How anatomy influences measurements of snakes.

Anatomy compromises the precision and accuracy of measurements made of the body length and head size of live snakes. Body measures (snout-vent length, SVL) incorporate many synovial intervertebral joints, each allowing flexion and limited extension and compression. Radiographs of the trunk in 14 phylogenetically diverse species in resting and stretched conditions combined with dissections and histological analysis of intervertebral joints show that the synovial nature of these joints underlies the variance in SVL measures.

Dorsal root ganglia, neural crest migration, and spinal cord form in snakes.

The classic view of the vertebrate dorsal root ganglion is that it arises from trunk neural crest cells that migrate to positions lateral to the spinal cord, sending axons dorsally into the spinal cord and dendrites ventrally to meet with motor axons in the ventral root to form spinal nerves. As a result, the ganglion ends up lying in the dorsal root of the spinal nerve. Serial histological sections of parts of the trunk of juveniles of different snake species revealed that the ganglia lie distal to the junction of dorsal and ventral roots of spinal nerves and outside the neural canal.

Interrogating Genomic-Scale Data for Squamata (Lizards, Snakes, and Amphisbaenians) Shows no Support for Key Traditional Morphological Relationships.

Genomics is narrowing uncertainty in the phylogenetic structure for many amniote groups. For one of the most diverse and species-rich groups, the squamate reptiles (lizards, snakes, and amphisbaenians), an inverse correlation between the number of taxa and loci sampled still persists across all publications using DNA sequence data and reaching a consensus on the relationships among them has been highly problematic. In this study, we use high-throughput sequence data from 289 samples covering 75 families of squamates to address phylogenetic affinities, estimate divergence times, and characterize residual topological uncertainty in the presence of genome-scale data.

The suction mechanism of the pipid frog, Pipa pipa (Linnaeus, 1758).

Most suction-feeding, aquatic vertebrates create suction by rapidly enlarging the oral cavity and pharynx. Forceful enlargement of the pharynx is powered by longitudinal muscles that retract skeletal elements of the hyoid, more caudal branchial arches, and, in many fish, the pectoral girdle. This arrangement was thought to characterize all suction-feeding vertebrates.

Highly extensible skeletal muscle in snakes.

Many snakes swallow large prey whole, and this process requires large displacements of the unfused tips of the mandibles and passive stretching of the soft tissues connecting them. Under these conditions, the intermandibular muscles are highly stretched but subsequently recover normal function. In the highly stretched condition we observed in snakes, sarcomere length (SL) increased 210% its resting value (SL0), and actin and myosin filaments no longer overlapped.

Snake lower jaw skin: extension and recovery of a hyperextensible keratinized integument.

The skin of squamates consists of a keratinized epidermis divided into thick scale and thinner, folded interscale regions underlain by a dermis containing a complex array of fibrous connective tissues. We examined the skin of the lower jaw of watersnakes (Nerodia sipedon) to determine how skin morphology changes when highly stretched during ingestion of large prey. Video records of skin behavior in the lower jaws of watersnakes feeding on fish or anesthetized watersnakes being stretched on an Instron machine showed that most skin extension involves the interscale skin.

A model of the anterior esophagus in snakes, with functional and developmental implications.

The gross anatomy of the mouth of snakes has always been interpreted as an evolutionary response to feeding demands. In most alethinophidian species, their anatomy allows limited functional independence of right and left sides and the roof and floor of the mouth as well as wide separation of the tips of the mandibles. However, locations of the tongue and glottis in snakes suggest extraordinary rearrangement of pharyngeal structures characteristic of all vertebrates.

Mammals as prey: estimating ingestible size.

Most mammals have deformable bodies, making it difficult to measure the size of living or freshly killed ones accurately. Because small rodents are common prey of many snakes, and because nearly all snakes swallow their prey whole, we explored four methods for determining the ingestible size (the smallest cross-sectional area that the largest part of the rodent can be made into without breaking bones or dislocating joints) of 100 intact rodents, including 50 Musmusculus and 50 Rattus norvegicus. Cross-sectional areas derived from maximal height and width of specimens at rest or the same specimens wrapped snout to pelvic girdle are roughly 1.

Drinking in snakes: resolving a biomechanical puzzle.

Snakes have long been thought to drink with a two-phase buccal-pump mechanism, but observations that some snakes can drink without sealing the margins of their mouths suggest that buccal pumping may not be the only drinking mechanism used by snakes. Here, we report that some snakes appear to drink using sponge-like qualities of specific regions of the oropharyngeal and esophageal mucosa and sponge-compressing functions of certain muscles and bones of the head. The resulting mechanism allows them to transport water upward against the effects of gravity using movements much slower than those of many other vertebrates.

Functional morphology of the palato-maxillary apparatus in "Palatine dragging" snakes (Serpentes: Elapidae: Acanthophis, Oxyuranus).

Elapid snakes have previously been divided into two groups (palatine erectors and palatine draggers) based on the morphology and inferred movements of their palatine bone during prey transport (swallowing). We investigated the morphology and the functioning of the feeding apparatus of several palatine draggers (Acanthophis antarcticus, Oxyuranus scutellatus, Pseudechis australis) and compared them to published records of palatine erectors. We found that the palatine in draggers does not move as a straight extension of the pterygoid as originally proposed.

Viper fangs: functional limitations of extreme teeth.

The fangs of vipers are extremely long, rotating, hollow teeth. Analysis of video records of more than 750 strikes recorded at 60 or 250 frames per second for 285 individuals representing 86 species in 31 genera shows that vipers reposition fangs after initial contact with prey in more than a third of the strikes. Repositioning resulted when fangs missed prey entirely or hit prey regions that did not permit adequate penetration.

Dynamic changes in body form during swimming in the water snake Nerodia sipedon.

Video records of swimming water snakes show that during moderate to rapid swimming, the rear half to two-thirds of the trunk is compressed laterally, approaching the body form of some sea snakes. Body form of swimming snakes differed significantly from their shape when resting on a flat surface or when anesthetized and suspended in water. The extent of lateral flattening is positively correlated with swimming speed, a relationship generally supported by tests of trunk models in a flow tank.

Kinematic analysis of an appetitive food-handling behavior: the functional morphology of Syrian hamster cheek pouches.

Prodigious food hoarding in Syrian hamsters Mesocricetus auratus Waterhouse is strongly linked to appetite and is made possible by large internal cheek pouches. We provide a functional analysis of the cheek pouch and its associated retractor muscle. Frame-by-frame analysis of videotaped pouch-filling behavior revealed multiple jaw cycles for each food item pouched and the use of more jaw cycles to pouch large food items ( approximately 2.

Feeding in Atractaspis (Serpentes: Atractaspididae): a study in conflicting functional constraints.

African fossorial colubroid snakes of the genus Atractaspis have relatively long fangs on short maxillae, a gap separating the pterygoid and palatine bones, a toothless pterygoid, and a snout tightly attached to the rest of the skull. They envenomate prey with a unilateral backward stab of one fang projected from a closed mouth. We combined structural reanalysis of the feeding apparatus, video records of prey envenomation and transport, and manipulations of live and dead Atractaspis to determine how structure relates to function in this unusual genus of snakes.

Prey transport in "palatine-erecting" elapid snakes.

Cobras and mambas are members of a group of elapid snakes supposedly united by the morphology and inferred behavior of their palatine bone during prey transport (palatine erectors). The palatine erectors investigated (Dendroaspis polylepis, Naja pallida, Ophiophagus hannah, Aspidelaps scutatus, A. lubricus) show differences in the morphology of their feeding apparatus that do not affect the overall behavior of the system.

Rhinokinetic snout of thamnophiine snakes.

Radiographic and cinegraphic behavioral data, combined with anatomical evidence, indicate that the snout in Nerodia and Thamnophis consists of four movable elements (1, premaxilla; 2, paired nasals; 3, right septomaxilla and vomer; and 4, left septomaxilla and vomer), a condition we refer to as rhinokinetic. In thamnophiine snakes, movements of the snout bones allow the teeth of the right and left sides to separate further and increase the effective stroke distance of each palatomaxillary cycle during swallowing. Histological and microdissectional analyses suggest that snout movement is keyed to the placement of the cartilaginous nasal septum and associated nasal capsules relative to the surrounding bones.

Morphology and behavior of the feeding apparatus in Cryptobranchus alleganiensis (Amphibia: Caudata).

Cine and high-speed videographic analyses of feeding in Cryptobranchus alleganiensis demonstrate that prey are captured by either inertial suction or a strike combined with suction. Movements of cephalic elements during capture are generally similar to those of other suction-feeding vertebrates but more variable than those of most aquatic salamanders. Following capture, prey frequently are manipulated and transported into and out of the buccal cavity across the teeth.

Variations of the cephalic muscles in the colubrid snake genera Entechinus, Opheodrys, and Symphimus.

The cephalic muscles in three species of Entechinus, two species of Opheodrys, and Symphimus mayae display patterns of interspecific variation that are largely congruent with patterns of variation previously described for the skulls of these species. This congruence does not stem from direct correlation between the shapes of associated bones and muscles. In these colubrid snakes, most interspecific variations in muscle form involve changes in the shape or relative position of attachment points that appear unrelated to changes in the gross form of the bony surfaces forming the attachment points and produce no major changes in the architectural array of fibers in the muscle.

The feeding apparatus of the salamander Amphiuma tridactylum: Morphology and behavior.

Anatomical studies of cephalic bones and muscles combined with cine and high-speed videographic analyses of feeding demonstrate that Amphiuma tridactylum uses two distinct types of suction feeding. Small or relatively immobile prey generally elicit a stationary capture mode in which mouth opening precedes buccal expansion and there is no forward movement of the head of the salamander. Actively moving prey are captured by a rapid strike during which mouth opening and buccal expansion are synchronous and the extent of buccal expansion is greater than in stationary feeding.