Sexual dimorphism is a common phenomenon among animals. The usual cause cited for sexual dimorphism in animals is sexual selection acting through female choice or male-male combat. Natural selection acting to reduce resource competition between the sexes, however, is an important alternative evolutionary scenario, but this possibility has received little empirical study. Here this issue is addressed by examining the relationships among body size, head shape and the functional aspects of diet in the adult male and female cottonmouth snake Agkistrodon piscivorus. In this species, males are larger in overall body and head size. Whereas an analysis of gross head measurements (simple linear head dimensions) shows little dimorphism in head shape, a more detailed analysis of head shape (using digital images of the snakes' heads) revealed some subtle, yet functionally significant, differences in head shape between adult male and female cottonmouths. Specifically, male cottonmouths have longer quadrate bones, and have greater lateral surface areas than females. Male cottonmouths also consumed relatively taller prey (prey size relative to snake body size) than conspecific females, and the sexes consumed significantly different proportions of prey. Because the size of the quadrate bone is a strong determinant of maximum gape in snakes, we suggest that the observed shape differences may reflect functional differences in maximum gape between similarly sized male and female cottonmouths. In turn, such differences in maximum gape width may explain why males consume taller prey than similarly sized females.
Despite repeated acquisitions of aquatic or semi‐aquatic lifestyles revolving around piscivory, snakes have not evolved suction feeding. Instead, snakes use frontally or laterally directed strikes to capture prey under water. If the aquatic medium constrains strike performance because of its physical properties, we predict morphological and functional convergence in snakes that use similar strike behaviours. Here we use natricine snakes to test for such patterns of convergence in morphology and function. Our data show that frontal strikers have converged on a similar morphology characterized by narrow elongate heads with a reduced projected frontal surface area. Moreover, simple computational fluid dynamics models show that the observed morphological differences are likely biologically relevant as they affect the flow of water around the head. In general, our data suggest that the direction of evolution may be predictable if constraints are strong and evolutionary solutions limited.
The evolutionary success of macrostomatan (enlarged-gape) snakes has been attributed to their ability to consume large prey, in turn made possible by their highly kinetic skulls. However, prey can be "large" in several ways, and we have little insight into which aspects of prey size and shape affect skull function during feeding. We used X-ray videos of broad-banded water snakes (Nerodia fasciata) feeding on both frogs and fish to quantify movements of the jaw elements during prey transport, and of the anterior vertebral column during post-cranial swallowing. In a sample of additional individuals feeding on both frogs and fish, we measured the time and the number of jaw protractions needed to transport prey through the buccal cavity. Prey type (fish vs. frog) did not influence transport kinematics, but did influence transport performance. Furthermore, wider and taller prey induced greater movements of most cranial elements, but wider prey were transported with significantly less anterior vertebral bending. In the performance trials, heavier, shorter, and wider prey took significantly more time and a greater number of jaw protractions to ingest. Thus, the functional challenges involved in prey transport depend not only upon prey mass, but also prey type (fish vs. frog) and prey shape (relative height, width and length), suggesting that from the perspective of a gape-limited predator, the difficulty of prey ingestion depends upon multiple aspects of prey size.
SUMMARY The effects of size on animal behaviour, ecology, and physiology are widespread. Theoretical models have been developed to predict how animal form,function, and performance should change with increasing size. Yet, numerous animals undergo dramatic shifts in ecology (e.g. habitat use, diet) that may directly influence the functioning and presumably the scaling of the musculoskeletal system. For example, previous studies have shown that banded watersnakes (Nerodia fasciata) switch from fish prey as juveniles to frog prey as adults, and that fish and frogs represent functionally distinct prey types to watersnakes. We therefore tested whether this ontogenetic shift in diet was coupled to changes in the scaling patterns of the cranial musculoskeletal system in an ontogenetic size series (70–600 mm snout–vent length) of banded watersnakes. We found that all cranial bones and gape size exhibited significant negative allometry, whereas the muscle physiological cross-sectional area (pCSAs) scaled either isometrically or with positive allometry against snout–vent length. By contrast, we found that gape size, most cranial bones, and muscle pCSAs exhibited highly significant positive allometry against head length. Furthermore, the mechanical advantage of the jaw-closing lever system remained constant over ontogeny. Overall, these cranial allometries should enable watersnakes to meet the functional requirements of switching from fusiform fish to bulky frog prey. However, recent studies have reported highly similar allometries in a wide diversity of vertebrate taxa, suggesting that positive allometry within the cranial musculoskeletal system may actually be a general characteristic of vertebrates.
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