Embryonic development and the evolution of fossoriality in lizards from the tribe Gymnophthalmini (Squamata, Gymnophthalmidae)

A história evolutiva da tribo Gymnophthalmini (família Gymnophthalmidae) é caracterizada por modificações morfológicas relacionadas à evolução de planos corpóreos serpentiformes adaptados à vida fossorial. As adaptações apresentadas por estas espécies (especialmente o alongamento do corpo e a redução dos membros) são frequentemente observadas em diversos outros grupos de Squamata. A formação do padrão corpóreo é coordenada por uma rede de regulação complexa que atua durante o desenvolvimento embrionário e, muitas vezes, mudanças sutis nesta rede podem resultar em alterações fenotípicas drásticas, levando à mudanças evolutivas nas formas corpóreas. Neste contexto, o grupo Squamata é um ótimo modelo de pesquisa para estudos sobre a evolução das formas, já que diversas espécies apresentam morfologias fossoriais semelhantes (e independentes, do ponto de vista filogenético) que podem ter se originado a partir de mecanismos de desenvolvimento comuns, mostrando uma certa direcionalidade das pressões seletivas que agem sob os possíveis caminhos de desenvolvimento. Este trabalho analisa o desenvolvimento dos embriões de cinco espécies da tribo Gymno-phthalmini (Procellosaurinus tetradactylus, Vanzosaura rubricauda, Psilophthalmus paeminosus, Nothobachia ablephara e Calyptommatus sinebrachiatus), estabelecendo critérios para a determinação de estágios embrionário a partir da morfologia externa e analisando o desenvolvimento dos elementos de sustentação do crânio e dos esqueletos axial e apendicular de forma comparada, relacionando a morfologia ao hábito de vida. O hábito fossorial impõe pressões significativas ao corpo do animal, que apresenta modificações osteológicas para suportar tais pressões, como um crânio robusto, com elementos organizados de forma a proteger o encéfalo dos impactos da locomoção subterrânea, corpo alongado adaptado à locomoção por movimentos ondulatórios e membros reduzidos que não participam ativamente da movimentação. Também são apresentados dados preliminares sobre os possíveis processos responsáveis pela redução dos membros e pelo alongamento do corpo nas espécies fossoriais, relacionando as modificações do desenvolvimento à evolução do plano corpóreo serpentiforme.The evolutionary history of the Gymnophthalmini (Gymnophthalmidae) is characterized by morphological modifications related to the evolution of a snake-like body plan adapted to fossorial habits. Fossorial snake-like species show adaptations, especially body lengthening and limb reduction, that are frequently observed throughout Squamate lineages. Pattern formation is coordinated by a complex regulation network that acts during embryonic development, and subtle changes to this network may result in drastic phenotypic modifica-tions, leading to evolutionary variation in body plans. In this context, Squamates represent an excellent research model since several distant related species show similar (and phylogenetically independent) adaptations to fossoriality, which may have originated trhough common developmental mechanisms, and can reflect, to some extent, a predominance of certain selective pressures acting upon developmental pathways. This work analyses the morphological aspect of embryonic development of five Gymnophthalmini species (Procellosaurinus tetradactylus, Vanzosaura rubricauda, Psilophthalmus paeminosus, Nothobachia ablephara e Calyptommatus sinebrachiatus) establishing criteria for the embryonic staging through external morphology, and analysing the development of cartilage and bone in the skull, and axial and appendicular systems in a comparative background, associating morphology to life habits. Fossorial life exerts great pressure upon the body, which demands an adaptative response in order to overcome this sort of impact. Thus, fossorial animals shows a robust skull, with bones tightly articulated as to protect the brain and sense organs, an elongated body for ondulatory locomotion and reduced limbs that do not actively participate in locomotion. Preliminary data on the possible developmental processes responsible for limb reduction and body elongation in the fossorial lineage are shown, and are discussed on the light of evolutionary developmental biology

Skeletal development in the fossorial gymnophthalmids Calyptommatus sinebrachiatus and Nothobachia ablephara

The development of the cartilaginous and bony elements that form the skull and axial and appendicular skeleton is described in detail for the post-ovipositional embryonic development of the fossorial gymnophthalmid species Calyptommatus sinebrachiatus and Nothobachia ablephara. Both species have a snake-like morphology, showing an elongated body and reduced or absent limbs, as well as modifications in skull bones for burrowing, such as complex articulation surfaces and development of bony extensions that enclose and protect the brain. Similar morphological changes have originated independently in several squamate groups, including the one that led to the snake radiation. This study characterizes the patterns of chondrogenesis and osteogenesis, with special emphasis on the features associated with the burrowing habit, and may be used for future comparative analyses of the developmental patterns involved in the origin of the convergent serpentiform morphologies. (C) 2012 Elsevier GmbH. All rights reserved.Fundacao de Amparo a Pesquisa do Estado de Sao Paulo (FAPESP)Fundacao de Amparo a Pesquisa do Estado de Sao Paulo (FAPESP)Conselho Nacional de Desenvolvimento Cientifico e Tecnologico (CNPq)Conselho Nacional de Desenvolvimento Cientifico e Tecnologico (CNPq

Embryonic development of the fossorial gymnophthalmid lizards Nothobachia ablephara and Calyptommatus sinebrachiatus

The evolutionary history of the lizard family Gymnophthalmidae is characterized by several independent events of morphological modifications to a snake-like body plan, such as limb reduction, body elongation, loss of external ear openings, and modifications in skull bones, as adaptive responses to a burrowing and fossorial lifestyle. The origins of such morphological modifications from an ancestral lizard-like condition can be traced back to evolutionary changes in the developmental processes that coordinate the building of the organism. Thus, the characterization of the embryonic development of gymnophthalmid lizards is an essential step because it lays the foundation for future studies aiming to understand the exact nature of these changes and the developmental mechanisms that could have been responsible for the evolution of a serpentiform (snake-like) from a lacertiform (lizard-like) body form. Here we describe the post-ovipositional embryonic development of the fossorial species Nothobachia ablephara and Calyptommatus sinebrachiatus, presenting a detailed staging system for each one, with special focus on the development of the reduced limbs, and comparing their development to that of other lizard species. The data provided by the staging series are essential for future experimental studies addressing the genetic basis of the evolutionary and developmental variation of the Gymnophthalmidae. (C) 2012 Elsevier GmbH. All rights reserved.Fundacao de Ampparo a Pesquisa do Estado de Sao Paulo (FAPESP)Fundacao de Ampparo a Pesquisa do Estado de Sao Paulo (FAPESP)Conselho Nacional de Desenvolvimento Cientifico e Tecnologico (CNPq)Conselho Nacional de Desenvolvimento Cientifico e Tecnologico (CNPq

Are Hemipenial Spines Related to Limb Reduction? A Spiny Discussion Focused on Gymnophthalmid Lizards (Squamata: Gymnophthalmidae)

Calcified spines in the hemipenial surface occur convergently in several gymnophthalmid lizard species and in advanced snakes. Based on the pronounced degrees of limb reduction in these distantly related lineages, such hemipenial structures were suggested to play a functional role in couple-anchoring during copulation, partly assuming the function of the limbs during mating. Herein, we assessed the hemipenial morphology of virtually all the valid genera of the family Gymnophthalmidae to test for a phylogenetic correlation between limb reduction and the presence of calcified hemipenial spines. The occurrence of calcified structures was mapped on the two most comprehensive phylogenies of the family. We concluded that spiny hemipenes are by no means necessarily associated with reduction of limbs. Conversely, the presence of well-developed hemipenial spines in specific limb-reduced taxa does not allow one to disregard the possibility that in some instances such structures might indeed be functionally associated with couple-anchoring, improving the success of matin

Through the Looking Glass: The Spectacle in Gymnophthalmid Lizards

The anatomy and development of the eyelids in squamate reptiles are still relatively unknown, considering its variation within the group. The neotropical Gymnophthalmini are traditionally characterized by having lost the eyelids, but their structure is not well described. In this study, the embryonic development and the adult morphology of the gymnophthalmid eye, with special attention to the eyelids, the nictitating membrane, and the spectacle are described. The eye in some Gymnophthalmini is covered by a spectacle, formed by the embryonic fusion of the dorsal and ventral eyelids, a character possibly synapomorphic to the tribe. The genus Tretioscincus, which floats either as sister to all other Gymnophthalmini, or is nested within the group, is unique in showing functional and movable eyelids. Thus, the presence of functional eyelids can be either considered as the primitive condition for the gymnophthalmini or as a re-acquisition of the character, showing the importance of a well-established phylogenetic hypothesis for understanding morphological evolution

Rediscovery of the Earless Microteiid Lizard Anotosaura collaris Amaral, 1933 (Squamata: Gymnophthalmidae): A redescription complemented by osteological, hemipenial, molecular, karyological, physiological and ecological data

Rodrigues, Miguel Trefaut, Jr, Mauro Teixeira, Vechio, Francisco Dal, Amaro, Renata Cecília, Nisa, Carolina, Guerrero, Agustín Camacho, Damasceno, Roberta, Roscito, Juliana Gusson, Sales Nunes, Pedro M., Recoder, Renato Sousa (2013): Rediscovery of the Earless Microteiid Lizard Anotosaura collaris Amaral, 1933 (Squamata: Gymnophthalmidae): A redescription complemented by osteological, hemipenial, molecular, karyological, physiological and ecological data. Zootaxa 3731 (3): 345-370, DOI: http://dx.doi.org/10.11646/zootaxa.3731.3.

Psilops mucugensis Rodrigues, Recoder, Jr, Roscito, Guerrero, Nunes, Freitas, Fernandes, Bocchiglieri, Vechio, Leite & Nogueira, 2017, sp. nov.

<i>Psilops mucugensis</i> sp. nov. <p>(Figs. 4–5)</p> <p> <i>Psilophthalmus</i> sp.—Cassimiro and Rodrigues (2009: 49); Freitas <i>et al.</i> (2012: 18, 20, 22); Magalhães <i>et al.</i> (2015: 247, 249, 258, 261)</p> <p> <b>Holotype.</b> an adult male, MZUSP 106188, collected by M.A. Freitas and T.F. Santos Silva on October 5th 2005 (field number MTR 11787) from Fazenda Três Irmãos (12°53'06"S 41°31'41"W, 1075 m above sea level, hereafter asl), district of Guiné, municipality of Mucugê, Serra do Espinhaço (Chapada Diamantina), Bahia, Brazil.</p> <p> <b>Paratypes.</b> MZUSP 96918, 106187 (field number MTR 11520–21), collected on 8th October 2005 in the same locality and collectors as for the holotype.</p> <p> <b>Referred specimens.</b> AAG 6640, 6652, 6663, 6692, 6715–6716, 7026, 7040–7041, 7065, 7080, 7109, 7120, 7142, 7144, 7190, 7199, 7220, 7228, 7249 from Palmeiras (12°32'S, 41°29'W), Bahia, Brazil; collected by Adrian Garda <i>et al</i>. on 6th January to 6th February 2013. MZUSP 106196, 106208, 106209 (all from 11°22'24.17"S, 40°31'4.22"W, 1141 m asl), MZUSP 106197 (11°22'23.74"S, 40°31'4.12"W, 1141 m asl), MZUSP 106210 (11°21'32"S, 40°31'37"W), all from Parque Estadual das Sete Passagens, municipality of Miguel Calmon: Bahia: Brazil; collected by the authors between 29th May 2010 to 10th March 2011, field numbers MTR 19871, RPD 0 40, RPD 110, MTR 20012, RPD 260, respectively. MZUSP 106206, from Morro do Chapéu hill (11°35'34.45"S, 41°12'27.65"W, 1271 m asl), municipality of Morro do Chapéu, Bahia, Brazil; collected by the authors on 30th December 2011, field number MTR 22533.</p> <p> <b>Diagnosis.</b> <i>Psilops mucugensis</i> differs from <i>P. paeminosus</i> (data in parenthesis) by having a higher number of smooth subcaudal scales, 7–14 (3–7), a higher number of total pores, 16–20 (14–16), two conspicuous dorsolateral white stripes running from supraciliaries to the tail (absent), a bright red tail (brownish in adults) and longer forelimbs, 25.9% SVL (21.5% SVL). <i>Psilops mucugensis</i> differs from <i>P. seductus</i> <b>sp. nov.</b> (described below, data in parenthesis) by having lower number of scale rows around midbody, 17–21 (22), a higher number of subdigital lamellae under Finger IV, 16–19 (13–16) and Toe IV, 11–14 (9–11), a higher number of smooth subcaudal scales, 7–14 (2–5), a higher number of total pores, 16–20 (10–13), a larger foot, 17.6% SVL (14.8% SVL), and by the presence of calcified spines (absent) and by lacking papillae ornamenting the hemipenial flounces (present).</p> <p> <b>Description of the holotype.</b> Rostral broad, visible from above wider than high, contacting first supralabial, nasal and frontonasal. Frontonasal hexagonal, slightly wider than long, in broad contact with rostral, nasal, prefrontals, and frontal. Prefrontals hexagonal, as long as wide, contacting frontonasal, frontal, first supraocular, first supraciliary, loreal, and nasal; separate at midline by contact between frontonasal and frontal. Frontal hexagonal with posteriorly convergent lateral margins, approximately twice as long as broad, wider anteriorly; in broad contact with first supraocular, its suture with frontonasal as broad as that of interparietal. Frontoparietals absent. Interparietal hexagonal, posteriorly rounded, with strongly posteriorly convergent lateral margins, longer than wide, longer and wider than frontal and contacting frontal, first supraocular, parietals, and occipitals. Parietals irregularly hexagonal, as long as wide, and smaller and shorter than interparietal, contacting first and second supraoculars, temporals, occipitals, and interparietal. Two supraoculars, first the largest, longer than wide, occupying most of supraocular area, second very small, sub-squared, with the same approximate size of the adjacent temporal. Three supraciliaries, first slightly larger than second, third the smallest; first supraciliary expanding on the lateral face of head and in contact with loreal and preocular; second supraciliary in the center of eye. Nasal large, subrectangular, longer than wide, with the nostril in the center, contacting first and second supralabials. Loreal narrow, as high as nasal, diagonally oriented and contacting second and third supralabials, frenocular, first supraciliary, prefrontal and nasal. Frenocular as large as loreal with its postero-ventral corner divided in a small scute and followed posteriorly by a series of small and narrow irregular preoculars and suboculars. Seven supralabials, fourth the largest, under the eye, fifth the highest, separated from parietal by two postocular scutes; seventh supralabial contacting ear border. Eyelid absent. Eye large, pupil rounded. Anterior and lower margin of eye with an irregularly narrow, smooth, and elongate semidivided scute, thinner at lower margin of eye; posterior margin of eye with a series of juxtaposed smooth granules. Temporal region covered by enlarged, smooth, and imbricate cycloid scales. Tympanum recessed. Ear opening bordered by cycloid, smooth, and imbricate scales, those on posterior margin smaller, those above tympanum larger, higher than wide and covering anterior and superior margin of ear.</p> <p>Mental wider than long. Postmental single, as wide as long, contacting first and second infralabials. Two pairs of enlarged genials in medial contact and contacting infralabials; first pair the largest, longer than wide, second one wider than long. Six infralabials, second and third the largest. All head scales smooth with irregularly disposed sensorial pits. Gulars disposed in nine irregularly transverse rows, smooth, imbricate, posteriorly rounded; central ones wider than long, diagonally disposed anteriorly, becoming gradually transverse posteriorly. Lateral gular scales smaller, cycloid, smooth, imbricate. Interbrachial row with five enlarged imbricate scales, central one the largest, subtriangular.</p> <p>Anterior dorsal scales disposed in three longitudinal rows until the level of arm, dorsolateral ones wider than long, scales in central row smaller, as wide as long. Scales of the first transverse dorsal row smooth or slightly striate becoming gradually tri or pentacarinate. After arm level dorsal scales become progressively more elongate, mucronate and tricarinate, having the central keel sharper. Thirty-four transverse rows between interparietal and posterior level of hindlimbs. Lateral scales cycloid, smooth, imbricate, except for an area with small, flat, smooth and juxtaposed scales around arm insertion. Eighteen scales around midbody. Ventrals wide, smooth, imbricate, in four longitudinal rows from interbrachials to precloacals and in 24 transverse rows between interbrachials and precloacals. Central rows distinctively enlarged, wider than long. Posterior margin of vent with four scales; central ones smaller. Total pores 20, two precloacals and eight femoral on each side.</p> <p>Scales of dorsal and lateral portion of tail imbricate, strongly keeled, elongate, mucronate, in complete rings; those covering dorsal portion of base of tail tricarinate, becoming unicarinate more elongate and lanceolate posteriorly. Scales of ventral portion of tail smooth, larger than correspondent dorsals at the base of tail but becoming gradually identical to them in shape and ornamentation toward the tip; 15 irregularly transverse rows of smooth subcaudals.</p> <p>Forelimbs with large, smooth and imbricate scales except for those on ventral parts of arm and forearm which are smaller. Anterior and posterior parts of tights with large, smooth and imbricate scales; posterior part of tights with granular, juxtaposed, smooth scales which grade progressively to a posterior irregular row of enlarged, mucronate, and keeled scales, identical to those covering dorsal part of tibia. Ventral parts of tibia with smooth, enlarged and imbricate scales. Palmar and plantar surfaces with small, conical, juxtaposed granules. Subdigital lamellae single, 12 on finger IV and 19 on toe IV. Fingers and toes clawed; first finger absent externally. Toes and fingers with the following relative sizes, respectively: 2=5<3<4 and 1<2<5<3<4.</p> <p>Dorsal surface of body and tail olive brown with irregular and scattered dark brown reticulum. A dorsolateral white stripe runs from supraciliaries to the tail, becoming inconspicuous toward its extremity. Below it a dark brown lateral stripe extends from posterior margin of eye to tail. Between supraocular region and midbody the white stripe is also dorsally emarginated by an irregularly dark brown stripe which makes its anterior third highly conspicuous. Dorsal and lateral surfaces of head identical to corresponding parts of dorsum and flanks. Ventral parts of head, body, and tail creamy white with dark brown to black spots especially concentrated on external rows of ventral scales and in gular region which is mottled with dark spots. Limbs olive brown dorsally, mottled with darker pigment; ventrally creamy white, almost immaculate. Tail dorsal color identical to that of body proximally, becoming uniformly light brown posteriorly. Ventral portion of tail with dark spots proximally, creamily immaculate posteriorly.</p> <p> <b>Measurements.</b> SVL= 35.3 mm; TAL = 42.3 mm (regenerated); TRL = 20.2 mm; HW = 4.3 mm; HL = 6.7 mm; FEM = 4.0 mm; TIB = 3.5 mm; FTL = 6.3 mm; HUM = 2.7 mm; FAL = 5.4 mm.</p> <p> <b>Variation.</b> Individuals from southern (municipalities of Mucugê and Palmeiras) and northern (Morro do Chapéu and Miguel Calmon) populations, agree in most scale counts. Nevertheless, northern populations show fewer femoral pores (15–16, 16– 20 in south) and number of smooth subcaudals (6–8, 10– 14 in south) (Table 2). Males and females are sexually dimorphic in body size, with females having larger SVL than males (ANOVA; <i>F</i> 1,17 = 7,95; <i>P</i> <0,05). Differences in shape are also significant, with females showing proportionally larger TRL (ANCOVA, SVL as covariate; <i>F</i> 1,16 = 13.42; <i>P</i> <0,01), but shorter HL (<i>F</i> 1,16 = 13.67; <i>P</i> <0,01), FEM (<i>F</i> 1,16 = 12.37; <i>P</i> <0.01) and TIB (<i>F</i> 1,16 = 10.49; <i>P</i> <0.01).</p> <p> <b>Distribution and natural history.</b> Known from the municipalities of Mucugê, Palmeiras (Parque Nacional Chapada Diamantina), Morro do Chapéu and Miguel Calmon, state of Bahia, Brazil (Fig. 6). At Miguel Calmon it is found at high altitude open areas over quartzite sandy soils (1140 m asl), while at Morro do Chapéu, the single specimen was found within a semi-deciduous montane open forest (1270 m asl) (Fig. 7). At Mucugê it was found at a flat plateau (1100 m a.s.l.) covered by a type of low semi-deciduous arborescent vegetation. At Palmeiras it was found in areas of rocky grasslands <i>campos rupestres</i> (Magalhães <i>et al.</i> 2015).</p> <p> <b>Etymology.</b> Named after the type locality, Mucugê, at Chapada Diamantina, Bahia.</p>Published as part of <i>Rodrigues, Miguel Trefaut, Recoder, Renato, Jr, Mauro Teixeira, Roscito, Juliana Gusson, Guerrero, Agustín Camacho, Sales Nunes, Pedro M., Freitas, Marco Antonio De, Fernandes, Daniel Silva, Bocchiglieri, Adriana, Vechio, Francisco Dal, Leite, Felipe Sá Fortes & Nogueira, Cristiano De Campos, 2017, A morphological and molecular study of Psilops, a replacement name for the Brazilian microteiid lizard genus Psilophthalmus Rodrigues 1991 (Squamata, Gymnophthalmidae), with the description of two new species, pp. 451-482 in Zootaxa 4286 (4)</i> on pages 462-466, DOI: 10.11646/zootaxa.4286.4.1, <a href="http://zenodo.org/record/828661">http://zenodo.org/record/828661</a&gt

FIGURE 11 in A morphological and molecular study of Psilops, a replacement name for the Brazilian microteiid lizard genus Psilophthalmus Rodrigues 1991 (Squamata, Gymnophthalmidae), with the description of two new species

FIGURE 11. Hemipenes of Psilops paeminosus (MZUSP 106186, from Gameleira do Assuruá, Bahia) (Clade B—type lineage), in its sulcate, lateral and asulcate faces (A), P. mucugensis (MZUSP 106196, from Miguel Calmon, Bahia) (Clade E), in its sulcate and asulcate faces (B) and, P. seductus (MNRJ 19099, from Jaborandi, Bahia) (Clade A) (C). Scale bars = 1 mm.Published as part of Rodrigues, Miguel Trefaut, Recoder, Renato, Jr, Mauro Teixeira, Roscito, Juliana Gusson, Guerrero, Agustín Camacho, Sales Nunes, Pedro M., Freitas, Marco Antonio De, Fernandes, Daniel Silva, Bocchiglieri, Adriana, Vechio, Francisco Dal, Leite, Felipe Sá Fortes & Nogueira, Cristiano De Campos, 2017, A morphological and molecular study of Psilops, a replacement name for the Brazilian microteiid lizard genus Psilophthalmus Rodrigues 1991 (Squamata, Gymnophthalmidae), with the description of two new species, pp. 451-482 in Zootaxa 4286 (4) on page 471, DOI: 10.11646/zootaxa.4286.4.1, http://zenodo.org/record/82866

A morphological and molecular study of Psilops, a replacement name for the Brazilian microteiid lizard genus Psilophthalmus Rodrigues 1991 (Squamata, Gymnophthalmidae), with the description of two new species

The lizard genus Psilophthalmus was originally described from the sandy deposits at the northern end of Serra do Espinhaco, in Santo Inacio, state of Bahia, but since then it has been recorded in other Brazilian localities of the states of Bahia, Minas Gerais, and Sergipe. Here, we review the collected specimens based on molecular markers (mitochondrial 12S, 16S, ND4 and cyt b, and nuclear C-mos and NT3) and morphological evidence (external, hemipenial and osteological morphologies). In the course of our revision we find out that Psilophthalmus Rodrigues 1991 was preoccupied by Psilophthalmus Szepligeti 1902 (Hymenoptera, Braconidae). The replacement name Psilops is proposed for the genus for which we recognize three species, with Psilops paeminosus as type species. One of the new species is found along the high elevation areas of the Chapada Diamantina plateaus, state of Bahia, while the other occurs in the cerrados of "Serra Geral", in the occidental plateaus of that state. Psilops paeminosus comprises three distinct allopatric clades that, based on current evidence, cannot be diagnosed morphologically: one from the vicinities of the type locality, one from the lower Sao Francisco River, and a third from the uplands of Minas Gerais and southern inland Bahia. We keep the latter two as candidate species but defer their formal description until further evidence allows robust diagnosis. Derived clades of Psilops with shorter limbs have invaded hotter and drier environments, while mostly used sandy soils along their evolution.Fundacao de Amparo a Pesquisa do Estado de Sao Paulo (FAPESP)Fundacao de Amparo a Pesquisa do Estado do Rio de Janeiro (FAPERJ)Conselho Nacional de Desenvolvimento Cientifico e Tecnologico (CNPq)Instituto Chico Mendes de Biodiversidade (ICMBIO)Instituto Brasileiro do Meio Ambiente e dos Recursos Naturais Renovaveis (IBAMA)Secretaria do Meio Ambiente do Estado da BahiaUniv Sao Paulo, Inst Biociencias, Dept Zool, BR-05508090 Sao Paulo, SP, BrazilUniv Fed Pernambuco, Ctr Biociencias, Dept Zool, BR-50670901 Recife, PE, BrazilInst Chico Mendes Conservacao Biodiversidade, PARNA Catimbau, BR-56537000 Buique, PE, BrazilUniv Fed Rio de Janeiro, Inst Biol, Dept Zool, BR-21941902 Rio De Janeiro, RJ, BrazilUniv Fed Sergipe, Ctr Ciencias Biol & Saude, Dept Ecol, BR-49100000 Sao Cristovao, SE, BrazilUniv Fed Vicosa, Campus Florestal, BR-35690000 Florestal, MG, BrazilUniv Sao Paulo, Inst Biociencias, Dept Ecol Geral, BR-05508090 Sao Paulo, SP, BrazilUniv Fed Sao Paulo, Dept Ciencias Biol, BR-09972270 Diadema, SP, BrazilUniv Estadual Santa Cruz, Programa Posgrad Zool, BR-45662900 Ilheus, BA, BrazilUniv Fed Sao Paulo, Dept Ciencias Biol, BR-09972270 Diadema, SP, BrazilFAPESPFAPERJCNPqICMBIO: 11596ICMBIO: 14555ICMBIO: 30309IBAMA: 2001.007793/00-31]Secretaria do Meio Ambiente do Estado da Bahia: 13/2010Web of Scienc

A morphological and molecular study of Psilops, a replacement name for the Brazilian microteiid lizard genus Psilophthalmus Rodrigues 1991 (Squamata, Gymnophthalmidae), with the description of two new species

The lizard genus Psilophthalmus was originally described from the sandy deposits at the northern end of Serra do Espinhaco, in Santo Inacio, state of Bahia, but since then it has been recorded in other Brazilian localities of the states of Bahia, Minas Gerais, and Sergipe. Here, we review the collected specimens based on molecular markers (mitochondrial 12S, 16S, ND4 and cyt b, and nuclear C-mos and NT3) and morphological evidence (external, hemipenial and osteological morphologies). In the course of our revision we find out that Psilophthalmus Rodrigues 1991 was preoccupied by Psilophthalmus Szepligeti 1902 (Hymenoptera, Braconidae). The replacement name Psilops is proposed for the genus for which we recognize three species, with Psilops paeminosus as type species. One of the new species is found along the high elevation areas of the Chapada Diamantina plateaus, state of Bahia, while the other occurs in the cerrados of "Serra Geral", in the occidental plateaus of that state. Psilops paeminosus comprises three distinct allopatric clades that, based on current evidence, cannot be diagnosed morphologically: one from the vicinities of the type locality, one from the lower Sao Francisco River, and a third from the uplands of Minas Gerais and southern inland Bahia. We keep the latter two as candidate species but defer their formal description until further evidence allows robust diagnosis. Derived clades of Psilops with shorter limbs have invaded hotter and drier environments, while mostly used sandy soils along their evolution.Fundacao de Amparo a Pesquisa do Estado de Sao Paulo (FAPESP)Fundacao de Amparo a Pesquisa do Estado do Rio de Janeiro (FAPERJ)Conselho Nacional de Desenvolvimento Cientifico e Tecnologico (CNPq)Instituto Chico Mendes de Biodiversidade (ICMBIO)Instituto Brasileiro do Meio Ambiente e dos Recursos Naturais Renovaveis (IBAMA)Secretaria do Meio Ambiente do Estado da BahiaUniv Sao Paulo, Inst Biociencias, Dept Zool, BR-05508090 Sao Paulo, SP, BrazilUniv Fed Pernambuco, Ctr Biociencias, Dept Zool, BR-50670901 Recife, PE, BrazilInst Chico Mendes Conservacao Biodiversidade, PARNA Catimbau, BR-56537000 Buique, PE, BrazilUniv Fed Rio de Janeiro, Inst Biol, Dept Zool, BR-21941902 Rio De Janeiro, RJ, BrazilUniv Fed Sergipe, Ctr Ciencias Biol & Saude, Dept Ecol, BR-49100000 Sao Cristovao, SE, BrazilUniv Fed Vicosa, Campus Florestal, BR-35690000 Florestal, MG, BrazilUniv Sao Paulo, Inst Biociencias, Dept Ecol Geral, BR-05508090 Sao Paulo, SP, BrazilUniv Fed Sao Paulo, Dept Ciencias Biol, BR-09972270 Diadema, SP, BrazilUniv Estadual Santa Cruz, Programa Posgrad Zool, BR-45662900 Ilheus, BA, BrazilUniv Fed Sao Paulo, Dept Ciencias Biol, BR-09972270 Diadema, SP, BrazilFAPESPFAPERJCNPqICMBIO: 11596ICMBIO: 14555ICMBIO: 30309IBAMA: 2001.007793/00-31]Secretaria do Meio Ambiente do Estado da Bahia: 13/2010Web of Scienc