.2- Etymology. The specific name “xacriaba” 9.9 mm (8.8 ± 0.5 mm). Foot length 17.5- honors the Xakriabás indians. A remaining 21.5 mm (18.7 ± 1.2 mm). Hindlimb 2.0 times longer than foot length. Hindlimb length community that occupies an Indian Territory in 35.2-42.8 mm (38.0 ± 2.3). Tail length 104- the vicinities of Peruaçu region and along the 134.6 mm (117.6 ± 12.2 mm), 1.9 times longer left bank of the river São Francisco, in the mu- than SVL. Supralabials scales 6-7 (6.3 ± 0.5); nicipalities of São João das Missões, state of infralabials scales 5-6 (5.2 ± 0.4). The speci- Minas Gerais, Brazil. men MZUSP 99685 has the frontonasal divided. Distribution and ecology. The new species is Variations in other meristic characters are sum- apparently endemic to the Planalto dos Gerais marized in table 2. (fig. 4), a large, eroded sandstone plateau that An evident sexual dichromatism is present, reaches 900 m in some points and extends especially during the reproductive season. In mainly along all the western region of Bahia adult males the flank color (between forelimbs state, in the eastern portion of the Brazilian Cer- and hindlimbs) is bright green, while in adult rado (Teixeira et al., 2013). The type series was females the lateroventral areas are bright yellow. all collected at Parque Nacional Cavernas do In the juveniles of both sexes the lateroventral Peruaçu, and it is the most common lizard in areas are light green, as for the ocelli (fig. 2). this region (Teixeira Jr., 2010). The new species Hemipenial morphology. Description is based was found only in savannas, despite significant on the fully everted and expanded right organ of sampling efforts involving pitfall traps and ac- MZUSP 99677 (fig. 3). Bilobed hemipenis. The tive searches in all vegetation types known in sulcus spermaticus is a broad channel, originat- the area (fig. 5). It is a Cerrado area that was ing at the base of the organ, reaching the apex. partially converted into pasturelands in the past, At the distal tip of the body the sulcus is di- but which is regenerating since 1997, and it vided in two lobes, each lobe has papillae at the is now protected. Except for some areas with outer edges, bellow them there are three sacs in bare sandy soil, the savanna vegetation in the each lobe. In the distal region there are 4-5 lam- region has an extensive herbaceous cover with inae, laterally located, separated by two longi- a sparse arboreal stratum that is predominantly tudinal septa on each side. The central zone of composed by porcada trees (Copaifera martii) distal region and proximal region of the organ and pequi trees (Caryocar brasiliense). are completely nude, in both faces (sulcate and Additional material from MZUSP confirms asulcate). the presence of Ameivula xacriaba sp. nov. in The hemipenial morphology of Ameivula Parque Nacional Grande Sertão Veredas, Minas xacriaba sp. nov. show features that are shared Gerais state, and in the municipalities of Jab- by other members of A. ocellifera group, such orandi and São Desiderio, state of Bahia (see as A. pyrrogularis, A. confusioniba, A. nigrigula Appendix). At Parque Nacional Grande Sertão (Silva and Ávila-Pires, 2013), and A. cipoensis Veredas Ameivula xacriaba sp. nov. is the sec- (Arias et al., 2014). The distal tip of the hemipe- ond most abundant species, after Vanzosaura nis of these species are papillate at the outer rubricauda, being sympatric with at least 25 edges, with two or three sacs at the edge of each other species of lizards, some of them typical of lobe. The hemipenis of the A. littoralis species the Serra Geral region (e.g., Stenocercus quinar- New Ameivula from eastern Brazilian Cerrado 235 12.0 0 128) ± ± 1.6 (23-33) 1.4 (16-22) 0.8 (26-31) 2.0 (24-34) 1.0 (13-19) = 7.1 (87-124) n ± ± ± ± ± ± A. pyrrhogularis ∗ standard deviation, with ± 25) ? 13 222.9 30 389.5 08 25) ( = ± ± (15-20) 18.5 (26-29) 28.3 1 (30-32) 29.8 ± n = 2.1 (31-35) ? 2.7(27-32) 26.7 3.0 (97-105) 104 ± ± 0.8 (16-18) 16.1 ± n 4.3 (44.7-55.3) 46.7 ± ± ± ± ± 20) 72% ( 7.3 208.6 22.5 328.4 = 40) ( ± 0(8) 8 ± n 1 (17-20) 17.1 5 (85-100) 99.8 = 1.6 (25-29) 30 2.2 (30-38) 33.3 0.8 (28-30) 27.4 1.2 (21-24) 31.3 0.8 (12-14) 17 ± 2.8 (39-49.5) 48.3 ± n ± ± ± ± ± ± ± 30) 100% ( 8.5 213.6 25.5 347.6 = 08 30) ( ± ± n 1 (15-17) 12.5 ± = 2.0 (22-28) 26.7 4.2 (29-39) 33.7 1.8 (16-21) 17.7 0.9 (27-29) 29.1 2.1 (29-35) 22.5 2.6 (42-50.5) 43.4 6.8 (87-105) 92.6 ± n ± ± ± ± ± group from Cerrado and Caatinga. Values indicate mean ± ± 38) 100% ( = 30.5 381.4 6.7 201.6 38) ( n ± 1.2 (6-8) 8 ± = 6.6 (41-64) 34.1 2.3 (10-16) 17.5 0.8 (25-27) 28.2 1.3 (14-16) 15.6 4.7 (89-100) 95 1.4 (22-26) 24.9 1.5 (23-28) 30.4 2.7 (40.3-49) 47.1 Ameivula ocellifera ± n ± ± ± ± ± ± ± ± 35) 23.68% ( 9.2 218 = 34.9 406.1 07.33 35) ( n ± ± ± = 2.0 (22-28) 24 4.4 (38-49) 51.3 1.3 (16-20) 13.1 0.6 (26-28) 25.7 1.5 (25-29) 26 3.0 (95-102) 94.3 0.6 (15-17) 14.7 n 2.4 (43.2-50.5) 44.3 ± ± ± ± ± ± ± A. mumbuca A. jalapensis A. confusioniba A. cipoensis A. nigrigula ± 92) 62.86% ( = sp. nov. 27.5 395.7 8.3 211.5 08 92) ( n ± ± 4 (41-52) 44.9 ± = 2.3 (23-29) 25.1 0.6 (13-15) 16 1.7 (23-28) 26.8 4.0 (94-106) 98.8 1.2 (14-17) 18.3 0.5 (27-29) 27.1 2 (47.2-52.2) 46.8 ± n ± ± ± ( (191-216) (200-227) (208-226) (188-211) (202-225) (190-220) (192-255) ± ± ± ± (341.4-417.4) (351.8-454.6) (356-454.6) (332-408) (307-373) (309.4-363) A. xacriaba Meristic characters take from the original description (Silva and Ávila-Pires, 2013). ∗ 100 49.2 100 383.6 × supraocular × a Comparison of selected meristic characters among the members of the and 2 a tail (SAT) ventral scales midbody (SAM) ventral scales 1 Scale around 99.3 Dorsal 202 range in parentheses. Transversal rows of 28 Longitudinal rows of 8 Four finger lamellae 14.1 Fourth toe lamellae 25.4 Scale around 26.3 SAM/SAT Table 2. Characters Gular scales 45.7 Femoral pores 15 % of contact between 52.17% ( SAM/Dorsal 236 F.J. Arias et al.

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. The triangle with a dot indicates the type locality of Ameivula xacriaba sp. nov. This ﬁgure is published in color in the online version. New Ameivula from eastern Brazilian Cerrado 237

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

Table 3. Results of a discriminant function analysis on and FAL for females (table 3). The two groups morphometric variables of Ameivula species from Cerrado and Caatinga groups. Scores with highest correlation with can be differentiated on basis of the significantly each discriminant function are highlighted in bold. larger size (ANOVA, males: F1,34 = 65.67, Males Females P<0.01, females: F1,34 = 20.05, P< Variables Coeffi- Corre- Coeffi- Corre- 0.01) and longer hind limbs of the species from cients lation cients lation the Caatinga group (ANCOVA, males: F1,34 = SVL −0.175 0.879 −0.565 0.517 4.23, P<0.05, females: F1,34 = 18.96, TRL 0.139 0.677 −0.443 0.372 P<0.01). Based on the discriminant function, HL 0.434 0.865 0.398 0.558 HW −0.291 0.676 0.610 0.496 88.6% of the individuals were correctly classi- FAL 0.364 0.837 0.516 0.639 fied in each group for males and 91.4% for fe- HLL 0.605 0.954 0.720 0.830 males (table 4). Eigenvalue 2.573 2.351 The three species from Cerrado (A. xacri- Wilks’ Lambda 0.280 0.298 aba sp. nov., A. mumbuca and A. jalapensis) showed significant differences in size (ANOVA, ius, Bachia geralista; Nogueira and Rodrigues, F2,29 = 28.88, P<0.01) but sexes did 2006; Recoder and Nogueira, 2007; Teixeira et not differed (F1,29 = 0.27, P = 0.607) al., 2013). and did not interacted with species differences (F = 0.49, P = 0.617). In post hoc com- Quantitative analyses. Group differences 2,29 were significant (MANOVA, Wilk’s Lambda = parisons, A. xacriaba sp. nov. is significantly larger than A. jalapensis (Tukey’s HSD Test, 0.313, F6,61 = 22.33, P<0.01). Sexual dimor- = phism in morphometry was detected (Wilk’s F1,9 32.4, P<0.01) and A. mumbuca (F = 5.47, P<0.05). Species did not dif- Lambda = 0.288, F6,61 = 25.11, P<0.01) 1,9 and interacted significantly with group differ- fered in shape (MANOVA, Wilk’s Lambda = = = ences (Wilk’s Lambda = 0.762, F6,61 = 3.17, 0.477, F10,40 1.79, P 0.09) and sexual P<0.01). dimorphism in shape was not detected (Wilk’s The discriminant function presented high cor- Lambda = 0.937, F10,40 = 1.31, P = 0.98). relation with HLL and SVL for males, and HLL However, A. xacriaba sp. nov. presented shorter 238 F.J. Arias et al.

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 93.3141 93.3 Caatinga 20 3 17 85.0 2 18 90.0

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

Variables DF1 DF2

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

Eigenvalue 11.853 1.140 Figure 6. Scatter plot of results of a discriminant analysis on % of variance 91.2 8.8 meristic variables using individuals of Ameivula xacriaba Cumulative % 91.2 100 sp. nov. (triangles), A. jalapensis (circles) and A. mumbuca (squares). The percentage of variation explained by each = discriminant function is presented between brackets. Group tails than A. jalapensis (ANCOVA, F1,9 6.79, centroids are represented by black crosses. This figure is P<0.05). published in color in the online version. A discriminant analysis performed on meristic characters using individuals of A. xacriaba xacriaba sp. nov. is distinguished from A. mum- sp. nov., A. jalapensis and A. mumbuca showed buca in presenting significantly lower number = a differentiation among the species in the multi- of pores (F1,16 33.33, P<0.01) and dor- = variate space. The first discriminant function ac- sals scales (F1,16 6.18, P<0.05). Based counted by 91.2% of variation among the three on the discriminant function, 89.3% of the indi- species, and was mainly explained by the num- viduals were correctly classified in each species ber of transversal rows of ventral scales (VT) (table 6). and number of gular scales (GUL) (table 5). In this axis, A. jalapensis present negative loadings Discussion and is differentiated from A. xacriaba sp. nov. and A. mumbuca which present positive load- In the last 15 years, the taxonomy of lizards cur- ings (fig. 6). Thus, A. jalapensis present a sig- rently assigned to the genus Ameivula has un- nificantly higher number of gulars scales than dergone important changes. For a long time, a A. xacriaba sp. nov. (ANOVA, F1,16 = 20.81, single species (Cnemidophorus ocellifer)was P<0.01) and A. mumbuca (F1,17 = 11.76, admitted, which was widely distributed along P<0.01), and a lower number of transverse open habitats of Cis-Andean South Amer- rows of ventrals (F2,25 = 29.22, P<0.01). The ica with conspicuous geographical variation in second function accounted by the remaining color pattern and lepidosis (Rodrigues, 1987). 8.8% of variation among groups and was mainly Subsequent studies led to the description of sev- explained by the number of femoral pores (FP) eral new species: A. nativa, A. littoralis, and A. and dorsal scales (D) (table 5). In this axis, A. abaetensis from the coastal Restingas (Rocha New Ameivula from eastern Brazilian Cerrado 239

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 77.8 A. jalapensis 10 0 10 0 100 A. mumbuca 91 0 888.9 et al., 1997, 2000; Dias et al., 2002); A. vene- Our results, based on a multivariate analy- tacauda, A. confusioniba, A. nigrigula, A. cya- sis of morphometric characters, show that the nura, and A. pyrrhogularis from the Caatinga species of the Ameivula ocellifera group clus- and transitional zones between Caatinga and ter in two groups, the first one being formed Cerrado (Arias et al., 2011a, 2011b; Silva and by the Caatinga species (A. nigrigula, A. aff. Ávila-Pires, 2013); A. abalosi from the Chaco ocellifera, A. confusioniba, and A. cipoensis) (Cabrera, 2012); Cnemidophorus parecis, A. while the second one is composed by the Cer- mumbuca, and A. jalapensis from the Cer- rado species (A. mumbuca, A. jalapensis and A. rado (Colli et al., 2003a, 2003b, 2009). Cur- xacriaba sp. nov.), were the Caatinga species rently, Cnemidophorus parecis is not consid- are significantly larger than Cerrado species. ered a member of the genus, because molecular This shows that there are more than two mor- and morphological evidence (Guigliano et al., phological groups in Ameivula, but their mono- 2006, 2013; Arias et al., 2011a) indicates that phyly must be tested with phylogenetic analy- this species is a member of Ameiva genus. sis. Furthermore, the differences between Cer- Ameivula was recently proposed by Harvey rado and Caatinga species suggest that environ- et al. (2012) as a new genus to accommodate ment pressure has an important role in the body the species formerly attributed to the Cnemi- size of this group of lizards. Futures studies, dophorus ocellifer group that formed a mono- such as biogeographc, phylogeographic, phylo- phyletic unit in their phylogenetic analysis. The genetic (among other), are needed to understand genus was diagnosed as follows: (1) presence of the evolutionary history of this group of lizards. a long first supraciliary; (2) prefrontal usually in Our analysis also separates A. mumbuca, A. contact with nasal; (3) prefrontal separated from jalapensis, and A. xacriaba sp. nov. on the the first supraciliary; and (4) papillate catch- basis of meristic characters such as number ment folds and awns present in the hemipenis of transversal rows of ventral scales, number (Harvey et al., 2012). However, this diagnosis is somewhat problematical as these characters of gular scales, number of femoral pores, and show some degree of variation within the genus. dorsal scales. Additional characters diagnosing A long first supraciliary is actually absent in all A. xacriaba sp.nov.fromA. jalapensis and species of the A. littoralis group, A. abalosi and A. mumbuca are: (1) the morphology of the A. confusioniba, while the prefrontal does not subdigital lamellae, keeled in A. jalapensis and contact the nasal in A. abalosi and the prefrontal A. mumbuca, smooth in A. xacriaba sp. nov.; contacts the first supraciliary in all species of (2) the clutch size, with reproductive female of the A. littoralis group (except in the holotype A. mumbuca and A. jalapensis (n = 4 for each of A. littoralis) (Arias et al., 2014). Therefore, species) containing a single egg, confirming an improved phylogenetic analysis, including previous observations (Colli et al., 2003b; Colli species of both A. littoralis and A. ocellifera et al., 2009) while the reproductive females groups, based on additional evidence both tax- (n = 4) of A. xacriaba sp. nov. retaining two onomically and geographically accurate, is still eggs; and (3) the color pattern in male and needed in order to adequately define and diag- female (adult and juveniles) of A. xacriaba sp. nose the genus. nov. with ocelli on lateral fields, while in A. 240 F.J. Arias et al.

geralista, and other undescribed species. The fast loss of the original vegetation cover in the region, which were in the last decades largely replaced 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 Planalto dos Gerais.

Acknowledgements. We thank Consejo de Investigación de la Universidad Nacional de Salta (CIUNSa), Consejo Nacional 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 and the officers from Parque Nacional das Cavernas do Peruaçu for logistical support and help with fieldwork; and Pedro Nunes for help in the preparation of the hemipenis.

References

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