Specimen collection is essential for modern science.

Natural history museums are vital repositories of specimens, samples and data that inform about the natural world; this Formal Comment revisits a Perspective that advocated for the adoption of compassionate collection practices, querying whether it will ever be possible to completely do away with whole animal specimen collection.

Phylogeography and taxonomy of Coleonyx elegans Gray 1845 (Squamata: Eublepharidae) in Mesoamerica: The Isthmus of Tehuantepec as an environmental barrier.

Population divergence leading to speciation is often explained by physical barriers causing allopatric distributions of historically connected populations. Environmental barriers have increasingly been shown to cause population divergence through local adaptation to distinct ecological characteristics. In this study, we evaluate population structuring and phylogeographic history within the Yucatán banded gecko Coleonyx elegans Gray 1845 to assess the role of both physical and environmental barriers in shaping the spatio-genetic distribution of a Mesoamerican tropical forest taxon. We generated RADseq and multi-locus Sanger datasets that included sampling across the entire species' range. Results find support for two distinct evolutionary lineages that diverged during the late Pliocene and show recent population expansions. Furthermore, these genetic lineages largely align with subspecies boundaries defined by morphology. Several mountain ranges identified as phylogeographic barriers in other taxa act as physical barriers to gene flow between the two clades. Despite the absence of a physical barrier between lineages across the lowland Isthmus of Tehuantepec, no introgression was observed. Here, a steep environmental cline associated with seasonality of precipitation corresponds exactly with the distributional limits of the lineages, whose closest samples are only 30 km apart. The combination of molecular and environmental evidence, and in conjunction with previous morphological evidence, allows us to reassess the current taxonomy in an integrative framework. Based on our findings, we elevate the previously recognized subspecies from the Pacific versant, the Colima banded gecko C. nemoralis Klauber 1945, to full species status and comment on conservation implications.

Phylogenetic and morphological evidence reveals the association between diet and the evolution of the venom delivery system in Neotropical goo-eating snakes

Advanced endoglyptodont snakes share a complex but homologous venom delivery system associated with the upperjaw and its dentition. Recently, a remarkable novel lower jaw venom delivery system was described for the Neotropicaldipsadine radiation of goo-eating snakes. While most dipsadines are opistoglyphous and exhibit large, mainly serousvenom glands associated with the upper jaw and supralabial glands, goo-eating dipsadine snakes are aglyphous and lackserous upper labial venom glands. Here, we provide new morphological and histological information on the oral glandsand maxillary dentition of representatives of the major lineages of dipsadines that help trace the evolutionary steps thatshaped the venom delivery system of dipsadines. We performed a maximum likelihood analysis on a molecular datasetthat includes 443 terminals and seven loci. Our results show that goo-eating dipsadines form a monophyletic assemblagethat includes the genusAdelphicosfor the first time, along withGeophis,Atractus,Ninia,Chersodromus,Tropidodipsas,Sibon, andDipsas. We also provide the first evidence of a complete shift from an upper jaw to a lower jaw venomdelivery system associated with their specialized feeding behaviour. Unlike other dipsadines who exhibit typicalendoglyptodont anteroposteriorly ridged posterior maxillary teeth, goo-eating dipsadines have uniform lateromediallyridged teeth throughout their maxilla. Our results indicate that the loss of the endoglyptodont venom delivery systemoccurred in the most recent common ancestor of goo-eating dipsadines, probably resulting from the loss of theembryonic posterior maxillary lamina responsible for the development of the venom delivery system.

A new species of Sceloporus of the torquatus group (Reptilia: Phrynosomatidae) from West Mexico.

We describe Sceloporus huichol sp. nov., a new species of spiny lizard of the genus Sceloporus, that is found in the mountainous regions of Jalisco and Nayarit in western Mexico. The new species belongs to the torquatus species group and has previously been confused with Sceloporus melanogaster, from which it differs by its smaller size, head color and nuchal collar. Additionally, Sceloporus huichol sp. nov. is different from the rest of its congeners by relevant genetic and morphologicall data. Despite having a complete nuchal collar, characteristic of Sceloporus torquatus, S. huichol sp. nov. is more closely related phylogenetically to S. melanogaster. Recognition of this new species brings the number of species in the torquatus group to nine.

A new species of leaf-toed gecko (Phyllodactylidae, Phyllodactylus) from María Cleofas Island, Nayarit, Mexico.

We describe a new species of leaf-toed gecko of the genus Phyllodactylus from María Cleofas Island, the smallest island of Tres Marías Archipelago, Nayarit, México. Genomic, phylogenomic, and morphological evidence support that the new species presents a unique combination of diagnostic characters. Morphologically, the new species has a high number of tubercles, head to tail (mean 47), longitudinal ventral scales (mean 61), and third labial-snout scales (mean 26). Gene flow tests revealed the genetic isolation of insular populations from mainland counterparts. In addition, we confirmed the non-monophyly of P. homolepidurus and P. nolascoensis, and we show that the taxon P. t. saxatilis is a complex; therefore, we propose taxonomic changes within the saxatilis clade. The discovery of this new insular endemic species highlights the urgency of continued exploration of the biological diversity of island faunas of Mexico.

Phylogenomics of the Mesoamerican alligator-lizard genera Abronia and Mesaspis (Anguidae: Gerrhonotinae) reveals multiple independent clades of arboreal and terrestrial species.

Abronia and Mesaspis are two of the five anguid lizard genera in the subfamily Gerrhonotinae. Their members are restricted to Mesoamerica, and most have allopatric distributions. Species of Abronia are primarily arboreal and occur in cloud and seasonally dry pine-oak forests, whereas those of Mesaspis are terrestrial and inhabit mesic microhabitats of montane forests. Recent molecular studies suggest that although these genera together form a monophyletic group, neither genus is monophyletic. Here we performed a phylogenetic study of Abronia and Mesaspis based on the most comprehensive taxonomic sampling of these genera to date and double digest restriction site-associated (ddRADseq) data. Our reconstructed phylogeny differed considerably from all previously published topologies, consistently recovering multiple independent clades of arboreal and terrestrial species and Abronia and Mesaspis as non-monophyletic. Geography, rather than current taxonomy, provides the best explanation of their phylogenetic relationships. Our analyses consistently recovered two main clades, distributed on the highlands of Middle America east and west of the Isthmus of Tehuantepec, respectively, and each composed of subclades of Abronia and Mesaspis. In the former main clade, members of the subgenus Auriculabronia formed the sister taxon to the Mesaspis moreletii complex, whereas the main clade west of the Isthmus was composed of two clades with a subclade of Abronia and another of Mesaspis each (one clade on the Atlantic versant of the main mountain ranges of eastern Mexico and another one on the Sierra Madre del Sur exclusive of its Atlantic versant) and a third clade with species of the subgenera Abronia and Scopaeabronia. We discuss the taxonomic implications of our results for the classification of the examined taxa and list the morphological characters that diagnose the recovered clades. This study highlights the utility of ddRADseq data to reconstruct the evolutionary history of supraspecific vertebrate taxa.

Phylogenomics and molecular species delimitation reveals great cryptic diversity of leaf-toed geckos (Phyllodactylidae: Phyllodactylus), ancient origins, and diversification in Mexico.

We utilize the efficient GBS technique to obtain thousands of nuclear loci and SNPs to reconstruct the evolutionary history of Mexican leaf-toed geckos (Phyllodactylus). Through the incorporation of unprecedented sampling for this group of geckos, in combination with genomic data analysis, we generate mostly consistent phylogenetic hypotheses using two approaches: supermatrix and coalescent-based inference. All topologies depict three, mutually exclusive major clades. Clade I comprises P. bordai and all species closer to P. bordai than to any other Phyllodactylus. Clade II comprises P. nocticolus and all species closer to P. nocticolus than to any other Phyllodactylus. Clade III comprises P. tuberculosus and all species closer to P. tuberculosus than to any other Phyllodactylus. Analyses estimate the age for the most recent common ancestor of Phyllodactylus in the Eocene (~43 mya), and the ancestors of each major clade date to the Eocene-Oligocene transition (32-36 mya). This group includes one late-Eocene lineage (P. bordai), Oligocene lineages (P. paucituberculatus, P. delcampi), but also topological patterns that indicate a recent radiation occurred during the Pleistocene on islands in the Gulf of California. The wide spatial and temporal scale indicates a complex and unique biogeographic history for each major clade. The 33 species delimited by BPP and stepping-stone BFD*coalescent based genomic approaches reflect this history. This diversity delimited for Mexican leaf-toed geckos demonstrates a vast underestimation in the number of species based on morphological data alone.

The roles of vicariance and dispersal in the differentiation of two species of the Rhinella marina species complex.

The high levels of Neotropical biodiversity are commonly associated with the intense Neogene-Quaternary geological events and climate dynamics. Here, we investigate the evolutionary history of two species of Neotropical closely related amphibians (R. horribilis and R. marina). We combine published data with new mitochondrial DNA sequences and multiple nuclear markers, including 12 microsatellites. The phylogenetic analyses showed support for grouping the samples in two main clades; R. horribilis (Central America and Mexico) and R. marina (South America east of the Andes). However, the phylogenetic inferences also show an evident mito-nuclear discordance. We use Approximate Bayesian Computation (ABC) to test the role of different events in the diversification between the two groups recovered. We found that both species were affected primarily by a recent Pleistocene divergence, which was similar to the divergence estimate revealed by the Isolation-with-Migration model, under persistent bidirectional gene flow through time. We provide the first evidence that R. horribilis is differentiated from the South American R. marina at the nuclear level supporting the taxonomic status of R. horribilis, which has been controversial for more than a century.

Detection of cryptic diversity in lizards (Squamata) from two Biosphere Reserves in Mesoamerica.

A combined approach based on karyology and DNA taxonomy allowed us to characterize the taxonomic peculiarities in 10 Mesoamerican lizard species, belonging to six genera and five families, inhabiting two Biosphere Reserve in Chiapas, Mexico: La Sepultura Biosphere Reserve, and Montes Azules Biosphere. The karyotypes of four species, Phyllodactylus sp. 3 (P. tuberculosus species group) (2n = 38), Holcosus festivus (Lichtenstein et von Martens, 1856) (2n = 50), Anolis lemurinus Cope, 1861 (2n = 40), and A. uniformis Cope, 1885 (2n = 29-30) are described for the first time, the last one showing a particular X1X1X2X2/X1X2Y condition. In Aspidoscelis deppii (Wiegmann, 1834) (2n = 50) and Anolis capito Peters, 1863 (2n = 42), we found a different karyotype from the ones previously reported for these species. Moreover, in A. capito, the cytogenetic observation is concurrent with a considerable genetic divergence (9%) at the studied mtDNA marker (MT-ND2), which is indicative of a putative new cryptic species. The skink Scincella cherriei (Cope, 1893), showed high values of genetic divergence (5.2% at 16S gene) between the specimens from Montes Azules and those from Costa Rica and Nicaragua, comparable to the values typical of sister species in skinks. A lower level of genetic divergence, compatible with an intraspecific phylogeographic structure, has been identified in Lepidophyma flavimaculatum Duméril, 1851. These new data identify taxa that urgently require more in-depth taxonomic studies especially in these areas where habitat alteration is proceeding at an alarming rate.

Geographic variation in morphology in the Mohave Rattlesnake (Crotalus scutulatus Kennicott 1861) (Serpentes: Viperidae): implications for species boundaries.

The Mohave Rattlesnake (Crotalus scutulatus) is a highly venomous pitviper inhabiting the arid interior deserts, grasslands, and savannas of western North America. Currently two subspecies are recognized: the Northern Mohave Rattlesnake (C. s. scutulatus) ranging from southern California to the southern Central Mexican Plateau, and the Huamantla Rattlesnake (C. s. salvini) from the region of Tlaxcala, Veracruz, and Puebla in south-central Mexico. Although recent studies have demonstrated extensive geographic variation in venom composition and cryptic genetic diversity in this species, no modern studies have focused on geographic variation in morphology. Here we analyzed a series of qualitative, meristic, and morphometric traits from 347 specimens of C. scutulatus and show that this species is phenotypically cohesive without discrete subgroups, and that morphology follows a continuous cline in primarily color pattern and meristic traits across the major axis of its expansive distribution. Interpreted in the context of previously published molecular evidence, our morphological analyses suggest that multiple episodes of isolation and secondary contact among metapopulations during the Pleistocene were sufficient to produce distinctive genetic populations, which have since experienced gene flow to produce clinal variation in phenotypes without discrete or diagnosable distinctions among these original populations. For taxonomic purposes, we recommend that C. scutulatus be retained as a single species, although it is possible that C. s. salvini, which is morphologically the most distinctive population, could represent a peripheral isolate in the initial stages of speciation.

A new species of patch-nosed snake (Colubridae: Salvadora Baird and Girard, 1853) from Oaxaca, Mexico.

A new species of snake of the genus Salvadora from Oaxaca, Mexico, is described. This taxon was confused with S. intermedia in previous taxonomic descriptions. It is characterized by lacking a pale vertebral stripe and by having incomplete dorsolateral stripes that do not reach the posterior part of the body, which is typical of congeners; by having both dorsolateral stripes separated each other by five to six scale rows on anterior part of body; and other scalation characters as well as the number maxillary teeth. The importance of this snake and its conservation is discussed.

Identification of leopard frogs (Anura: Ranidae: Lithobates) distributed in some localities of the Southern Mexican Plateau using mitochondrial DNA sequences.

Species of the genus Lithobates are difficult to identify, especially on the 'Rana pipiens' complex, because the morphological differences among the species are often subtle. In this study, we used 12S ribosomal RNA gene partial sequences to identify specimens of leopard frogs from some new localities (not sampled on previous studies) of the Southern Mexican Plateau, to carry out a phylogenetic analysis, and also a morphometric analysis of some morphological features were analyzed to evaluate their morphological variation. A phylogenetic analysis using partial sequences of 12S rDNA mitochondrial gene for 31 samples was carried out using Bayesian Inference, Maximum Parsimony, and Maximum Likelihood. In addition, 20 morphological lineal measurements from 97 specimens were analyzed by principal component analysis (PCA) to evaluate if the frogs studied are conspecific. Partial sequences of the 12S rDNA obtained from frogs distributed in the Southern Mexican Plateau show two haplotypes with a divergence of 0.4%. Phylogenetic hypothesis shows an exclusive group with a previous sequence of Lithobates montezumae. The PCA indicates that variables are not linearly correlated and specimens belong to a single group. Evidence found, let us consider that specimens studied belong to one species of the Lithobates montezumae subgroup, distributed principally in the Southern Mexican Plateau.

Solitary ecology as a phenomenon extending beyond insular systems: exaptive evolution in Anolis lizards.

The mechanisms driving phenotypic evolution have been of interest to biologists since Darwin. Ecological release-wherein adaptive evolution occurs following relaxation of constraining selective pressures-and environmental filtering-wherein exaptive traits allow colonization of a new area-have been studied in several insular cases. Anolis lizards, which may exist in solitude or sympatry with multiple congeners, are an excellent system for evaluating whether ecological release and environmental filtering are associated with phenotypic shifts across phylogenetic and geographical scales. Insular solitary Anolis exhibit phenotypic differentiation in body size and sexual size dimorphism-SSD-through exaptive and adaptive evolution, respectively. But, the generality of these effects has not yet been addressed. Here, we analyse the evolution of body size and SSD relative to sympatry in mainland Anolis. We found that mainland species co-occurring with few congeners exhibit uniform body size and greater SSD relative to other random mainland assemblages, consistent with the insular solitary pattern. The locations of evolutionary shifts for both traits do not coincide with evolutionary transitions to decreased levels of sympatry. These results are consistent with exaptive environmental filtering but not adaptive ecological release. Future studies should be conducted at local scales to evaluate the role of these factors in the evolution of solitary existence in mainland and island species.

Environmental heterogeneity explains coarse-scale ß-diversity of terrestrial vertebrates in Mexico.

We explored the hypothesis that high ß-diversity of terrestrial vertebrates of Mexico is associated with a high environmental heterogeneity (HEH) and identify the drivers of ß-diversity at different spatial scales. We used distribution range maps of 2,513 species of amphibians, reptiles, mammals, and birds occurring in Mexico. We estimated ß-diversity for each taxon at four spatial scales (grid cells of 2°, 1°, 0.5° and 0.25°) using the multiplicative formula of Whittaker ßw. For each spatial scale, we derived 10 variables of environmental heterogeneity among cells based on raw data of temperature, precipitation, elevation, vegetation and soil. We applied conditional autoregressive models (CAR) to identify the drivers of ß-diversity for each taxon at each spatial scale. CARs increased in explanatory power from fine-to-coarse spatial scales in amphibians, reptiles and mammals. The heterogeneity in precipitation including both, coefficient of variation (CV) and range of values (ROV), resulted in the most important drivers of ß-diversity of amphibians; the heterogeneity in temperature (CV) and elevation (ROV) were the most important drivers of ß-diversity for reptiles; the heterogeneity in temperature (ROV) resulted in the most important driver in ß-diversity for mammals. For birds, CARs resulted significant at fine scales (grid cells of 0.5° and 0.25°), and the precipitation (ROV and CV), temperature (ROV), and vegetation (H) and soil (H) were heterogeneity variables retained in the model. We found support for the hypothesis of environmental heterogeneity (HEH) for terrestrial vertebrates at coarse scales (grid cell of 2°). Different variables of heterogeneity, mainly abiotic, were significant for each taxon, reflecting physiological differences among terrestrial vertebrate groups. Our study revealed the importance of mountain areas in the geographic patterns of ß-diversity of terrestrial vertebrates in Mexico. At a coarse scale, specific variables of heterogeneity can be used as a proxy of ß-diversity for amphibians and reptiles.

Evidence for divergent patterns of local selection driving venom variation in Mojave Rattlesnakes (Crotalus scutulatus).

Snake venoms represent an enriched system for investigating the evolutionary processes that lead to complex and dynamic trophic adaptations. It has long been hypothesized that natural selection may drive geographic variation in venom composition, yet previous studies have lacked the population genetic context to examine these patterns. We leverage range-wide sampling of Mojave Rattlesnakes (Crotalus scutulatus) and use a combination of venom, morphological, phylogenetic, population genetic, and environmental data to characterize the striking dichotomy of neurotoxic (Type A) and hemorrhagic (Type B) venoms throughout the range of this species. We find that three of the four previously identified major lineages within C. scutulatus possess a combination of Type A, Type B, and a 'mixed' Type A + B venom phenotypes, and that fixation of the two main venom phenotypes occurs on a more fine geographic scale than previously appreciated. We also find that Type A + B individuals occur in regions of inferred introgression, and that this mixed phenotype is comparatively rare. Our results support strong directional local selection leading to fixation of alternative venom phenotypes on a fine geographic scale, and are inconsistent with balancing selection to maintain both phenotypes within a single population. Our comparisons to biotic and abiotic factors further indicate that venom phenotype correlates with fang morphology and climatic variables. We hypothesize that links to fang morphology may be indicative of co-evolution of venom and other trophic adaptations, and that climatic variables may be linked to prey distributions and/or physiology, which in turn impose selection pressures on snake venoms.

Systematics of the frogs allocated to Sarcohyla bistincta sensu lato (Cope, 1877), with description of a new species from Western Mexico.

A new species of hylid frog is described from the southwestern edge of the Mexican Plateau from the states of Morelos and Mexico through Michoacán and Jalisco, reaching the Sierra Madre Occidental in Sinaloa and western Durango. The new species is part of the widespread Mexican hylid Sarcohyla bistincta (sensu amplo) complex, comprised of S. bistincta, S. pentheter, S. calthula, and S. ephemera. One subspecies of S. bistincta (labeculata) was proposed for an isolated population in Oaxaca. We restrict the group's nominal species, S. bistincta (sensu stricto), to the Sierra Madre Oriental of Mexico and southward into the Sierra Madre del Sur of Guerrero and Oaxaca. Examination of type material places S. calthula and S. ephemera in the synonymy of S. labeculata (new combination). The species allied to S. bistincta, namely, S. bistincta, S. labeculata, S. pentheter, and the new species described herein, are diagnosed and described following recent suggested taxonomic changes and new available material.

An updated checklist of the herpetofauna from Guerrero, Mexico.

We present an updated checklist of the amphibians and reptiles for the state of Guerrero, Mexico. This list is based on bibliographic records, the revision of scientific collections and the authors' field notes. The herpetofauna of the state includes 259 species (78 amphibians, 181 reptiles), of which three are introduced. The conservation status of all native species was assessed using three independent systems (NOM-059, IUCN and EVS). Finally, the geographic distribution of amphibians and reptiles is discussed in the context of both the biogeographic provinces within the state and the influence of road location on specimen collection patterns.

Cryptic genetic diversity, population structure, and gene flow in the Mojave rattlesnake (Crotalus scutulatus).

The Mojave rattlesnake (Crotalus scutulatus) inhabits deserts and arid grasslands of the western United States and Mexico. Despite considerable interest in its highly toxic venom and the recognition of two subspecies, no molecular studies have characterized range-wide genetic diversity and population structure or tested species limits within C. scutulatus. We used mitochondrial DNA and thousands of nuclear loci from double-digest restriction site associated DNA sequencing to infer population genetic structure throughout the range of C. scutulatus, and to evaluate divergence times and gene flow between populations. We find strong support for several divergent mitochondrial and nuclear clades of C. scutulatus, including splits coincident with two major phylogeographic barriers: the Continental Divide and the elevational increase associated with the Central Mexican Plateau. We apply Bayesian clustering, phylogenetic inference, and coalescent-based species delimitation to our nuclear genetic data to test hypotheses of population structure. We also performed demographic analyses to test hypotheses relating to population divergence and gene flow. Collectively, our results support the existence of four distinct lineages within C. scutulatus, and genetically defined populations do not correspond with currently recognized subspecies ranges. Finally, we use approximate Bayesian computation to test hypotheses of divergence among multiple rattlesnake species groups distributed across the Continental Divide, and find evidence for co-divergence at this boundary during the mid-Pleistocene.

Taxonomic changes and description of two new species for the Phyllodactylus lanei complex (Gekkota: Phyllodactylidae) in Mexico.

An integrative taxonomy approach was implemented based on analysis of genetic, phylogenetic, morphological and ecological data to identify the cryptic diversity within the Phyllodactylus lanei complex. At least six species can be identified, of which four are currently considered subspecies: Phyllodactylus lanei, Phyllodactylus rupinus, Phyllodactylus isabelae, Phyllodactylus lupitae and two corresponding to undescribed taxa, which are identified and described in this contribution. These differ from other Mexican geckos in several characters: genetic distance (DNAmt), position in molecular phylogeny (concatened data DNAmt+DNAnu), species tree, morphological characters such as snout-vent length, longitudinal scales, tubercles from head to tail, interorbital scales, scales across venter, third labial-snout scales and rows of tubercles across dorsum; there are also differences in their bioclimatic profiles (temperature and precipitation) and geographical distribution. The most recent studies on taxonomy and evolution of Mexican geckos (Phyllodactylus) show that the diversity of this group of reptiles is currently underestimated, suggesting that more research and conservation efforts are should be addressed at these lizards.

Phylogenomic insights into the diversification of salamanders in the Isthmura bellii group across the Mexican highlands.

Mountain formation in Mexico has played an important role in the diversification of many Mexican taxa. The Trans-Mexican Volcanic Belt in particular has served as both a cradle of diversification and conduit for dispersal. We investigated the evolutionary history of the Isthmura bellii group of salamanders, a widespread amphibian across the Mexican highlands, using sequence capture of ultraconserved elements. Results suggest that the I. bellii group probably originated in southeastern Mexico in the late Miocene and later dispersed across the Trans-Mexican Volcanic Belt and into the Sierra Madre Occidental. Pre-Pleistocene uplift of the Trans-Volcanic Belt likely promoted early diversification by serving as a mesic land-bridge across central Mexico. These findings highlight the importance of the Trans-Volcanic Belt in generating Mexico's rich biodiversity.