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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.

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., (© 2024 Anatomical Society.)

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., (© The Author(s) 2023. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology.)

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., (© 2022 The Authors. Journal of Evolutionary Biology published by John Wiley & Sons Ltd on behalf of European Society for Evolutionary Biology.)

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.

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.

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.

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., (© 2020 Wiley Periodicals LLC.)

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.

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., Competing Interests: No. The authors have declared that no competing interests exist.

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., (© The Author(s) 2020. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. All rights reserved. For permissions please email: journals.permissions@oup.com.)

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.

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., (© 2020 International Society of Zoological Sciences, Institute of Zoology/Chinese Academy of Sciences and John Wiley & Sons Australia, Ltd.)

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., (© 2020 Wiley Periodicals, Inc.)

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 (F 1,131  = 11.05; r 2 adj  = 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.

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., (© 2020 Wiley Periodicals, Inc.)

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 mass 0.15 and acceleration scaled with mass 0.17 . 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., (© 2019 Wiley Periodicals, Inc.)

Most colubrid snake venoms have been poorly studied, despite the fact that they represent a great resource for biological, ecological, toxinological and pharmacological research. Herein, we explore the venom delivery system of the Aesculapian False Coral Snake Erythrolamprus aesculapii as well as some biochemical and toxicological properties of its venom. Its Duvernoy's venom gland is composed of serous secretory cells arranged in densely packed secretory tubules, and the most striking feature of its fang is their double-curved shape, exhibiting a beveled bladelike appearance near the tips. Although E. aesculapii resembles elapid snakes of the genus Micrurus in color pattern, this species produces a venom reminiscent of viperid venoms, containing mainly tissue-damaging toxins such as proteinases. Prominent hemorrhage developed both locally and systemically in mice injected with the venom, and the minimum hemorrhagic dose was found to be 18.8 μg/mouse; the lethal dose, determined in mice, was 9.5 ± 3.7 μg/g body weight. This work has toxicological implications that bites to humans by E. aesculapii could result in moderately severe local (and perhaps systemic) hemorrhage and gives insight into future directions for research on the venom of this species., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

We analyzed some aspects of reproduction and sexual dimorphism of the semi-aquatic dipsadid snake Erythrolamprus miliaris in the state of Rio de Janeiro, southeastern Brazil. We detected sexual dimorphism in body size (snout-vent length), with females averaging larger than males, but no sexual dimorphism in the relative length of the tail. Oviductal eggs and secondary follicles were found in all seasons, suggesting that female reproductive cycles are continuous, in spite of the tropical seasonal climate in the region. Reproductive males were present throughout the year, suggesting a continuous cycle for males as well. Clutch size averaged 10.3 ± 4.8 (range 4-21) and there was a positive and significant relationship between clutch size and female size. Compared to conspecific populations previously studied in other Atlantic Rainforest areas, populations of E. miliaris from the state of Rio de Janeiro appear more similar overall in their reproductive traits to a more northern population from the state of Bahia than to populations from further south in the states of São Paulo and Paraná.

Herein we investigated diet, sexual dimorphism and reproductive biology of two sympatric congeneric species at the coastal Brazilian Pampa, Philodryas aestiva and P. patagoniensis. Analysis of the stomach content of the scansorial Philodryas aestiva revealed that it feeds mainly on Anura and small mammals, while it occasionally consumes Insecta, Squamata and Aves. The more terrestrial Philodryas patagoniensis presented a broader diet, composed mainly of Anura, Squamata and small mammals. Insecta and Actinopterygii were found in lower frequencies. In females of both species, snout-vent length was significantly larger than in males, while tail length/snout-vent length ratio was significantly higher in males. Males reach sexual maturity at smaller sizes than females in both species. Fecundity was higher in P. patagoniensis (3 to 24 eggs; 15 ± 8.15) than in P. aestiva (10 to 20 eggs; X = 14.50 ± 3.53). Females of P. aestiva showed secondary follicles in all seasons while eggs occurred from middle-winter to early-summer. Secondary follicles occurred throughout the year in P. patagoniensis, but concentrated in spring. Eggs occurred in late-summer, middle-autumn and spring. Both species presented positive correlations between SVL and clutch size. Differences in analyzed traits seem to reflect divergences in species morphology and use of habitat/microhabitat.

The Japanese rat snake (Elaphe climacophora) is a common species in Japan and is widely distributed across the Japanese islands. An albino mutant of the Japanese rat snake ("pet trade" albino) has been bred and traded by hobbyists for around two decades because of its remarkable light-yellowish coloration with red eyes, attributable to a lack of melanin. Another albino Japanese rat snake mutant found in a natural population of the Japanese rat snake at high frequency in Iwakuni City, Yamaguchi Prefecture is known as "Iwakuni no Shirohebi". It has been conserved by the government as a natural monument. The Iwakuni albino also lacks melanin, having light-yellowish body coloration and red eyes. Albino mutants of several organisms have been studied, and mutation of the tyrosinase gene (TYR) is responsible for this phenotype. By determining the sequence of the TYR coding region of the pet trade albino, we identified a nonsense mutation in the second exon. Furthermore, RT-PCR revealed that TYR transcripts were not detected in this snake. These findings suggest that mutation of TYR is responsible for the albino phenotype of the pet trade line of the Japanese rat snake. However, the Iwakuni albino did not share this TYR mutation; thus, these two albino lines differ in their origins.

Snakes are the major group of venomous vertebrates, and the rear-fanged snakes represent the vast majority of species and occur worldwide; however, relatively few studies have characterized their venoms and evaluated their potential hazards for humans. Herein we explore the protein composition and properties of the venom of the rear-fanged Green Parrot Snake, Leptophis ahaetulla marginatus, the most common snake found in the Iguazu National Park (Argentina), as well as the main features of its venom delivery system. This species has venom reminiscent of elapid venoms, composed mainly of components such as 3FTxs, CRiSPs and AChE, but it shows low toxicity toward mammals (LD 50  > 20 μg/g mouse). The histology of its Duvernoy's venom gland is similar to that of other colubrids, with serous secretory cells arranged in densely packed secretory tubules. The posterior end of its maxilla exhibits 1-3 blade-shaped and slightly recurved fangs but without grooves. This study provides an initial analysis of the biological role of venom in Leptophis, with implications for potential symptoms that might be anticipated from bites by this species., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

A snake's body represents an extreme degree of elongation with immense muscle complexity. Snakes have approximately 25 different muscles on each side of the body at each vertebra. These muscles serially repeat, overlap, interconnect, and rarely insert parallel to the vertebral column. The angled muscles mean that simple measurements of anatomical cross-sectional area (ACSA, perpendicular to the long-axis of the body) serve only as proxies for the primary determinant of muscle force, physiological cross-sectional area (PCSA, area perpendicular to the muscle fibers). Here, I describe and quantify the musculature of two intraguild constrictors: kingsnakes (Lampropeltis holbrooki) and ratsnakes (Pantherophis obsoletus) whose predation performance varies considerably. Kingsnakes can produce significantly higher constriction pressures compared with ratsnakes of similar size. In both snakes, I provide qualitative descriptions, detail previously undescribed complexity, identify a new lateral muscle, and provide some of the first quantitative measures of individual muscle and whole-body PCSA. Furthermore, I compare measurements of ACSA with measurements of PCSA. There was no significant difference in PCSA of muscles between kingsnakes and ratsnakes. There is, however, a strong relationship between ACSA and PCSA measurements. I could not identify a significant difference in musculature between kingsnakes and ratsnakes that explains their different levels of constriction performance. Unmeasured components of muscle function, such as endurance and force production, might account for differences in performance between two species with similar muscle structure., (© 2018 Anatomical Society.)

In this work, we performed a biogeographic analysis with Bayesian binary MCMC (BBM) statistical dispersal-vicariance analysis (S-DIVA) and species distribution models (SDM) on three phylogenetically closely related Mediterranean whipsnakes (Hierophis gemonensis, H. carbonarius, H. viridiflavus), to investigate the pathways of their geographical diversification and locate putative refugial areas in the last glacial maximum (LGM). Our analysis suggests that the diversification processes between the studied species overall followed an east-west route, from eastern Greece to the Iberian Peninsula and continental France, highlighting a significant role of dispersal and vicariance processes at both inter- and intraspecific levels. In particular, the main lineage-splitting events between H. gemonensis, H. carbonarius and H. viridiflavus coincide with two events of vicariance, involving respectively eastern Greece and eastern Italy, and eastern Italy and western Italy, Iberian Peninsula and continental France. SDM analyses highlight the occurrence of multiple putative glacial refugia in the Balkans, Italy and southern France, which represent well the occurrence of distinct "refugia within refugia" in the main Mediterranean peninsulas. Furthermore, our results suggest how these refugia may have played an important role during the Quaternary climatic oscillations in shaping the current haplotype distribution of European whipsnakes., (Copyright © 2018 Elsevier GmbH. All rights reserved.)

As body size increases, some predators eliminate small prey from their diet exhibiting an ontogenetic shift toward larger prey. In contrast, some predators show a telescoping pattern of prey size in which both large and small prey are consumed with increasing predator size. To explore a functional explanation for the two feeding patterns, I examined feeding effort as both handling time and number of upper jaw movements during ingestion of fish of consistent size. I used a range of body sizes from two snake species that exhibit ontogenetic shifts in prey size (Nerodia fasciata and N. rhombifer) and a species that exhibits telescoping prey size with increased body size (Thamnophis proximus). For the two Nerodia species, individuals with small or large heads exhibited greater difficulty in feeding effort compared to snakes of intermediate size. However, for T. proximus measures of feeding effort were negatively correlated with head length and snout-vent length (SVL). These data indicate that ontogenetic shifters of prey size develop trophic morphology large enough that feeding effort increases for disproportionately small prey. I also compared changes in body size among the two diet strategies for active foraging snake species using data gleaned from the literature to determine if increased change in body size and thereby feeding morphology is observable in snakes regardless of prey type or foraging habitat. Of the 30 species sampled from literature, snakes that exhibit ontogenetic shifts in prey size have a greater magnitude of change in SVL than species that have telescoping prey size patterns. Based upon the results of the two data sets above, I conclude that ontogenetic shifts away from small prey occur in snakes due, in part, to growth of body size and feeding structures beyond what is efficient for handling small prey., (Copyright © 2017. Published by Elsevier GmbH.)

The developmental mechanisms of color patterns formation and its evolution remain unclear in reptilian sauropsids. We, therefore, studied the pigment cell mechanisms of stripe pattern formation during embryonic development of the snake Elaphe quadrivirgata. We identified 10 post-ovipositional embryonic developmental stages based on external morphological characteristics. Examination for the temporal changes in differentiation, distribution, and density of pigment cells during embryonic development revealed that melanophores first appeared in myotome and body cavity but not in skin surface at Stage 5. Epidermal melanophores were first recognized at Stage 7, and dermal melanophores and iridophores appeared in Stage 9. Stripe pattern first appeared to establish at Stage 8 as a spatial density gradient of epidermal melanophores between the regions of future dark brown longitudinal stripes and light colored background. Our study, thus, provides a comprehensive pigment-cell-based understanding of stripe pattern formation during embryonic development. We briefly discuss the importance of the gene expression studies by considering the biologically relevant theoretical models with standard developmental staging for understanding reptilian color pattern evolution., (© 2017 Wiley Periodicals, Inc.)

A new species of natricine snake of the Southeast Asian genus Opisthotropis Gü nther, 1872 is described from western Hunan Province of China based on both mitochondrial DNA and morphological data. The new species is morphologically most similar and genetically most closely related to O. cheni Zhao, 1999 and O. latouchii (Boulenger, 1899), but possesses considerable genetic divergence ( p -distance 5.1%-16.7%) and can be differentiated from all other congeners by a combination of the following morphological characters: (1) body size large (total length 514-586 mm) and strongly built; (2) dorsal scale rows 17 throughout, feebly keeled anteriorly and moderately keeled posteriorly; (3) ventral scales 147-152, subcaudal scales 54-62; (4) preocular absent, loreal elongated and touching orbit; (5) supralabials 8-9, fifth and sixth entering obit; (6) anterior temporals short, length 1.74-2.04 times longer than width; (7) maxillary teeth subequal, 28-30; (8) dorsal surface of head with distinct irregular yellow stripes and markings edged with ochre; (9) body with clear black and yellow longitudinal streaks, partly fused to several lighter patches or thicker stripes anteriorly; and (10) venter pale yellow, with asymmetric blackish speckles along outer margin. We present an updated diagnostic key to all members of the genus Opisthotropis , and recommendations on the ecological study for the group are provided.

Background: Covering the eye of all snakes is a transparent integumental structure known as the spectacle. In order to determine variations in spectacle thickness among species, the spectacles of 217 alcohol-preserved museum specimens of 44 species belonging to 14 different families underwent optical coherence tomography (OCT) to measure spectacular thickness. Multivariable analyses were made to determine whether family, activity period (diurnal/nocturnal) and habitat (arboreal/terrestrial/fossorial/aquatic) influenced spectacle thickness., Results: The thinnest spectacles in absolute terms were found in the Usambara bush viper (Viperidae) with a thickness of 74 ± 9 μm and the absolute thickest spectacle was found in the red-tailed pipe snake (Cylindrophiidae) which had a spectacle thickness of 244 ± 57 μm. Fossorial and aquatic snakes had significantly thicker spectacles than arboreal and terrestrial snakes. When spectacle thickness was correlated to eye size (horizontal spectacle diameter), Gray's earth snake (Uropeltidae) had the lowest ratio (1:7) and the cottonmouth (Viperidae) had the highest ratio (1:65). Multivariable and phylogenetic analyses showed that spectacular thickness could be predicted by taxonomic family and habitat, but not activity period., Conclusion: This phylogenetically broad systematic study of the thickness of the snake spectacle showed that spectacular thickness varies greatly across snake species and may reflect evolutionary adaptation and development.

Synopsis: The morphology-performance-fitness paradigm for testing selection on morphological traits has seen decades of successful application. At the same time, life-history approaches using matrix methods and perturbation studies have also allowed the direct estimate of selection acting on vital rates and the traits that comprise them. Both methodologies have been successfully applied to the garter snakes of the long-term Eagle Lake research project to reveal selection on morphology, such as color pattern, number of vertebrae, and gape size; and life-history traits such as birth size, growth rates, and juvenile survival. Here we conduct a reciprocal transplant study in a common laboratory environment to study selection on morphology and life-history. To place our results in the ecomorphology paradigm, we measure performance outcomes (feeding rates, growth, insulin-like growth factor 1 titers) of morphological variation (body size, condition) and their fitness consequences for juvenile survival-a trait that has large fitness sensitivities in these garter snake populations, and therefore is thought to be subject to strong selection. To better merge these two complementary theories, we end by discussing our findings in a nexus of morphology-performance-fitness-life history to highlight what these approaches, when combined, can reveal about selection in the wild., (© The Author 2017. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. All rights reserved. For permissions please email: journals.permissions@oup.com.)

Non-avian reptiles commonly are assumed to be like birds in their overall patterns of development. However, colubrid corn snakes (Pantherophis guttatus) have mechanisms of yolk cellularization and processing that are entirely different from the avian pattern. In birds, a vascular "yolk sac" surrounds and digests the liquid yolk. In contrast, in corn snakes, the yolk material is converted into vascularized cords of yolk-filled cells. In this study, we used stereomicroscopy, histology, and scanning electron microscopy to analyze this unusual developmental pattern in corn snakes. Our observations reveal that the yolk sac cavity is invaded by endodermal cells that proliferate, absorb yolk spheres, and form aggregates of interconnected cells within the liquid yolk mass. As development proceeds, small blood vessels arise from the yolk sac omphalopleure, penetrate into the yolk mass, and become tightly encased in the endodermal cells. The entire vitellus ultimately becomes converted into a mass of vascularized, "spaghetti-like" strands of yolk-laden cells. The resulting arrangement allows yolk to be digested intracellularly and yolk products to be transported to the developing embryo. Indirect evidence for this pattern in other species raises the possibility that it is ancestral for squamates and quite possibly Reptilia in general., (© 2017 Wiley Periodicals, Inc.)

Venoms can deleteriously affect any physiological system reachable by the bloodstream, including directly interfering with the coagulation cascade. Such coagulopathic toxins may be anticoagulants or procoagulants. Snake venoms are unique in their use of procoagulant toxins for predatory purposes. The boomslang ( Dispholidus typus ) and the twig snakes ( Thelotornis species) are iconic African snakes belonging to the family Colubridae. Both species produce strikingly similar lethal procoagulant pathologies. Despite these similarities, antivenom is only produced for treating bites by D. typus , and the mechanisms of action of both venoms have been understudied. In this study, we investigated the venom of D. typus and T. mossambicanus utilising a range of proteomic and bioactivity approaches, including determining the procoagulant properties of both venoms in relation to the human coagulation pathways. In doing so, we developed a novel procoagulant assay, utilising a Stago STA-R Max analyser, to accurately detect real time clotting in plasma at varying concentrations of venom. This approach was used to assess the clotting capabilities of the two venoms both with and without calcium and phospholipid co-factors. We found that T. mossambicanus produced a significantly stronger coagulation response compared to D. typus . Functional enzyme assays showed that T. mossambicanus also exhibited a higher metalloprotease and phospholipase activity but had a much lower serine protease activity relative to D. typus venom. The neutralising capability of the available boomslang antivenom was also investigated on both species, with it being 11.3 times more effective upon D. typus venom than T. mossambicanus . In addition to being a faster clotting venom, T. mossambicanus was revealed to be a much more complex venom composition than D. typus . This is consistent with patterns seen for other snakes with venom complexity linked to dietary complexity. Consistent with the external morphological differences in head shape between the two species, CT and MRI analyses revealed significant internal structural differences in skull architecture and venom gland anatomy. This study increases our understanding of not only the biodiscovery potential of these medically important species but also increases our knowledge of the pathological relationship between venom and the human coagulation cascade., Competing Interests: The authors declare no conflict of interest. The founding sponsors had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript; or in the decision to publish the results.

As part of a broad survey of placental structure, function, and evolution in reptilian sauropsids paraffin-section histology was used to study microscopic anatomy of the uterus and fetal membranes of three species of North American watersnakes (Nerodia: Colubridae). The pre-ovulatory uterus is poorly vascularized with inactive shell glands. These shell glands are activated during vitellogenesis but regress during pregnancy. Two placentas develop through apposition of the uterine lining to the chorioallantois and the yolk sac omphalopleure. Fetal and maternal components of the chorioallantoic placenta are progressively vascularized during development. Their epithelia are attenuated, but (contrary to a previous report), epithelia of neither the uterus nor the chorion are eroded. The fetal portion of the yolk sac placenta is an omphalallantois, formed of avascular omphalopleure, isolated yolk mass, and allantois. This placenta is progressively replaced by chorioallantoic placenta during mid- to late-development through depletion of the isolated yolk mass. The chorioallantoic placenta is anatomically specialized for maternal-fetal gas exchange, and its expansion during development reflects the growing needs of the fetus for gas exchange. The yolk sac placenta is morphologically unsuited for gas exchange, but may serve other functions in maternal-fetal exchange., (© 2017 Wiley Periodicals, Inc.)

Across ecosystems and trophic levels, predators are usually larger than their prey, and when trophic morphology converges, predators typically avoid predation on intraguild competitors unless the prey is notably smaller in size. However, a currently unexplained exception occurs in kingsnakes in the genus Lampropeltis Kingsnakes are able to capture, constrict and consume other snakes that are not only larger than themselves but that are also powerful constrictors (such as ratsnakes in the genus Pantherophis ). Their mechanisms of success as intraguild predators on other constrictors remain unknown. To begin addressing these mechanisms, we studied the scaling of muscle cross-sectional area, pulling force and constriction pressure across the ontogeny of six species of snakes ( Lampropeltis californiae , L. getula , L. holbrooki , Pantherophis alleghaniensis , P. guttatus and P. obsoletus ). Muscle cross-sectional area is an indicator of potential force production, pulling force is an indicator of escape performance, and constriction pressure is a measure of prey-handling performance. Muscle cross-sectional area scaled similarly for all snakes, and there was no significant difference in maximum pulling force among species. However, kingsnakes exerted significantly higher pressures on their prey than ratsnakes. The similar escape performance among species indicates that kingsnakes win in predatory encounters because of their superior constriction performance, not because ratsnakes have inferior escape performance. The superior constriction performance by kingsnakes results from their consistent and distinctive coil posture and perhaps from additional aspects of muscle structure and function that need to be tested in future research., (© 2017. Published by The Company of Biologists Ltd.)

Across the diversity of vertebrates, bite force has been studied and suggested to have important ecological and evolutionary consequences. However, there is a notable lineage of vertebrates that use this performance trait yet are missing from the bite-force literature: the snakes. Snakes often rely on biting during prey subjugation and handling. Many snakes bite and hold prey while a constriction coil is formed or while venom is being delivered, or both. Others use biting exclusively without employing any additional prey-handling behaviors. In addition to biting, constriction is an important predation mechanism. Here, I quantify bite force and constriction pressure in kingsnakes (Lampropeltis getula). Furthermore, I explore the proximate determinants of bite force as well as the relationship between biting and constriction performance. Bite force increased linearly with all head and body measures. Of these, head height was the best predictor of bite force. Bite force in kingsnakes was within the range of values reported for lizards, but their relative performance was lower for their head size compared to lizards. Peak constriction pressure also increased with all body measures. Biting and constricting use 2 different parts of the musculoskeletal system and are positively and significantly correlated with one another. Future work targeting a greater diversity of snakes that rely more heavily on biting may reveal a greater range of bite performance in this diverse and successful vertebrate group., (© 2016 International Society of Zoological Sciences, Institute of Zoology/Chinese Academy of Sciences and John Wiley & Sons Australia, Ltd.)

A new species of Dendrelaphis, Dendrelaphis sinharajensis sp. nov. is described, the sixth species of the genus known from Sri Lanka. The new species is readily distinguished from all other congeners by its colour pattern and scalation. The species is a canopy-dweller known only from the Sinharaja World Heritage Site and its vicinity, in the lowland wet zone of Sri Lanka.

Hydrodynastes bicinctus was described with no type material or locality and it has two subspecies currently recognized that are not taxonomically well defined. We tested the validity of the two subspecies through meristic, morphometric, and color pattern characters. Two apparently distinct color patterns of H. bicinctus were noticed, one from the Cerrado open formations and the other from the Amazon rainforest. These aforementioned patterns, however, exhibited a high degree of geographic overlap and many specimens showed a blended pattern. Based on these results we propose synonymizing H. bicinctus schultzi with the nominal taxon. Furthermore, we designate a neotype for the species, present data on geographic distribution, and provide morphological descriptions of the hemipenis, cephalic glands, and skull.

We describe a new kukri snake, Oligodon condaoensis sp. nov., from Con Dao Islands, southern Vietnam based on the morphological characters of four specimens. It differs from other congeners by a combination of the following characters: medium size in adults (total length up to 552 mm); 17-17-15 dorsal scale rows; deeply forked hemipenes without spines and papillae, extending to subcaudal 13 or 14; 11-13 maxillary teeth, the posterior three being enlarged; cloacal plate undivided; head scalation complete; nasal divided; presubocular absent; 168-176 ventrals; 33-37 subcaudals; overall dorsal coloration dark gray, faint body stripes present or absent; and ventral coloration cream to dark gray without rectangular blotches. We also provide a list of 43 species of amphibians and terrestrial reptiles recorded from Con Dao Islands.

Indigo Snakes (genus Drymarchon) occur from northern Argentina northward into to the United States, where they inhabit southern Texas and disjunct populations in Mississippi, Florida and Georgia. Based on allopatry and morphological differences Collins (1991) hypothesized that the two United States taxa-the Western Indigo Snake, D. melanurus erebennus (Cope, 1860), and the Eastern Indigo Snake, D. couperi (Holbrook, 1842)-deserved full species recognition. Building upon this hypothesis with molecular and morphological analyses we illustrate that D. couperi is split into two distinct lineages. Based on the General Lineage Concept of Species, we describe the lineage that occurs along the Gulf coast of Florida and Mississippi as a new species, Drymarchon kolpobasileus. The new species is distinguished from D. couperi by a suite of morphological features, including a shorter and shallower head, deeper and shorter 7 th infralabial scales, and shorter temporal scales. Overall, the presence of a deep 7 th infralabial scale provides the best univariate identifier of D. kolpobasileus sp. nov. This study illustrates the usefulness of using both morphological and genetic data in refining accurate descriptions of geographical distributions.

This study aimed to describe the reproductive biology of populations of Thamnodynastes hypoconia in subtemperate wetlands based on macroscopic analyses of their gonads. We analyzed 101 specimens from the southernmost regions of Brazil. The males had a greater snout-vent length, but the females reached sexual maturity with a greater body size. The reproductive cycle of the females was seasonal, with secondary vitellogenesis occurring between the winter and spring (May-October). Based on macroscopic analysis of gonads, data suggests that males have a continuous reproductive cycle. Parturition occurs between the late summer and early fall (January-April). The clutch size ranged between 4 and 16 embryos and showed no relationship with the female's body size. The recorded reproductive frequency of T. hypoconia is low (38 %) compared to other phylogenetically related species. Conversely, the studied populations appear to have a high reproductive success because it is an abundant species in the study site.

The colubrid snake species Pseudorabdion torquatum is resurrected from the synonymy of P. longiceps. Out of the three syntypes, only two were located and examined at Muséum national d'Histoire naturelle, Paris, France (MNHN-RA). Pseudorabdion torquatum differs from P. longiceps in having a larger SVL (227.0 mm vs. 132.0-193.0 mm in males); higher number of ventral in males, 141 (vs. 127-139); lower number of ventrals in females, 137 (vs. 139-145); lower number of subcaudals in both sexes, 12-17 (vs. 17-30); and six supralabials (vs. five). Pseudorabdion torquatum is thus regarded as a valid and endemic species to Sulawesi. The species is here redescribed and a lectotype is designated. Furthermore we provide an English translation of the original description of Rabdion torquatum which was originally written in French. Thus, besides unconfirmed records of P. longiceps from Sulawesi, there is a single known congener from that island, P. sarasinorum.

We describe a new species of Rhadinella from the Sierra Madre del Sur of Guerrero, Mexico, a region where the genus was previously unknown. This diminutive species is a member of a group of snakes previously allocated in the Rhadinaea godmani group, and more recently transferred to the genus Rhadinella. These snakes may have conspicuous dark longitudinal striping on a pale brown to orange background or may have dark brown to blackish dorsal ground coloration, which mostly or completely obfuscates a pattern of longitudinal striping. The new species is mostly dark with barely discernible slightly paler or darker striping (depending on how striping is interpreted). The closest relative of the new species, on the basis of morphological similarities and biogeography, appears to be Rhadinella donaji which occurs to the east in the Sierra Madre del Sur of Oaxaca about 275 km from the type-locality of the new species.

The colubrid snake genus Tantilla currently comprises 62 valid species distributed from the southern United States of America to southern Bolivia in the west, and Uruguay and northern Argentina in the east (Wilson & Mata-Silva 2014, 2015). Of these 62 species, 13 are found in South America (Wilson & Mata-Silva 2014, 2015). The most recently described of these Southamerican species is Tantilla marcovani Lema, 2004. While we were working on a checklist and key to the members of the Tantilla clade (sensu Holm 2008, including the genera Geagras Cope 1876, Scolecophis Fitzinger 1843, Tantilla Baird & Girard 1853, and Tantillita Smith 1941; Wilson & Mata-Silva 2015), it became apparent that the diagnosis of T. marcovani entirely falls to distinguish this nominal taxon from the widespread and extensively variable T. melanocephala Linnaeus 1758. Tantilla melanocephala, as presently envisioned, is the most broadly distributed member of its genus (Wilson & Mena 1980; Savage 2002; Greenbaum et al. 2004), occurring from Panama southward into the major portion of the South American continent, apart from Chile and southern Argentina.

Snakes are a highly successful group of vertebrates, within great diversity in habitat, diet, and morphology. The unique adaptations for the snake skull for ingesting large prey in more primitive macrostomatan snakes have been well documented. However, subsequent diversification in snake cranial shape in relation to dietary specializations has rarely been studied (e.g. piscivory in natricine snakes). Here we examine a large clade of snakes with a broad spectrum of diet preferences to test if diet preferences are correlated to shape variation in snake skulls. Specifically, we studied the Xenodontinae snakes, a speciose clade of South American snakes, which show a broad range of diets including invertebrates, amphibians, snakes, lizards, and small mammals. We characterized the skull morphology of 19 species of xenodontine snakes using geometric morphometric techniques, and used phylogenetic comparative methods to test the association between diet and skull morphology. Using phylogenetic partial least squares analysis (PPLS) we show that skull morphology is highly associated with diet preferences in xenodontine snakes.

Postembryonic changes in the dermal and epidermal pigment cell architecture of the striped and nonstriped morph of the Japanese four-lined snake Elaphe quadrivirgata were examined to reveal stripe pattern formation after hatching. The striped and nonstriped morphs were distinguishable at the hatching, suggesting that the basis of stripe pattern was formed during embryonic development. In the striped morph, the color of stripes changed from red-brown in juveniles to vivid dark-brown in adults, and density of dermal melanophore increased much more in the stripe than background dorsal scales with growth. This increase in density of dermal melanophore was accompanied not only by the increased epidermal melanophore density but also by the change in vertical structures of dermal melanophore. By contrast, the density of epidermal and dermal melanophore evenly increased over the dorsal scales in the nonstriped morph. Thus, the increased vividness of the stripe pattern after hatching is achieved through localized increase of melanophore density particularly in the stripe region but not over the whole dorsal scales., (© 2015 Wiley Periodicals, Inc.)

A new species of the genus Oligodon from the coastal area of Binh Chau-Phuoc Buu Nature Reserve, Ba Ria-Vung Tau Province, southern Vietnam, is described. Oligodon arenarius sp. nov. is distinguishable from all other species by the unique combination of the following characters: medium size; 17 dorsal scale rows; 6-8 maxillary teeth, the posterior three being enlarged; head scalation lacking a loreal but usually including a presubocular; divided nasal; two postoculars; 131-144 ventrals; 36-60 subcaudals; unforked hemipenis, without spines or obvious papillae; sexual dimorphism displayed in the number of subcaudals (36-40 in females, 58-60 in males) and a relative tail length, tails being quite long in males (TaL/TL = 0.26-0.28) and moderate in females (TaL/TL = 0.13-0.17); head coloration pattern including ocular band, temporal bands and chevron-shaped mark on nape; dorsal coloration without distinct pattern, uniform or with dark speckling; ventrals pinkish in life, immaculate. To date, this species appear to be endemic to Vietnam.

In reptilian sauropsids, fetal (extraembryonic) membranes that line the eggshell sustain developing embryos by providing for gas exchange and uptake of water and eggshell calcium. However, a scarcity of morphological studies hinders an understanding of functional specializations and their evolution. In kingsnakes (Lampropeltis getula), scanning electron microscopy reveals two major fetal membranes: the chorioallantois and yolk sac omphalopleure. In early development, the chorioallantois contains tall chorionic epithelial cells, avascular connective tissue, and enlarged allantoic epithelial cells. During its maturation, the chorionic and allantoic epithelia thin dramatically and become underlain by a rich network of allantoic capillaries, yielding a membrane ideally suited for respiratory gas exchange. Yolk sac development initially is like that of typical lizards and snakes, forming an avascular omphalopleure, isolated yolk mass (IYM), and yolk cleft. However, unlike the situation in most squamates studied, the omphalopleure becomes transformed into a "secondary chorioallantois" via three asynchronous events: flattening of the epithelium, regression of the IYM, and vascularization by the allantois. Progressive expansion of chorioallantois parallels growing embryonic needs for gas exchange. In early through mid-development, external surfaces of both the chorionic and omphalopleure epithelium show an abundance of irregular surface protrusions that possibly increase surface area for water absorption. We postulate that the hypertrophied allantoic epithelial cells produce allantoic fluid, a viscous substance that facilitates water uptake and storage. Our findings are consistent with a previous study on the corn snake Pantherophis guttatus, but include new observations and novel functional hypotheses relevant to a reconstruction of basal squamate patterns., (© 2015 Wiley Periodicals, Inc.)