A new dwarf gecko of the genus Pseudogonatodes (Squamata: Sphaerodactylidae) from the eastern slope of the Andean Cordillera de Mrida in northern South America.

Pseudogonatodes is a poorly known genus of small bodied, diurnal, ground-dwelling geckos widely distributed in northern South America. No additional species have been described in over two decades. Herein we describe a new species from the eastern slope of the Cordillera de Mrida in the Venezuelan Andes. The new species is readily diagnosable morphologically from the other seven recognized species of Pseudogonatodes by having a single postnasal scalea putative autapomorphy. Furthermore, it is characterized by a unique combination of phenotypic characters that includes granular dorsal scales, three large postrostrals, five to six loreals, mental U-shaped, four to six postmentals, 2629 ventrals between anterior levels of fore- and hind limbs, third lamella under fourth toe not distinctly enlarged, and subcaudal pattern 11. In addition to characterizing the external morphology, we present a description of the skull, based on 3D digital models reconstructed from high resolution computed microtomography scans. The discovery of this new species highlights the still underestimated diversity of this group of Neotropical dwarf geckos and underscores the need for further studies on its systematics and taxonomy.

Unveiling the evolutionary relationships and the high cryptic diversity in Andean rainfrogs (Craugastoridae: Pristimantis myersi group).

Background: Pristimantis is the most diverse genus of terrestrial frogs. Historically, it has been divided into several phenetic groups in order to facilitate species identification. However, in light of phylogenetic analysis, many of these groups have been shown to be non-monophyletic, denoting a high degree of morphological convergence and limited number of diagnostic traits. In this study, we focus on the Pristimantis myersi group, an assemblage of small rainfrogs distributed throughout the Andes of Ecuador and Colombia, whose external morphology is highly conserved, and its species diversity and evolutionary relationships largely unknown. Methods: We inferred a new phylogenetic hypothesis for the frog genus Pristimantis, including all available sequences of the mtDNA 16S rRNA, as well as new DNA sequences from 175 specimens. Our sampling included 19 of the 24 species currently recognized as part of the Pristimantis myersi group. Results: Our new evolutionary hypothesis recovered the P. myersi group as non-monophyletic and composed of 16 species. Therefore, we exclude P. albujai, P. bicantus, P. sambalan, and P. nelsongalloi in order to preserve the monophyly of the group. We discovered at least eight candidate species, most of them hidden under the names of P. leoni, P. hectus, P. festae, P. gladiator, and P. ocreatus. Discussion: Our results reveal the occurrence of a high level of cryptic diversity to the species level within the P. myersi group and highlight the need to redefine some of its species and reassess their conservation status. We suggest that the conservation status of six species within the group need to be re-evaluated because they exhibit smaller distributions than previously thought; these species are: P. festae, P. gladiator, P. hectus, P. leoni, P. ocreatus, and P. pyrrhomerus. Finally, given that the Pristimantis myersi group, as defined in this work, is monophyletic and morphologically diagnosable, and that Trachyphrynus is an available name for the clade containing P. myersi, we implement Trachyphrynus as a formal subgenus name for the Pristimantis myersi group.

The performance of protected-area expansions in representing tropical Andean species: past trends and climate change prospects.

Protected area (PA) extent has increased significantly over the last 150 years globally, but it is yet unclear whether progress in expanding coverage has been accompanied by improved performance in ecological representation. Here, we explore temporal trends in the performance of PA networks in representing > 16,000 vertebrate and plant species in tropical Andean countries based on species bioclimatic niche modelling. We use a randomization analysis to assess whether representation gains over time (1937-2015) are the expected consequence of increasing the overall area of the network or the result of better designed networks. We also explore the impact of climate change on protected-area representation based on projected species distributions in 2070. We found that PAs added in the last three to four decades were better at representing species diversity than random additions overall. Threatened species, amphibians and reptiles are the exception. Species representation is projected to decrease across PAs under climate change, although PA expansions over the last decade (2006-2015) better represented species' future bioclimatic niches than did sites selected at random for most evaluated groups. These findings indicate an unbalanced representation across taxa, and raises concern over under-represented groups, including threatened species, and species' representation under climate change scenarios. However, they also suggest that decisions related to locating protected areas have become more strategic in recent decades and illustrate that indicators tracking representativeness of networks are crucial in PA monitoring frameworks.

Synthesis of geological data and comparative phylogeography of lowland tetrapods suggests recent dispersal through lowland portals crossing the Eastern Andean Cordillera.

Vicariance is the simplest explanation for divergence between sister lineages separated by a potential barrier, and the northern Andes would seem to provide an ideal example of a vicariant driver of divergence. We evaluated the potential role of the uplift of the Eastern Cordillera (EC) of the Colombian Andes and the Mérida Andes (MA) of Venezuela as drivers of vicariance between lowland populations co-distributed on both flanks. We synthesized published geological data and provided a new reconstruction showing that the EC-MA grew from north to south, reaching significant heights and separating drainages and changing sediment composition by 38-33 million years ago (Ma). A few lowland passes across the EC-MA may have reached their current heights (~1,900 m a.s.l.) at 3-5 Ma. We created a comparative phylogeographic data set for 37 lineages of lowland tetrapods. Based on molecular phylogenetic analyses, most divergences between sister populations or species across the EC-MA occurred during Pliocene and the Quaternary and a few during the latest Miocene, and coalescent simulations rejected synchronous divergence for most groups. Divergence times were on average slightly but significantly more recent in homeotherms relative to poikilotherms. Because divergence ages are mostly too recent relative to the geological history and too asynchronous relative to each other, divergence across the northern Andes may be better explained by organism-environment interactions concomitant with climate oscillations during the Pleistocene, and/or dispersal across portals through the Andes.

The taxonomic status of Cryptobatrachus frogs (Anura: Hemiphractidae) from the Serrana del Perij.

The Neotropical frog genus Cryptobatrachus includes six currently recognized species distributed throughout the northern Andes in Colombia and Venezuela. Cryptobatrachus conditus, C. pedroruizi, and C. remotus were described from the Colombian and Venezuelan slopes of the Serrana del Perij. Due to the great morphological similarity among these species, we re-assess their taxonomic status based on morphological and molecular data from types, topotypes, and specimens from localities referred to in the original descriptions of these species. Morphometric comparisons showed that all these species are virtually indistinguishable, although some subtle qualitative differences in morphological traits distinguish C. conditus from the other species. Phylogenetic analyses of DNA sequences found that C. remotus and C. pedroruizi are not reciprocally monophyletic and exhibit low genetic divergence ( 1 %). Therefore, C. remotus should be considered a junior synonym of C. pedroruizi. This work stresses the importance of comprehensive taxon sampling in poorly explored areas, especially between neighbouring countries.

A new species of nurse-frog (Aromobatidae, Allobates) from the Amazonian forest of Loreto, Peru.

We describe a new species of nurse-frog (Aromobatidae, Allobates) from the Amazonian forest of Loreto, Peru using morphological, acoustic and genetic data. Our phylogenetic analysis placed Allobates sieggreenae sp. nov. as the sister species of A. trilineatus, the most similar-looking species and with which it was previously confused. However, the new species has a brown dorsum, solid dark brown lateral dark stripe not fading towards groin, adult males with few and sparse melanophores over a cream background on chin, chest, and belly, dark transverse bars absent on thighs, and an advertisement call formed by a trill of single notes (in A. trilineatus dorsum dark brown, blackish brown lateral dark stripe, paler from mid-body to groin, adult males with a dark background color on chin, chest, and belly due to a dense layer of melanophores, dark transverse bar present on dorsal surface of thighs, and trills of paired notes). Allobates sieggreenae is known from two localities of Amazonian white-sand forest ecosystems east of the Ucayali River.

Total evidence and sensitivity phylogenetic analyses of egg-brooding frogs (Anura: Hemiphractidae).

We study the phylogenetic relationships of egg-brooding frogs, a group of 118 neotropical species, unique among anurans by having embryos with large bell-shaped gills and females carrying their eggs on the dorsum, exposed or inside a pouch. We assembled a total evidence dataset of published and newly generated data containing 51 phenotypic characters and DNA sequences of 20 loci for 143 hemiphractids and 127 outgroup terminals. We performed six analytical strategies combining different optimality criteria (parsimony and maximum likelihood), alignment methods (tree- and similarity-alignment), and three different indel coding schemes (fifth character state, unknown nucleotide, and presence/absence characters matrix). Furthermore, we analyzed a subset of the total evidence dataset to evaluate the impact of phenotypic characters on hemiphractid phylogenetic relationships. Our main results include: (i) monophyly of Hemiphractidae and its six genera for all our analyses, novel relationships among hemiphractid genera, and non-monophyly of Hemiphractinae according to our preferred phylogenetic hypothesis; (ii) non-monophyly of current supraspecific taxonomies of Gastrotheca, an updated taxonomy is provided; (iii) previous differences among studies were mainly caused by differences in analytical factors, not by differences in character/taxon sampling; (iv) optimality criteria, alignment method, and indel coding caused differences among optimal topologies, in that order of degree; (v) in most cases, parsimony analyses are more sensitive to the addition of phenotypic data than maximum likelihood analyses; (vi) adding phenotypic data resulted in an increase of shared clades for most analyses.

Integrative taxonomy reveals a new but common Neotropical treefrog, hidden under the name Boana xerophylla.

Boana xerophylla is a common treefrog widely distributed in northern Brazil, Colombia, French Guiana, Guyana, Panama, Suriname, Trinidad and Tobago, and Venezuela. A recent study found molecular, acoustic, and morphometric differences between the populations located on opposite sides of the Orinoco River. Here, we carry out an updated molecular phylogenetic analysis, including new samples from all the countries along the distribution area, and analyzed additional call recordings from Brazil, Colombia, French Guiana, Panama, Trinidad and Tobago, and Venezuela. Our phylogenetic inference reveals three geographically restricted lineages: one in the eastern Guiana Shield (corresponding to B. xerophylla sensu stricto), another in the western Guiana Shield, and a third one north of the Orinoco River. Morphological and acoustic data agree with the differentiation between the populations north of the Orinoco River and the eastern Guiana Shield despite the low genetic p-distances observed (16S rRNA: 0.72.2 %). We argue that the populations north of the Orinoco River correspond to a new species, sister of B. xerophylla. We name and describe Boana platanera sp. nov. from the southern versant of the Cordillera de Mérida (08º48'26'' N, 70º30'46'' W, WGS 84; 947 m asl), Venezuela, and refer all the populations north of the Orinoco River currently identified as B. xerophylla to this species. The new species can be readily diagnosed from B. xerophylla (characters of the latter in parentheses) by a pale orange-yellow or light brown dorsal coloration (dark brown to green), palpebral membrane with dark pigments (pigments absent); pericloacal region dark brown (cream), advertisement call with shorter first note length than B. xerophylla. This study represents an empirical example regarding false negatives behind genetic thresholds for species discovery, appraising the use of integrative taxonomic approaches.

Discovery of an additional piece of the large gymnophthalmid puzzle: a new genus and species of stream spiny lizard (Squamata: Gymnophthalmidae: Cercosaurinae) from the western Guiana Shield in Venezuela.

Gymnophthalmids are a highly diverse group of Neotropical lizards and its species richness is still in process of discovery. The incorporation of molecular evidence and a noticeable increase in taxon and geographic sampling in systematic studies has led to the description of numerous new genera and species of gymnophthalmids (particularly in Cercosaurinae) in recent years. Herein we describe a new genus and species of cercosaurine lizard with crocodile-like morphology, from the Venezuelan Guiana Shield on the basis of molecular phylogenetic and morphological evidence. Kataphraktosaurus gen. nov. can be readily distinguished from all other genera of Cercosaurinae by a unique combination of morphological characters that includes heterogeneous dorsal scalation with enlarged and strongly keeled scales forming two paravertebral rows, ventral and subcaudal scales imbricated and strongly keeled, large and symmetrical cephalic scales, absence of postmental scale, palpebral disc translucent and divided, tail slightly compressed, all digits clawed, and only six femoral pores (three at each hindlimb) inserted in a clump of small scales. This genus is described as monotypic and only contains Kataphraktosaurus ungerhamiltoni sp. nov., which is known from one specimen and diagnosed by the same set of aforementioned characters. The secretive habits of this species and the remoteness of the locality may explain its singleton situation. Following the International Union for Conservation of Nature's criteria, we categorized the new species as Data Deficient.

Tachiramantis lassoalcalai (Barrio-Amorós, Rojas-Runjaic amp; Barros, 2010) (Anura, Craugastoridae): a new combination revealed by molecular evidence, with a description of its advertisement call.

Pristimantis lassoalcalai Barrio-Amorós, Rojas-Runjaic Barros, 2010 is a poorly known terrarana, endemic to the eastern slope of Sierra de Perijá in Venezuela. Although a close relationship of this species with Tachiramantis has been suspected based on its overall morphological similarity, this relationship had not been tested so far. On the basis of molecular data (two fragments of the 12S and 16S mtDNA genes) obtained from the type series, we reconstruct its evolutionary relationships and establish its phylogenetic position as a member of Tachiramantis. Based on this phylogenetic hypothesis, we transfer Pristimantis lassoalcalai to Tachiramantis as Tachiramantis lassoalcalai comb. nov. In addition, we describe its advertisement call. This is the fourth known species of the genus and the second to which its vocalization is described.

Resolving the taxonomic puzzle of Boana cinerascens (Spix, 1824), with resurrection of Hyla granosa gracilis Melin, 1941 (Anura: Hylidae).

Expeditions to unexplored or little explored places are important for discovering new species and also for collecting new samples (including specimens and tissues for DNA sequencing ) that may help resolve a plethora of taxonomic problems. In the 19th century, several naturalists explored a number of localities in Amazonia, describing species for which type material was deposited, mostly, in European museums of natural history. Some of these types were lost or destroyed in World War II and recent expeditions have focused on sampling new material from the type localities. material from Boana cinerascens, which allowed us to infer phylogenetic relationships of the Boana punctata group (i.e., green Boana), based on DNA sequence data, and to revaluate the status of B. cinerascens and its synonyms. We designate, redescribe and illustrate a neotype for B. cinerascens, which was described by Spix in 1824, from the Municipality of Tefé, State of Amazonas, Brazil. We revalidate, redescribe, and illustrate Hyla granosa gracilis Melin, 1941(= Boana gracilis). Corroborating previous studies, the green Boana were not recovered as a monophyletic group. Boana cinerascens is sister of B. gracilis plus a clade containing B. atlantica + B. punctata (both species not recovered as monophyletic).

Unveiling species diversity in collared frogs through morphological and bioacoustic evidence: a new Mannophryne (Amphibia, Aromobatidae) from Sierra de Aroa, northwestern Venezuela, and an amended definition and call description of M. herminae (Boettger, 1893).

Species diversity in collared frogs of the genus Mannophryne is presumed to be underestimated due to the paucity of external morphology characters, but combining morphology with bioacoustics and other lines of evidence has shown to be useful in delimiting species of this group. Herein we describe a new species of Mannophryne from Sierra de Aroa in northwestern Venezuela. The new species is morphologically similar to M. herminae but is readily recognized by its strikingly different advertisement call. It also can be distinguished from all its congeners by the unique combination of its small body size, general color pattern, basal toe webbing, and advertisement call consisting of long trills of single tonal notes emitted at a rate of 2-3 notes/s. Additionally, to facilitate future diagnosis of undescribed species related to M. herminae, we amend the definition of the latter, describe in detail its advertisement call, and redefine its known distribution range. The new species increases the number of described species of Mannophryne to 20.

A new species of Hyloscirtus (Anura, Hylidae) from the Colombian and Venezuelan slopes of Sierra de Perijá, and the phylogenetic position of Hyloscirtus jahni (Rivero, 1961).

A new species of Hyloscirtus, belonging to the H. bogotensis species Group, is described from the Venezuelan and Colombian slopes of the Sierra de Perijá. The new species can be readily distinguished from its congeners by the combination of the following characters: mental gland present, disc-shaped and small; ulnar, outer, and inner tarsal folds present; calcar tubercle absent; whitish stripes on external border of upper eyelids and supratympanic folds, longitudinally on the mid-dorsum, on supracloacal fold, outer ulnar folds, inner and outer tarsal folds, and also on dorsal internal surface of shanks. We estimate phylogenetic relationships based on mtDNA (spanning fragments of 12S rRNA, tRNA-Val and 16S rRNA), of all Hyloscirtus species available in Genbank, as well as the new species described herein, H. callipeza, H. jahni, and H. platydactylus, all of which have not been previously sequenced. Our molecular data support the hypothesis of the new species as sister species of H. callipeza and indicates that H. jahni does not belong to the H. bogotensis species Group, but rather is sister species of all other Hyloscirtus (sensu Faivovich et al. 2005). Based on this last result we propose a new species group for H. jahni and the synonymy of Colomascirtus in Hyloscirtus. We also provide the first description of the advertisement call of H. callipeza. With the new species described herein, the number of Hyloscirtus species increases to 37.

A new species of nurse-frog (Aromobatidae, Allobates) from the Juami River basin, northwestern Brazilian Amazonia.

We describe a new species of nurse-frog (Aromobatidae, Allobates) from northwestern Brazilian Amazon. Allobates juami sp. nov. is distinguished from similar congeneric species by the combination of the following characters: body-size range (snout-to-vent length 17.5-18.5 mm), lack of dark pigments on ventral surfaces of male specimens, dorsal color pattern (predominantly solid dark brown, but conspicuously light brown over snout and urostyle regions), presence of conspicuous pale dorsolateral and ventrolateral stripes, and presence of a diffuse pale paracloacal mark. The advertisement call of the new species lasts 2.5-5.1 s, contains 60-73 short notes (trills), and is emitted at an average rate of 13 notes per second within trills. Duration of silent intervals between notes ranges between 0.020-0.050 s, and the peak frequency of notes ranges from 4.59 to 5.47 kHz. The new species is currently known only from the type locality at Estação Ecológica Juami-Japurá (1.96455° S, 67.93579° W; ~ 87 m a.s.l.).

A new species of Phyzelaphryne Heyer, 1977 (Anura: Eleutherodactylidae) from the Japurá River basin, with a discussion of the diversity and distribution of the genus.

We describe and name the second species of Phyzelaphryne (Brachycephaloidea, Eleutherodactylidae), from northwestern Brazilian Amazonia. Phyzelaphryne nimio sp. nov. is distinguished from its only congener, Phyzelaphryne miriamae, by its smaller body size and the anatomy of the carpal and metacarpal regions, with relatively larger (sometimes fused) supernumerary carpal and metacarpal tubercles. Molecular phylogenetic analyses based on fragments of the mitochondrial genes 16S rRNA and COI suggest that the currently known distribution of the species is restricted to its type locality and other areas within Estação Ecológica Juami-Japurá, state of Amazonas, Brazil. Based on molecular, morphological and bioacoustic evidence, we assigned other specimens recently collected in Parque Nacional do Jaú, state of Amazonas, Brazil, to P. miriamae, extending the species' known geographic distribution north of the Amazon River.

Phylogenetic systematics of egg-brooding frogs (Anura: Hemiphractidae) and the evolution of direct development.

Egg-brooding frogs (Hemiphractidae) are a group of 105 currently recognized Neotropical species, with a remarkable diversity of developmental modes, from direct development to free-living and exotrophic tadpoles. Females carry their eggs on the back and embryos have unique bell-shaped gills. We inferred the evolutionary relationships of these frogs and used the resulting phylogeny to review their taxonomy and test hypotheses on the evolution of developmental modes and bell-shaped gills. Our inferences relied on a total evidence parsimony analysis of DNA sequences of up to 20 mitochondrial and nuclear genes (analyzed under tree-alignment), and 51 phenotypic characters sampled for 83% of currently valid hemiphractid species. Our analyses rendered a well-resolved phylogeny, with both Hemiphractidae (sister of Athesphatanura) and its six recognized genera being monophyletic. We also inferred novel intergeneric relationships [((Cryptobatrachus, Flectonotus), (Stefania, (Fritziana, (Hemiphractus, Gastrotheca))))], the non-monophyly of all species groups previously proposed within Gastrotheca and Stefania, and the existence of several putative new species within Fritziana and Hemiphractus. Contrary to previous hypotheses, our results support the most recent common ancestor of hemiphractids as a direct-developer. Free-living aquatic tadpoles apparently evolved from direct-developing ancestors three to eight times. Embryos of the sister taxa Cryptobatrachus and Flectonotus share a pair of single gills derived from branchial arch I, while embryos of the clade including the other four genera have two pairs of gills derived from branchial arches I and II respectively. Furthermore, in Gastrotheca the fusion of the two pairs of gills is a putative synapomorphy. We propose a revised taxonomy concordant with our optimal topologies.

High levels of diversity uncovered in a widespread nominal taxon: continental phylogeography of the neotropical tree frog Dendropsophus minutus.

Species distributed across vast continental areas and across major biomes provide unique model systems for studies of biotic diversification, yet also constitute daunting financial, logistic and political challenges for data collection across such regions. The tree frog Dendropsophus minutus (Anura: Hylidae) is a nominal species, continentally distributed in South America, that may represent a complex of multiple species, each with a more limited distribution. To understand the spatial pattern of molecular diversity throughout the range of this species complex, we obtained DNA sequence data from two mitochondrial genes, cytochrome oxidase I (COI) and the 16S rhibosomal gene (16S) for 407 samples of D. minutus and closely related species distributed across eleven countries, effectively comprising the entire range of the group. We performed phylogenetic and spatially explicit phylogeographic analyses to assess the genetic structure of lineages and infer ancestral areas. We found 43 statistically supported, deep mitochondrial lineages, several of which may represent currently unrecognized distinct species. One major clade, containing 25 divergent lineages, includes samples from the type locality of D. minutus. We defined that clade as the D. minutus complex. The remaining lineages together with the D. minutus complex constitute the D. minutus species group. Historical analyses support an Amazonian origin for the D. minutus species group with a subsequent dispersal to eastern Brazil where the D. minutus complex originated. According to our dataset, a total of eight mtDNA lineages have ranges >100,000 km2. One of them occupies an area of almost one million km2 encompassing multiple biomes. Our results, at a spatial scale and resolution unprecedented for a Neotropical vertebrate, confirm that widespread amphibian species occur in lowland South America, yet at the same time a large proportion of cryptic diversity still remains to be discovered.

A new rainfrog of the Pristimantis myersi Group (Amphibia, Craugastoridae) from Volcán Pichincha, Ecuador.

A new frog of the Pristimantis myersi Group is described from a bamboo patch within the Reserva Ecológica Verdecocha (0°5'46.9"S, 78°36'15.3"W; 2851 m), located at northwestern flank of the Volcán Pichincha, in the vicinities of Quito, Ecuador. The new species is known from eight adult males, whereas the females remain unknown; it can be readily distinguished from all species of the P. myersi Group that inhabit the highlands of the Ecuadorian Andes by the unique combination of the following characters: body small (adult male SVL 14.9-19.7 mm; females unknown); dorsal skin shagreen, with a barely visible middorsal raphe, scapular and dorsolateral folds; tympanum small but well-defined; upper eyelid with one enlarged tubercle; males with prominent vocal slits, but without nuptial pads on thumbs; fold-like tarsal tubercles. With this new species, the number of Pristimantis assigned to the P. myersi Group raises to 16, of which, 12 are in Ecuador. We provide notes on morphology and color variation, advertisement call, and natural history of the new species.

Terraranans of the Lost World: a new species of Pristimantis (Amphibia, Craugastoridae) from Abakapá-tepui in the Chimantá massif, Venezuelan Guayana, and additions to the knowledge of P. muchimuk.

A new frog of the genus Pristimantis is named and described from the summit of Abakapá-tepui in the Chimantd massif, south-eastern Venezuela. The new species is known from two adult specimens and is the second craugastorid species described from this massif. It can be readily distinguished from all congeners inhabiting the highlands of the Guiana Shield by the unique combination of the following characters: dorsal skin shagreen and ventral skin coarsely areolate, tympanum small and ill-defined, vocal slits absent in males, finger I shorter than II, thumbs with two whitish and non-spinous nuptial pads in adult males, fingers and toes with broad lateral fringes, basal webbing between all toes, throat and chest nacreous white in life. Also, based on five specimens of Pristimantis muchimuk recently collected from Churi-tepui, we provide new information on this little known species, including an amended diagnosis, notes on morphology, color variation, advertisement calls, and natural history.