Relicts in the mist: Two new frog families, genera and species highlight the role of Pantepui as a biodiversity museum throughout the Cenozoic.

The iconic mountains of the Pantepui biogeographical region host many early-diverging endemic animal and plant lineages, concurring with Conan Doyle's novel about an ancient "Lost World". While this is the case of several frog lineages, others appear to have more recent origins, adding to the controversy around the diversification processes in this region. Due to its remoteness, Pantepui is challenging for biological surveys, and only a glimpse of its biodiversity has been described, which hampers comprehensive evolutionary studies in many groups. During a recent expedition to the Neblina massif on the Brazil-Venezuela border, we sampled two new frog species that could not be assigned to any known genus. Here, we perform phylogenetic analyses of mitogenomic and nuclear loci to infer the evolutionary relationships of the new taxa and support their description. We find that both species represent single lineages deeply nested within Brachycephaloidea, a major Neotropical clade of direct-developing frogs. Both species diverged >45 Ma from their closest relatives: the first is sister to all other Brachycephaloidea except for Ceuthomantis, another Pantepui endemic, and the second is sister to Brachycephalidae, endemic to the Brazilian Atlantic Forest. In addition to these considerable phylogenetic and biogeographic divergences, external morphology and osteological features support the proposition of two new family and genus-level taxa to accommodate these new branches of the amphibian tree of life. These findings add to other recently described ancient vertebrate lineages from the Neblina massif, providing a bewildering reminder that our perception of the Pantepui's biodiversity remains vastly incomplete. It also provides insights into how these mountains acted as "museums" during the diversification of Brachycephaloidea and of Neotropical biotas more broadly, in line with the influential "Plateau theory". Finally, these discoveries point at the yet unknown branches of the tree of life that may go extinct, due to global climate change and zoonotic diseases, before we even learn about their existence, amphibians living at higher elevations being particularly at risk.

A new Crossodactylodes Cochran, 1938 (Anura: Leptodactylidae: Paratelmatobiinae) from the highlands of the Atlantic Forests of southern Bahia, Brazil.

A new Crossodactylodes is described from Serra das Lontras, in the highlands of the Atlantic Forests of southern Bahia. The new species can be distinguished from all other Crossodactylodes by having Finger I ending in an acute tip, a larger body size, by cranial features, and by molecular data. Like their congeners, the new species live in bromeliads but is widely geographically disjunct, being apparently restricted to the summit of a mountain range in Northeastern Brazil.

A new species of Leposoma (Squamata: Gymnophthalmidae) with four fingers from the Atlantic Forest central corridor in Bahia, Brazil.

Leposoma sinepollex sp. nov., a new species of the scincoides group, is described from a mountain region in the Atlantic Forest central corridor in state of Bahia, Brazil. The new species is characterized by elongate dorsal and lanceolate ventral scales arranged in diagonal rows, a single and smooth frontonasal, five supraoculars, absence of pollex, third toe as long as or longer than fourth, absence of striations in lower part of head, parietals longer than wide and as long as interparietal, 27-29 dorsals, 25-29 scales around body, 17-19 ventrals, 12-14 total pores in the male (absent in females), 9-10 and 9-11 subdigital lamellae respectively under IV Finger and IV Toe, and strong sexual color dichromatism with a black pigmentation in the ventral parts of males, creamy in females. The new species is morphologically similar to Leposoma nanodactylus, sharing with it among other features the synapomorphic division of the first supraocular. Phylogenetic analyses of 981 bp of combined sequences (cyt b+ ND4) recovered also a strongly supported (PP = 1,0; BP = 100) sister relationship between both species. The new species and Leposoma nanodactylus are placed sister to all the other Atlantic Forest species, with L. baturitensis being the first to diverge in this radiation. We discuss the distribution of the Atlantic Forest Leposoma, as well as possible scenarios for the origin of the new species.

A new species of Bachia Gray, 1845 (Squamata: Gymnophthalmidae) from the Eastern Brazilian Cerrado, and data on its ecology, physiology and behavior.

A new species of Bachia of the bresslaui group, Bachia geralista sp. nov., is described from Planalto dos Gerais, an old and partially dissected plateau extending along the Cerrados of Bahia, Minas Gerais and Tocantins states, Brazil. The new species is morphologically similar to B. bresslaui, with which it has been confused; however head scalation resembles other species from sandy spots within the Cerrado (B. psamophila and B. oxyrhina). Like in B. psamophila and B. oxyrhina, the shovel-shaped snout of the new species is highly prominent, a typical trait of psammophilous habits in other gymnophthalmids. The examination of specimens of B. bresslaui from several populations within the Cerrado revealed great variation among localities, leading to the reidentification of a specimen from Utiariti, Mato Grosso, previously referred to in the literature as the second record of B. bresslaui, as the recently described B. didactyla, suggesting that cryptic diversity might remain still undiscovered within this genus in the Cerrado. Despite occurring in a relatively open Cerrado, thermal physiology of Bachia geralista sp. nov. restricts its occurrence to shaded microhabitats within this habitat.

Rediscovery of the earless microteiid lizard Anotosaura collaris Amaral, 1933 (Squamata: Gymnophthalmidae): a redescription complemented by osteological, hemipenial, molecular, karyological, physiological and ecological data.

More than a century after its discovery by Ernest Garbe, and almost 80 years after its original description, we obtained a series of specimens of the earless gymnophthalmid Anotosaura collaris, the type species of the genus, up to now known only by a single specimen. On the basis of the material obtained at and close to the type locality we redescribe the species, adding information about the external and hemipenial morphology, osteology and karytoype. Molecular data confirm its sister relationship with Anotosaura vanzolinia as well as the close relationship of Anotosaura with the Ecpleopodini Colobosauroides and Dryadosaura. We supplement this information with thermophysiological, ecogeographical, karyotypic and ecological data.

Morphological variation and genealogical discordance in Caatinga sand lizards Calyptommatus Rodrigues 1991 (Squamata: Gymnophthalmidae) with the description of a new species.

In the last decades, a remarkable fauna of psammophilous and fossorial squamates was discovered in sandy habitats of the semiarid Caatinga of northeast Brazil. Despite the increasing accumulation of genetic data from this unique fauna, an incomplete knowledge of its diversity still hampers a better understanding of its origins and diversification. The fossorial lizard genus Calyptommatus (Gymnophthalmidae) is endemic to sandy habitats of the Caatinga, being currently represented by four allopatric species. In this study, we used morphological and molecular data to assess population-level variation in Calyptommatus. We found a new morphotype of Calyptommatus from the state of Bahia, Brazil, readily distinguished from congeners by the presence of a frontal scale. Morphological, nuDNA and geographic data support the recognition of a new species herein described as Calyptommatus frontalis sp. nov. Nevertheless, genetic data revealed mito-nuclear+morphology discordance, with populations with frontal scales distributed in three distantly related mtDNA clades, suggesting either potential historical and/or current introgressions or incomplete lineage sorting. Further data are needed to clarify the status of the two other mtDNA clades displaying a frontal scale.

A new large canopy-dwelling species of Phyllodytes Wagler, 1930 (Anura, Hylidae) from the Atlantic Forest of the state of Bahia, Northeastern Brazil.

The known diversity of treefrogs of the genus Phyllodytes has rapidly increased in recent years, currently comprising 14 species. Recent field work in the Atlantic Rainforest of the state of Bahia lead to the discovery of a new large species of Phyllodytes which is herein described based on multiple evidence including morphological, acoustical and genetic data. Phyllodytes sp. nov. is one of the largest species within the genus and presents immaculate yellowish dorsum and limbs. The advertisement call of the species is composed of 7-31 notes (half pulsed/pulsatile-half harmonic) with frequency-modulated harmonics. Phyllodytes sp. nov. has a karyotype of 2n = 22 chromosomes, as also found in other species of the genus. Genetic distance values of the 16S mitochondrial rRNA among Phyllodytes sp. nov. and its congeners range between 6.4 to 10.2%. The description of another new species for this state reinforces the need for further taxonomic work with Phyllodytes in this region that has been revealed as a priority area for research and conservation of this genus.

Geographic variation and taxonomy of red-tailed Gymnophthalmus (Squamata: Gymnophthalmidae) from Amazonian Savannas.

Gymnophthalmus represents a challenging group for systematics of microteiid lizards. Due to scarcity of exemplars and molecular data, the taxonomy of the genus has been unstable, and six of its eight species are considered part of a poorly-delimited complex of unisexual and bisexual species. Unnamed populations of red-tailed Gymnophthalmus from savanna enclaves in Amazon, similar to G. vanzoi but with differences in color pattern, have been noted in the literature. Here, we used molecular and morphological data to test the taxonomic status of the Central Amazonian red-tailed Gymnophthalmus (G. sp.). Our molecular analysis recovered a close relationship between G. vanzoi and G. sp. from Central Amazon, with minimal divergence. Samples were similar in scale counts but presented significant variation in color pattern and morphometry. Despite the geographic isolation, individuals of both populations cannot be fully differentiated based on morphology. Thus, our results suggest that G. vanzoi has a wider distribution, including the Central Amazonian red-tailed Gymnophthalmus and likely morphologically similar populations from other enclaves of Amazonian savannas, which may have been isolated recently. Based on our comparative analysis, we highlight some characters proved useful for differentiating Brazilian Gymnophthalmus and discuss some prospects for the taxonomy of the genus.

The herpetofauna of Parque Nacional da Serra das Confusões, state of Piauí, Brazil, with a regional species list from an ecotonal area of Cerrado and Caatinga / A herpetofauna do Parque Nacional da Serra das Confusões, Piauí, Brasil, com uma lista regional para uma área ecotonal entre o Cerrado e a Caatinga

Abstract Different physiognomies at Parque Nacional da Serra das Confusões (PNSCo) were intensively sampled aiming to access the distribution pattern of its herpetofauna. Sixty six species were found in the park (47 reptiles and 19 amphibians); the rarefaction curve for lizards, although not fully stabilized in an asymptote, indicates that the sampling effort was enough to reveal most lizard species occurring in the area; and richness estimators recovered values close to observed. For amphibians, the curve shows a weak tendency to stabilization with richness estimators indicating that additional records could be done. Field work carried out at PNSCo has highlighted an unique herpetofauna: five new species were described and there are three candidates as new species. The regional list including Cerrados's units - Estação Ecológica Serra Geral do Tocantins (EESGT) and Estação Ecológica de Uruçuí-Una (EEUU) with Caatinga's ones - PNSCo and Parque Nacional da Serra da Capivara (PNSCa), shows a high herpetofaunal diversity (191 species) to the region. The cluster analysis recovered the Cerrados's units and Caatinga's ones, in separate clusters evidencing a species turnover between domains, despite its geographical proximity. Thus, although there is widespread fauna throughout region shared by the units, each reserve holds its own faunal identity, harboring a singular assemblage of species.

Resumo Diferentes fisionomias foram intensivamente amostradas no Parque Nacional da Serra das Confusões (PNSCo) com o objetivo de acessar o padrão de distribuição da sua herpetofauna. Sessenta e seis espécies foram encontradas no parque (47 répteis e 19 anfíbios); a curva de rarefação de lagartos, embora não totalmente estabilizada em uma assíntota, indica que o esforço amostral foi suficiente para revelar a maioria das espécies de lagartos que ocorrem na área; e estimadores de riqueza recuperaram valores perto dos observados. Para os anfíbios, a curva mostra uma fraca tendência para a estabilização com estimadores de riqueza indicando que registros adicionais podem ser feitos. O trabalho de campo realizado no PNSCo destacou uma herpetofauna única: cinco novas espécies foram descritas e há três candidatos como novas espécies. A lista regional, incluindo unidades de Cerrado - Estação Ecológica Serra Geral do Tocantins (EESGT) e Estação Ecológica Uruçuí-Una (EEUU) com aqueles da Caatinga - PNSCo e Parque Nacional da Serra da Capivara (PNSCa), mostra uma alta diversidade da herpetofauna (191 espécies) para a região. A análise de agrupamento recupera as unidades do Cerrado e as da Caatinga em grupos separados, evidenciando uma substituição de espécies entre os domínios, apesar da sua proximidade geográfica. Assim, embora haja fauna ocorrente em toda a região e compartilhada pelas unidades, cada uma das reservas mantém a sua própria identidade, abrigando um conjunto singular de espécies.

Anfíbios da Estação Ecológica Serra Geral do Tocantins, região do Jalapão, Estados do Tocantins e Bahia / Amphibians from Estação Ecológica Serra Geral do Tocantins, Jalapão region, Tocantins and Bahia States

A porção norte do domínio do Cerrado é uma das áreas historicamente menos conhecidas com relação à sua biodiversidade. Recentemente, alguns estudos tem revelado valores de riqueza comparáveis a outras regiões dentro do domínio. A Estação Ecológica Serra Geral do Tocantins (EESGT) está localizada na região do Jalapão, porção Nordeste do Cerrado, e faz parte do maior bloco de áreas protegidas neste domínio. Neste estudo descrevemos a riqueza e composição de espécies de anfíbios da EESGT, discutindo-as em um contexto biogeográfico, e caracterizamos o uso de sítios reprodutivos pelas espécies de anfíbios registradas em relação às fitofisionomias e aos tipos de corpos d'água. Utilizamos os métodos de busca ativa e armadilhas de queda, no período considerado como o auge da estação reprodutiva para a maior parte das espécies do Cerrado. Foram registradas 36 espécies de anfíbios na EESGT, totalizando 39 espécies conhecidas para a região do Jalapão. Aplicando o estimador Jackknife, sugerimos uma riqueza potencial de 42 espécies para a EESGT. A maior parte das espécies registradas é endêmica ou fortemente associada ao Cerrado, seguidas pelas espécies de ampla distribuição no Brasil ou América do Sul. A maior parte da espécies se reproduz em poças temporárias localizadas em áreas abertas, embora existam espécies que ocorrem exclusivamente em matas de galeria e utilizem corpos d'água lóticos para se reproduzir.

The Northern part of Cerrado is one of the biologically poorest known areas in the domain. Recent studies revealed richness values that are as high as those from other sites in the domain. The Estação Ecológica Serra Geral do Tocantins is located in a region known as Jalapão, in Northeastern Cerrado, and it is included in the largest set of protected areas in the domain. We describe amphibian richness and species composition in EESGT, and discuss it in a biogeographic context. We also describe breeding sites focusing on phytophysiognomy and types of water bodies. We sampled amphibians through active search and pitfall traps, during the peak of breeding season for most of the anuran species in Cerrado. We registered 36 species, which coupled with former studies results in a regional richness of 39 species known for Jalapão. After applying Jackknife estimator, we suggest a potential richness of 42 species for the EESGT. Most registered species are endemic or strongly associated to Cerrado, followed by species widespread in Brazil or South America, and those associated with Caatinga. Most species breed in temporary ponds located in open areas, although there are some forest specialists, and stream-breeding species.

Répteis da Estação Ecológica Serra Geral do Tocantins, Brasil Central / Reptiles of Serra Geral do Tocantins Ecological Station, Central Brazil

Os Cerrados sul-americanos abrigam alta diversidade de répteis, incluindo elevado número de endemismos. No entanto, o conhecimento desta diversidade é ainda incompleto frente à acelerada transformação das paisagens naturais no Brasil central. Constituem, portanto, uma das regiões prioritárias para estudo e conservação da biodiversidade mundial. Estudos intensivos sobre a fauna de répteis do Cerrado são necessários e urgentes para melhor compreensão dos processos que levaram à sua origem e distribuição e para subsidiar ações de conservação. Por meio de métodos padronizados, amostramos duas regiões ainda inexploradas da Estação Ecológica Serra Geral do Tocantins, situada na região do Jalapão. Registramos 45 espécies de répteis para a EESGT e entorno, o que representa uma riqueza alta e comparável à de outras regiões bem amostradas do Cerrado. Curvas de acumulação e estimadores indicam que a riqueza local de lagartos e anfisbenídeos aproxima-se da riqueza real enquanto a de serpentes é subestimada. A distribuição não-aleatória das espécies na paisagem concorda com evidências anteriores sugerindo utilização diferencial dos hábitats pelos répteis. Reunindo os resultados do presente estudo com os de levantamentos prévios realizados na região, registramos 88 espécies de répteis para o Jalapão sendo oito registros novos que incluem Bachia oxyrhina uma espécie recém descrita da região. As espécies da área apresentam três padrões gerais de distribuição: (1) espécies endêmicas do Cerrado, (2) espécies compartilhadas com domínios da diagonal de formações abertas sul-americanas, e (3) espécies de ampla ocorrência, compartilhadas também com ecossistemas florestais. Prevalecem espécies de ampla distribuição, porém é grande o número de espécies típicas do Cerrado, incluindo cinco possivelmente endêmicas do Jalapão, e há contribuição importante da fauna da Caatinga. A distribuição dos répteis em escala local e regional demonstra a necessidade de considerar a heterogeneidade paisagística para o planejamento de diretrizes visando à conservação em regiões do Cerrado. Por sua grande extensão, posição biogeográfica e complexidade de relevo e tipos de hábitat, a EESGT tem papel fundamental para a preservação e conhecimento da diversidade de répteis do Cerrado.

The South American Cerrado harbors a rich reptilian fauna with high number of endemics. However, knowledge of this diversity is still incipient in front of accelerated transformation of natural landscapes, which makes the domain a "hotspot" for study and conservation of global biodiversity. Studies on the reptile fauna of Cerrado are urgently needed for a better understanding of the processes that led to their origin and distribution and to support conservation planning. We used standard methods to sample two unexplored regions of the Serra Geral do Tocantins Ecological Station (EESGT), located in the Jalapão region of the Cerrado domain. We recorded 45 species of reptiles for the EESGT and surroundings, which represents high species richness, comparable to other well-sampled regions of the Cerrado. Accumulation curves and estimators indicate that sampled richness of lizards and amphisbaenas are close to real values. However, current sampling underestimates snake richness. The non-random distribution of species in the landscape agrees with previous evidence suggesting differential use of habitats by Cerrado reptiles. Combining the results of this study with those of previous surveys conducted in the region, we recorded 88 species of reptiles for the Jalapão region with eight new records, including Bachia oxyrhina, a recent described species of this region. We found three general patterns of distribution: (1) species endemic of the Cerrado domain, (2) species shared with the domains of the South American diagonal of tropical open formations, (3) species widely distributed, also shared with forest ecosystems. Species of wide distribution prevails in the region, but there is large number of species typical of the Cerrado, possibly including five endemics to Jalapão, and there is an important contribution of the Caatinga fauna. The distribution of reptile species in local and regional scales shows the need to consider the landscape heterogeneity for adequate conservation planning in the Cerrado region. Because of the physical and biotic characteristics, EESGT has a key role in the preservation and knowledge of the diversity of Cerrado reptiles.

Vertebrados da Estação Ecológica Serra Geral do Tocantins: faunística, biodiversidade e conservação no Cerrado brasileiro / Vertebrate fauna of Estação Ecológica Serra Geral do Tocantins: biodiversity and conservation in the Brazilian Cerrado hotspot

Inventários e estudos faunísticos detalhados sobre vertebrados são uma das fontes mais relevantes de dados para interpretações de padrões detalhados de diversidade biológica. Dados básicos e de boa qualidade sobre faunística são ainda mais urgentes em regiões pouco estudadas e sob intensa ameaça antrópica, tais como a região do Cerrado, um dos 34 hotspots globais para a conservação da biodiversidade. Apresentamos aqui uma síntese dos resultados dos inventários de vertebrados na Estação Ecológica Serra Geral do Tocantins (~716.000 ha), a segunda maior unidade de conservação em todo o Cerrado. Foram registradas 450 espécies de vertebrados na EESGT e entorno imediato, incluindo 17 espécies ameaçadas, 50 espécies endêmicas do Cerrado e 11 espécies com distribuição potencialmente restrita. Do total de espécies amostradas, 180 são novos registros para a região do Jalapão. Ao menos 12 espécies amostradas foram consideradas potenciais espécies novas, das quais quatro foram descritas recentemente, a partir do material obtido no inventário. Os resultados evidenciam que a EESGT é uma das mais importantes áreas protegidas no Brasil central, contribuindo para a persistência de espécies ameaçadas, dependentes dos últimos grandes blocos contínuos de vegetação nativa de Cerrado. Nossos resultados indicam ainda que a conservação da EESGT e suas principais subunidades é crucial para a representatividade do sistema de áreas protegidas do Cerrado, protegendo potenciais endemismos restritos que aliam alta vulnerabilidade intrínseca e valor como indicadores de padrões e processos biogeográficos formadores da rica e cada vez mais ameaçada fauna Neotropical.

Basic taxonomic and distributional data on vertebrates are one of the most useful and reliable sources of information for conservation planning. Biological data are even more relevant in rich and highly threatened regions such as the Brazilian Cerrado, one of the least studied global biodiversity hotspots. Herein we provide a summary of the results of a vertebrate survey at Estação Ecológica Serra Geral do Tocantins (~716.000 ha), the second largest protected area in the Cerrado region. We recorded 450 species in EESGT and surroundings, including 17 threatened species, 50 Cerrado endemics and 11 potential restricted-range species. Our results also added 180 new vertebrate records for the Jalapão region. At least 12 species were considered potential undescribed taxa; four of these were recently described based on specimens obtained in the present study. Our results indicate that EESGT is among the most biologically relevant protected areas in the Cerrado. Proper management will favor the persistence of threatened vertebrates dependent on the last remaining large blocks of pristine Cerrado savannas. Moreover, EESGT and its major biological subunits contribute decisively to the representativeness of the reserve system in the Cerrado, conserving presumed narrow endemics with high intrinsic vulnerability and high potential value as indicators of biogeographic processes of diversification in rich and complex Neotropical biotas.