Functional morphology of terrestrial prey capture in salamandrid salamanders.

Salamanders use the hyobranchial apparatus and its associated musculature for tongue projection on land and for suction feeding in water. Hyobranchial apparatus composition and morphology vary across species, and different morphologies are better suited for feeding in aquatic versus terrestrial environments. We hypothesize that differences in hyobranchial morphology result in functional trade-offs in feeding performance. We predict that semi-aquatic and aquatic salamandrids with hyobranchial morphology suited for aquatic feeding will have lower performance, in terms of tongue-projection distance, velocity, acceleration and power, compared with terrestrial salamandrids when feeding in a terrestrial environment. We found that semi-aquatic and aquatic newts had lower velocity, acceleration and muscle-mass-specific power of tongue projection when compared with the terrestrial salamanders Chioglossa lusitanica and Salamandra salamandra The fully aquatic newt, Paramesotriton labiatus, has a robust, heavily mineralized hyobranchial apparatus and was unable to project its tongue during terrestrial feeding, and instead exhibited suction-feeding movements better suited for aquatic feeding. Conversely, terrestrial species have slender, cartilaginous hyobranchial apparatus and enlarged tongue pads that coincided with greater tongue-projection distance, velocity, acceleration and power. Chioglossalusitanica exhibited extreme tongue-projection performance, similar to that seen in elastically projecting plethodontid salamanders; muscle-mass-specific power of tongue projection exceeded 2200 W kg-1, more than 350 times that of the next highest performer, Ssalamandra, which reached 6.3 W kg-1 These findings reveal that two fully terrestrial salamandrids have morphological specializations that yield greater tongue-projection performance compared with species that naturally feed in both aquatic and terrestrial environments.

Functional trade-offs in the aquatic feeding performance of salamanders.

During aquatic feeding salamanders use the hyobranchial apparatus to capture prey. The hyobranchial apparatus depresses the floor of the mouth, effectively expanding the oropharyngeal cavity and generating suction. Within the family Salamandridae, there is a wide range of ecological diversity, with salamanders being terrestrial, semi-aquatic, or aquatic as adults. The purpose of this research was to quantify the diverse morphology and suction feeding performance of aquatically feeding salamandrids. We hypothesized that a more robust hyobranchial apparatus morphology would yield increased aquatic feeding performance. When compared to semi-aquatic newts, the fully aquatic species Paramesotriton labiatus had greater mineralization of the hyobranchial apparatus, as well as relatively more narrow basibranchial and wider ceratobranchial I+II complexes. These morphological differences coincide with greater aquatic feeding performance. Kinematics from high-speed videography revealed that maximum mouth opening velocity and acceleration were approximately two and five times greater, respectively, in Paramesotriton, and hyobranchial depression acceleration was found to be approximately three times greater than in the semi-aquatic species Pleurodeles waltl, Notophthalmus viridescens, Triturus dobrogicus, and Cynops cyanurus. Using digital particle image velocimetry, peak and average fluid velocity generated in Paramesotriton during suction feeding events were found to be 0.5ms-1 and 0.2ms-1, respectively, doubling that of all semi-aquatic species. These findings reveal that specialized morphology increases aquatic feeding performance in a fully aquatic newt.

Extreme Performance and Functional Robustness of Movement are Linked to Muscle Architecture: Comparing Elastic and Nonelastic Feeding Movements in Salamanders.

Muscle-powered movements are limited by the contractile properties of muscles and are sensitive to temperature changes. Elastic-recoil mechanisms can both increase performance and mitigate the effects of temperature on performance. Here, we compare feeding movements in two species of plethodontid salamanders, Bolitoglossa franklini and Desmognathus quadramaculatus, across a range of body temperatures (5-25°C) to better understand the mechanism of elastically powered, thermally robust movements. Bolitoglossa exhibited ballistic, elastically powered tongue projection with a maximum muscle mass specific power of 4,642 W kg(-1) while Desmognathus demonstrated nonballistic, muscle-powered tongue projection with a maximum power of 359 W kg(-1) . Tongue-projection performance in Bolitoglossa was more thermally robust than that of Desmognathus, especially below 15°C. The improved performance and thermal robustness of Bolitoglossa was associated with morphological changes in the projector muscle, including elaborated collagen aponeuroses and the absence of myofibers attaching directly to the tongue skeleton. The elongated aponeuroses likely increase the capacity for elastic energy storage, and the lack of myofibers inserting on the tongue skeleton permits ballistic projection. These results suggest that relatively simple changes in myofiber architecture and the amount of connective tissue can improve the performance and functional robustness of movements in the face of environmental challenges such as variable temperature.

Mechanical properties of sand tiger shark (Carcharias taurus) vertebrae in relation to spinal deformity.

Approximately 35% of sand tiger sharks (Carcharias taurus) in public aquaria exhibit spinal deformities ranging from compressed vertebrae and loss of intervertebral space to dislocated spines with vertebral degeneration and massive spondylosis caused by excessive mineralization both within vertebrae and outside the notochordal sheath. To identify the mechanical basis of these deformities, vertebral centra from affected (N=12) and non-affected (N=9) C. taurus were subjected to axial compression tests on an MTS 858 Bionix material testing system, after which mineral content was determined. Vertebral centra from affected sharks had significantly lower mineral content and material behavior in nearly all variables characterizing elasticity, plasticity and failure. These mechanical deficiencies are correlated with size at capture, capture method, vitamin C and zinc deficiency, aquarium size and swimming behavior in public aquaria. Non-affected C. taurus had greater stiffness and toughness even though these properties are generally incompatible in mineralized structures, suggesting that the biphasic (mineralized, unmineralized phases) nature of chondrichthyan vertebrae yields material behavior not otherwise observed in vertebrate skeletons. However, vertebral centra from non-affected sharks had lower mineral content (33%), stiffness (167 MPa), yield strain (14%) and ultimate strength (16 MPa) than other species of sharks and bony vertebrates, indicating that biomechanical precautions must be taken in the husbandry of this species.