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

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Whiptail Lizards in South America: a New Ameivula (Squamata, Teiidae) from Planalto Dos Gerais, Eastern Brazilian Cerrado 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 morpholo- gical 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- © Koninklijke Brill NV, Leiden, 2014. DOI:10.1163/15685381-00002948 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, 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 ex- cluding 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 addi- tion, we took the following measurements using a digital
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