Perils of ingesting harmful prey by advanced snakes.

The advanced snakes (Alethinophidia) include the extant snakes with a highly evolved head morphology providing increased gape and jaw flexibility. Along with other physiological and morphological adaptations, this allows them to immobilize, ingest, and transport prey that may be disproportionately large or presents danger to the predator from bites, teeth, horns, or spines. Reported incidents of snakes failing to consume prey and being injured or killed during feeding mostly reflect information in the form of natural-history notes. Here we provide the first extensive review of such incidents, including 101 publications describing at least 143 cases of mortality (including six of 'multiple individuals') caused by ingestion or attempted consumption of injurious prey. We also report on 15 previously unpublished injurious feeding incidents from the USA, Austria, and Bulgaria, including mortality of five juvenile piscivorous dice snakes (Natrix tessellata) from a single location. Occurrences are spread across taxa, with mortality documented for at least 73 species from eight families and 45 genera. Incidents were generally well represented within each of three major categories: oversized prey (40.6%), potentially harmful prey (40.6%), and predator's behavioural/mechanical errors (18.9%). Reptile (33%) and fish (26%) prey caused disproportionately high mortality compared to mammals (16%). Feeding can be dangerous throughout a snake's life, with the later stages of feeding likely being more perilous. The number of reports has increased over time, and the data seem biased towards localities with a higher number of field-working herpetologists. We propose a standardized framework, comprising a set of basic information that should ideally be collected and published, and which could be useful as a template for future data collection, reporting, and analyses. We conclude that incidents of mortality during feeding are likely to be more common than previously assumed, and this hypothesis has implications for the ecology of persistence where populations are impacted by changing trophic environments.

Analyses of Skin Secretions of Vipera ammodytes (Linnaeus, 1758) (Reptilia: Serpentes), with Focus on the Complex Compounds and Their Possible Role in the Chemical Communication.

Snakes rely heavily on chemical cues when foraging, searching for mates, etc. Snakes' sex attractiveness pheromones comprise mainly heavy, semi-volatile compounds such as ketones. Here we investigated the composition of skin secretions of adult Vipera ammodytes (Linnaeus, 1758) individuals. The samples were analyzed by gas chromatography/mass spectrometry and the identification of the compounds was performed using commercial mass spectral libraries and retention times. The relative concentrations of all detected compounds were tested for significant differences between (1) male vs. female live individuals, (2) shed skin vs. live individuals, and (3) pre-reproductive vs. reproductive live individuals. We detected fifty-nine compounds of which six were ketones. Two ketones (2-pentacosanone and 2-heptacosanone) were present in many of the samples and thus may have an important role in the V. ammodytes chemical communication. We did not find significant differences between the relative concentrations of the compounds between male and female individuals (only three compounds are exceptions). Significant differences were found between extracts from shed skins and live individuals and between live pre-reproductive individuals and live reproductive individuals. The results of the study suggest that chemical communication in V. ammodytes involves less compounds in comparison to the known literature data for other species.

Physiological and behavioral responses to salinity in coastal Dice snakes.

Secondarily marine tetrapods have evolved adaptations to maintain their osmotic balance in a hyperosmotic environment. During the transition to a marine habitat, the evolution of a euryhaline physiology likely encompassed successive changes in behavior and physiology that released organisms from regular access to fresh water. Deciphering these key steps is a complicated task. In this study, we investigated a species of freshwater natricine snake in which some populations are known to use marine environments. We experimentally subjected 30 adult Dice snakes (Natrix tessellata) from a population inhabiting the Black Sea coast to three salinities corresponding to freshwater (~0.1‰), brackish water (~15.0‰), and full-strength seawater (~34.0‰) in order to investigate their physiological (variation of body mass, osmolality) and behavioral (activity, drinking behavior) responses to salinity. Our results show that coastal Dice snakes from the study population are relatively tolerant to salinity close to that recorded in the Black Sea, but that prolonged exposure to full-strength seawater increases osmolality, stimulates thirst, decreases the activity of snakes and may ultimately jeopardize survival. Collectively with previously published data, our results strongly suggest specific physiological adaptations to withstand hyperosmolality rather than to reduce intake of salt, in coastal populations or species of semi-aquatic snakes. Future comparative investigations of Dice snakes from populations restricted to freshwater environment might reveal the functional traits and the behavioral and physiological responses of coastal N. tessellata to life in water with elevated salinity.

Does Sexual Selection Influence Ornamentation of Hemipenes in Old World Snakes?

In the present study, we investigated and documented the morphology of the male copulatory organs (hemipenes) in fifteen wide-ranging snake species. The species represent four families (Boidae, Colubridae, Lamprophiidae, and Viperidae) and ten genera. We applied the same preparation techniques for all species, successfully everting and expanding the organs completely. The detailed description of the general morphology of the male copulatory organs was based on 31 specimens. Our data were compared with published observations and we point out some incorrectly described details in previous investigations. We provide the first description of the hemipenial morphology for three ophidian species (Elaphe sauromates, Telescopus fallax, and Malpolon insignitus). In addition to the morphological characteristics of the hemipenes presented in the research, we propose the adoption of a standardized index describing the hemipenial proportions. The immense variation in hemipenial morphology presupposes its dynamic evolution, but we suggest that many of the significant structures observed here may have escaped previous researchers due to differing methodologies. Some of the highly ornamented morphologies that we describe are consistent with a locking mechanism during copulation. However, other morphologies may relate to the variety of mating behaviors observed. As a result, we propose that sexual selection is the major driver affecting the hemipenial ornamentation in snakes. Anat Rec, 300:1680-1694, 2017. © 2017 Wiley Periodicals, Inc.

Hypernatremia in Dice snakes (Natrix tessellata) from a coastal population: implications for osmoregulation in marine snake prototypes.

The widespread relationship between salt excreting structures (e.g., salt glands) and marine life strongly suggests that the ability to regulate salt balance has been crucial during the transition to marine life in tetrapods. Elevated natremia (plasma sodium) recorded in several marine snakes species suggests that the development of a tolerance toward hypernatremia, in addition to salt gland development, has been a critical feature in the evolution of marine snakes. However, data from intermediate stage (species lacking salt glands but occasionally using salty environments) are lacking to draw a comprehensive picture of the evolution of an euryhaline physiology in these organisms. In this study, we assessed natremia of free-ranging Dice snakes (Natrix tessellata, a predominantly fresh water natricine lacking salt glands) from a coastal population in Bulgaria. Our results show that coastal N. tessellata can display hypernatremia (up to 195.5 mmol x l(-1)) without any apparent effect on several physiological and behavioural traits (e.g., hematocrit, body condition, foraging). More generally, a review of natremia in species situated along a continuum of habitat use between fresh- and seawater shows that snake species display a concomitant tolerance toward hypernatremia, even in species lacking salt glands. Collectively, these data suggest that a physiological tolerance toward hypernatremia has been critical during the evolution of an euryhaline physiology, and may well have preceded the evolution of salt glands.