Corallus hortulanus testes histology: morphological and reproductive aspects.

In general snakes show differentiate anatomical, biological and behavioral particularities compared to other species. Basic information about the snakes anatomy, physiology and reproductive biology is scarce in several species, making the reproduction a challenge. Thus, the present work aims to evaluate morphological aspects of the Corallus hortulanus testes, correlating these findings with environmental factors and reproductive aspects. The testes of three specimens of Corallus hortulanus were cut to a thickness of 3µm in microtome, stained with 1% toluidine blue, photo documented and described. Seasonality was observed in the sperm production of Corallus hortulanus, with the presence of mature spermatozoa in the wettest and hottest periods of the year, as well as the largest testicular volume in these periods.

Comparative morphology of oral glands in snakes of the family Homalopsidae reveals substantial variation and additional independent origins of salt glands within Serpentes.

Using diffusible iodine-based contrast-enhanced computed tomography (diceCT), we examined the morphology of the oral glands of 12 species of the family Homalopsidae. Snakes of this family exhibit substantial interspecific morphological variation in their oral glands. Particular variables are the venom glands, ranging from large (e.g., Subsessor bocourti) to small (e.g., Erpeton tentaculatum). The supra- and infralabial glands are more uniform in morphology, being the second most developed in almost all the sampled species. Premaxillary glands distinct from the supralabial glands were observed in five species (Myron richardsonii, Bitia hydroides, Cantoria violacea, Fordonia leucobalia, and Gerarda prevostiana), in addition to Cerberus rynchops, the only species in which this condition was previously documented associated with the excretion of salt. In the three species of the saltwater group of homalopsids (C. violacea, F. leucobalia, and G. prevostiana), the premaxillary glands also extend posteriorly, occupying a large area above the supralabial gland, a condition not observed in any other species of snake studied thus far. Character evolution analyses indicate that premaxillary glands differentiated from the supralabial gland and evolved independently three or four times in the family, always in lineages that invaded marine habitats. Our results suggest that the differentiated premaxillary glands are likely salt glands, as is the case in C. rynchops. If corroborated, this increases to six or seven the number of independent evolutionary origins of salt glands in snakes that have undergone an evolutionary transition to marine life.

When Food Fights Back: Skull Morphology and Feeding Behavior of Centipede-Eating Snakes.

Feeding is a complex process that involves an integrated response of multiple functional systems. Animals evolve phenotypic integration of complex morphological traits to covary and maximize performance of feeding behaviors. Specialization, such as feeding on dangerous prey, can further shape the integration of behavior and morphology as traits are expected to evolve and maintain function in parallel. Feeding on centipedes, with their powerful forcipules that pinch and inject venom, has evolved multiple times within snakes, including the genus Tantilla. However, the behavioral and morphological adaptations used to consume this dangerous prey are poorly understood. By studying snakes with varying degrees of dietary specialization, we can test the integration of diet, morphology, and behavior to better understand the evolution of consuming difficult prey. We studied the prey preference and feeding behavior of Tantilla using the flat-headed snake (T. gracilis) and the crowned snake (T. coronata), which differ in the percentage of centipedes in their diet. We then quantified cranial anatomy using geometric morphometric data from CT scans. To test prey preference, we offered multiple types of prey and recorded snake behavior. Both species of snakes showed interest in multiple prey types, but only struck or consumed centipedes. To subdue centipedes, crowned snakes used coiling and holding (envenomation) immediately after striking, while flat-headed snakes used the novel behavior of pausing and holding onto centipedes for a prolonged time prior to the completion of swallowing. Each skull element differed in shape after removing the effects of size, position, and orientation. The rear fang was larger in crowned snakes, but the mechanical advantage of the lower jaw was greater in flat-headed snakes. Our results suggest that the integration of behavioral and morphological adaptations is important for the success of subduing and consuming dangerous prey.

Diet and habit explain head-shape convergences in natricine snakes.

The concept of ecomorphs, whereby species with similar ecologies have similar phenotypes regardless of their phylogenetic relatedness, is often central to discussions regarding the relationship between ecology and phenotype. However, some aspects of the concept have been questioned, and sometimes species have been grouped as ecomorphs based on phenotypic similarity without demonstrating ecological similarity. Within snakes, similar head shapes have convergently evolved in species living in comparable environments and/or with similar diets. Therefore, ecomorphs could exist in some snake lineages, but this assertion has rarely been tested for a wide-ranging group within a single framework. Natricine snakes (Natricinae) are ecomorphologically diverse and currently distributed in Asia, Africa, Europe and north-central America. They are primarily semiaquatic or ground-dwelling terrestrial snakes, but some are aquatic, burrowing or aquatic and burrowing in habit and may be generalist or specialist in diet. Thus, natricines present an interesting system to test whether snakes from different major habit categories represent ecomorphs. We quantify morphological similarity and disparity in head shape among 191 of the ca. 250 currently recognized natricine species and apply phylogenetic comparative methods to test for convergence. Natricine head shape is largely correlated with habit, but in some burrowers is better explained by dietary specialism. Convergence in head shape is especially strong for aquatic burrowing, semiaquatic and terrestrial ecomorphs and less strong for aquatic and burrowing ecomorphs. The ecomorph concept is useful for understanding natricine diversity and evolution, though would benefit from further refinement, especially for aquatic and burrowing taxa.

Molecular phylogeny and morphological comparisons of the genus Hebius Thompson, 1913 (Reptilia: Squamata: Colubridae) uncover a new taxon from Yunnan Province, China, and support revalidation of Hebius septemlineatus (Schmidt, 1925).

We describe a new species of the genus Hebius and provide evidence for the validity of H. septemlineatuscomb. nov. Morphological and molecular analyses of Hebius specimens collected in Yunnan Province, China, revealed three distinct lineages, namely the newly described Hebius weixiensissp. nov., as well as H. octolineatus (Boulenger, 1904), and H. septemlineatuscomb. nov. (Schmidt 1925), which is removed from synonymy with H. octolineatus. Based on mitochondrial genealogy, Hebius weixiensissp. nov. is sister to H. septemlineatuscomb. nov., while H. octolineatus is sister to H. bitaeniatus. The new species and H. septemlineatuscomb. nov. showed considerable genetic divergence from their recognized congeners (uncorrected P-distance ≥3.9%). Furthermore, the new species and H. septemlineatuscomb. nov. can be diagnosed from closely related congeners by a combination of pholidosis characters.

A new snake species of the genus Gonyosoma Wagler, 1828 (Serpentes: Colubridae) from Hainan Island, China.

A new species of the genus Gonyosoma Wagler, 1828 is described herein based on six specimens from the Diaoluoshan Mountains, Hainan Island, Hainan Province, China. The new species, Gonyosoma hainanensesp. nov., is most similar to its continental sister species, Gonyosoma boulengeri (Mocquard, 1897). Both taxa have a scaled protrusion on the anterior portion of the rostrum, distinct from other congeners. However, Gonyosoma hainanensesp. nov. can be distinguished from G. boulengeri by two significant morphological characters: (1) black orbital stripe absent in adults (vs. present in G. boulengeri); and (2) two loreals (vs. one loreal in G. boulengeri). The new species is also genetically divergent and forms a unique clade from its sister species and all other congeners based on sequences of the mitochondrial gene cytochrome b (cyt b).

A new species of Ahaetulla Link, 1807 (Squamata: Colubridae) from the Mekong Delta, Vietnam.

A new vine snake, genus Ahaetulla, from Soc Trang Province, southern Vietnam is described based on morphological data and nucleotide sequences from COI and Cytb. Ahaetulla rufusoculara sp. nov. is diagnosed by the following morphological characters: bright red eyes; snout without dermal appendage; internasal separated from supralabial by loreal; body scale rows 15-15-13; ventrals 186190 in males and 182185 in females; subcaudals 143153 in males and 128 or 129 in females; 8 (rarely 9) supralabials, fourth and fifth or fourth to sixth entering orbit; 1214 maxillary teeth; hemipenis short, reaching 6th or 7th SC; cloacal plate divided; dorsum bright green; and yellow or white stripe along the lower flank. The new species differs from its congeners by an uncorrected p-distance in COI and Cytb sequences of at least 7.7% and 7.5%, respectively.

A new species of Boiga (Serpentes: Colubridae) from the Southern Western Ghats of India with a molecular phylogeny and expanded characterisation of related species.

We conducted a molecular phylogenetic study on the Boiga ceylonensis group from the Western Ghats of India, building on a recent morphology-based study. Our analysis supports previous work in elucidating the phylogenetic position of B. nuchalis and B. beddomei s. str. (of Matheran, Western Ghats) as clustering closely with B. ceylonensis, while B. flaviviridis clustered with the Sri Lankan taxon close to B. ranawanei. Additionally, our phylogenetic study revealed the presence of an undescribed taxon in the Southern Western Ghats, more closely related to B. ceylonensis than to any other sampled taxon, including sympatric congeners. This new taxon is described here as a new species Boiga whitakeri sp. nov. from the Devar MalaiAnaimalai hill complex. Based on molecular and morphometric studies, we expand the description of the recently described B. thackerayi as inhabiting almost the entire stretch of the Western Ghats. We also complement the diagnoses of B. nuchalis and B. flaviviridis, and determine the phylogenetic position of B. thackerayi with molecular data.

Night stalkers from above: A monograph of Toxicodryas/ tree snakes (Squamata: Colubridae) with descriptions of two new cryptic species from Central Africa.

The genus Toxicodryas, historically included with the renowned Australasian cat-eyed snakes of the colubrid genus Boiga, currently includes two widespread species (T. blandingii and T. pulverulenta) in western, central, and eastern Africa. We leverage findings from a recent phylogenomic and historical demographic analysis of this genus (based on 2848-4471 Rad-seq loci from across the genome), with robust sampling from throughout the ranges of both species, to define two additional taxonomic units, with species boundaries corresponding to river barriers. Additional morphometric data from scores of examined museum specimens and literature records bolster the recognition of these two new cryptic species. We hypothesize that T. blandingii occurs west of the confluence of the Congo and Ubangi rivers, whereas a cryptic new species that is found east of this biogeographic barrier has significantly higher numbers of ventral scale counts in both sexes, additional significant differences in several scale counts, and lower venom toxicity. Toxicodryas pulverulenta occurs west of the Niger Delta in West Africa, whereas a cryptic new species that is found east of this biogeographic barrier has significantly higher numbers of subcaudal scale counts in both sexes. A review of published information regarding morphological variation, ecology, natural history, habitat, and venom is summarized for these four Toxicodryas species.

Structural and ultrastructural studies on the developing vomeronasal sensory epithelium in the grass snake Natrix natrix (Squamata: Colubroidea).

The sensory olfactory epithelium and the vomeronasal sensory epithelium (VSE) are characterized by continuous turnover of the receptor cells during postnatal life and are capable of regeneration after injury. The VSE, like the entire vomeronasal organ, is generally well developed in squamates and is crucial for detection of pheromones and prey odors. Despite the numerous studies on embryonic development of the VSE in squamates, especially in snakes, an ultrastructural analysis, as far as we know, has never been performed. Therefore, we investigated the embryology of the VSE of the grass snake (Natrix natrix) using electron microscopy (SEM and TEM) and light microscopy. As was shown for adult snakes, the hypertrophied ophidian VSE may provide great resolution of changes in neuron morphology located at various epithelial levels. The results of this study suggest that different populations of stem/progenitor cells occur at the base of the ophidian VSE during embryonic development. One of them may be radial glia-like cells, described previously in mouse. The various structure and ultrastructure of neurons located at different parts of the VSE provide evidence for neuronal maturation and aging. Based on these results, a few nonmutually exclusive hypotheses explaining the formation of the peculiar columnar organization of the VSE in snakes were proposed.

Contribution to the taxonomy of the genus Lycodon H. Boie in Fitzinger, 1827 (Reptilia: Squamata: Colubridae) in China, with description of two new species and resurrection and elevation of Dinodon septentrionale chapaense Angel, Bourret, 1933.

While considerable progress has been made in the taxonomic studies of the genus Lycodon in Asia, questions remain to be clarified regarding the taxonomy of certain groups, particularly those containing species in China. Not only do many regions in China remain overlooked by herpetologists, resulting in the possibility of undiscovered new species, but the surveyed areas also have suspicious records of recognized congeners that require taxonomic confirmations. Combining both morphological and genetic data, we tackle these outstanding issues in the taxonomy of Lycodon in China. In particular, we discover two new species of Lycodon: one from the previously neglected hot-dry valley in the northern Hengduan Mountain Region close to Tibet, and another recluse and cryptic species from the L. fasciatus complex in the downtown park of a major city in southern Sichuan Province. Additionally, we clarify the distribution of L. septentrionalis in China and resurrect and elevate its junior synonym subspecies, Dinodon septentrionale chapaense, as a full, valid species, and we synonymize the recently described L. namdongensis to the resurrected L. chapaensis comb. nov.. Lycodon chapaensis comb. nov. thus represents a new national record of reptilian fauna of China. Lastly, based on literature review, we also correct some of the erroneous records of L. fasciatus and L. ruhstrati in China, point out remaining taxonomic issues of the genus for future research, and update the dichotomous key and distribution of the 20 species of Lycodon currently recorded from China.

Taxonomic revision of the genus Xenopholis Peters, 1869 (Serpentes: Dipsadidae): Integrating morphology with ecological niche.

A reliable identification and delimitation of species is an essential pre-requisite for many fields of science and conservation. The Neotropical herpetofauna is the world's most diverse, including many taxa of uncertain or debated taxonomy. Here we tackle one such species complex, by evaluating the taxonomic status of species currently allocated in the snake genus Xenopholis (X. scalaris, X. undulatus, and X. werdingorum). We base our conclusions on concordance between quantitative (meristic and morphometric) and qualitative (color pattern, hemipenes and skull features) analyses of morphological characters, in combination with ecological niche modeling. We recognize all three taxa as valid species and improve their respective diagnosis, including new data on color in life, pholidosis, bony morphology, and male genitalia. We find low overlap among the niches of each species, corroborating the independent source of phenotypic evidence. Even though all three species occur in the leaf litter of distinct forested habitats, Xenopholis undulatus is found in the elevated areas of the Brazilian Shield (Caatinga, Cerrado and Chaco), whereas X. scalaris occurs in the Amazon and Atlantic rainforests, and X. werdingorum in the Chiquitanos forest and Pantanal wetlands. We discuss the disjunct distribution between Amazonian and Atlantic Forest snake species in the light of available natural history and ecological aspects. This study shows the advantages of combining multiple data sources for reliable identification and circumscription of ecologically similar species.

Mandibular sawing in a snail-eating snake.

The jaws of vertebrates display a striking diversity in form and function, but they typically open and close like a trapdoor rather than sliding like a saw. Here, we report unique feeding behaviour in the blunt-headed snail-eating snake, Aplopeltura boa (family Pareidae), where the snake cuts off and circumvents the indigestible part (the operculum) of its prey in the mouth using long sliding excursions of one side of the mandible, while the upper jaws and the mandible on the other side maintain a stable grasp on the prey. This behaviour, which we call 'mandibular sawing', is made possible by extraordinarily independent movements of the jaw elements and is a surprising departure from usual feeding behaviour in vertebrates.

Skull osteology and osteological phylogeny of the Western whip snake Hierophis viridiflavus (Squamata, Colubridae).

The skull osteology of Hierophis viridiflavus is here described and figured in detail on the basis of 18 specimens. The sample includes specimens from the ranges of both H. viridiflavus viridiflavus and H. viridiflavus carbonarius as well as specimens not identified at sub-specific level. The main characters that define H. viridiflavus in comparison to the parapatric congeneric species Hierophis gemonensis are wide maxillary diastema, basioccipital crest well distinct in three lobes and basioccipital process well marked. The foramina of the otoccipital and prootic, and the basioccipital process of the basioccipital are among the most ontogenetically variable characters, as indicated by two juvenile specimens included in the sample. A specimen-level phylogenetic analysis including H. gemonensis and other outgroups (overall 6 species, 26 specimens, 64 skull characters) recovered all H. viridiflavus specimens in one clade, indicating the presence of a clear phylogenetic signal in the applied characters. However, the resolution within the H. viridiflavus clade is poor the monophyly of H. viridiflavus carbonarius was retrieved, but not that of Hierophis v. viridiflavus. Probably due to the relatively high variability, the skull morphology does not support the recently proposed specific status of the two subspecies.

Captivity Affects Head Morphology and Allometry in Headstarted Garter Snakes, Thamnophis sirtalis.

In response to the growing number of amphibian and reptiles species in decline, many conservation managers have implemented captive breeding and headstarting programs in an effort to restore these populations. However, many of these programs suffer from low survival success, and it is often unclear as to why some individuals do not survive after reintroduction. Here I document changes to head morphology in the eastern garter snake, Thamnophis sirtalis, in response to time spent in captivity. Thamnophis raised on three diet treatments all differed in head size from wild individuals, and head size differed between the three treatments. Overall, head size was smaller in all three diet treatments than in wild snakes, potentially limiting the available prey for the captive garter snakes. Allometric patterns of growth in head size were also different for each diet treatment. Several potential implications of these changes in morphology are discussed, and what these changes may mean for other species that are part of headstarting and reintroduction programs.

Relative tail length correlates with body condition in male but not in female crowned leafnose snakes (Lytorhynchus diadema).

Reproductive success is the ultimate measure of individual quality; however, it is difficult to determine in free-living animals. Therefore, indirect measures that are related to reproduction are generally employed. In snakes, males typically possess longer tails than females and this sexual size dimorphism in tail length (TL) has generally been attributed to the importance of the tail in mating and reproduction. Thus, intra-sexual differences in tail length, specifically within males, were hypothesized to reflect individual quality. We used a body condition index (BCI) as a measure of quality in snakes and predicted that tail length would be correlated with BCI in males. We tested our prediction by determining BCI in the free-ranging adult male and female crowned leafnose snake (Lytorhynchus diadema), a colubrid species that inhabits mainly desert sand dunes. The relative TL was correlated positively and significantly to BCI in males (F1,131 = 11.05; r2adj = 0.07; P < 0.01) but not in females, thus supporting our prediction. This is the first time that the relationship between TL and body condition was tested in a free-ranging species. In addition, sexual size dimorphism of TL increased intra-specifically with body size, which was also found in interspecific analyses following Rensch's rule.

Role of phenotypic plasticity in morphological differentiation between watersnake populations.

An individual's morphology is shaped by the environmental pressures it experiences, and the resulting morphological response is the culmination of both genetic factors and environmental (non-genetic) conditions experienced early in its life (i.e. phenotypic plasticity). The role that phenotypic plasticity plays in shaping phenotypes is important, but evidence for its influence is often mixed. We exposed female neonate diamond-backed watersnakes (Nerodia rhombifer) from populations experiencing different prey-size regimes to different feeding treatments to test the influence of phenotypic plasticity in shaping trophic morphology. We found that snakes in a large-prey treatment from a population frequently encountering large prey exhibited a higher growth rate in body size than individuals in a small-prey treatment from the same population. This pattern was not observed in snakes from a population that regularly encounters small prey. We also found that regardless of treatment, snakes from the small-prey population were smaller at birth than snakes from the large-prey population and remained so throughout the study. These results suggest that the ability to plastically respond to environmental pressures may be population-specific. These results also indicate a genetic predisposition towards larger body sizes in a population where large prey items are more common.

Variation in the formation of crista sellaris and basisphenoid in the skull of the grass snake Natrix natrix embryos (Serpentes, Colubridae).

The crista sellaris is one of the basic structures of the snake chondrocranium. In embryogenesis it ossifies, forming the basisphenoid. This article describes variations of the crista sellaris, which, in turn, leads to variation in the formation of the basisphenoid in the skull of grass snake Natrix natrix (Serpentes, Colubridae) embryos. Several embryos at different developmental stages are investigated. Embryos from one developmental stage are taken simultaneously from one clutch. Thus, variation of the crista sellaris and basisphenoid in the skull of embryos is discovered, including those from one clutch. In several embryos, the crista sellaris is absent or partially formed. The absence of the crista sellaris does not preclude the formation of the basisphenoid. Based on the observed variations of the crista sellaris, the basisphenoid may have a different origin. In embryos with formed crista sellaris, the basisphenoid develops as a complex endochondral bone from three ossification centers (paired-in the polar cartilages and unpaired in the crista sellaris). In embryos without fully formed crista sellaris the basisphenoid develops as a paired bone of mixed origin: endochondral ossification appears in the polar cartilages, but where the crista sellaris is absent, it is formed by dermal ossification. In general, we observed the absence of the crista sellaris in 15% of studied embryos. We assume that it may be due to a reduction of the orbital-temporal region of the chondrocranium in N. natrix embryos. RESEARCH HIGHLIGHTS: The variation of the crista sellaris and basisphenoid formation in grass snake embryos is revealed. Depending on the presence or absence of the crista sellaris, basisphenoid is formed as a complex chondral bone or as a paired bone of mixed origin.

The scaling of terrestrial striking performance in western ratsnakes (Pantherophis obsoletus).

In many organisms, juveniles have performance capabilities that partly offset their disadvantageous sizes. Using high-speed video recordings and imaging software, we measured the scaling of head morphology, axial morphology, and defensive strike performance of Pantherophis obsoletus across their ontogeny to understand how size and morphology affect performance. Head measurements were negatively allometric whereas the cross-sectional area (CSA) of epaxial muscles displayed positive allometry. The greater relative muscle CSA of larger ratsnakes allows them to produce higher forces relative to their mass, and those forces act on a relatively smaller head mass when it is thrust forward during striking. Maximum strike accelerations of 70-273.8 ms-2 and velocities of 1.08-3.39 ms-1 scaled positively with body mass but differed from the geometric predictions. Velocity scaled with mass0.15 and acceleration scaled with mass0.17 . Larger snakes struck from greater distances (range = 4.1-26 cm), but all snakes covered the strike distances with similarly short durations (84 ± 3 ms). The negatively allometric head size, isometry of anterior mass, and positively allometric muscle CSA enable larger P. obsoletus to strike with higher velocities and accelerations than smaller individuals. Our results contrast with the scaling of strike performance in an arboreal viper, whose strike distance and velocity were independent of body mass. When strike distance is modulated, all other performance capacities are affected because of the interdependence of acceleration, velocity, duration, and distance.

Morphology-based intraspecific taxonomy of Oreocryptophis porphyraceus (Cantor, 1839) in mainland China (Serpentes: Colubridae).

In this study, a total of 106 individuals of Oreocryptophis porphyraceus from mainland China were morphologically examined and recorded. Differences between populations were compared by combining data from this study and other published research. The skulls of three specimens representing three proposed subspecies (i.e., O. p. pulchra, O. p. vaillanti, and O. p. hainana) were examined by computed tomography (CT) scanning. Both external morphological characters and skull comparisons consistently showed significant differences between the studied populations. Based on these data, we suggest that at least four subspecies of O. porphyraceus should be recognized in mainland China: i.e., O. p. porphyraceus, O. p. pulchra, O. p. vaillanti, and O. p. hainana. However, the taxonomical arrangement of the central Chinese populations with intermediate morphology remain unresolved.