Zootaxa, a New Microteiid Lizard of the Genus Acratosaura

A new microteiid lizard of the genus Acratosaura (Squamata: Gymnophthalmidae) from Serra do Sincorá, State of Bahia, Brazil

MIGUEL TREFAUT RODRIGUES1, JOSÉ CASSIMIRO1,3, MARCO ANTONIO DE FREITAS2 & THAÍS FIGUEIREDO SANTOS SILVA2 1Universidade de São Paulo, Instituto de Biociências, Departamento de Zoologia, Caixa Postal 11.461, CEP 05422–970, São Paulo, Brasil. E-mail: [email protected] 2Universidade Estadual Santa Cruz, Programa de Pós-Graduação em Zoologia, Rodovia Ilhéus/Itabuna km 16, 45.650–000, Ilhéus, BA, Brasil. E-mail: [email protected] (MAF) 3Corresponding author. E-mail: [email protected]

Abstract

A new species of the genus Acratosaura is described on the basis of two specimens obtained at the “campos rupestres” (rocky meadows) near Mucugê municipality, state of Bahia, Brazil. Acratosaura spinosa sp. nov., has ear openings and eyelids, pentadactyl limbs lacking a claw on first toe, a single frontonasal, prefrontals, frontoparietals, parietals, interparietals, occipitals, no collar fold, three pairs of genials, three supraoculars and three superciliaries and is further characterized by presenting 28–31 rows of strongly keeled, lanceolate and imbricate dorsal scales, sides of the neck with keeled scales, striate temporal scales, four longitudinal and 17–18 transverse rows of smooth ventral scales, 28–30 scales around the body, 14 and 19–20 infradigital lamellae under finger IV and toe IV, respectively, 13 total preanal and femoral pores in male, absent in female. The new species differs strikingly in color pattern from its only congener A. mentalis. The two species occur sympatricaly in the high altitude open habitats near the type locality.

Key words: Acratosaura spinosa sp. nov., Acratosaura mentalis, Lizards, Squamates, Reptiles, Distribution, Campos rupestres, Serra do Sincorá, Espinhaço range

Resumo

Uma nova espécie do gênero Acratosaura é descrita a partir de dois espécimes obtidos nos campos rupestres da vizinhança do município de Mucugê, estado da Bahia, nordeste do Brasil. Acratosaura spinosa sp. nov., apresenta ouvido externo, pálpebras, membros pentadáctilos sem unha no primeiro dedo, uma única frontonasal, prefrontais, frontoparietais, parietais, interparietais, occipitais, ausência de dobra colar, três pares de geniais, três supraoculares e três superciliares. A nova espécie é ainda caracterizada por apresentar de 28 a 31 fileiras de escamas dorsais fortemente quilhadas, lanceoladas e imbricadas, laterais do pescoço com escamas quilhadas e escamas temporais estriadas, quatro longitudinais e 17 a 18 fileiras transversais de escamas ventrais lisas, 28 a 30 escamas ao redor do corpo, 14 lamelas infradigitais sob o quarto dedo, 19 a 20 lamelas infradigitais sob o quarto artelho; 13 poros preanais e femorais no macho, ausentes na fêmea. A nova espécie difere marcadamente no padrão de colorido de sua única espécie congenérica, A. mentalis. As duas espécies ocorrem simpatricamente nos campos rupestres de altitude das proximidades da localidade- tipo.

Palavras-chave: Acratosaura spinosa sp. nov., Acratosaura mentalis, Lagartos, Squamata, Répteis, Distribuição, Campos rupestres, Serra do Sincorá, Cadeia do Espinhaço

Accepted by S. Carranza: 12 Dec. 2008; published: 16 Feb. 2009 17 Introduction

The gymnophthalmid lizard genus Acratosaura was recently created to allocate the former Colobosaura mentalis Amaral, 1933 (Rodrigues et al. 2007). Although the most evident external difference between Acratosaura and Colobosaura is the presence of three pairs of chin-chields in Acratosaura, this decision resulted from extensive phylogenetic studies of Gymnophthalmidae involving molecular (Castoe et al. 2004; Pellegrino et al. 2001) and both morphological and molecular characters (Rodrigues et al. 2007). According to the most recent phylogenetic studies, the sister genus of Acratosaura is Stenolepis and both are the sister group of a clade comprising the genera Alexandresaurus, Colobosaura and Iphisa. All these relationships were highly supported as was the monophyly of these five genera, which are presently being referred to as the Iphisiinae (Rodrigues et al. in press). Acratosaura is unique among them in having three pairs of chin-shields, all the other genera having two pairs. The Iphisiinae together with the Heterodactylini sensu stricto [i.e. Heterodactylus, Colobodactylus and a new genus under description (Rodrigues et al. in press)] and the Gymnophthalmini compose the subfamily Gymnophthalminae, which in turn together with the Rachisaurinae and the Alopoglossinae compose the Gymnophthalmidae (Pellegrino et al. 2001). Much of these results, which are progressively leading to a phylogenetic classification of Gymnophthalmidae, were possible thanks to the use of molecular characters in recent studies but also to the discovery of newly collected material, which increased substantially microteiid diversity. In recent surveys conducted in the northern part of highlands of the Espinhaço mountain range in state of Bahia, a region referred to as Chapada Diamantina, we obtained two specimens of a new species of Acratosaura herein described.

Material and methods

Snout-vent length (SVL) and tail length (TL) were measured to the nearest mm with a rule; scale counts were taken with the aid of a stereomicroscope. Scale counts and scale nomenclature are according to Rodrigues et al. (2007). Sex was determined by the presence/absence of preanal/femoral pores. All data were taken from preserved specimens housed at MZUSP (Museu de Zoologia, Universidade de São Paulo) and MNRJ (Museu Nacional, Rio de Janeiro) collections.

Acratosaura spinosa sp. nov. (Figs. 1, 2A, 3, 4, 5A)

Holotype: MZUSP 98088, an adult male obtained at 1,250 m at Fazenda Caraíbas (13º06'59"S 41º22'58"W), Serra do Sincorá, Espinhaço range, district of Cascavel, municipality of Mucugê, state of Bahia, Brazil, collected by Marco Antonio de Freitas and Thaís Figueiredo Santos Silva on 8 December 2005, field number MTR 13999. Paratype: MZUSP 98188, a female obtained at 997 m at Mucugê (13°01'38"S, 41°21'04"W), state of Bahia, Brazil, collected by José Cassimiro on 11 March 2005, field number JC 1211. Etymology: the specific epithet is given in reference to the spinose aspect of the body scales. Diagnosis: a gymnophthalmid with ear openings and eyelids, and slender pentadactyl limbs lacking the claw on the first toe. Frontonasal single; prefrontals, frontoparietals, parietals, interparietals, and occipitals present. Parietals longer than wide. Collar fold absent. Three pairs of genials; three supraoculars. Dorsal scales in 28–31 rows, strongly keeled, lanceolate, imbricate. Ventrals smooth, in four regular longitudinal and 17–18 transverse rows; external rows wider than long, midventral rows longer than wide. Scales around body 28–30; 14 and 19–20 infradigital lamellae under finger IV and toe IV respectively. Male with a continuous series of 13 total very conspicuous pores without gap between preanal and femoral (Fig. 3); pores absent in female.

NEW ACRATOSAURA FROM BRAZIL Zootaxa 2013 © 2009 Magnolia Press · 19 FIGURE 2. Acratosaura spinosa sp. nov. (holotype, MZUSP 98088), adult male (A), from Serra do Sincorá, Mucugê municipality, Bahia State; and Acratosaura mentalis (B) from Santo Inácio, Gentio do Ouro municipality, Bahia State, Brazil. Photographs: 2.A by Marco Antonio de Freitas and 2.B by Miguel T. Rodrigues.

20 · Zootaxa 2013 © 2009 Magnolia Press RODRIGUES ET AL. FIGURE 3. Preanal region of Acratosaura spinosa sp. nov. (holotype, MZUSP 98088), from Serra do Sincorá, Mucugê municipality, Bahia State. Photograph by Flávio Uemori Yamamoto.

Description of the holotype (Fig. 1, 2A, 3, 4, 5A): rostral broad, wider than high, contacting first supralabial, nasal and frontonasal. Frontonasal heptagonal, twice as wide as long, contacting rostral, nasal, loreal and prefrontals. Prefrontals hexagonal, as wide as long, in broad contact at midline. Frontal hexagonal, with almost parallel lateral margins, twice as long as wide, anteriorly indenting the prefrontal and posteriorly the frontoparietal sutures. Frontoparietals pentagonal, slightly larger than prefrontals, in broad contact, strongly indented by the interparietal. Interparietal longer than wide, longer and narrower than frontal, as long as and narrower than parietals. Parietals heptagonal, bordered laterally by three temporal scales, anteriorly by the third supraocular and frontoparietal, medially by the interparietal, and posteriorly by an occipital. Two pairs of distinctively enlarged occipitals separated at midline by a smaller scale. Three supraoculars, first the smallest, second the largest and in broad contact with frontal, third smaller, in broad contact with frontoparietal. Nasal above first supralabial, large, longer than high, with the nostril in the middle lower part of the scale, indenting the suture with the labial. Loreal posterior to nasal, diagonally oriented; contacting posteriorly first supraocular, first superciliary, preocular, and frenocular. Frenocular below preocular, followed posteriorly by a series of four suboculars, the first with the same approximate size of frenocular, second and third longer and narrower. Six supralabials, third and fourth under the eye, fifth the highest and contacting three smaller scales following posteriorly the subocular series. An elongate scale following sixth supralabial contacts the granules surrounding anterior margin of ear. Three superciliaries, first largest, wider anteriorly, longer than first supraocular, contacting preocular, loreal, first and second supraoculars, second superciliary and upper eyelid. Third superciliary followed by a smaller, higher than wide scale which contacts its posterior part, third supraocular and a small postocular. Central part of eyelid with a semitransparent undivided disc surrounded by small and slightly pigmented granular smooth scales. Lower eyelid with twelve strongly pigmented palpebrals. Temporal region with juxtaposed scales, irregular in size and shape; posterior ones

NEW ACRATOSAURA FROM BRAZIL Zootaxa 2013 © 2009 Magnolia Press · 21 largest, longitudinally striated. Ear opening surrounded by a series of juxtaposed granules, those from anterior margin elongate; external auditory meatus large, tympanum distinct, subovoid, recessed. Lateral surface of neck with keeled, imbricate scales which mostly are longer than wide. Except for striated temporals, all head scales smooth and juxtaposed with scattered sensorial organs.

FIGURE 4. Gular region of Acratosaura spinosa sp. nov. (holotype, MZUSP 98088), from Serra do Sincorá, Mucugê municipality, Bahia State. Photograph by Flávio Uemori Yamamoto.

22 · Zootaxa 2013 © 2009 Magnolia Press RODRIGUES ET AL. FIGURE 5. Details of the dorsal scales at midbody of (A) Acratosaura spinosa sp. nov. (holotype, MZUSP 98088), from Serra do Sincorá, Mucugê municipality, Bahia State; and (B) Acratosaura mentalis (MZUSP 98183) from Santo Inácio, Gentio do Ouro municipality, Bahia state, Brazil. Photographs by Pedro M. S. Nunes.

NEW ACRATOSAURA FROM BRAZIL Zootaxa 2013 © 2009 Magnolia Press · 23 Mental broad, wider than high. Postmental heptagonal, wider than long. Three pairs of genials, all in contact with infralabials and at midline; first smallest, third largest. An enlarged pair of symmetric flat and chevron-like scutes follows third pair of genials near the midline. Six infralabials, third the largest. Gulars smooth, imbricate, rounded posteriorly, in eight transverse rows; those from first two rows smaller with the same approximate size, posterior rows with enlarged scales, wider than long, the two central ones wider. Gulars followed by a distinct interbrachial region with seven much larger and elongate scales, about twice as long as gulars; central intrabrachial triangular (Fig. 4). Collar fold absent. Dorsal scales imbricate, strongly keeled and arranged in regular transverse rows; anterior ones wider, becoming progressively lanceolate, mucronate, more strongly keeled, tricarinate. Twenty eight transverse rows between interparietal and the posterior level of hind limbs. Lateral scales identical to dorsals except those closer to ventrals that are larger and scales of paraventral row that are smooth. Scales near the arm insertion smaller, almost granular. A distinctive area with granular scales surrounds the area of arm insertion. Twenty eight scales around midbody. Ventral scales smooth, imbricate, rounded posteriorly; 17 transverse rows from interbrachials (excluded) to preanals. Four longitudinal rows of ventrals, external ones wider than long, central ones longer than wide. Six scales in precloacal region, central and paramedials the largest. Total pores 13, continuous, with no gap between femoral and preanal ones. Scales of tail identical to the dorsals at midbody, keeled, lanceolate, strongly imbricate, except those from ventral part of base of tail which are slightly larger and smooth, but become gradually identical to other scales around tail towards the extremity. Fore limbs with large, smooth and imbricate scales; those on ventral part of brachium much smaller. Anterior and ventral parts of hind limbs with irregularly large, smooth and imbricate scales, identical to the correspondent parts of fore limbs. Posterior part of thighs with granular, juxtaposed scales; larger, imbricate and keeled scales on dorsal part of tibia. Carpal and tarsal scales large, imbricate with some striated scales on tarsus; supradigital lamellae smooth, imbricate. Palmar and plantar surfaces with smooth, small granules; infradigital lamellae single, 14 on Finger IV and 19 on Toe IV. Toes and fingers, except for Finger I, clawed, and respectively having the following relative sizes: 1 < 2 = 5 < 3 < 4 and, 1 < 2 < 5 < 3 < 4. Dorsal surfaces of body, flanks and tail and lateral part of tail dark brown to black with a series of longitudinal and transversely arranged yellow round spots about one scale size. Irregularly distributed darker punctuation smaller than one fourth of the scale size. Head light brown dorsally with scattered dark brown marks irregular in size and shape. Labials cream yellowish with wide irregular vertical dark bars on central part of scales. Ventral parts of head, body and tail creamy yellow with a series of irregular dark blotches. Regenerated part of tail uniform dark brown. Limbs dark brown irregularly mottled with yellow. Measurements of the holotype: snout-vent length: 53 mm; tail length (regenerated): 75 mm. Variation: the paratype and only other specimen known is a female 53 mm SVL, tail length 65 mm (regenerated) with 31 transverse dorsal scale rows, 18 transverse ventral rows, 30 scales around midbody, 14 and 20 subdigital lamellae respectively under the fourth finger and fourth toe, 8 transverse rows of gulars, 7 supralabials, 7 infralabials, 3 supraoculars, and 3 superciliaries. Except for the dorsal scales that are less intensively keeled, the absence of pores, and the absence of dark blotches in the venter, the female is very similar to the holotype. Comparisons: Among the Heterodactylini (sensu Rodrigues et al., 2007) the presence of three pairs of genials is a synapomorphy of Acratosaura; in all other genera there are two pairs. Acratosaura spinosa and Acratosaura mentalis are very similar in number of transverse dorsal scale rows (28–31 vs 30–32, respectively), ventral scale rows (17–18 vs 17–19), scales around midbody (28–30 vs 27–30), fourth finger and fourth toe subdigital lamellae (respectively 14 vs 12–14 and 19–20 vs 16–23), number of gular scale rows (8 vs 7–9), and number of femoral pores (13 vs 13–18). They are identical with respect to the number of supralabials (7), infralabials (7), supraoculars (3), superciliaries (3), pairs of chin-shields (3), and have similar maximum body size (53 mm vs 59 mm). Nevertheless there are several important differences in scalation and color pattern. In Acratosaura spinosa anterior dorsal scales are strongly keeled and mucronate becoming more strongly keeled, strongly mucronate and lanceolate at midbody; in Acratosaura mentalis they are smooth and

24 · Zootaxa 2013 © 2009 Magnolia Press RODRIGUES ET AL. rounded anteriorly becoming progressively hexagonal and slightly keeled at midbody (Fig. 5). Temporal scales are striate in Acratosaura spinosa and smooth in Acratosaura mentalis. On the side of neck scales are keeled and slightly imbricate in Acratosaura spinosa but smooth and juxtaposed in A. mentalis. A small, narrow and elongate scute follows the sixth supralabial in Acratosaura spinosa; in A. mentalis it is followed by a large scute, identical to the precedent supralabial. There are two distinctive transversely enlarged rows of gulars in Acratosaura spinosa, in A. mentalis these scales are much smaller and never so conspicuously enlarged. In Acratosaura spinosa there is a series of enlarged and strongly keeled scales in the dorsal part of tight separating the posterior granular area and the anterior part recovered by enlarged smooth scales; in A. mentalis scales from this area are very small and slightly keeled. Tibia scales are also more pronouncedly keeled in Acratosaura spinosa than in its congener. There are also striking differences in color pattern between these two species (Fig. 2). The dorsal color pattern of Acratosaura spinosa consists basically of regular series of transversely and longitudinally arranged yellow spots, one scale wide on a dark brown background. In Acratosaura mentalis instead of this pattern and on an olive gray background there is a characteristic, wide middorsal yellow stripe, 2–3 dorsals wide, which extends from the frontal scale to the level of the arm insertion.

FIGURE 6. Campos rupestres (rocky meadows), the habitat of Acratosaura spinosa sp. nov. at the type locality, near Mucugê municipality, state of Bahia, Brazil. Photograph by Marco Antonio de Freitas.

Distribution and natural history. Serra do Sincorá comprises a series of elevated plateaus and mountain ridges reaching up to about 1,600 m above sea level. It is a segment of Chapada Diamantina, the local name attributed to the northern portion of the extensive mountain ridge referred to as Serra do Espinhaço. The entire region is located in the municipality of Mucugê, state of Bahia, Brazil. Part of this area is now protected by the

NEW ACRATOSAURA FROM BRAZIL Zootaxa 2013 © 2009 Magnolia Press · 25 FIGURE 7. Type-locality of Acratosaura spinosa sp. nov. (red star), Mucugê municipality, State of Bahia, Brazil, and distribution of Acratosaura mentalis (yellow circles) in eastern Brazil. Localities: 1. Fechados, Santana de Pirapama, MG; 2. Guinda, Diamantina, MG; 3. Peixe Cru, Turmalina, MG; 4. Cristália, MG; 5. Grão Mogol, MG; 6. Parque Nacional do Peruaçu, Januária, MG; 7. Maracás, BA; 8. Mucugê, BA; 9. Santo Inácio, Gentio do Ouro, BA; 10. Senhor do Bonfim, BA; 11. UHE de Xingo, Delmiro Gouveia, AL; 12. Serra Negra, Mata Grande, AL; 13. Serra do Triunfo, Garanhuns, PE; 14. Parque Nacional do Catimbau, Buíque, PE; 15. Brejo da Madre de Deus, PE; 16. Fazenda Bravo, Barra de São Miguel, PB; 17. Cabaceiras, PB; 18. RPPN Fazenda Almas, São José dos Cordeiros and Sumé municipalities, PB; and 19. Cacimba de Dentro, PB. For localities see Appendix, and Arzabe et al. (2005), Delfin (2007), Queiroz & Lema (1996), and Rodrigues (1986). State Abbreviations: AL = Alagoas; BA = Bahia; MG = Minas Gerais; PE = Pernambuco; PB = Paraíba.

26 · Zootaxa 2013 © 2009 Magnolia Press RODRIGUES ET AL. Parque Nacional da Chapada Diamantina. The entire area is dominated by “campos rupestres” (rocky meadows, see Giulietti & Pirani 1988; Rodrigues 1988) the typical open vegetation from altitudinal areas in the Espinhaço range where quartzitic sandy soils and rocky outcrops are characteristic (Fig. 6). Vegetation in the region also includes several types of Cerrado like vegetation, semidecidual seasonal forest (carrasco), and gallery forest along the river valleys. Specimens were obtained at Mucugê, near the village, and at Fazenda Caraíbas, a farm situated on a relatively flat upland plateau at around 1,100 m a.s.l. on the west side of the escarpments of Serra do Sincorá. The holotype, was found under a rock in typical campos rupestres; the paratype was exposed on the side of a road in a disturbed area in campos rupestres. Distance between the two localities in a straight line is about 12 km.

Discussion

As far as we know Acratosaura spinosa is restricted to and supposed to be endemic from Serra do Sincorá. In the last years several new taxa have been described from the highlands of the northern portion of Espinhaço range (Cassimiro et al. 2008; Heyer 2002; Lugli & Haddad 2006a; Lugli & Haddad 2006b; Napoli & Juncá 2006; Pinna 1992; Rodrigues 1987; Rodrigues et al. 2006). These new discoveries reinforced previous ideas suggesting a high level of endemism at Chapada Diamantina, and especially in the Mucugê region (Rodrigues 1987). Endemic species from this area are generally associated with altitude and restricted to a small extension of the Espinhaço ridge. This seems to be the case of Acratosaura spinosa, Tropidurus mucujensis (Rodrigues 1987), Enyalius erythroceneus (Rodrigues et al. 2006), undescribed species of Eurolophosaurus and Mabuya (Rodrigues et al. 2006), Rupirana cardosoi (Heyer 2002), Bokermanohyla spp. (Lugli & Haddad 2006a; Lugli & Haddad 2006b; Napoli & Juncá 2006) and Strabomantis aramunha (Cassimiro et al. 2008), and even of the fish subfamily Copionodontinae (Pinna 1992). It is a relieve to know that most of these species are now under the protected shelter of Parque Nacional da Chapada Diamantina, the largest conservation area in the region with about 150,000 ha. Acratosaura mentalis, the only other congener of A. spinosa, is sympatric with it at Mucugê and was also found in the campos rupestres but unlike the new species here described it is more widely distributed (Fig. 7). It occurs at low and moderate elevations (500–1,000 m) in mesic habitats and relictual forests in the Caatingas, at moderate elevations (700 m) in semidecidual forests in the Cerrado Domain, and up to 1,400 m in the highlands of Serra do Espinhaço. The discovery of Acratosaura spinosa adds a new species to a genus up to now considered monotypic, and reinforces the idea of its generic distinction from Colobosaura where until recently only Colobosaura mentalis was allocated (Rodrigues et al. 2007). In fact, the most recent evidence based on morphological and molecular data indicates that the sister taxon of Acratosaura is Stenolepis ridleyi, a species restricted to continuous and relict patches of Atlantic Forest of Northeastern Brazil.

Acknowledgments

We thank Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP), Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), and IBAMA for collection permits (185/2005), Agroinvest Kayatani S/A, Felipe S. F. Leite, L. E. Lopes, Mitsuru Origushi, Eduardo Varejão, Mauro Teixeira Jr., and Renato S. Recoder for their valuable help in the field, Luciana Lobo and Mauro Teixeira Jr. for the drawings, Pedro M. S. Nunes for stereomicroscopic photographs, Flávio ‘Japonês’ Uemori Yamamoto for figures 3 and 4, Marinus S. Hoogmoed for their critical review of the manuscript, and Hussam Zaher and Carolina Castro-Mello for access to specimens.

Arzabe, C., Skuk, G., Santana, G.G., Delfin, F.R., Lima, Y.C.C. & Abrantes, S.H.F. (2005) Herpetofauna da área do Curimataú, Paraíba. In: Araújo, F.S., Rodal, M.J.N. & Barbosa, M.R.V., Análise das variações da biodiversidade do Bioma Caatinga: suporte a estratégias regionais de conservação. Ministério do Meio Ambiente: Brasília, D.F., 264–280. Cassimiro, J., Verdade, V.K. & Rodrigues, M.T. (2008) A large and enigmatic new eleutherodactyline frog (Anura, Strabomantidae) from Serra do Sincorá, Espinhaço range, Northeastern Brazil. Zootaxa, 1761, 59–68. Castoe, T.A., Doan, T.M. & Parkinson, C.L. (2004) Data partitions and complex models in Bayesian analysis: the phylogeny of gymnophthalmid lizards. Systematic Biology, 53, 1–22. Delfin, F.R. & Freire, E.M.X. (2007) Os lagartos gimnoftalmídeos (Squamata: Gymnophthalmidae) do Cariri Paraibano e do Seridó do Rio Grande do Norte, Nordeste do Brasil: Considerações acerca da distribuição geográfica e ecologia. Oecologia Brasileira, 11, 365–382. Giulietti, A.M. & Pirani, J.R. (1988) Patterns of geographic distribution of some plant species from the Espinhaço range, Minas Gerais and Bahia, Brazil. In: Heyer, W.R. & Vanzolini, P.E., Proceedings of a Workshop on Neotropical Distribution Patterns. Academia Brasileira de Ciências: Rio de Janeiro, 39–69. Heyer, W.R. (2002) A new genus and species of frog from Bahia, Brazil (Amphibia: Anura: Leptodactylidae) with comments on the zoogeography of the brazilian campos rupestres. Proceedings of the Biological Society of Washington, 112, 19–39. Lugli, L. & Haddad, C.F.B. (2006a) A new species of Bokermannohyla pseudopseudis group from Central Bahia, Brazil (Amphibia, Hylidae). Herpetologica, 62, 453–462. Lugli, L. & Haddad, C.F.B. (2006b) A new species of Bokermannohyla (Anura, Hylidae) from central Bahia, Brazil. Journal of Herpetology, 40, 7–15. Napoli, M.F. & Juncá, F.A. (2006) A new species of the Bokermannohyla circumdata group (Amphibia: Anura: Hylidae) from Chapada Diamantina, State of Bahia, Brazil. Zootaxa, 1244, 57–68. Pellegrino, K.C.M., Rodrigues, M.T., Yonenaga-Yassuda, Y. & Sites Jr, J.W. (2001) A molecular perspective on the evolution of microteiid lizards (Squamata, Gymnophthalmidae), and a new classification for the family. Biological Journal of the Linnean Society, 74, 315–338. Pinna, M.C.C. (1992) A new subfamily of Trichomycteridae (Teleostei, Siluriformes), lower loricaroid relationships and a discussion on the impact of additional taxa for phylogenetic analysis. Zoological Journal of the Linnean Society, 106, 175–229. Queiroz, A.N. & Lema, T. (1996) Novo registro de Colobosaura mentalis Amaral, 1933, (Sauria; Teiidae) para o nordeste do Brasil. Biociências, Porto Alegre, 4, 87–90. Rodrigues, M.T. (1986) Uma nova espécie do gênero Phyllopezus de Cabaceiras: Paraíba: Brasil; com comentários sobre a fauna de lagartos da área (Sauria Gekkonidae). Papéis Avulsos de Zoologia, São Paulo, 36, 237–250. Rodrigues, M.T. (1987) Sistemática, ecologia e zoogeografia dos Tropidurus do grupo torquatus ao sul do rio Amazonas (Sauria, Iguanidae). Arquivos de Zoologia, São Paulo, 31, 105–230. Rodrigues, M.T. (1988) Distribution of lizards of the genus Tropidurus in Brazil (Sauria, Iguanidae). Academia Brasileira de Ciências: Rio de Janeiro, 305–315. Rodrigues, M.T., Cassimiro, J., Pellegrino, K.C.M., Verdade, V.K., Pavan, D. & Curcio, F.F. (In press) A new genus of microteiid lizard from the Caparaó mountains, Southeastern Brazil, with a discussion of relationships among Gymnophthalminae (Squamata). American Museum Novitates. Rodrigues, M.T., Freitas, M.A., Silva, T.F.S. & Bertolotto, C.E.V. (2006) A new species of lizard genus Enyalius (Squamata, Leiosauridae) from the highlands of Chapada Diamantina, state of Bahia, Brazil, with a key to species. Phyllomedusa, 5, 11–24. Rodrigues, M.T., Pellegrino, K.C.M., Dixo, M., Verdade, V.K., Pavan, D., Argôlo, A.J.S. & Sites Jr, J.W. (2007) A new genus of microteiid lizard from the Atlantic Forests of state of Bahia, Brazil, with a new generic name for Colobosaura mentalis, and a discussion of relationships among the Heterodactylini (Squamata, Gymnophthalmidae). American Museum Novitates, 3565, 1–27.

Acratosaura mentalis: BRAZIL: Alagoas/Sergipe: UHE Xingó: MZUSP 80152–3, MNRJ 9964. Bahia: Morro do Chapéu: MZUSP 74224; Mucugê: MZUSP 74223, 96894; Santo Inácio: MZUSP 79743, 98183–5; Senhor do Bonfim: MZUSP 40084–5; Central: Toca dos Pilões: MZUSP 89290. Minas Gerais: Diamantina: Guinda MZUSP 98187; Grão Mogol: MZUSP 56933, 56983, 98186; Santana do Pirapama: Fechados: JC 1062; Turmalina: MNRJ 9965. Paraíba: Cabaceiras: Fazenda Bravo: MZUSP 60796, 66231. Pernambuco: Buíque: Serra do Catimbau: VC 44; Garanhuns: MZUSP 65675; João Alfredo: MZUSP 7076. VC = field number of Universidade Rural de Pernambuco, Serra Talhada; JC = field number of José Cassimiro.