Sexual divergence in diets and morphology in Fijian sea snakes Laticauda colubrina (Laticaudinae)

In the Fiji Islands, female yellow‐lipped sea kraits (Laticauda colubrina) grow much larger than males, and have longer and wider heads than do conspecific males of the same body length. This morphological divergence is accompanied by (and may be adaptive to) a marked sex divergence in dietary habits. Adult female sea kraits feed primarily on large conger eels, and take only a single prey item per foraging bout. In contrast, adult males feed upon smaller moray eels, and frequently take multiple prey items. Prey size increases with snake body size in both males and females, but the sexes follow different trajectories in this respect. Female sea kraits consume larger eels relative to predator head size and body length than do males. Thus, the larger relative head size of female sea kraits is interpreted as an adaptation to consuming larger prey items. Our results are similar to those of previous studies on American water snakes (natricines) and Australian file snakes (acrochordids), indicating that similar patterns of sex divergence in dietary habits and feeding structures have evolved convergently in at least three separate lineages of aquatic snakes.     

Morphological adaptations to marine life in snakes

We investigated morphological adaptations to aquatic life within animals that exhibit a structurally simple, elongate body form, i.e., snakes. This linear body plan should impose different biomechanical constraints than the classical streamlined body shape associated with propulsion by fins, feet, or wings. Our measurements of general body shape of terrestrial, amphibious, and marine snakes (all from the same phylogenetic lineage, the Elapidae) show that seasnakes display specialized morphological attributes for life in water. Most notably, the cross‐sectional body shape is circular in terrestrial snakes but dorso‐ventrally elongated in seasnakes (due to a prominent ventral keel); amphibious species (sea kraits) exhibit an intermediate shape. The tail of amphibious and marine species (a major propulsive structure during swimming) is higher and thinner than in terrestrial snakes (i.e., paddle‐shaped) but shorter relative to body length. The evolution of a laterally compressed shape has been achieved by an increase in body height rather than a decrease in body width, possibly reflecting selection for more effective propulsive thrust, and for an ability to maintain hydrodynamic efficiency despite the minor bodily distension inevitably caused by prey items and developing offspring. J. Morphol., 2011. © 2011 Wiley‐Liss, Inc     

Terrestrial locomotion in sea snakes: the effects of sex and species on cliff-climbing ability in sea kraits (Serpentes, Elapidae, Laticauda)

Ecomorphological theory predicts a match between an organism's environment and its locomotor abilities, such that animals function most effectively under the conditions they experience in nature. However, amphibious species must simultaneously optimize performance in two different habitats posing incompatible demands on locomotor morphology and physiology. This situation may generate a mismatch between environment and locomotor function, with performance optimized only for the more important habitat type; alternatively, selection may fine-tune locomotor abilities for both types of challenges. Two species of sea kraits in New Caledonia offer an opportunity to examine this question: Laticauda laticaudata is more highly aquatic than L. colubrina , and males are more terrestrial than females within each taxon. We examined an aspect of locomotor performance that is critical to coming ashore on steep-walled rocky islets: the ability to climb steep cliffs. We also measured the muscular strength of these animals, a character that is likely critical to climbing performance. Laticauda colubrina was heavier-bodied and stronger (even relative to its body mass) than the more aquatic L. laticaudata ; and within each species, males were heavier-bodied and stronger than females. The same patterns were evident in cliff-climbing ability. Thus, the ability of different species and sexes of sea kraits to climb steep cliffs correlates with their body shape even though these primarily aquatic animals use terrestrial habitats only rarely.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 433–441.     

Evolution of alternative foraging tactics driven by water temperature and physiological constraints in an amphibious snake

Amphibious predatory ectotherms live and forage in two environments (aquatic and terrestrial) that can drastically differ in temperature means and variance across space and time. The locomotor performance of ectotherms is known to be strongly affected by temperature. However, how differences in water temperature may drive the evolution of alternative foraging tactics in amphibious animals remains poorly understood. Fish‐eating Viperine snakes Natrix maura occur from high altitude cold water streams to warm shallow lakes, and employ two main feeding strategies: sentinel foraging (underwater sit‐and‐wait behaviour) and active foraging (fish chasing). Using 272 juvenile snakes we measured: the performance kinetics of diving and swimming in a wide range of water temperatures; basal metabolic levels in relation to body temperature; and the type of foraging mode expressed in water‐temperature‐acclimated snakes. Individual swimming performances increased with testing temperature (10, 15, 20, 25 or 30 °C). Apnoea time followed an opposite trend however, plausibly reflecting the fact that oxygen demands are related to the metabolic rate of ectotherms. That is, snake heart rates increased with body temperature. Snakes acclimated to 10 °C water mostly displayed sentinel foraging. By contrast, 20 °C and 30 °C water‐acclimated snakes were extremely active fish chasers. Individual apnoea times at the various testing temperatures were all correlated; as were individual swimming speeds. There was however no clear relationship between an individual's ability to hold its breath and its ability to swim, suggesting that both performance traits may be the target of different selective pressures. Fast swimming speed and long breath holding abilities are likely key determinants of both foraging success and predatory evasion, although in a context dependent manner. Active swimming foraging is likely to be advantageous in warm water (> 20 °C), while sentinel foraging appears better suited to cold water (< 14 °C). The physiological aspects of foraging tactics of amphibious snakes combined with field and laboratory observations support the idea that physiological and environmental constraints may generate shifts in habitat use and associated foraging tactics in amphibious ectotherms. Avenues for further research are discussed. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 411–422.     

Conflicts between feeding and reproduction in amphibious snakes (sea kraits,Laticauda spp.)

If reproduction impairs an organism's ability to perform other fitness‐related activities, natural selection may favour behavioural adjustments to minimize these conflicts. This is presumably the reason why many animals are anorexic during the breeding season. We studied amphibious sea snakes, a group whose ecology facilitates teasing apart the causal links between reproduction and feeding. In both Laticauda laticaudata and L. saintgironsi in New Caledonia, adult females cease feeding as their eggs develop. The advantages of foregoing feeding do not relate to thermoregulation (because foraging does not entail lower body temperatures), nor are they attributable to physical constraints on abdominal volume (because in a snake's linear body, there is little overlap between the stomach and the oviducts). Instead, female sea kraits appear to cease feeding because their bodily distension impedes locomotor ability, rendering them less effective at foraging and more vulnerable to aquatic predators.     

Isotopic signatures, foraging habitats and trophic relationships between fish and seasnakes on the coral reefs of New Caledonia

A predator’s species, sex and body size can influence the types of prey that it consumes, but why? Do such dietary divergences result from differences in foraging habitats, or reflect differential ability to locate, capture or ingest different types of prey? That question is difficult to answer if foraging occurs in places that preclude direct observation. In New Caledonia, amphibious sea kraits (Laticauda laticaudata and L. saintgironsi) mostly eat eels—but the species consumed differ between snake species and vary with snake body size and sex. Because the snakes capture eels within crevices on the sea floor, it is not possible to observe snake foraging on any quantitative basis. We used stable isotopes to investigate habitat-divergence and ontogenetic shifts in feeding habits of sympatric species of sea kraits. Similarities in δ15 N (~10.5‰) values suggest that the two snake species occupy similar trophic levels in the coral-reef foodweb. However, δ13C values differed among the eight eel species consumed by snakes, as well as between the two snake species, and were linked to habitat types. Specifically, δ13C differed between soft- vs. hard-substrate eel species, and consistently differed between the soft-bottom forager L. laticaudata (~ −14.7‰) and the hard-bottom forager L. saintgironsi (~ −12.5‰). Differences in isotopic signatures within and between the two sea krait species and their prey were consistent with the hypothesis of habitat-based dietary divergence. Isotopic composition varied with body size within each of the snake species and varied with body size within some eel species, reflecting ontogenetic shifts in feeding habits of both the sea kraits and their prey. Our results support the findings of previous studies based on snake stomach contents, indicating that further studies could usefully expand these isotopic analyses to a broader range of trophic levels, fish species and spatial scales.     

Dehydration and drinking responses in a pelagic sea snake

Recent investigations of water balance in sea snakes demonstrated that amphibious sea kraits (Laticauda spp.) dehydrate in seawater and require fresh water to restore deficits in body water. Here, we report similar findings for Pelamis platurus, a viviparous, pelagic, entirely marine species of hydrophiine ("true") sea snake. We sampled snakes at Golfo de Papagayo, Guanacaste, Costa Rica and demonstrated they do not drink seawater but fresh water at variable deficits of body water incurred by dehydration. The threshold dehydration at which snakes first drink fresh water is -18.3 ± 1.1 % (mean ± SE) loss of body mass, which is roughly twice the magnitude of mass deficit at which sea kraits drink fresh water. Compared to sea kraits, Pelamis drink relatively larger volumes of water and make up a larger percentage of the dehydration deficit. Some dehydrated Pelamis also were shown to drink brackish water up to 50% seawater, but most drank at lower brackish values and 20% of the snakes tested did not drink at all. Like sea kraits, Pelamis dehydrate when kept in seawater in the laboratory. Moreover, some individuals drank fresh water immediately following capture, providing preliminary evidence that Pelamis dehydrate at sea. Thus, this widely distributed pelagic species remains subject to dehydration in marine environments where it retains a capacity to sense and to drink fresh water. In comparison with sea kraits, however, Pelamis represents a more advanced stage in the evolutionary transition to a fully marine life and appears to be less dependent on fresh water.     

CONSTRAINTS ON REPRODUCTIVE INVESTMENT: A COMPARISON BETWEEN AQUATIC AND TERRESTRIAL SNAKES

Life-history theory predicts that “costs” of reproduction may be important evolutionary determinants of reproductive investment; previous studies on reptiles indicate that decrements to maternal mobility may be among the most important components of such costs. Biomechanical models suggest that reproductive investment in aquatic snakes may be constrained by the important locomotory role of the posterior part of the body during swimming: carrying eggs or offspring in this region would more seriously impair locomotory efficiency in swimming than in terrestrial lateral undulation. If this constraint is important, aquatic snakes would be expected to have lower clutch masses relative to body mass than terrestrial species and to carry the clutch in a more anterior position (commencing at the same proportion of maternal body length anteriorly, but not extending as far posteriorly). Comparisons between aquatic and terrestrial snakes of several families confirm these predictions. Phylogenetic analysis suggests that this pattern of reduced reproductive investment has evolved independently in each of the four ophidian lineages that contain marine species (acrochordids, homalopsine colubrids, laticaudid sea snakes, and hydrophiid sea snakes). Although it thus seems likely that these patterns represent adaptations to aquatic versus terrestrial life, the nature of the selective forces involved remains speculative. The hypothesis based on locomotory impairment of gravid females has better empirical support than any alternative hypothesis, as it successfully predicts modifications in the position of the clutch within the female's body, as well as overall reduced reproductive investment.     

Comparison of freshwater discrimination ability in three species of sea kraits (Laticauda semifasciata, L. laticaudata and L. colubrina)

Three species of amphibious sea kraits (Laticauda spp.) require drinking freshwater to regulate water balance. The extent of terrestriality is known to differ among them. Species with higher extent of terrestriality would drink freshwater accumulated on land, whereas less terrestrial species would rely totally on freshwater that runs into the sea. Consequently, we predicted that the latter species might have a better ability to follow the flow of freshwater or lower salinity water in the sea than the former. We investigated the freshwater discrimination ability of three sea krait species, using a Y-maze apparatus. We found that Laticauda semifasciata and Laticauda laticaudata, less terrestrial species, followed freshwater significantly more frequently than seawater, whereas Laticauda colubrina, more terrestrial species, unbiasedly selected freshwater and seawater. This result supports our prediction and suggests that less terrestrial sea kraits more efficiently access freshwater sources in the sea than highly terrestrial sea kraits. It is likely that behavioral rehydration systems vary among sea kraits in relation to their terrestrial tendency.     

Limb movements and locomotor function in the California sea lion (Zalophus californianus)

The peculiar amphibious mode of life of California sea lions suggests that their locomotor systems may contain adaptations both to life on land and in the water. Previous studies of their locomotor behaviour have been either superficial or based on inferences which were derived from limb structure. Limb movements associated with locomotor behaviour in California sea lions are described on the basis of frame-by-frame analysis of slow motion cinematography of typical aquatic and terrestrial locomotor sequences. Results are compared to reports of terrestrial and aquatic locomotor behaviour in fissiped carnivores, whose locomotor behaviour is presumed to reflect the framework from which the locomotor behaviour of sea lions was derived. The major distinction between sea lions and fissipeds in terms of aquatic locomotor behaviour involves the use of the forelimb in sea lions. Propulsive thrust is generated by medial rotation, adduction and retraction of the forelimbs in sea lions, in contrast to nearly pure limb retraction in fissipeds. The major features which distinguish terrestrial locomotor behaviour in sea lions from that of fissipeds are use of the manus as a transverse rather than sagittal propulsive lever and extensive use of posterior axial and head and neck movements rather than hindlimb movements. The biomechanical implications of these movements are used to elucidate their potentially adaptive features.     

Thermal biology of sea snakes and sea kraits1

Temperature probably had no direct effect on the evolution of sea kraits within their center of origin, a geologically stable thermal zone straddling the equator, but may have indirectly affected expansions and contractions in distributions beyond that zone through global fluctuations that caused alternation of higher and lower sea levels. The northern limit of the Laticauda colubrina complex seems to be the 20°C isotherm; in the south, the range does not reach that isotherm because there is no land (also a habitat requirement of sea kraits) within the zone of suitable temperature. The relationship of temperature to the pattern of geographic variation in morphology supports either the hypothesis of peripheral convergence or the developmental hypothesis but does not distinguish between them. Quadratic surfaces relating cumulative scores for coloration and morphological characters to global position showed a strong latitudinal component and an even stronger longitudinal one in which the direction of the latitudinal effect was reversed between east and west. A multivariate analysis revealed that while morphological characters vary significantly by location and climate when tested separately, when the influence of location on morphology is taken into account, no residual relationship between climate and morphology remains. Most marine snakes have mean upper temperature tolerances between 39°C and 40°C and operate at temperatures much nearer their upper thermal limits than their lower limits but still avoid deleterious extremes by diving from excessively hot water to deeper, cooler strata, and by surfacing when water is cold. At the surface in still water in sunlight, Pelamis can maintain its body temperature slightly above that of the water, but whether this is significant in nature is questionable. As temperature falls below 18-20°C, survival time is progressively reduced, accompanied by the successive occurrence of cessation of feeding, cessation of swimming, and failure to orient. Acclimation does not seem to be in this species' repertoire. In the water column, marine snakes track water temperature; on land, sea kraits can thermoregulate by basking, selecting favorable locations, and by kleptothermy. Laticauda colubrina adjusts its reproductive cycle geographically in ways that avoid breeding in the coldest months. Mean voluntary diving time is not temperature-dependent within the normal range of temperatures experienced by marine snakes in the field, but is reduced in water colder than 20°C. On land, much as while diving in the sea, sea kraits maintain long periods of apnea; intervals between breaths are inversely related to temperature.     

Comparative limb bone scaling in turtles: Phylogenetic transitions with changes in functional demands?

Several terrestrial vertebrate clades include lineages that have evolved nearly exclusive use of aquatic habitats. In many cases, such transitions are associated with the evolution of flattened limbs that are used to swim via dorsoventral flapping. Such changes in shape may have been facilitated by changes in limb bone loading in novel aquatic environments. Studies on limb bone loading in turtles found that torsion is high relative to bending loads on land, but reduced compared to bending during aquatic rowing. Release from torsion among rowers could have facilitated the evolution of hydrodynamically advantageous flattened limbs among aquatic species. Because rowing is regarded as an intermediate locomotor stage between walking and flapping, rowing species might show limb bone flattening intermediate between the tubular shapes of walkers and the flattened shapes of flappers. We collected measurements of humeri and femora from specimens representing four functionally divergent turtle clades: sea turtles (marine flappers), softshells (specialized freshwater rowers), emydids (generalist semiaquatic rowers), and tortoises (terrestrial walkers). Patterns of limb bone scaling with size were compared across lineages using phylogenetic comparative methods. Although rowing taxa did not show the intermediate scaling patterns we predicted, our data provide other functional insights. For example, flattening of sea turtle humeri was associated with positive allometry (relative to body mass) for the limb bone diameter perpendicular to the flexion-extension plane of the elbow. Moreover, softshell limb bones exhibit positive allometry of femoral diameters relative to body mass, potentially helping them maintain their typical benthic position in water by providing additional weight to compensate for shell reduction. Tortoise limb bones showed positive allometry of diameters, as well as long humeri, relative to body mass, potentially reflecting specializations for resisting loads associated with digging. Overall, scaling patterns of many turtle lineages appear to correlate with distinctive behaviors or locomotor habits.     

The influence of natural selection and sexual selection on the tails of sea-snakes (Laticauda colubrina)

Many phenotypic traits perform more than one function, and so can influence organismal fitness in more than one way. Sexually dimorphic traits offer an exceptional opportunity to clarify such complexity, especially if the trait involved is subject to natural as well as sexual selection, and if the sexes differ in ecology as well as reproductive behaviour. Relative tail length in sea-snakes fulfils these conditions. Our field studies on a Fijian population of yellow-lipped sea kraits ( Laticauda colubrina ) show that relative tail lengths in male sea kraits have strong consequences for individual fitness, both via natural and sexual selection. Males have much longer tails (relative to snout-vent length) than do females. Mark-recapture studies revealed a trade-off between growth and survival: males with relatively longer tails grew more slowly, but were more likely to survive, than were shorter-tailed males. A male snake's tail length relative to body length influenced not only his growth rate and probability of survival, but also his locomotor ability and mating success. Relative tail length in male sea kraits was thus under a complex combination of selective forces. These forces included directional natural selection (through effects on survival, growth and swimming speed) as well as stabilizing natural selection (males with average-length tails swam faster) and stabilizing sexual selection (males with average-length tails obtained more matings). In contrast, our study did not detect significant selection on relative tail length in females. This sex difference may reflect the fact that females use their tails primarily for swimming, whereas males also must frequently use the tail in terrestrial locomotion and in courtship as well as for swimming.     

Triturus newts defy the running-swimming dilemma

Conflicts between structural requirements for carrying out different ecologically relevant functions may result in a compromise phenotype that maximizes neither function. Identifying and evaluating functional trade-offs may therefore aid in understanding the evolution of organismal performance. We examined the possibility of an evolutionary trade-off between aquatic and terrestrial locomotion in females of European species of the newt genus Triturus. Biomechanical models suggest a conflict between the requirements for aquatic and terrestrial locomotion. For instance, having an elongate, slender body, a large tail, and reduced limbs should benefit undulatory swimming, but at the cost of reduced running capacity. To test the prediction of an evolutionary trade-off between swimming and running capacity, we investigated relationships between size-corrected morphology and maximum locomotor performance in females of ten species of newts. Phylogenetic comparative analyses revealed that an evolutionary trend of body elongation (increasing axilla-groin distance) is associated with a reduction in head width and forelimb length. Body elongation resulted in reduced maximum running speed, but, surprisingly, also led to a reduction in swimming speed. The evolution of longer tails was associated with an increase in maximal swimming speed. We found no evidence for an evolutionary trade-off between aquatic and terrestrial locomotor performance, probably because of the unexpected negative effect of body elongation on swimming speed. We conclude that the idea of a design conflict between aquatic and terrestrial locomotion, mediated through antagonistic effects of body elongation, does not apply to our model system.     

Thirsty sea snakes forsake refuge during rainfall

Abstract Vertebrates living in dry or salty habitats spend significant amounts of time and energy maintaining their hydro‐mineral balance; any opportunity to drink fresh water should reduce such expenses. However, to find fresh water, individuals are generally forced to leave the safe and buffered conditions that prevail in their refuges. Therefore, they must face harsh environmental conditions and increased predation risks. We examined this trade‐off in the field, using a very large dataset gathered on amphibious sea snakes that confront periodic droughts. The study was carried out on different islets in the lagoon of New Caledonia. As expected, sea kraits remain concealed most of the time under rocks or within burrows during dry weather. They select precise time windows to move rapidly between the sea and the land, thereby minimizing the time spent in the open. Rainfall triggered massive disruptions of this rhythm: many snakes quickly forsook their shelter to drink in the open, and remained almost motionless until satiety. Interestingly, they adopted specific and unusual postures to drink. They perched on the top of rocks that were benefiting from running and less salty water during downpours. They also drank the droplets attached to vegetation during drizzle.     

Long-term field study of sea kraits in New Caledonia: fundamental issues and conservation

This short review focuses on the findings associated with a long-term field study on two species of sea kraits in New Caledonia. Since 2002, more than 30 sites in the lagoon have been sampled, and in most places mark-recapture was implemented. We collected detailed data on more than 14,000 marked individuals (>6000 recaptures) and used different techniques (stable isotopes, bio-logging, analyses of diet). The objective was fundamental: to examine how amphibious snakes cope with both terrestrial and aquatic environments. As access to abundant food is likely the main evolutionary driver for the return transition toward the sea in marine tetrapods, foraging ecology was an important part of the research and novel information was obtained on this subject. Rapidly however, field observations revealed the potential interest of sea kraits for conservation issues. Our results show that these snakes are useful bio-indicators of marine biodiversity; they also provide a useful signal to monitor levels of contamination by heavy metals in the lagoon, and more generally as a means of studying the functioning of reef ecosystems. Importantly, anecdotal observations (e.g., a krait drinking during rain) provided unsuspected physiological insights of general importance to fundamental problems and conservation. One of the lessons of this long-term study is that key results emerged in an unexpected way, but all were dependent on intensive field work.     

An interspecific comparison between morphology and swimming performance in cyprinids

Flow regimes are believed to be of major evolutionary significance in fish. The flow regimes inhabited by cyprinids vary extensively from still flow regimes to riptide flow regimes. To test (i) whether flow‐driven swimming performance and relevant morphological differentiation are present among fish species and (ii) whether evolutionary shifts between high‐flow and low‐flow habitats in cyprinids are associated with evolutionary trade‐offs in locomotor performance, we obtained data on both steady and unsteady swimming performance and external body shape for 19 species of cyprinids that typically occur in different flow regimes (still, intermediate and riptide). We also measured the routine energy expenditure (RMR) and maximum metabolic rate (MMR) and calculated the optimal swimming speed. Our results showed that fish species from riptide groups tend to have a higher critical swimming speed (U_crit), maximum linear velocity (V_max) and fineness ratio (FR) than fish from the other two groups. However, there was no correlation between the reconstructed changes in the steady and unsteady swimming performance of the 19 species. According to the phylogenetically independent contrast (PIC) method, the U_crit was actively correlated with the MMR. These results indicated that selection will favour both higher steady and unsteady swimming performance and a more streamlined body shape in environments with high water velocities. The results suggested that steady swimming performance was more sensitive to the flow regime and that for this reason, changes in body shape resulted more from selective pressure on steady swimming performance than on unsteady swimming performance. No evolutionary trade‐off was observed between steady and unsteady swimming performance, although U_crit and MMR were found to have coevolved. However, a further analysis within each typically occurring habitat group suggested that the trade‐off that may exist between steady and unsteady swimming performance may be concealed by the effect of habitat.     

Effects of oceanic salinity on body condition in sea snakes

Since the transition from terrestrial to marine environments poses strong osmoregulatory and energetic challenges, temporal and spatial fluctuations in oceanic salinity might influence salt and water balance (and hence, body condition) in marine tetrapods. We assessed the effects of salinity on three species of sea snakes studied by mark-recapture in coral-reef habitats in the Neo-Caledonian Lagoon. These three species include one fully aquatic hydrophiine (Emydocephalus annulatus), one primarily aquatic laticaudine (Laticauda laticaudata), and one frequently terrestrial laticaudine (Laticauda saintgironsi). We explored how oceanic salinity affected the snakes' body condition across various temporal and spatial scales relevant to each species' ecology, using linear mixed models and multimodel inference. Mean annual salinity exerted a consistent and negative effect on the body condition of all three snake species. The most terrestrial taxon (L. saintgironsi) was sensitive to salinity over a short temporal scale, corresponding to the duration of a typical marine foraging trip for this species. In contrast, links between oceanic salinity and body condition in the fully aquatic E. annulatus and the highly aquatic L. laticaudata were strongest at a long-term (annual) scale. The sophisticated salt-excreting systems of sea snakes allow them to exploit marine environments, but do not completely overcome the osmoregulatory challenges posed by oceanic conditions. Future studies could usefully explore such effects in other secondarily marine taxa such as seabirds, turtles, and marine mammals.     

Vertebral evolution and ontogenetic allometry: The developmental basis of extreme body shape divergence in microcephalic sea snakes

Snakes exhibit a diverse array of body shapes despite their characteristically simplified morphology. The most extreme shape changes along the precloacal axis are seen in fully aquatic sea snakes (Hydrophiinae): “microcephalic” sea snakes have tiny heads and dramatically reduced forebody girths that can be less than a third of the hindbody girth. This morphology has evolved repeatedly in sea snakes that specialize in hunting eels in burrows, but its developmental basis has not previously been examined. Here, we infer the developmental mechanisms underlying body shape changes in sea snakes by examining evolutionary patterns of changes in vertebral number and postnatal ontogenetic growth. Our results show that microcephalic species develop their characteristic shape via changes in both the embryonic and postnatal stages. Ontogenetic changes cause the hindbodies of microcephalic species to reach greater sizes relative to their forebodies in adulthood, suggesting heterochronic shifts that may be linked to homeotic effects (axial regionalization). However, microcephalic species also have greater numbers of vertebrae, especially in their forebodies, indicating that somitogenetic effects also contribute to evolutionary changes in body shape. Our findings highlight sea snakes as an excellent system for studying the development of segment number and regional identity in the snake precloacal axial skeleton.