Publicaciones Torres Carvajal Lenin Omar

Thermal biology of two tropical lizards from the ecuadorian andes and their vulnerability to climate change
REVISTA
PLOS ONE

Publicación
2020-01-24
This study aims to analyze the thermal biology and climatic vulnerability of two closely related lizard species (Stenocercus festae and S. guentheri) inhabiting the Ecuadorian Andes at high altitudes. Four physiological parameters—body temperature (Tb), preferred temperature (Tpref), critical thermal maximum (CTmax), and critical thermal minimum (CTmin)—were evaluated to analyze the variation of thermophysiological traits among these populations that inhabit different environmental and altitudinal conditions. We also evaluate the availability of operative temperatures, warming tolerance, and thermal safety margin of each population to estimate their possible risks in the face of future raising temperatures. Similar to previous studies, our results suggest that some physiological traits (CTmax and Tb) are influenced by environmental heterogeneity, which brings changes on the thermoregulatory behavior. Other parameters (Tpref and CTmin), may be also influenced by phylogenetic constraints. Moreover, the fluctuating air temperature (Tair) as well as the operative temperatures (Te) showed that these lizards exploit a variety of thermal microenvironments, which may facilitate behavioral thermoregulation. Warming tolerance and thermal safety margin analyses suggest that both species find thermal refugia and remain active without reducing their performance or undergoing thermal stress within their habitats. We suggest that studies on the thermal biology of tropical Andean lizards living at high altitudes are extremely important as these environments exhibit a unique diversity of microclimates, which consequently result on particular thermophysiological adaptations.

A Revision And Key For The Tribe Diaphorolepidini (Serpentes: Dipsadidae) And Checklist For The Genus Synophis.
REVISTA
ZOOTAXA

Publicación
2016-09-28
The genus Synophis contains a number of enigmatic species, distributed primarily in the Andean highlands of northern South America. Their extreme crypsis and rarity has precluded detailed study of most species. A recent flurry of collection activity resulted in the accession of many new specimens, and the description of 4 new species in 2015, doubling the number of described taxa. However, lingering questions remain regarding the assignment of many new and historical specimens, the morphological limits and geographical ranges of the species, and their phylogenetic relationships. We analyze new and existing morphological and molecular data to produce a new molecular phylogeny and revised morphological descriptions. We validate the previously unavailable tribe name Diaphorolepidini Jenner, Pyron, Arteaga, Echevarría, & Torres-Carvajal tribe nov., describe a 9th species Synophis niceforomariae Pyron, Arteaga, Echevarría, & Torres-Carvajal sp. nov., and offer new Standard Names in English and Spanish for the group: Andean Shadow Snakes and Culebras Andinas de la Sombra, respectively. A variety of features such as vertebrae and hemipenes show an interesting range of variation in the group, which should be evaluated in future studies, to refine species limits and diagnoses further. Cryptic and undiscovered diversity undoubtedly remains, and we hope this summary provides a robust basis for future work.

A Phylogenetic, Biogeographic, And Taxonomic Study Of All Extant Species Of Anolis (Squamata; Iguanidae)
REVISTA
SYSTEMATIC BIOLOGY

Publicación
2017-02-27
Anolis lizards (anoles) are textbook study organisms in evolution and ecology. Although several topics in evolutionary biology have been elucidated by the study of anoles, progress in some areas has been hampered by limited phylogenetic information on this group. Here, we present a phylogenetic analysis of all 379 extant species of Anolis, with new phylogenetic data for 139 species including new DNA data for 101 species. We use the resulting estimates as a basis for defining anole clade names under the principles of phylogenetic nomenclature and to examine the biogeographic history of anoles. Our new taxonomic treatment achieves the supposed advantages of recent subdivisions of anoles that employed ranked Linnaean-based nomenclature while avoiding the pitfalls of those approaches regarding artificial constraints imposed by ranks. Our biogeographic analyses demonstrate complexity in the dispersal history of anoles, including multiple crossings of the Isthmus of Panama, two invasions of the Caribbean, single invasions to Jamaica and Cuba, and a single evolutionary dispersal from the Caribbean to the mainland that resulted in substantial anole diversity. Our comprehensive phylogenetic estimate of anoles should prove useful for rigorous testing of many comparative evolutionary hypotheses. [Anoles; biogeography; lizards; Neotropics; phylogeny; taxonomy]

A New Species Of Phenacosaur Anolis (Squamata; Iguanidae) From Peru And A Comprehensive Phylogeny Of Dactyloa-Clade Anolis Based On New Dna Sequences And Morphology.
REVISTA
COPEIA

Publicación
2015-09-10
We describe a new species of Anolis from the Andes of northern Peru. This form is similar to species formerly assigned to the genus Phenacosaurus and to Ernest Williams’ tigrinus series. That is, the new species possesses large smooth headscales, cryptic coloration, and short limbs and tail. We present new DNA and morphological data and perform a comprehensive phylogenetic analysis of the Dactyloa clade of Anolis. The new species is estimated to be close to phenotypically similar species from Peru and Venezuela. We revise the taxonomy of Dactyloa based on our estimate. Most previously recognized phylogenetically defined groups are retained with restricted species contents. Several species currently scorable only for morphological data could not be placed in any group with confidence. The species status of the enigmatic mainland form Anolis deltae should be re-examined with reference to the roquet group of southern Lesser Antillean Anolis.

Connectivity, Population Structure, And Conservation Of Ecuadorian Green Sea Turtles.
REVISTA
ENDANGERED SPECIES RESEARCH

Publicación
2017-03-14
Studies of highly migratory species that increase our understanding of the dynamics of genetic diversity, migratory routes, and genetic connectivity are essential for informing conservation actions. Genetic data for green turtles Chelonia mydas from Ecuador have only been available from Galápagos Islands (GPS) rookeries, but not from foraging aggregations. Furthermore, green turtles from habitats associated with mainland Ecuador (Machalilla National Park; MNP) have not been sampled. To assess the genetic relationships between nesting and foraging aggregations from these 2 regions and other regional populations, the mitochondrial DNA (mtDNA) control region was sequenced from 133 turtles. Conventional FST (haplotype frequency) and ΦST (sequence-based) values were low and non-significant between Ecuadorian rookeries, suggesting high connectivity between these sites located ca. 1000 km apart. Mixed stock analysis (MSA) indicated a dominant (>94%) GPS-MNP contribution to both foraging grounds, with small and nearly negligible contributions from other rookeries in the region (e.g. Costa Rica and Mexico). While orphan haplotypes were not included in the MSA because their rookery of origin is not known, their close genetic relationships to Western and Central Pacific mtDNA clades suggests that a relatively large percentage of turtles at the combined foraging sites (>10%) have been involved in transoceanic migration events. The genetic links between GPS and MNP C. mydas nesting populations revealed by our study highlight the need to incorporate the nesting populations from coastal Ecuador in more comprehensive future conservation planning.

Description And Phylogeny Of Three New Species Of Synophis (Colubridae, Dipsadinae) From The Tropical Andes In Ecuador And Perú.
REVISTA
ZOOKEYS

Publicación
2015-12-15
Se reporta el descubrimiento de tres especies nuevas de serpientes Synophis de las estribaciones orientales de los Andes tropicales en Ecuador y Perú. Todos los registros previos de S. bogerti del oriente ecuatoriano corresponden a S. bogerti sp. n., la cual ocurre entre 1000–1750 m a lo largo de gran parte de las estribaciones amazónicas de los Andes ecuatorianos. En contraste, Synophis zamora sp. n. se restringe al suroriente de Ecuador, incluyendo la Cordillera del Cóndor, entre 1543–1843 m. Synophis insulomontanus sp. n. es de las estribaciones orientales de los Andes del centro y norte del Perú, entre 1122–1798 m, y representa el primer registro de Synophis para este país. Todas las tres especies nuevas comparten en común una espina lateral larga en la base del cuerpo del hemipene. Un árbol molecular filogenético, basado en tres genes mitocondriales es presentado, incluyendo muestras de Diaphorolepis wagneri. Nuestro árbol apoya fuertemente a Synophis y Diaphorolepis como taxa hermanos, así como la monofilia de las tres especies descritas y de S. calamitus. La inclusión de Synophis y Diaphorolepis dentro de Dipsadinae, como hermanas a un clado que contiene a Imantodes, Dipsas, Ninia, Hypsiglena y Pseudoleptodeira también es apoyada.

First Record Of Macropholidus Ruthveni Noble 1921 (Squamata: Gymnophthalmidae) From Ecuador.
REVISTA
HERPETOLOGY NOTES

Publicación
2015-03-10
The gymnophthalmid lizard clade Macropholidus, traditionally ranked as a genus, was recently defined by Torres-Carvajal and Mafla-Endara (2013) as the largest crown clade containing Macropholidus ruthveni Noble1921, but not Pholidobolus montium Peters 1863. This phylogenetic definition (de Queiroz and Gauthier, 1994) is based on a phylogenetic tree obtained from analyses of mitochondrial DNA nucleotide sequence data (Torres-Carvajal and Mafla-Endara, 2013), and is in conflict with previous non-phylogenetic definitions of both Pholidobolus and Macropholidus (Montanucci, 1973; Reeder, 1996) based on morphological data. In contrast to Pholidobolus, members of Macropholidus have a single transparent palpebral disc in the lower eyelid and lack a lateral fold between fore and hind limbs (Torres-Carvajal and Mafla-Endara, 2013). Macropholidus lizards occur between 800 and 3000 m within a region of relatively low-elevation mountains in the Andes of southern Ecuador and northern Peru, known as the Huancabamba Depression (Torres-Carvajal and Mafla-Endara, 2013). As defined by Torres-Carvajal and Mafla-Endara (2013), Macropholidus contains four species (M. annectens, M. ataktolepis, M. huancabambae, M. ruthveni) of which only M. annectens has been reported for Ecuador. Macropholidus ruthveni is currently known from both humid and dry Andean montane forests in northern Peru at elevations between 800-2500 m (Cadle and Chuna, 1995; P. Venegas pers. comm.). Herein, we present the first record of M. ruthveni for Ecuador, based on a male collected at Reserva Natural Tumbesia - La Ceiba, Zapotillo, Province of Loja in southwestern Ecuador (Fig. 1; S 4.2746, W 80.3158; WGS84; 477 m).

First Record Of The Invasive Brown Anole, Anolis Sagrei Duméril & Bibron, 1837 (Squamata: Iguanidae: Dactyloinae), In South América.
REVISTA
CHECK LIST

Publicación
2017-04-03
Introduced species are known to be potentially harmful and cause negative effects on native species and ecosystems. In addition, economic losses are common because of the occurrence and persistence of introduced species (Pimentel et al. 2001). Events of species introduction constitute one of the main drivers of biodiversity loss (Chapin et al. 2000; Didham et al. 2005). In some cases, non-native species constitute the cause for local or global changes in abundance, and of species composition (Kraus2015; Nuñez & Pauchard 2010). Thus, it becomes a big challenge to counteract the advance of introduced species, especially in megadiverse countries that are undergoing intensive economic development (Lövei et al. 2012).

Independent Demographic Responses To Climate Change Among Temperate And Tropical Milksnakes (Colubridae: Genus Lampropeltis).
REVISTA
PLOS ONE

Publicación
2015-06-17
The effects of Late Quaternary climate change have been examined for many temperate New World taxa, but the impact of Pleistocene glacial cycles on Neotropical taxa is less well understood, specifically with respect to changes in population demography. Here, we examine historical demographic trends for six species of milksnake with representatives in both the temperate and tropical Americas to determine if species share responses to climate change as a taxon or by area (i.e., temperate versus tropical environments). Using a multilocus dataset, we test for the demographic signature of population expansion and decline using non-genealogical summary statistics, as well as coalescent-based methods. In addition, we determine whether range sizes are correlated with effective population sizes for milksnakes. Results indicate that there are no identifiable trends with respect to demographic response based on location, and that species responded to changing climates independently, with tropical taxa showing greater instability. There is also no correlation between range size and effective population size, with the largest population size belonging to the species with the smallest geographic distribution. Our study highlights the importance of not generalizing the demographic histories of taxa by region and further illustrates that the New World tropics may not have been a stable refuge during the Pleistocene.

Phylogeny Of Riama (Squamata: Gymnophthalmidae), Impact Of Phenotypic Evidence On Molecular Datasets, And The Origin Of The Sierra Nevada De Santa Marta Endemic Fauna.
REVISTA
CLADISTICS

Publicación
2017-04-23
Riama is the most speciose genus of the Neotropical lizard family Gymnophthalmidae. Its more than 30 montane species occur throughout the northern Andes, the Cordillera de la Costa (CC) in Venezuela, and Trinidad. We present the most comprehensive phylogenetic analysis of Riama to date based on a total evidence (TE) approach and direct optimization of molecular and morphological evidence. Analyses use DNA sequences from four loci and 35 phenotypic characters. The dataset consists of 55 ingroup terminals representing 25 of the 30 currently recognized species of Riama plus five undescribed taxa, including an endemic species from the Sierra Nevada de Santa Marta (SNSM) in Colombia, and 66 outgroup terminals of 47 species. Analysis results in a well-supported hypothesis in which Riama is polyphyletic, with its species falling into three clades. The Tepuian Anadia mcdiarmidi nests within one clade of Riama, and the recently resurrected Pantodactylus nests within Cercosaura. Accordingly, we propose a monophyletic taxonomy that reflects historical relationships. Analysis of character evolution indicates that the presence/absence of prefrontals—a cornerstone of the early genus-level taxonomy of cercosaurines—is optimally explained as having been plesiomorphically present in the most recent common ancestor of Cercosaurinae and lost in that of the immediately less inclusive clade. Multiple independent reversals to present and subsequent returns to absent occur within this clade.

Photography-Based Taxonomy Is Inadequate, Unnecessary, And Potentially Harmful For Biological Sciences.
REVISTA
ZOOTAXA

Publicación
2016-11-23
The question whether taxonomic descriptions naming new animal species without type specimen(s) deposited in collections should be accepted for publication by scientific journals and allowed by the Code has already been discussed in Zootaxa (Dubois & Nemésio 2007; Donegan 2008, 2009; Nemésio 2009a–b; Dubois 2009; Gentile & Snell 2009; Minelli 2009; Cianferoni & Bartolozzi 2016; Amorim et al. 2016). This question was again raised in a letter supported by 35 signatories published in the journal Nature (Pape et al. 2016) on 15 September 2016. On 25 September 2016, the following rebuttal (strictly limited to 300 words as per the editorial rules of Nature) was submitted to Nature, which on 18 October 2016 refused to publish it. As we think this problem is a very important one for zoological taxonomy, this text is published here exactly as submitted to Nature, followed by the list of the 493 taxonomists and collection-based researchers who signed it in the short time span from 20 September to 6 October 2016.

Phylogeny And Biogeography Of The Most Diverse Clade Of South American Gymnophthalmid Lizards (Squamata, Gymnophthalmidae, Cercosaurinae).
REVISTA
MOLECULAR PHYLOGENETICS AND EVOLUTION

Publicación
2016-03-11
Nearly 50% of the diversity of the speciose Neotropical lizard clade Gymnophthalmidae is nested within the subclade Cercosaurinae. The taxonomy of ercosaurinae lizards has been historically confusing because many diagnostic characters of those clades traditionally ranked as genera do not represent true diagnostic apomorphies. Even though molecular phylogenies of several ‘genera’ have been presented in the last few years, some of them remain poorly sampled (e.g., Anadia, Echinosaura, Potamites, Riama). In this paper we present a more comprehensive phylogeny of Cercosaurinae lizards with emphasis on Andean taxa from Ecuador and Peru, as well as a time-calibrated phylogeny with reconstruction of ancestral areas. Our analysis includes 52% of all recognized species of Cercosaurinae (67 species) and 1914 characters including three mitochondrial and one nuclear gene. We find that Anadia, Echinosaura, Euspondylus, Potamites, Proctoporus, and Riama are not monophyletic: the Tepuian Anadia mcdiarmidi is not sister to Andean species of Anadia; Echinosaura sulcarostrum is not included in the same clade formed by other species of Echinosaura and their more recent common ancestor; Teuchocercus is nested within Echinosaura; species of Euspondylus included in this study are nested within Proctoporus; Riama laudahnae is included in Proctoporus; and Potamites is paraphyletic and split in two separate clades, one of which we name Gelanesaurus, also a new genus-group name. Within Potamites, P. ecpleopus is paraphyletic, and P. strangulatus strangulatus and P. strangulatus trachodus are recognized as two distinct species.

Present Diversity Of Galápagos Leaf-Toed Geckos (Phyllodactylidae: Phyllodactylus) Stems From Three Independent Colonization Events.
REVISTA
MOLECULAR PHYLOGENETICS AND EVOLUTION

Publicación
2016-07-09
We re-examined the biogeography of the leaf-toed geckos (Phyllodactylus) endemic to the Galápagos Islands by analyzing for the first time samples of P. gilberti, a species endemic to Wolf island, in a phylogenetic framework. Our aim was to test the three-colonizations scenario previously proposed for these lizards and estimate the age of each colonization event. To achieve this we estimated simultaneously a species tree and divergence times with Bayesian methods. Our results supported the three-colonizations scenario. Similar to a previous hypothesis, the species tree obtained here showed that most species of Phyllodactylus are nested in a single clade with an age between 5.49 and 13.8 Ma, whereas a second independent colonization corresponding to P. darwini from San Cristóbal island occurred 3.03 Ma ago. The species from Wolf island, P. gilberti, stems from a more recent colonization event (0.69 Ma). Thus, present diversity of Galápagos leaf-toed geckos stems from three independent, asynchronous colonization events. As with other Galápagos organisms, the Pacific coast of South America seems to be the source for the founders of P. gilberti.

Short-Term Predicted Extinction Of Andean Populations Of The Lizard Stenocercus Guentheri (Iguanidae: Tropidurinae).
REVISTA
JOURNAL OF THERMAL BIOLOGY

Publicación
2016-12-01
We studied the thermal physiology of the Andean lizard Stenocercus guentheri in order to evaluate the possible effects of global warming on this species. We determined the preferred body temperature (Tpref), critical thermals (CTmin, CTmax), and hours of restriction and activity. Tpref was 32.14±1.83°C; CTmin was 8.31°C in adults and 9.14°C in juveniles, whereas CTmax was 43.28°C in adults and 41.68°C in juveniles. To assess extinction risk, we used the model created by Sinervo et al. (2010) and predicted that 16.7% of populations will have a high risk of extinction by 2020, with an increase to 26.7% by 2050. These results suggest that this species, despite being able to maintain its Tpref through behavioral thermoregulation and habitat selection, could be physiologically sensitive to climate warming; thus, the potential for local adaptation may be limited under a warmer climate. Further studies focusing on the ability of S. guentheri to evolve higher Tpref and thermal tolerances are needed to understand the ability of this species to respond to climate change.

Priority Areas For Biodiversity Conservation In Mainland Ecuador
REVISTA
NEOTROPICAL BIODIVERSITY

Publicación
2017-08-07
Ecuador’s territory harbors a unique set of species and ecosystems, many of them endemic to the countries’ territory and subject to different sources of threat of anthropogenic origin. Despite national and subnational conservation strategies developed in Ecuador to conserve its biodiversity in the long run, including the National System of Protected Areas (PANE) and the forest conservation incentive program SocioBosque (PSB), further actions are needed to mitigate and reverse the effects of threats for the persistence of biodiversity. This study was designed to identify the most important areas for biodiversity conservation in mainland Ecuador that can contribute to preserving key species (i.e. endemic, threatened) and ecosystems in the wider landscape, thus complementing current conservation efforts (i.e. PANE). Species distribution models and recent maps were used to identify a set of 744 species and 87 ecosystems as surrogates of the country’s biodiversity. Marxan, a systematic reserve selection algorithm was used to identify important biodiversity areas that could represent between 10% and 20% of the remnant distribution of the surrogates. The optimized solution generated by Marxan included 24% (3.64 million ha) of Ecuador′s remnant vegetation, of which 35% is within the current national protected area system and 13% (456 000 ha) are included within SocioBosque communal and private conservation agreements. Major conservation shortfalls of the PANE were concentrated in the Southern Andes, Central Amazonia, and the Central and Southern portions of the Coastal plain.

Extinct, Obscure Or Imaginary: The Lizard Species With The Smallest Ranges
REVISTA
DIVERSITY AND DISTRIBUTIONS

Publicación
2017-11-23
Aim: Small geographic ranges make species especially prone to extinction from anthropogenic disturbances or natural stochastic events. We assemble and analyse a comprehensive dataset of all the world's lizard species and identify the species with the smallest ranges—those known only from their type localities. We compare them to wide-ranging species to infer whether specific geographic regions or biological traits predispose species to have small ranges. Location: Global. Methods: We extensively surveyed museum collections, the primary literature and our own field records to identify all the species of lizards with a maximum linear geographic extent of <10 km. We compared their biogeography, key biological traits and threat status to those of all other lizards. Results: One in seven lizards (927 of the 6,568 currently recognized species) are known only from their type localities. These include 213 species known only from a single specimen. Compared to more wide-ranging taxa, they mostly inhabit relatively inaccessible regions at lower, mostly tropical, latitudes. Surprisingly, we found that burrowing lifestyle is a relatively unimportant driver of small range size. Geckos are especially prone to having tiny ranges, and skinks dominate lists of such species not seen for over 50 years, as well as of species known only from their holotype.

Systematics And Ecology Of Anolis Biporcatus (Squamata: Iguanidae)
REVISTA
SALAMANDRA

Publicación
2017-05-15
Anolis biporcatus is a large green anole found in southern Mexico, Central America and northern South America. We examined morphological, molecular, and ecological aspects of specimens from localities throughout the range of A. biporcatus and found evidence that warrants elevating A. biporcatus parvauritus to species status. A. parvauritus differs from A. biporcatus in its dewlap colour pattern, overall morphology, and mitochondrial DNA. A deep mitochondrial split exists between northern and central Costa Rican samples of A. biporcatus, and substantial morphological variation occurs within this form. These patterns suggest potential species-level splits within currently recognized A. biporcatus. More extensive sampling of DNA data is needed to evaluate potential species boundaries within A. biporcatus.

Use Of A Rostral Appendage During Social Interactions In The Ecuadorian Anolis Proboscis
REVISTA
JOURNAL OF NATURAL HISTORY

Publicación
2017-06-02
The use of sexually selected characters in inter- and intra-sexual interactions has long been of interest to evolutionary biologists. Recently, a distinction between sexually selected traits as ornaments versus weapons has been advanced. We investigated the behaviour of an enigmatic lizard with a prominent sexually dimorphic trait in an effort to describe whether the trait was the product of sexual selection and further whether it functioned as a weapon or an ornament. The subject of our study was the Ecuadorian proboscis anole (Anolis proboscis), a slow-moving cryptic species endemic to the north-western slopes of the Andes in Ecuador. Males, but not females, of this species bear a rostral appendage that has been described as an exaggerated trait resulting from sexual selection. However, a thorough description of the use of the rostral appendage in social interactions is lacking. Here, we describe social interactions of this species during 11 male–female courtships and mating interactions, as well as three male–male agonistic interactions. We describe four types of displays by males, many involving the rostral appendage. We found that the rostral appendage is used as an ornament in social displays but not as a weapon in combat. We also show that, unlike other lizards with rostral appendages, male A. proboscis hatch with this structure already developed.

Diversity, Threat, And Conservation Of Reptiles From Continental Ecuador
REVISTA
AMPHIBIAN & REPTILE CONSERVATION

Publicación
2017-12-21
Ecuador is one of the most reptile-diverse countries in the world, with 464 currently recognized species. Similar to other taxa, reptiles in Ecuador face important conservation challenges because of anthropogenic activities. Using distribution data of nearly 90% of the species of reptiles from continental Ecuador, as well as information on ecosystem protection status and anthropogenic activities, we present the first comprehensive quantitative study of reptile conservation in Ecuador. While species richness is higher in northwestern Ecuador and the central-northern Amazon, the conservation priority areas identified in this study also include the central Pacific coast, southwestern Ecuador, and the central-southern Amazon. Similar areas have been identified by previous studies as conservation gaps. Thus, our study reinforces the idea of protecting those areas to improve the conservation of biodiversity in continental Ecuador.

The Global Distribution Of Tetrapods Reveals A Need For Targeted Reptile Conservation
REVISTA
NATURE ECOLOGY AND EVOLUTION

Publicación
2017-10-09
The distributions of amphibians, birds and mammals have underpinned global and local conservation priorities, and have been fundamental to our understanding of the determinants of global biodiversity. In contrast, the global distributions of reptiles, representing a third of terrestrial vertebrate diversity, have been unavailable. This prevented the incorporation of reptiles into conservation planning and biased our understanding of the underlying processes governing global vertebrate biodiversity. Here, we present and analyse the global distribution of 10,064 reptile species (99% of extant terrestrial species). We show that richness patterns of the other three tetrapod classes are good spatial surrogates for species richness of all reptiles combined and of snakes, but characterize diversity patterns of lizards and turtles poorly. Hotspots of total and endemic lizard richness overlap very little with those of other taxa. Moreover, existing protected areas, sites of biodiversity significance and global conservation schemes represent birds and mammals better than reptiles. We show that additional conservation actions are needed to effectively protect reptiles, particularly lizards and turtles. Adding reptile knowledge to a global complementarity conservation priority scheme identifies many locations that consequently become important. Notably, investing resources in some of the world’s arid, grassland and savannah habitats might be necessary to represent all terrestrial vertebrates efficiently.

Phylogeny And Diversity Of Neotropical Monkey Lizards (Iguanidae: Polychrus Cuvier, 1817)
REVISTA
PLOS ONE

Publicación
2017-06-01
Neotropical monkey lizards (Polychrus) are arboreal lizards with compressed bodies, partially fused eyelids and strikingly long, whip-like tails. The eight currently recognized species occur in the lowlands of South and Central America. Based on the largest taxon and character sampling to date, we analyze three mitochondrial and one nuclear gene using Bayesian methods to (1) infer the phylogeny of Polychrus under both concatenated-tree and speciestree methods; (2) identify lineages that could represent putative undescribed species; and (3) estimate divergence times. Our species tree places P. acutirostris as the sister taxon to all other species of Polychrus. While the phylogenetic position of P. gutturosus and P. peruvianus is poorly resolved, P. marmoratus and P. femoralis are strongly supported as sister to P. liogaster and P. jacquelinae, respectively. Recognition of P. auduboni and P. marmoratus sensu stricto as distinct species indicates that the populations of "P. marmoratus" from the Amazon and the Atlantic coast in Brazil represent separate species. Similarly, populations of P. femoralis from the Tumbes region might belong to a cryptic undescribed species. Relative divergence times and published age estimates suggest that the orogeny of the Andes did not play a significant role in the early evolution of Polychrus.

A New Species Of Cat-Eyed Snake (Serpentes: Dipsadinae: Leptodeirini) From The Andes Of Southern Ecuador
REVISTA
ZOOTAXA

Publicación
2020-12-16
Leptodeira is one of the most widespread and taxonomically problematic snake taxa in the Americas. Here we describe a new species of Leptodeira from the Andes of southern Ecuador based on morphological and molecular data. The new species is geographically close and morphologically similar to L. ornata and L. larcorum, from which it can be distinguished by having smaller dorsal body blotches, a longer tail, and shorter spines on the hemipenial body. The shortest genetic distances between the new species and its congeners are 0.02 (16S), 0.05 (cytb), and 0.18 (ND4). The new species is restricted to the Jubones River Basin in southern Ecuador, an area of endemism for other reptile species. Our phylogenetic analysis based on mitochondrial and nuclear DNA sequence data also supports recognition of the names L. larcorum (restricted to Peru) for “L. septentrionalis larcorum”, and L. ornata for populations of “L. s. ornata” from central and eastern Panama, western Colombia, and western Ecuador. However, some samples of “L. s. ornata” from Panama and Costa Rica, as well as the new species described herein, are not included within or more closely related to L. ornata, which is sister to the clade (L. bakeri, L. ashmeadii

Thermal Biology Of Two Tropical Lizards From The Ecuadorian Andes And Their Vulnerability To Climate Change
REVISTA
PLOS ONE

Publicación
2020-01-24
This study aims to analyze the thermal biology and climatic vulnerability of two closely related lizard species (Stenocercus festae and S. guentheri) inhabiting the Ecuadorian Andes at high altitudes. Four physiological parameters—body temperature (Tb), preferred temperature (Tpref), critical thermal maximum (CTmax), and critical thermal minimum (CTmin)—were evaluated to analyze the variation of thermophysiological traits among these populations that inhabit different environmental and altitudinal conditions. We also evaluate the availability of operative temperatures, warming tolerance, and thermal safety margin of each population to estimate their possible risks in the face of future raising temperatures. Similar to previous studies, our results suggest that some physiological traits (CTmax and Tb) are influenced by environmental heterogeneity, which brings changes on the thermoregulatory behavior. Other parameters (Tpref and CTmin), may be also influenced by phylogenetic constraints. Moreover, the fluctuating air temperature (Tair) as well as the operative temperatures (Te) showed that these lizards exploit a variety of thermal microenvironments, which may facilitate behavioral thermoregulation. Warming tolerance and thermal safety margin analyses suggest that both species find thermal refugia and remain active without reducing their performance or undergoing thermal stress within their habitats. We suggest that studies on the thermal biology of tropical Andean lizards living at high altitudes are extremely important as these environments exhibit a unique diversity of microclimates, which consequently result on particular thermophysiological adaptations.

Alternative Reproductive Adaptations Predict Asymmetric Responses To Climate Change In Lizards
REVISTA
SCIENTIFIC REPORTS

Publicación
2019-03-25
Anthropogenic climate change ranks among the major global-scale threats to modern biodiversity. Extinction risks are known to increase via the interactions between rapid climatic alterations and environmentally-sensitive species traits that fail to adapt to those changes. Accumulating evidence reveals the influence of Eco physiological, ecological and phenological factors as drivers underlying demographic collapses that lead to population extinctions. However, the extent to which life-history traits influence population responses to climate change remains largely unexplored. The emerging ‘cul-de-sac hypothesis’ predicts that reptilian viviparity (‘live-bearing’ reproduction), a ‘key innovation’ facilitating historical invasions of cold climates, increases extinction risks under progressively warming climates compared to oviparous reproduction – as warming advances poleward/mountain wards, historically cold-climates shrink, leading viviparous species to face demographic collapses. We present the first large-scale test of this prediction based on multiple lizard radiations and on future projections of climate-based ecological niche models. Viviparous species were found to experience stronger elevational range shifts (and potentially increased extinctions) in coming decades, compared to oviparous lizards. Therefore, our analyses support the hypothesis’s fundamental prediction that elevational shifts are more severe in viviparous species, and highlight the role that life-history adaptations play in the responses of biodiversity to ongoing climate change.

Divergence Of Tropical Pitvipers Promoted By Independent Colonization Events Of Dry Montane Andean Habitats
REVISTA
JOURNAL OF BIOGEOGRAPHY

Publicación
2019-07-17
Aim: A poorly explored feature of the origin and maintenance of Neotropical bio‐ diversity is how the evolutionary dynamics of colonization and differentiation in relation to lowland and highland habitats has impacted lineage formation. Most spe‐ ciation models for this region have focused on vicariant events, whereas the need to assess the influence of demographic processes has been recognized only recently. We evaluate if the origin of Andean montane lineages of terciopelo pitvipers is ex‐ plained by either of two historical processes that represent distinct phylogeographic mechanisms: differentiation by isolation within the highlands or different dispersal events from the lowlands. Location: Western Ecuador. Taxon: Terciopelo pitvipers (Bothrops asper species complex). Methods: We use genomic data and genetic clustering analyses, evaluation of histori‐ cal migration between genetic clusters and demographic model selection to investi‐ gate recent diversification events in South America using a vertebrate group rarely explored in phylogeographic studies: tropical Andean snakes. Specifically, the ori‐ gin of two Ecuadorian montane lineages of terciopelo pitvipers was evaluated given ambiguous phylogenetic relationships with the presumably ancestral Pacific lowland lineage. Results: Discrepancies of evolutionary relationships previously obtained with tree‐ like methods are resolved through the use of modelling approaches. We found strong support for the independent origin of montane lineages based on topologies inferred by maximum‐likelihood trees and modelling approaches that take into account pos‐ sible gene flow. This suggests dispersal rather than in‐situ differentiation as the most likely mechanism by which the montane linages originated.

Systematic Review Of Atractus Schach (Serpentes: Dipsadidae) Species Complex From The Guiana Shield With Description Of Three New Species
REVISTA
SYSTEMATICS AND BIODIVERSITY

Publicación
2019-04-03
The Guiana Shield harbours one of the best preserved and largest extents of tropical forest on Earth and an immense biodiversity. The herpetofauna of this region remains poorly known. The species-rich snake genus Atractus contains +-140 species, many with complicated taxonomic histories, including A. schach. Examination of specimens in museums and newly collected material from French Guiana has allowed the illustration of hemipenial morphology for the first time and an expanded diagnosis. Concatenated molecular phylogenetic (mitochondrial and nuclear genes) and phenotypic (morphometrics, external and hemipenial morphology) analyses confirm non-monophyly of the A. flammigerus group and indicate that A. schach is a species complex with three new species described here. The geographic distribution of A. schach sensu stricto is restricted to Guiana, Surinam, and French Guiana north of Tumucumaque massif. Populations tentatively assigned to A. schach from the east from French Guiana in the Roura lowlands to Almeirim, and from central Amazonia between the Negro and Trombetas rivers in Brazil are also recognized as new species. Our results suggest that populations from south of the Amazon River are not conspecific with those from the Guiana Shield

Phylogeny, Diversity And Biogeography Of Neotropical Sipo Snakes (Serpentes: Colubrinae: Chironius)
REVISTA
MOLECULAR PHYLOGENETICS AND EVOLUTION

Publicación
2019-01-01
Neotropical sipo snakes (Chironius) are large diurnal snakes with a long tail and big eyes that differ from other Neotropical snakes in having 10 or 12 dorsal scale rows at midbody. The 22 currently recognized species occur from Central America south to Uruguay and northeastern Argentina. Based on the largest geographical sampling to date including ∼90% of all species, we analyzed one nuclear and three mitochondrial genes using phylogenetic methods to (1) test the monophyly of Chironius and some of its widely distributed species; (2) identify lineages that could represent undescribed species; and (3) reconstruct ancestral distributions. Our best hypothesis placed C. grandisquamis (Chocoan Rainforest) + C. challenger (Pantepui) as sister to all other species. Based on phylogeny and geographic distribution, we identified 14 subclades as putative species within Chironius fuscus, C. multiventris (including C. foveatus and C. laurenti), C. monticola, and C. exoletus. Under current taxonomy, these species show nearly twice as much genetic diversity as other species of Chironius for ND4. Biogeographical analyses using BioGeoBEARS suggest that current distribution patterns of Chironius species across South America resulted from multiple range expansions. The MRCA of the clade C. challenger + C. grandisquamis was most likely distributed over the Pantepui region, the Andes, and the Chocoan Rainforest, whereas the remaining lineages probably evolved from an Amazonian ancestor.

Reptiles Of Ecuador: A Resource-Rich Online Portal, With Dynamic Checklists And Photographic Guides
REVISTA
AMPHIBIAN & REPTILE CONSERVATION

Publicación
2019-05-18
Con 477 especies de reptiles (excluyendo aves) en un área de 283,561 km2, Ecuador tiene la mayor densidad de riqueza de especies de reptiles de los países megadiversos del mundo. Esta riqueza está representada por 35 especies de tortugas, cinco cocodrilos y 437 escamosos, incluyendo tres anfisbénidos, 197 lagartijas y 237 serpientes. De estas, 45 especies son endémicas de las Islas Galápagos y 111 son endémicas del Ecuador continental. La alta tasa de descripciones de especies durante las últimas décadas, junto con los frecuentes cambios taxonómicos, han impedido que las listas de especies y los libros impresos mantengan un registro razonablemente actualizado de las especies de reptiles del Ecuador. Aquí presentamos Reptiles del Ecuador (http://bioweb.bio/faunaweb/reptiliaweb), un portal en línea gratuito y rico en recursos con información actualizada sobre los reptiles ecuatorianos. Este portal interactivo incluye información enciclopédica sobre todas las especies, presentaciones multimedia, mapas de distribución, modelos de aptitud de hábitat y guías dinámicas en PDF. Incluimos además una lista de especies actualizada, con información sobre distribución, endemismo y estado de conservación, así como una guía fotográfica de los reptiles del Ecuador.

The Global Biogeography Of Lizard Functional Groups
REVISTA
JOURNAL OF BIOGEOGRAPHY

Publicación
2019-07-19
Aim: Understanding the mechanisms determining species richness is a primary goal of biogeography. Richness patterns of sub‐groups within a taxon are usually assumed to be driven by similar processes. However, if richness of distinct ecological strategies responds differently to the same processes, inferences made for an entire taxon may be misleading. We deconstruct the global lizard assemblage into functional groups and examine the congruence among richness patterns between them. We further examine the species richness – functional richness relationship to elucidate the way functional diversity contributes to the overall species richness patterns. Location: Global. Methods: Using comprehensive biological trait databases we classified the global lizard assemblage into ecological strategies based on body size, diet, activity times and microhabitat preferences, using Archetypal Analysis. We then examined spatial gradients in the richness of each strategy at the one‐degree grid cell, biome, and realm scales. Results: We found that lizards can best be characterized by seven “ecological strategies”: scansorial, terrestrial, nocturnal, herbivorous, fossorial, large, and semi‐aquatic. There are large differences among the global richness patterns of these strategies. While the major richness hotspot for lizards in general is in Australia, several strategies exhibit highest richness in the Amazon Basin. Importantly, the global maximum in lizard species richness is achieved at intermediate values of functional diversity and increasing functional diversity further result in a slow decline of species richness.

Sistemática De Los Saurios Anadia Gray, 1845 (Squamata: Gymnophthalmidae) De Ecuador: Límite De Especies, Distribución Geográfica Y Descripción De Una Especie Nueva
REVISTA
NEOTROPICAL BIODIVERSITY

Publicación
2018-07-20
Realizamos una revisión sistemática del género Anadia en Ecuador para delimitar especies basándonos en varias líneas de evidencia: morfología externa, hemipenes, coloración, secuencias de ADN y distribución geográfica. Describimos una especie nueva, Anadia buenaventura sp. nov., que se distingue de otras especies de Anadia por presentar un patrón de coloración con bandas dorsolaterales claras e hileras paravertebrales de puntos oscuros, y menos escamas alrededor del cuerpo. También presentamos nuevos datos sobre la distribución de Anadia petersi Oftedal y una filogenia molecular que sugiere que Anadia rhombifera (Günther) constituye un complejo de especies que requiere más trabajo taxonómico que debería incluir material de Colombia.