The phylogenetic position of ridley's worm lizard reveals the complex biogeographic history of New World insular amphisbaenids.

The archipelago of Fernando de Noronha (FN) is located in the Equatorial South Atlantic Ocean, at 375 km off the northeastern coast of Brazil. Its endemic vertebrate land fauna is restricted to only six species, and three main hypotheses have been proposed to explain their presence in the archipelago. These hypotheses suggest FN had alternative biogeographic connections with: 1) the West Indies; 2) the South America mainland; or 3) Africa. Here, we evaluate for the first time the phylogenetic position of Amphisbaena ridleyi within the diversity of Amphisbaenia, and we infer the biogeographic processes that explain its presence in FN and its relationship with amphisbaenids from the West Indies. We analyzed a comprehensive multilocus dataset for Amphisbaenidae using maximum likelihood and time-calibrated Bayesian phylogenetic approaches. Based on our time-calibrated tree, we tested different biogeographic scenarios through historical biogeographic analyses. Our phylogenetic results for the high-level relationships of Amphisbaenia can be parenthetically summarized as (Rhineuridae, (Blanidae, (Bipedidae, (Cadeidae, (Trogonophidae, (Amphisbaenidae))))). Nine highly supported groups of species were recovered among the mainland South American amphisbaenids (SAA), whereas two phylogenetically distant groups of species were inferred for the West Indies: 1) WIC01, an Oligocene lineage present in Cuba and Hispaniola, which is the sister group of all other SAA groups; and 2) WIC02, a Miocene lineage that is restricted to southern Hispaniola and is closely related to Am. ridleyi. We estimated two events of transatlantic dispersal of amphisbaenians from Africa to West Indies: the dispersal of Cadeidae during the Middle Eocene, and the dispersal of the ancestor of Amphisbaena during the transition Eocene/Oligocene. These events were likely affected by the North Equatorial and South Equatorial currents, respectively, which have been flowing westwards since the Paleocene. The ancestral cladogenesis of Amphisbaena during the Late Oligocene is likely related to overwater dispersal events, or alternatively can be associated with the fragmentation of GAARlandia, when WIC01 was isolated in the West Indies, while the remaining groups of Amphisbaena diversified throughout the South American continent. During the Late Miocene, the ancestor of WIC02 dispersed from northern South America to the West Indies, while Am. ridleyi dispersed from the same region to FN. The overwater dispersal of WIC02 was driven by the North Brazilian Current and the dispersal of Am. ridleyi was likely influenced by the periodic shifts in direction and strength estimated for the North Equatorial Counter-Current during the Late Miocene.

Revealing the cryptic diversity of the widespread and poorly known South American blind snake genus Amerotyphlops (Typhlopidae: Scolecophidia) through integrative taxonomy

Morphological stasis is generally associated with relative constancy in ecological pressures throughout time, producing strong stabilizing selection that retains similar shared morphology. Although climate and vegetation are commonly the main key factors driving diversity and phenotypic diversification in terrestrial vertebrates, fossorial organisms have their morphology mostly defined by their fossorial lifestyle. Among these secretive fossorial organisms, blind snakes of the South American genus Amerotyphlops are considered poorly studied when compared to other taxa. Here, we evaluate the cryptic diversity of Amerotyphlops using phylogenetic and multivariate approaches. We based our phylogenetic analysis on a molecular dataset composed of 12 gene fragments (eight nuclear and four mitochondrial) for 109 species of Typhlopidae. The multivariate analysis was implemented using 36 morphological variables for 377 specimens of Amerotyphlops. Additionally, we contrast our phylogenetic result with the morphological variation found in cranial, external and hemipenial traits. Our phylogenetic results recovered with strong support the following monophyletic groups within Amerotyphlops: (1) a clade formed by A. tasymicris and A. minuisquamus; (2) a clade composed of A. reticulatus; (3) a north-eastern Brazilian clade including A. yonenagae, A. arenensis, A. paucisquamus and A. amoipira; and (4) a clade composed of A. brongersmianus and a complex of cryptic species. Based on these results we describe four new species of Amerotyphlops from north-eastern and south-eastern Brazil, which can be distinguished from the morphologically similar species, A. brongersmianus and A. arenensis.

The phylogenetic position of Ridley's worm lizard reveals the complex biogeographic history of new world insular amphisbaenids

The archipelago of Fernando de Noronha (FN) is located in the Equatorial South Atlantic Ocean, at 375 km off the northeastern coast of Brazil. Its endemic vertebrate land fauna is restricted to only six species, and three main hypotheses have been proposed to explain their presence in the archipelago. These hypotheses suggest FN had alternative biogeographic connections with: 1) the West Indies; 2) the South America mainland; or 3) Africa. Here, we evaluate for the first time the phylogenetic position of Amphisbaena ridleyi within the diversity of Amphisbaenia, and we infer the biogeographic processes that explain its presence in FN and its relationship with amphisbaenids from the West Indies. We analyzed a comprehensive multilocus dataset for Amphisbaenidae using maximum likelihood and time-calibrated Bayesian phylogenetic approaches. Based on our time-calibrated tree, we tested different biogeographic scenarios through historical biogeographic analyses. Our phylogenetic results for the high-level relationships of Amphisbaenia can be parenthetically summarized as (Rhineuridae, (Blanidae, (Bipedidae, (Cadeidae, (Trogonophidae, (Amphisbaenidae))))). Nine highly supported groups of species were recovered among the mainland South American amphisbaenids (SAA), whereas two phylogenetically distant groups of species were inferred for the West Indies: 1) WIC01, an Oligocene lineage present in Cuba and Hispaniola, which is the sister group of all other SAA groups; and 2) WIC02, a Miocene lineage that is restricted to southern Hispaniola and is closely related to Am. ridleyi. We estimated two events of transatlantic dispersal of amphisbaenians from Africa to West Indies: the dispersal of Cadeidae during the Middle Eocene, and the dispersal of the ancestor of Amphisbaena during the transition Eocene/Oligocene. These events were likely affected by the North Equatorial and South Equatorial currents, respectively, which have been flowing westwards since the Paleocene. The ancestral cladogenesis of Amphisbaena during the Late Oligocene is likely related to overwater dispersal events, or alternatively can be associated with the fragmentation of GAARlandia, when WIC01 was isolated in the West Indies, while the remaining groups of Amphisbaena diversified throughout the South American continent. During the Late Miocene, the ancestor of WIC02 dispersed from northern South America to the West Indies, while Am. ridleyi dispersed from the same region to FN. The overwater dispersal of WIC02 was driven by the North Brazilian Current and the dispersal of Am. ridleyi was likely influenced by the periodic shifts in direction and strength estimated for the North Equatorial Counter-Current during the Late Miocene.

Reassessing the systematics of Leptodeira (Serpentes, Dipsadidae) with emphasis in the South American species

Within the diverse subfamily Dipsadinae, Imantodini represents one of the few groups distributed in North, Central and South America. The tribe comprises the genera Leptodeira and Imantodes, from which Leptodeira is the most diverse, including 15 species and 11 subspecies, distributed from southern USA to central Argentina. Taxonomy and affinities among these taxa are poorly resolved, and the phylogenetic relationships among the South American diversity were never properly assessed before. Here, we investigate the phylogenetic relationships and the taxonomic status of Leptodeira spp. based on a comprehensive multilocus dataset with emphasis in the South American radiation. Besides assessing the phylogenetic relationship and species cohesion, we also evaluate the morphological variation among the South American diversity of Leptodeira. Our results support the monophyly of Imatodini and Leptodeira, while indicating that several individuals classified as Leptodeira annulata and L. septentrionalis do not cluster together within their respective species. Moreover, specimens identified as belonging to the subspecies L. a. annulata, L. a. cussiliris, L. s. ornata and L. s. polysticta do not group together suggesting the current classification includes non-natural groups. The analysis of morphological evidence also supports the phylogenetics results, indicating that several clades can be recognized as evolutionary units presenting distinct phenotypes. To equate the taxonomy to our results, we propose a new taxonomic arrangement for Leptodeira in which we are: (1) redefining the composition of L. annulata and L. septentrionalis; (2) elevating five subspecies to species level; (3) revalidating one species; (4) recognizing four species complexes; and (5) indicating the presence of hidden diversity (probably four undescribed species). Finally, we describe a new species (Leptodeira tarairiu sp. nov.) from the open formations of South America (Cerrado and Caatinga), and we provide detailed redescriptions for all South American species of Leptodeira.

Molecular and phenotypic data reveal a new Amazonian species of pit vipers (Serpentes: Viperidae: Bothrops)

Recent genetic studies have found unclear species boundaries and evidence of undescribed diversity in the poorly studied jararacussu species group within Bothrops. In this contribution, we investigate phenotypic and genetic diversity in the Amazonian snake Bothrops brazili to test previous assertions of unrecognised species diversity within this taxon. Our phylogenetic results and inferences of independently evolving lineages based on molecular data recover two divergent clades within B. brazili, one restricted to areas north and another to areas south of the Amazon River. Phylogenetic relationships between these lineages and other species in the jararacussu species group reveal B. brazili to be paraphyletic, with the northern clade inferred as sister to a clade composed of Atlantic Forest taxa (B. jararacussu, B. muriciensis, B. pirajai). External morphology (number of ventral and subcaudal scales) and colouration patterns (lateral trapezoidal marks) consistently separate the two lineages of B. brazili. We therefore recognise and describe the northern lineage as a new species of Bothrops, improving our knowledge of species diversity within a medically important clade of venomous South American snakes.

Rain forest shifts through time and riverine barriers shaped the diversification of South American terrestrial pit vipers (Bothrops jararacussu species group)

Aim To investigate (a) historical biogeographical connections and species interchange among rain forest habitats and (b) the role of riverine barriers on population divergence and speciation in the Neotropical region. Location Amazonia and Atlantic Forest in South America. Taxon Bothrops jararacussu species group (Serpentes: Viperidae). Methods We inferred phylogenetic relationships within Bothrops with an emphasis on the jararacussu species group under a Bayesian framework based on six molecular loci. We also used genetic coalescent simulations and approximate Bayesian computation to infer historical demography within the jararacussu group based on tests of alternative scenarios. Results We found the jararacussu species group to be monophyletic. The Atlantic Forest species B. pirajai and B. muriciensis were inferred nested within this group, closely related to B. jararacussu, confirming that Atlantic Forest species form a clade. The historical demographic analyses support vicariant separation between populations of B. brazili north and south of the Amazon River during the Miocene–Pliocene border, as well as colonization of the Atlantic Forest by a northern Amazonian ancestor in the Pleistocene. Main Conclusion The evolutionary history of the jararacussu species group sheds light on the dynamism of Neotropical rain forests over time, with at least one event of forest expansion leading to faunal interchange between Amazonian and Atlantic forests in the Pleistocene. Moreover, tests of alternative demographic scenarios suggest that the populations of B. brazili from north and south of the Amazon River originated from a vicariant event during the Miocene–Pliocene border, in agreement with the proposed age of establishment of the modern Amazon River drainage. Our results also have taxonomic implications for these medically important venomous snakes, supporting unrecognized diversity at the species level.

Rain forest shifts through time and riverine barriers shaped the diversification of South American terrestrial pit vipers (Bothrops jararacussu species group)

Aim To investigate (a) historical biogeographical connections and species interchange among rain forest habitats and (b) the role of riverine barriers on population divergence and speciation in the Neotropical region. Location Amazonia and Atlantic Forest in South America. Taxon Bothrops jararacussu species group (Serpentes: Viperidae). Methods We inferred phylogenetic relationships within Bothrops with an emphasis on the jararacussu species group under a Bayesian framework based on six molecular loci. We also used genetic coalescent simulations and approximate Bayesian computation to infer historical demography within the jararacussu group based on tests of alternative scenarios. Results We found the jararacussu species group to be monophyletic. The Atlantic Forest species B. pirajai and B. muriciensis were inferred nested within this group, closely related to B. jararacussu, confirming that Atlantic Forest species form a clade. The historical demographic analyses support vicariant separation between populations of B. brazili north and south of the Amazon River during the Miocene–Pliocene border, as well as colonization of the Atlantic Forest by a northern Amazonian ancestor in the Pleistocene. Main Conclusion The evolutionary history of the jararacussu species group sheds light on the dynamism of Neotropical rain forests over time, with at least one event of forest expansion leading to faunal interchange between Amazonian and Atlantic forests in the Pleistocene. Moreover, tests of alternative demographic scenarios suggest that the populations of B. brazili from north and south of the Amazon River originated from a vicariant event during the Miocene–Pliocene border, in agreement with the proposed age of establishment of the modern Amazon River drainage. Our results also have taxonomic implications for these medically important venomous snakes, supporting unrecognized diversity at the species level.

Correction: Large-scale molecular phylogeny, morphology, divergence-time estimation, and the fossil record of advanced caenophidian snakes (Squamata: Serpentes).

[This corrects the article DOI: 10.1371/journal.pone.0216148.].

Large-scale molecular phylogeny, morphology, divergence-time estimation, and the fossil record of advanced caenophidian snakes (Squamata: Serpentes).

Caenophidian snakes include the file snake genus Acrochordus and advanced colubroidean snakes that radiated mainly during the Neogene. Although caenophidian snakes are a well-supported clade, their inferred affinities, based either on molecular or morphological data, remain poorly known or controversial. Here, we provide an expanded molecular phylogenetic analysis of Caenophidia and use three non-parametric measures of support-Shimodaira-Hasegawa-Like test (SHL), Felsentein (FBP) and transfer (TBE) bootstrap measures-to evaluate the robustness of each clade in the molecular tree. That very different alternative support values are common suggests that results based on only one support value should be viewed with caution. Using a scheme to combine support values, we find 20.9% of the 1265 clades comprising the inferred caenophidian tree are unambiguously supported by both SHL and FBP values, while almost 37% are unsupported or ambiguously supported, revealing the substantial extent of phylogenetic problems within Caenophidia. Combined FBP/TBE support values show similar results, while SHL/TBE result in slightly higher combined values. We consider key morphological attributes of colubroidean cranial, vertebral and hemipenial anatomy and provide additional morphological evidence supporting the clades Colubroides, Colubriformes, and Endoglyptodonta. We review and revise the relevant caenophidian fossil record and provide a time-calibrated tree derived from our molecular data to discuss the main cladogenetic events that resulted in present-day patterns of caenophidian diversification. Our results suggest that all extant families of Colubroidea and Elapoidea composing the present-day endoglyptodont fauna originated rapidly within the early Oligocene-between approximately 33 and 28 Mya-following the major terrestrial faunal turnover known as the "Grande Coupure" and associated with the overall climate shift at the Eocene-Oligocene boundary. Our results further suggest that the caenophidian radiation originated within the Caenozoic, with the divergence between Colubroides and Acrochordidae occurring in the early Eocene, at ~ 56 Mya.

A new species of the Amphisbaena (Squamata, Amphisbaenidae) from the Brazilian Cerrado with a key for the two-pored species.

Here, we describe a new species of Amphisbaena with two precloacal pores from open Cerrado areas of the municipality of Arenópolis, in the Brazilian state of Goiás. The new species differs from other South American amphisbaenids by the folllowing combination of characters: (1) snout rounded in dorsal view and slightly convex in lateral view; (2) two precloacal pores; (3) 161-176 dorsal half-annuli; and (4) 12-15 tail annuli. Our molecular phylogenetic analysis retrieved a monophyletic Amphisbaena silvestrii group, with A. silvestrii positioned as the sister-group of a clade formed by Amphisbaena anaemariae and the new species described herein. Members of the A. silvestrii group including A. neglecta and A. crisae not added in our phylogenetic analysis are characterized by a relatively small body, two precloacal pores, body coloration with dark and light areas, and lack of specializations on the cephalic or caudal shields. We present a key for two-pored species of Amphisbaena.

Amphisbaena mensae Catro-Mello, 2000 is a synonym of Amphisbaena talisiae Vanzolini, 1995 (Squamata: Amphisbaenia: Amphisbaenidae).

Amphisbaena talisiae and A. mensae, two worm lizard species endemic to the Cerrado ecoregion, in central Brazil, are considered synonyms based on morphological characters. With the proposed synonymy, the name A. talisiae has priority of use over A. mensae. Amphisbaena talisiae can be distinguished from its congeners by a series of morphological characters, including a round head, three supralabial and three infralabial scales, postmalar row absent, four precloacal pores without a median hiatus, 205-234 body annuli, 17-29 caudal annuli, 10-14 dorsal and 14-18 ventral segments in a midbody annulus. The species is recorded from southeastern Mato Grosso, eastern Minas Gerais and central Tocantins states in central Brazil, and its conservation status should be changed from Data Deficient to Least Concern.

Large-scale molecular phylogeny, morphology, divergence-time estimation, and the fossil record of advanced caenophidian snakes (Squamata: Serpentes)

Caenophidian snakes include the file snake genus Acrochordus and advanced colubroidean snakes that radiated mainly during the Neogene. Although caenophidian snakes are a well-supported clade, their inferred affinities, based either on molecular or morphological data, remain poorly known or controversial. Here, we provide an expanded molecular phylogenetic analysis of Caenophidia and use three non-parametric measures of support–Shimodaira-Hasegawa-Like test (SHL), Felsentein (FBP) and transfer (TBE) bootstrap measures–to evaluate the robustness of each clade in the molecular tree. That very different alternative support values are common suggests that results based on only one support value should be viewed with caution. Using a scheme to combine support values, we find 20.9% of the 1265 clades comprising the inferred caenophidian tree are unambiguously supported by both SHL and FBP values, while almost 37% are unsupported or ambiguously supported, revealing the substantial extent of phylogenetic problems within Caenophidia. Combined FBP/TBE support values show similar results, while SHL/TBE result in slightly higher combined values. We consider key morphological attributes of colubroidean cranial, vertebral and hemipenial anatomy and provide additional morphological evidence supporting the clades Colubroides, Colubriformes, and Endoglyptodonta. We review and revise the relevant caenophidian fossil record and provide a time-calibrated tree derived from our molecular data to discuss the main cladogenetic events that resulted in present-day patterns of caenophidian diversification. Our results suggest that all extant families of Colubroidea and Elapoidea composing the present-day endoglyptodont fauna originated rapidly within the early Oligocene–between approximately 33 and 28 Mya–following the major terrestrial faunal turnover known as the "Grande Coupure" and associated with the overall climate shift at the Eocene-Oligocene boundary. Our results further suggest that the caenophidian radiation originated within the Caenozoic, with the divergence between Colubroides and Acrochordidae occurring in the early Eocene, at ~ 56 Mya.

Molecular phylogeny and hemipenial diversity of South American species of Amerotyphlops (Typhlopidae, Scolecophidia)

Typhlopidae is the most diverse family of Scolecophidia, with 269 species. Amerotyphlops was recently erected within subfamily Typhlopinae and comprises fifteen species distributed from Mexico to Argentina and the southern Lesser Antilles. Despite recent advances, affinities among typhlopines remain poorly explored, and the phylogenetic relationships and morphology of the South American (SA) species were never accessed before. Here, we performed a phylogenetic analysis including 106 species of Typhlopidae and ten genes. Our dataset represents the most comprehensive for SA species, containing seven of eight recognized species. Corroborating previous studies, we recovered the main groups of Typhlopoidea, and for typhlopines, we recovered with strong support two clades: (a) the Greater Antilles radiation, and the (b) Lesser Antilles and SA radiation. Within the SA radiation, we recovered four main lineages: (a) a clade formed by A. tasymicris and A. minuisquamus; (b) a clade composed by A. reticulatus as the sister group of all other SA species; (c) a clade composed by A. brongersmianus as the sister group of a clade comprising all Northeast Brazilian Species (NBS); and (d) a clade of the NBS, including A. yonenagae, A. arenensis, A. paucisquamus, and A. amoipira. We supplemented our phylogenetic result with the description of hemipenial morphology for seven SA species and comment their relevance to the systematics of Typhlopinae. Hemipenes of SA Amerotyphlops follow the general pattern in scolecophidians (single organ with undivided sulcus). Only A. reticulatus and A. minuisquamus have organs with calcified spines. According to our results, hemipenial ornamentation have shown highly informative and could represent a potential source of systematic and taxonomic characters in that group. We also present an extensive review of the geographical distribution for all SA species. Our study represents the first integrative analysis of a poorly known evolutionary radiation of one of the most widespread SA fossorial snakes.

Large-scale molecular phylogeny, morphology, divergence-time estimation, and the fossil record of advanced caenophidian snakes (Squamata: Serpentes)

Caenophidian snakes include the file snake genus Acrochordus and advanced colubroidean snakes that radiated mainly during the Neogene. Although caenophidian snakes are a well-supported clade, their inferred affinities, based either on molecular or morphological data, remain poorly known or controversial. Here, we provide an expanded molecular phylogenetic analysis of Caenophidia and use three non-parametric measures of support–Shimodaira-Hasegawa-Like test (SHL), Felsentein (FBP) and transfer (TBE) bootstrap measures–to evaluate the robustness of each clade in the molecular tree. That very different alternative support values are common suggests that results based on only one support value should be viewed with caution. Using a scheme to combine support values, we find 20.9% of the 1265 clades comprising the inferred caenophidian tree are unambiguously supported by both SHL and FBP values, while almost 37% are unsupported or ambiguously supported, revealing the substantial extent of phylogenetic problems within Caenophidia. Combined FBP/TBE support values show similar results, while SHL/TBE result in slightly higher combined values. We consider key morphological attributes of colubroidean cranial, vertebral and hemipenial anatomy and provide additional morphological evidence supporting the clades Colubroides, Colubriformes, and Endoglyptodonta. We review and revise the relevant caenophidian fossil record and provide a time-calibrated tree derived from our molecular data to discuss the main cladogenetic events that resulted in present-day patterns of caenophidian diversification. Our results suggest that all extant families of Colubroidea and Elapoidea composing the present-day endoglyptodont fauna originated rapidly within the early Oligocene–between approximately 33 and 28 Mya–following the major terrestrial faunal turnover known as the "Grande Coupure" and associated with the overall climate shift at the Eocene-Oligocene boundary. Our results further suggest that the caenophidian radiation originated within the Caenozoic, with the divergence between Colubroides and Acrochordidae occurring in the early Eocene, at ~ 56 Mya.

Molecular phylogeny and hemipenial diversity of South American species of Amerotyphlops (Typhlopidae, Scolecophidia)

Typhlopidae is the most diverse family of Scolecophidia, with 269 species. Amerotyphlops was recently erected within subfamily Typhlopinae and comprises fifteen species distributed from Mexico to Argentina and the southern Lesser Antilles. Despite recent advances, affinities among typhlopines remain poorly explored, and the phylogenetic relationships and morphology of the South American (SA) species were never accessed before. Here, we performed a phylogenetic analysis including 106 species of Typhlopidae and ten genes. Our dataset represents the most comprehensive for SA species, containing seven of eight recognized species. Corroborating previous studies, we recovered the main groups of Typhlopoidea, and for typhlopines, we recovered with strong support two clades: (a) the Greater Antilles radiation, and the (b) Lesser Antilles and SA radiation. Within the SA radiation, we recovered four main lineages: (a) a clade formed by A. tasymicris and A. minuisquamus; (b) a clade composed by A. reticulatus as the sister group of all other SA species; (c) a clade composed by A. brongersmianus as the sister group of a clade comprising all Northeast Brazilian Species (NBS); and (d) a clade of the NBS, including A. yonenagae, A. arenensis, A. paucisquamus, and A. amoipira. We supplemented our phylogenetic result with the description of hemipenial morphology for seven SA species and comment their relevance to the systematics of Typhlopinae. Hemipenes of SA Amerotyphlops follow the general pattern in scolecophidians (single organ with undivided sulcus). Only A. reticulatus and A. minuisquamus have organs with calcified spines. According to our results, hemipenial ornamentation have shown highly informative and could represent a potential source of systematic and taxonomic characters in that group. We also present an extensive review of the geographical distribution for all SA species. Our study represents the first integrative analysis of a poorly known evolutionary radiation of one of the most widespread SA fossorial snakes.

A new four-pored Amphisbaena Linnaeus, 1758 (Amphisbaenia, Amphisbaenidae) from Brazilian Amazon.

A new species of Amphisbaena is described from the Brazilian Amazon, within the area impacted by the Teles Pires hydroelectric power plant, Jacareacanga municipality, State of Pará. Amphisbaena hoogmoedi sp. nov. can be diagnosed from its congeners by the following combination of characters: snout convex in profile view, sligthly compressed not keeled; pectoral scales arranged in regular annuli; conspicuous autotomic site between 7th-8th caudal annuli; 247-252 dorsal half-annuli; 27 caudal annuli; tail length 9.5-10.4% of snoutvent length; four precloacal pores arranged in sequence; three supralabials; a rounded tail; 22-24 dorsal segments in midbody annulus; postmalar row absent; head length 2.1-2.9% of snout-vent length; prefrontals length 46.6-49.5% of head length; prefrontals suture length 38-44.6% of head length; small malar length 10.6-13.4% of ventral length of head ; second infralabial length 33.8-38.5% of head length; ventral length of head 2.7-2.9% of snout-vent length; mouth length 80.2-81.8% of head length; third infralabial length 16.4-19.6% of head length; snout length 62.5-78.6% of head length; ocular length 23.4-26.2% of head length; mental length 23.2-25.4% of ventral length of head; postmental length 27.2-31.3% of ventral length of head; frontals suture length 23.4-32.3% of head length; postocular width 25-31.9% of maximun width of head; first supralabial length 24.9-30.6% of head length; second supralabial length 27.7-30% of head length and second supralabial height 26.9-28.8% of maximun head height. The hemipenis is bilobed, capitate and with lateral lamellae on the lobes; with a centrally-positioned spermatic groove, bifurcated at the base of the lobes, and with each branch extending to the tip of organ.

Origin and hidden diversity within the poorly known Galápagos snake radiation (Serpentes: Dipsadidae)

Galápagos snakes are among the least studied terrestrial vertebrates of the Archipelago. Here, we provide a phylogenetic analysis and a time calibrated tree for the group, based on a sampling of the major populations known to occur in the Archipelago. Our study revealed the presence of two previously unknown species from Santiago and Rábida Islands, and one from Tortuga, Isabela, and Fernandina. We also recognize six additional species of Pseudalsophis in the Galápagos Archipelago (Pseudalsophis biserialis from San Cristobal, Floreana and adjacent islets; Pseudalsophis hoodensis from Española and adjacent islets; Pseudalsophis dorsalis from Santa Cruz, Baltra, Santa Fé, and adjacent islets; Pseudalsophis occidentalis from Fernandina, Isabela, and Tortuga; Pseudalsophis slevini from Pinzon, and Pseudalsophis steindachneri from Baltra, Santa Cruz and adjacent islets). Our time calibrated tree suggests that the genus Pseudalsophis colonized the Galápagos Archipelago through a single event of oceanic dispersion from the coast of South America that occurred at approximately between 6.9?Ma and 4.4?Ma, near the Miocene/Pliocene boundary.

Origin and hidden diversity within the poorly known Galápagos snake radiation (Serpentes: Dipsadidae)

Galápagos snakes are among the least studied terrestrial vertebrates of the Archipelago. Here, we provide a phylogenetic analysis and a time calibrated tree for the group, based on a sampling of the major populations known to occur in the Archipelago. Our study revealed the presence of two previously unknown species from Santiago and Rábida Islands, and one from Tortuga, Isabela, and Fernandina. We also recognize six additional species of Pseudalsophis in the Galápagos Archipelago (Pseudalsophis biserialis from San Cristobal, Floreana and adjacent islets; Pseudalsophis hoodensis from Española and adjacent islets; Pseudalsophis dorsalis from Santa Cruz, Baltra, Santa Fé, and adjacent islets; Pseudalsophis occidentalis from Fernandina, Isabela, and Tortuga; Pseudalsophis slevini from Pinzon, and Pseudalsophis steindachneri from Baltra, Santa Cruz and adjacent islets). Our time calibrated tree suggests that the genus Pseudalsophis colonized the Galápagos Archipelago through a single event of oceanic dispersion from the coast of South America that occurred at approximately between 6.9?Ma and 4.4?Ma, near the Miocene/Pliocene boundary.

A new species of Amerotyphlops from Northeastern Brazil, with comments on distribution of related species.

We describe a new species of Amerotyphlops from an upland forest enclave in the state of Paraíba, Northeastern Brazil. The new species is distinguished from the other seven South American species of Amerotyphlops by the combination of the following characters: nasal suture incomplete; rostral scale oval and yellowish cream with some dark brown spots; four supralabial scales; three infralabial scales; rows of scales around the body 18/18/18; middorsal scales from 204 to 225; dorsum with twelve to thirteen rows of scales dark brown and belly with four to five rows of scales immaculate yellowish cream; caudal spine dark brown; subcaudal scales 8-10 in female and 11-13 in males; maximum total length 233 mm. The new species is morphologically similar to A. amoipira and A. paucisquamus, sharing 18/18/18 rows of scales around the body and a small overlap of counts of middorsal scales.