A new species of Leposoma (Squamata, Gymnophthalmidae) from the remnant Atlantic forests of the state of Bahia, Brazil

A new species of Leposoma (Squamata, Gymnophthalmidae) from the remnant atlantic forests of the state of Bahia, Brazil

Miguel Trefaut Rodrigues^I; Marianna Dixo^II; Dante Pavan^I; Vanessa Kruth Verdade^I

^IUniversidade de São Paulo, Instituto de Biociências, Departamento de Zoologia, Caixa Postal 11.461, CEP 05422-970, São Paulo, SP, Brasil

^IIUniversidade de São Paulo, Instituto de Biociências, Departamento de Ecologia, Caixa Postal 11.461, CEP 05422-970, São Paulo, Brasil

ABSTRACT

Leposoma puk, a new species of the scincoides group is described from the Atlantic forests of southern state of Bahia, Brazil. The new species is characterized by a single frontonasal; elongate dorsal and lanceolate ventral scales arranged in diagonal rows; weakly striated or smooth scales on top of head; a longer than wide interparietal, with almost straight lateral margins; 17-18 pores; third supraocular the largest one, wider than long and clearly rectangular in shape; a suture between frontal and frontoparietal scales coincident with the middle of the third supraocular; and, a black pigmentation in the venter or along venter margins in males. The discovery of this new species, sympatric with Leposoma annectans, L. nanodactylus, and L. scincoides ranks the Atlantic Rainforest region of Southern Bahia first in species diversity for the genus. Zoogeography and speciation of Leposoma of the scincoides group are discussed.

Keywords: Leposoma puk, new species, Brazil, Squamata, Gymnophthalmidae.

RESUMO

Leposoma puk, uma nova espécie do grupo scincoides é descrita para a região da mata Atlântica do sul da Bahia, Brasil. A nova espécie é caracterizada por apresentar: frontonasal simples; escamas dorsais alongadas e ventrais lanceoladas dispostas em fileiras diagonais; escamas do topo da cabeça lisas ou levemente estriadas; interparietal mais longa que larga, com margens laterais quase paralelas; 17-18 poros; terceira supraocular mais larga que longa e a maior da área supraocular; sutura entre a frontal e frontoparietal coincidente com a região mediana da terceira supraocular; e machos com pigmentação negra no ventre ou ao longo da região paraventral. A descoberta desta nova espécie, simpátrica com Leposoma annectans, L. nanodactylus, e L. scincoides torna a região da mata atlântica do sul da Bahia a área com a maior diversidade local de espécies do gênero. Discutem-se alguns aspectos sobre a zoogeografia e a especiação de Leposoma do grupo scincoides.

Palavras-chave: Leposoma puk, espécie nova, Brazil, Squamata, Gymnophthalmidae.

INTRODUCTION

The Atlantic rainforests and their highly diverse and endemic biota are one of the top five threatened hotspots of the world (Mittermeier et al., 1997, 1999; Myers et al., 2000). Originally extending along the eastern coast of Brazil for about 5.500 km from near 6° to 30° S and covering a total area of approximatelly 1.000.000 km², less than 6% of the original coverage remains today. Despite the enormous amount of forests that have been cut down, species surveys are far from complete and several new species have been discovered during the past few years. Under the present regime of forest disturbance, discovery of new species contributes to phylogenetic understanding of particular groups and consequently the history of these forests and their relationships with other habitats in South America.

The gymnophthalmid lizards of genus Leposoma are a conspicuous component of the forest litter herpetofauna of Amazonian and Atlantic forests (Pellegrino et. al., 1999; Rodrigues, 1997; Rodrigues and Borges, 1997; Rodrigues et al., 2002). Two assemblages of species are presently known: the parietale and scincoides species groups. Until now, the highest species diversity in Leposoma was concentrated in the species of parietale group which ranges from Amazonia to Costa Rica. Nine species are presently recognized in parietale group and a maximum of two species have been reported to occur sympatrically (Avila-Pires, 1995; Rodrigues, 1997). The scincoides species group of Leposoma is restricted to the Atlantic Forest and until recently the only species admitted in the group was Leposoma scincoides. More recent surveys in the remaining patches of the Atlantic forests revealed a much higher diversity in this group: Leposoma baturitensis (Rodrigues and Borges, 1997) and Leposoma nanodactylus (Rodrigues, 1997) were described and Leposoma annectans resurrected from synonymy of Leposoma scincoides (Rodrigues et al., 2002).The revalidation of Leposoma annectans resulted from the study of a large series of specimens obtained during fieldwork at Una, state of Bahia, in the context of a broad project (RestaUna), designed to study the comparative response of several biological groups facing forest fragmentation in northern Atlantic forests (Rodrigues et. al., 2002). Although not describing it, Rodrigues et al. (2002) made reference to an adult specimen of a still undescribed Leposoma collected more than 10 years ago in the Atlantic forests of Bahia (see also Rodrigues, 1990). Fortunately, in the last field campaign of the RestaUna project another specimen of that undescribed species was obtained and subsequently, an additional specimen was collected on an adjacent mountain. Therefore, the description presented here is based on three specimens. The discovery of this new species raises to five the number of species of Leposoma known from the Atlantic forests and to four the number of species in the genus that occur sympatrically in the forests of southern Bahia, placing the area to that of with the highest local diversity for the genus.

SPECIES DESCRIPTION

Leposoma puk sp.nov.

Holotype: MZUSP 87.959, an adult male from Una: state of Bahia: Brazil (15°10'S, 39°03'W), collected by Marianna Dixo on 25 February 2000; field number MD 2592.

Paratypes: MZUSP 66.475, an adult male from São José do Macuco, presently São José da Vitória (Fazenda Unacau): state of Bahia: Brazil (15°09'S, 39°18'W); collected by Miguel Trefaut Rodrigues on 15. x. 1986, field number 86.7231, and MZUSP 89334, an adult male from Serra do Teimoso: Jussari: state of Bahia: Brazil (15°09'S, 39°31'W); collected by M. T. Rodrigues, D. Pavan, M. Dixo and V. K. Verdade on 20.iii.2001, field number MRT 6160.

Etymology: puk derives from the vocabulary of the now extinct Botocudo indians who formerly lived in the Atlantic forests of State of Bahia, Brazil. It means "the one who cries", and here is intended to reflect the loss of the Botocudo culture, as well as the highly threatened and dwindling present condition of the Atlantic Rainforests, the remaining habitat of these little forest lizards.

Diagnosis: A Leposoma of the scincoides group characterized by a single frontonasal, elongate dorsal and lanceolate ventral scales arranged in diagonal rows and having: 1) weakly striated to smooth scales on top of head; 2) interparietal longer than wide, with almost straight lateral margins; 3) parietals longer than wide; 4) 17-18 pores; 4) third supraocular the largest one, wider than long and clearly rectangular in shape; 5) a suture between frontal and frontoparietal scales that coincides with the middle of the third supraocular, and, 6) a black pigmentation in the venter or along venter margins in males.

Description of the holotype: (Figs. 1-2). Snout normal, sligthly acuminate. Rostral broad, wider than high, contacting first supralabial, nasal and frontonasal. Frontonasal entire, as wide as long; in broad contact with rostral, nasal, loreal, and prefrontal, always indenting posteriorly the latter; separating or just touching first supraocular. Prefrontals as long as large, in broad contact at midline and with first supraocular; indented anteriorly by the frontonasal and posteriorly by the frontal. Frontal hexagonal, with almost straight lateral margins; approximately twice as long as broad; indenting posteriorly a pair of frontoparietals. Frontoparietals pentagonal, larger than prefrontals, in broad contact; their midline suture approximatelly the same size as suture between prefrontals. Interparietal wider and longer than frontal, with almost straight lateral margins, longer than wide; almost straight posteriorly. Parietals irregularly hexagonal, longer than wide; as large as and as long as interparietal, always reaching the level of its posterior margin. Supraoculars four; first smallest, triangular, longer than wide and contacting loreal. Third supraocular the largest, roughly rectangular, wider than second and fourth supraoculars that are equivalent in size; all supraoculars contacting superciliaries. Nasal above first supralabial, large, longer than wide, with the nostril in the center. Nasal semidivided superiorly or divided at the level of posterior margin of nostril. Ventral margin of nasal slightly convex, dorsal margin straight. Loreal narrow, posterior to nasal, diagonally and ventro-posteriorly oriented; contacting nasal, internasal, first supraocular and first superciliar, a small preocular granule, and frenocular. Frenocular contacting nasal and followed by a series of four infraorbital elongate scutes; the first one quadrangular, smaller than frenocular, the second, third and fourth elongate, the fifth enlarged, quadrangular, and followed by a temporal scute with about the same size. Six supralabials, first and third the longest, fifth highest, third and fourth bellow eye. Sixth supralabial followed by a granule much smaller than last supralabial and separated from the tympanum by two similar granules. Five superciliaries, first largest, sligthly larger than first supraocular, expanded both on lateral and dorsal surface of head and in broad contact with loreal and a small granule placed between the latter and frenocular. Other superciliaries elongated, the last being the largest one. Eyelid with semitransparent disc formed by two enlarged scales. Temporal region almost all covered with smooth or slightly keeled and juxtaposed scales that are larger around tympanum, around parietals and surrounding postocular area. Lateral surface of neck with conical and juxtaposed keeled granules, irregularly arranged in transversal rows. Ear opening bordered by a series of very small, smooth, granules; tympanum distinct, subovoid. Scales of the top of head with very slight, low relief irregular longitudinal striations; almost smooth when compared to most of other Leposoma.

Mental broad, wider than long. Postmental single, contacting first infralabial. Three pairs of enlarged genials, all contacting infralabials; first pair the smallest one, second largest, both in broad contact at middline, and in contact with infralabials. Third pair of genials wider than long, separated medially by two elongate granules. An additional series of relatively large, diagonally oriented and flat chin-shield like scutes follows the genials. Mental, postmental and genials smooth. Infralabials five; first largest, followed by second in size. Scales immediately posterior and lateral to third pair of genials, juxtaposed, smooth, flat. From there they gradually change to lanceolate, imbricate and keeled, like posterior gulars. Gulars variable in shape, in nine transverse rows; anterior rows smooth, rounded and juxtaposed, changing progressively to lanceolate, imbricate, keeled and mucronate. Collar fold indistinct.

Dorsal scales large, strongly keeled, mucronate, lanceolate, and imbricate anteriorly, becoming wider and laterally juxtaposed with almost straight lateral sides at midbody; 28 regular transverse rows between interparietal and posterior level of hind limbs. Lateral scales resembling dorsals but more lanceolate; changing progressively to ventrals except for an area with small, rounded, almost smooth and juxtaposed conical granules around arm level. Groin scales identical to flank scales except for their smaller size. Twenty seven scales around midbody. Ventrals leaf-shaped, keeled, mucronate, imbricate, in 21 regular transverse and diagonal rows from interbrachials (included) to preanals. Posterior margin of the vent with five scales; central and lateral scales the smallest, paramedials the largest. Total pores 18; 4 of them preanal.

Scales of tail imbricate, keeled, in complete rings that became more regular ventrally. Near the base of tail dorsal and lateral tail scales are more strongly keeled and wider than ventral tail scales, becoming gradually identical towards the extremity.

Limb scales keeled and imbricate, except on ventral surface of brachium and on posterior surface of thigh which are mostly subimbricate or juxtaposed, sometimes granular. Palmar and plantar surfaces with small, conical granules. Subdigital lamellae double, 13 on Finger IV and 17 on Toe IV. Fingers and toes clawed, with the following relative sizes: 1 < 2 = 5 < 3 < 4 and 1 < 2 < 5 < 3 < 4, respectively.

Dorsal surface of body and tail light brown with irregular dark brown or black spots. Dorsal surface of head lighter. A dorsolateral light stripe one to one and half scale wide extends from lateral part of parietal scale, converge to occipital region and run to third part of tail. The dorsolateral stripe is more conspicuous anteriorly, fading progressively towards midbody. An irregular longitudinal dark brown to black area one to three scales size marginate inferiorly the dorsolateral light stripe. The dorsolateral stripes delimit a midorsal light brown area of about 4 scales wide with the characteristic dark brown reticulate background. Flanks and lateral surface of tail slightly darker than back with a few small light spots more conspicuous on neck. Dorsal surface of head lighter with scattered dark brown reticulation. Lateral surface of head darker, coloured as flanks. Ventral parts of head and body cream with a very conspicuous reticulate pattern of irregularly disposed dark spots. The dark reticulation is more concentrated in the ventrolateral area. In the ventral part of tail only the anterior third remains with some light pigmentation, rest of tail almost entirely black. Limbs dark brown dorsally, mottled with cream; ventrally yellowish cream, like in the ventral parts of body and tail.

Left hemipenis everted at preservation; bilobed, with short lobes. Sulcus spermaticus medial, bifurcating centripetally towards the apex. Right and left lobes symetric with two longitudinal rows of 20-22 enlarged spines on the apex of naked W-shaped flounces.

Measurements of the holotype: Snout-vent lenght: 38 mm; tail lenght 53 mm.

Comparisons: species of genus Leposoma are separated in two distinctive groups: parietale and scincoides (Rodrigues, 1997; Rodrigues et al., 2002). Since originally proposed by Ruibal (1952) their content varied due to the description of new species but their similarity based definition stands until now. Species of the scincoides group (annectans, scincoides, nanodactylus and baturitensis) have elongate dorsal and lanceolate ventral scales that are characteristically arranged in diagonal rather than in longitudinal rows. In the parietale group, dorsal scales are wider and shorter than those present in species of scincoides group, and ventral scales are arranged in regular longitudinal rows. Leposoma puk, is a member of the scincoides group. Leposoma puk, Leposoma nanodactylus and Leposoma baturitensis are unique in the scincoides group in presenting an entire and smooth frontonasal and weakly striated or smooth scales on head (longitudinally divided and highly striated in L. annectans and L. scincoides). Leposoma puk, Leposoma nanodactylus and Leposoma baturitensis are also the only species in the scincoides group presenting a characteristically black pigmentation in the venter or along venter margins in males (venter cream, immaculate in L. annectans and L. scincoides). Leposoma puk, nanodactylus, and baturitensis can be separated by the following list of characters.

1) Supraoculars: in Leposoma puk the third supraocular is the largest one, wider than long and clearly rectangular in shape; in L. baturitensis the second supraocular is the largest, which is squared. In L. nanodactylus, the scale corresponding to the first supraocular of baturitensis and puk is characteristically divided in two scales. This implies that the largest supraocular in L. nanodactylus (the fourth), which is also squared, is actually homologous to the third of nanodactylus and baturitensis.

2) Frontal: frontoparietal suture: in Leposoma puk the suture between frontal and frontoparietal scales coincides with the middle of the third supraocular. In L. baturitensis and L. nanodactylus the suture always reaches the anterior third of the third supraocular (actually the fourth in L. nanodactylus)

3) Shape of dorsal scales: in Leposoma puk and L. baturitensis midbody dorsal scales are wider than the anterior ones and are laterally juxtaposed with almost straight lateral sides. Dorsals in L. nanodactylus are always elongate, lanceolate, imbricate.

4) In Leposoma puk, L. baturitensis and L. nanodactylus the fingers III and IV have approximately the same comparative size but in L. puk and baturitensis the size difference between the referred fingers corresponds to less than the equivalent size of the distal supradigital lamellae; in nanodactylus to two of those scales.

5) Leposoma puk has 17-18 total pores; there are 9-11 in baturitensis and 10 in nanodactylus.

6) Leposoma puk additionally differs from L. baturitensis by slight differences in the number of dorsal and ventral scales, higher counts of scales around midbody, and of fourth finger infradigital lamellae, and from nanodactylus by a lower number of dorsal scales and higher counts of IV finger and IV toe infradigital lamellae (Table 1). Data for L. nanodactylus includes data for two specimens (one female) recently obtained at Serra do Teimoso. Table 1 also presents comparative data for other species of the scincoides group.

DISCUSSION

As usual in the Atlantic Forest domain at this latitude, most of the original forest was disturbed for cacao plantations. The paratype and first specimen of Leposoma puk was collected at São José do Macuco. It was active in the litter of a cacao grove where only large trees from primary forest (up to 30 m tall and over approximately 50 cm diameter at breast height) had been preserved to shade the understory cacao plants. At the time, the mountainous area had been recently disturbed for cacao planting and was adjacent to a hilly region covered with primary forest. The holotype was obtained in a pit-fall trap located in cacao groves at the Una Reserve, in the same general area but characterized by lower reliefs and located a few kilometers away. The third specimen was also collected in the same area but at Serra do Teimoso, a mountain reaching approximatelly 1000 meters altitude. Most of the southern face of the mountain has been preserved as a private reserve (RPPN Serra do Teimoso) and is still covered with the original forest. At the base of the mountain the forest is less diverse and humid than at the peak which is dominated by large trees (up to 30 m tall and approximately 1.0 m DBH), and characterized by higher abundance of palms, ferns, bromeliads and more leaf litter than the forest situated at lower altitudes. The specimen was obtained in a pit-fall trap located at about 900 m altitude, near the top of the mountain dominated by primary rainforest. Maximum strait-line distances among the three localities is less than 30 kilometers.

The Una area is characterized by low hills dissected by medium to small size streams. This relatively homogeneous landscape is interrupted only by a series of isolated, higher mountains like Serra do Teimoso. In contrast to the lower altitude areas used for growing cacao, these mountains are still partially covered with the original primary forest (e.g. Serra das Trempes, Serra do Padre, Serra da Lontra or Serra do Teimoso). Additional data on habitat and miscellaneous information on the local herpetofauna was presented previously (Rodrigues et al., 1989; Rodrigues, 1997; Rodrigues et al., 2002).

Four species of Leposoma occur sympatrically in this region, the highest diversity locality so far reported for the genus. Most of the specimens were collected in pit-falls traps during field work of the RestaUna project. Table 2 shows capture data for the four species in three field campaigns during the year in a low relief area (about 100 m altitude) near the Una reserve (Rodrigues et al., 2002). Leposoma scincoides was the most abundant species, followed by L. annectans; Leposoma nanodactylus and Leposoma puk were extremely rare. Table 2 also includes comparative data for species of Leposoma collected at Serra do Teimoso. Leposoma scincoides is the only widespread species, distributed from the state of Bahia to the state of Rio de Janeiro; the other three species are restricted to the state of Bahia: L. nanodactylus and L. puk are found in the area around Una, with L. annectans probably extending to Salvador.

Why is Leposoma scincoides so widespread and locally abundant in Atlantic Forests, whereas L. annectans is locally common and the other two sympatric species are so rare? Why do four species of Leposoma occur sympatrically in the general area of Una and not elsewhere? We suggest that the distribution of these species probably reflects historical patterns.

South American tropical forests have fluctuated in time and space, especially during the Quaternary (Haffer, 1979; Vanzolini and Williams, 1970) giving rise to the "refuge hypothesis" of animal and plant distributions in the Amazon. Biological and geomorphological evidence for Quaternary forest fluctuations in the Atlantic forest domain is vast (Ab'Saber, 1977, 1982; Haffer, 1979, 2001; Heyer and Maxon, 1983; Jackson, 1978).

The "brejos", isolated forest islands located in the semiarid Caatingas of northeastern Brazil provide textbook examples of refuges (Vanzolini, 1981; Vanzolini and Williams, 1981; Rodrigues, 1990; 1996). Their relict forest-adapted fauna persists from eras when forest was continuous between Atlantic forest and the "brejos" of today. Presently, vestiges of similar forest patches not larger than 300 km² occur within the semiarid Caatingas; a region that spans approximately 800,000 km² (Ab'Saber, 1977). One example is Serra de Baturité, a mountain complex in the State of Ceará, and the type locality of Leposoma baturitensis the only extralimital species of the scincoides group of Leposoma known. From the top of the highest Baturité mountain one can see that forest is restricted to the mountainous area. Other physionomically similar scenarios exist in the Caatingas of northeastern Brazil as a testimony of times when, under more humid climates, these areas and Atlantic forest were possibly connected.

If we examined early periods in the Quaternary characterized by higher humidity than present day and look at the prevalent landscape physiognomy, the relictual condition dissappears. At those times, Serra de Baturité would have been connected by forest to the surrounding area. What would have been the distributional limits of Leposoma baturitensis in those times? Would it have been restricted to Baturité mountains or widespread around forest areas surrounding them? An affirmative answer to both questions is possible. Restriction of Leposoma baturitensis to the Baturité mountains could be explained by recent differentiation in isolation and/or by historical contact with another widespread generalist congeneric species that despite the continuous forest coverage prevented expansion of its distribution. As there are no vestiges of a putative relative we could hypothesize that it became extinct when, following climatic deterioration towards present, the forest dwindled and disappeared from surrounding areas. Under the second possibility, the same arguments could explain disappearance of a more widespread baturitensis from the adjacent areas today dominated by the semiarid Caatingas.

As in the Baturité example, we may use similar logic to explain the history of Atlantic forest in the study area. The aridity that progressively affected northeastern Brazil left a formerly continuous forest fauna isolated in hilly areas surrounding by Caatingas (Oliveira et al., 1999). Forested mountains like Serra das Trempes, Serra dos Padres, Serra do Teimoso, or Serra da Lontra, may have acted as forest refugia in the recent past. Presently, they are what we are here calling concealed refuges because they remain as yet unrecognized. This is because under the present humid climate the general green coverage of these mountains are simply recognized as part of the Atlantic forest region, a large core area described mainly on geomorphologic grounds, rather than faunal attributes.

Considering the rarity of Leposoma nanodactylus and Leposoma puk in the lowlands surrounding Una, we hypothesize that these species may have differentiated and remained isolated in some of these concealed refuge areas. Both the widespread and generalist Leposoma scincoides and the relatively common Leposoma annectans might outcompete these rare species in forest surrounding the concealed refuge areas.

Presently, even after climatic change and intensive exploration for wood, agriculture, mining, and cattle ranching, has dramatically altered the landscape, the contrast between open and formerly forested areas remains evident in the semiarid Caatingas of northeastern Brazil. Evergreen forested areas are visually recognized as islands in the grayish-brown dry Caatingas. This is why putative refugia or centers of differentiation of forest fauna in the semiarid Caatingas were first surveyed and should receive priority for conservation. Their insularity and endemisms are so obvious that they are obvious candidates for conservation priority (Conservation International, 1994). In contrast, several important probable refuges are overlooked because they are part of an otherwise homogeneous habitat. These concealed refuges and their limits can only be detected with intensive biogeographical study.

The highly endemic sand dune area of Rio São Francisco in the core area of semiarid Caatingas of State of Bahia is now recognized as a concealed refugium (Rodrigues, 1991a,b,c,d; 1993a,b; 1996). We suggest that the relictual mountains near Una in the Atlantic Forest of State of Bahia provide other examples of concealed refugia. Our suggestion could be tested with additional field work in the region complemented by molecular phylogeographic studies. If our hypothesis is confirmed, we hope there will be enough time left to take adequate conservation measures before the remaining forest habitats and their associated fauna are destroyed.

ACKNOWLEDGEMENTS

We thank Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP), Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), and Financiadora de Estudos e Projetos (FINEP) for support. For their invaluable help in the field we thank A. Langguth, E.X. Freire, C. Bonvicino, M.N. Stevaux, the staff of Fazenda Unacau and of RestaUna project, especially R. Pardini and D. Faria, and H. and L.V. Carvalho of the Reserva Particular do Patrimônio Natural (RPPN) Serra do Teimoso. We also thank L. Seky for valuable help with Botocudo vocabulary, Luciana Lobo for the drawing, and B. Zimmerman and A. Chek for reading the manuscript and making invaluable suggestions for its improvement.

Trabalho recebido para publicação em 18.III.2002 e aceito em 06.VIII.2002

ADDITIONAL SPECIMENS EXAMINED

Leposoma nanodactylus – Brazil: Bahia: Una: MZUSP 87955-87957, 87986-87987; Serra do Teimoso: Jussari: MZUSP 89332-89333.

Leposoma baturitensis – Brazil: Ceará: Serra de Baturité: Pacoti (Sítio Barbosa) MZUSP 79378; Pacoti (Sítio São Gonçalo do Freire) MZUSP 79379.

Leposoma annectans – Brazil: Bahia: Una: MZUSP 87792-87804; 87851-87874; 87958.

Leposoma scincoides – Brazil: Bahia: Una: MZUSP 87805-87050, 87875-87954; Serra do Teimoso: Jussari: MZUSP 89324-89329.

• Avila-Pires, T.C.S. 1995. Lizards of Brazilian Amazonia (Reptilia: Squamata). Zool. Verh. Leiden 299:1-706.
• Ab'Saber, A.N. 1977. Os domínios morfoclimáticos da América do Sul. Primeira aproximação. Geomorfologia 53:1-23.
• Ab'Saber, A.N. 1982. The paleoclimate and paleoecology of Brazilian Amazonia. Pp. 41-59. In G. T. Prance (Ed.), Biological Diversification in the Tropics. Columbia University Press, New York, New York.
• Conservation International, 1994. Prioridades para Conservação da Biodiversidade da Mata Atlântica do Nordeste. Map. Ministério do Meio Ambiente, dos Recursos Hídricos e da Amazônia Legal.
• Haffer, J. 1979. Quaternary Biogeography of Tropical Lowland South America. Pp. 107-140 In: W. E. Duellman (Ed.), The South American Herpetofauna: Its Origin, Evolution, and Dispersal. Monogr. Mus. Nat. Hist. Univ. Kansas (7):1-485.
• Haffer, J. 2001. Hypothesis to explain the origin of species in Amazonia. Pp. 45-118 In I.C.G. Vieira, J.M.C. Silva, D.C. Oren & M.A. d'Incao (eds.), Diversidade Biológica e Cultural da Amazônia. Museu Paraense Emílio Goeldi, Belém, Pará.
• Heyer, W.R. & L.R. Maxon. 1983. Relationships, zoogeography, and spetiation mechanisms of frogs of the genus Cycloramphus (Amphibia, Leptodactylidae). Arquivos Zool., S. Paulo 30(5):341-373.
• Jackson, J.J. 1978. Differentiation in the genera Enyalius and Strobilurus (Iguanidae): Implications for Pleistocene climatic changes in eastern Brazil. Arquivos Zool., S. Paulo 30(1):79 pp.
• Mittermeier, R.A. & C.G. Mittermeier. 1997. Megadiversity. Earth biologically wealthiest nations. CEMEX. Quebecor Printing. Canada. 501 pp.
• Mittermeier, R.A., N. Myers, P. Robles Gil & C.G.. Mittermeier. 1999. Hotspots: earth biologically richest and most endangered terrestrial ecoregions. CEMEX. Agrupación Sierra Madre, Mexico. 430 pp
• Myers, N., R.A. Mittermeier, C. Mittermeier, G.A.B. Fonseca & J. Kent. 2000. Biodiversity hotspots for conservation priorities. Nature 403:853-858.
• Oliveira, P.E.; A.M.F. Barreto & K. Suguio. 1999. Late Pleistocene/Holocene climatic and vegetational history of the Brazilian Caatinga: the fossil dunes of the middle São Francisco River. Palaeo 152:319-337.
• Pellegrino, K.C.M., M.T. Rodrigues & Y. Yonenaga-Yassuda. 1999. Chromosomal evolution in Brazilian lizards of genus Leposoma (Squamata, Gymnophthalmidae) from Amazon an Atlantic forest: evidences based on R-banding and FISH of telomeric sequences. Hereditas 131(1):15-21.
• Rodrigues, M.T. 1990. Os lagartos da floresta Atlântica brasileira: distribuição atual e pretérita e suas implicações para estudos futuros. Pp. 404-410 in II Simpósio sobre ecossistemas da costa sul e sudeste brasileira. Estrutura, Função e Manejo. Vol. 1. Academia de Ciências do Estado de São Paulo.
• Rodrigues, M.T. 1991a. Herpetofauna das dunas interiores do rio São Francisco: Bahia: Brasil. I. Introdução à área e descrição de um novo gênero de microteiideos (Calyptommatus) com notas sobre sua ecologia, distribuição e especiação (Sauria, Teiidae). Papéis Avulsos de Zoologia, S. Paulo 37(19):285-320.
• Rodrigues, M.T. 1991b. Herpetofauna das dunas interiores do rio São Francisco: Bahia: Brasil. II. Psilophthalmus: um novo gênero de microteiideos sem pálpebra (Sauria, Teiidae). Papéis Avulsos Zool., S. Paulo 37(20):321-327.
• Rodrigues, M.T. 1991c. Herpetofauna das dunas interiores do rio São Francisco: Bahia: Brasil. III. Procellosaurinus: um novo gênero de microteiideo sem pálpebra, com a redefinição do gênero Gymnophthalmus (Sauria, Teiidae). Papéis Avulsos Zool., S. Paulo 37(21):329-342.
• Rodrigues, M.T. 1991d. Herpetofauna das dunas interiores do rio São Francisco: Bahia: Brasil. IV. Uma nova espécie de Typhlops (Ophidia, Typhlopidae). Papéis Avulsos Zool., S. Paulo 37(22):343-346.
• Rodrigues, M.T. 1993a. Herpetofauna das dunas interiores do rio São Francisco: Bahia: Brasil. V. Duas novas espécies de Apostolepis (Serpentes, Colubridae). Mem. Inst. Butantan 54(2):53-59 (1992).
• Rodrigues, M.T. 1993b. Herpetofauna of palaeoquaternary sand dunes of the middle São Francisco river: Bahia: Brazil. VI. Two new species of Phimophis (Serpentes: Colubridae) with notes on the origin of psammophilic adaptations. Papéis Avulsos Zool., S. Paulo 38(11):187-198.
• Rodrigues, M.T. 1996. Lizards, snakes and amphisbaenians from the Quaternary sand dunes of the middle Rio São Francisco: Bahia: Brazil. Jour. Herpetol. 30(4):513-523.
• Rodrigues, M.T. 1997. A new species of Leposoma (Squamata: Gymnophthalmidae) from the Atlantic forest of Brazil. Herpetologica 53(3):383-389.
• Rodrigues, M.T. & D.M. Borges. 1997. A new species of Leposoma (Sauria, Gymnophthalmidae) from a relictual forest of semiarid northeastern Brazil. Herpetologica 53(1):1-6.
• Rodrigues, M.T., M. Dixo & G.M. Accacio. 2002. A large sample of Leposoma (Squamata, Gymnophthalmidae) from the Atlantic Forests of Bahia, the status of Leposoma annectans Ruibal, 1952, and notes on conservation. Papéis Avulsos Zool., S. Paulo 42(5):103-117.
• Rodrigues, M.T., Y. Yonenaga-Yassuda & S. Kasahara. 1989. Notes on the ecology and karyotypic description of Strobilurus torquatus (Sauria, Iguanidae). Ver. Brasil. Genet. 12:747-759.
• Ruibal, R. 1952. Revisionary studies of some South American Teiidae. Bull. Mus. Comp. Zool. 106:407-529.
• Vanzolini, P.E. 1981. A quasi-historical approach to the natural history of the differentiation of reptiles in tropical geographic isolates. Papéis Avulsos Zool., S. Paulo 39(19):189-204.
• Vanzolini, P.E. & E.E. Williams. 1981. The vanishing refuge: a mechanism for ecogeographic speciation. Papéis Avulsos Zool., S. Paulo, 34(23):251-255.

Publication Dates

• Publication in this collection
  28 Oct 2003
• Date of issue
  Nov 2002

History

• Received
  18 Mar 2002
• Accepted
  06 Aug 2002