Biomechanics of the jaws of spotted ratfish.

Elasmobranch fishes (sharks, skates and rays) consume prey of a variety of sizes and properties, and the feeding mechanism typically reflects diet. Spotted ratfish, Hydrolagus colliei (Holocephali, sister group of elasmobranchs), consume both hard and soft prey; however, the morphology of the jaws does not reflect the characteristics typical of durophagous elasmobranchs. This study investigated the mechanical properties and morphological characteristics of the jaws of spotted ratfish over ontogeny, including strain, stiffness and second moment of area, to evaluate the biomechanical function of the feeding structures. Compressive stiffness of the jaws (E=13.51-21.48 MPa) is similar to that of silicone rubber, a very flexible material. In Holocephali, the upper jaw is fused to the cranium; we show that this fusion reduces deformation experienced by the upper jaw during feeding. The lower jaw resists bending primarily in the posterior half of the jaw, which occludes with the region of the upper jaw that is wider and flatter, thus potentially providing an ideal location for the lower jaw to crush or crack prey. The mechanical properties and morphology of the feeding apparatus of spotted ratfish suggest that while the low compressive stiffness is a material limit of the jaw cartilage, spotted ratfish, and perhaps all holocephalans, evolved structural solutions (i.e. fused upper jaw, shape variation along lower jaw) to meet the demands of a durophagous diet.

Saws, Scissors, and Sharks: Late Paleozoic Experimentation with Symphyseal Dentition.

Sharks of Late Paleozoic oceans evolved unique dentitions for catching and eating soft bodied prey. A diverse but poorly preserved clade, edestoids are noted for developing biting teeth at the midline of their jaws. Helicoprion has a continuously growing root to accommodate >100 crowns that spiraled on top of one another to form a symphyseal whorl supported and laterally braced within the lower jaw. Reconstruction of jaw mechanics shows that individual serrated crowns grasped, sliced, and pulled prey items into the esophagus. A new description and interpretation of Edestus provides insight into the anatomy and functional morphology of another specialized edestoid. Edestus has opposing curved blades of teeth that are segmented and shed with growth of the animal. Set on a long jaw the lower blade closes with a posterior motion, effectively slicing prey across multiple opposing serrated crowns. Further examples of symphyseal whorls among Edestoidae are provided from previously undescribed North American examples of Toxoprion, Campyloprion, Agassizodus, and Sinohelicoprion. The symphyseal dentition in edestoids is associated with a rigid jaw suspension and may have arisen in response to an increase in pelagic cephalopod prey during the Late Paleozoic. Anat Rec, 2018. © 2018 Wiley Periodicals, Inc. Anat Rec, 303:363-376, 2020. © 2018 American Association for Anatomy.

Intra-oropharyngeal food transport and swallowing in white-spotted bamboo sharks.

Despite the importance of intraoral food transport and swallowing, relatively few studies have examined the biomechanics of these behaviors in non-tetrapods, which lack a muscular tongue. Studies show that elasmobranch and teleost fishes generate water currents as a 'hydrodynamic tongue' that presumably transports food towards and into the esophagus. However, it remains largely unknown how specific musculoskeletal motions during transport correspond to food motion. Previous studies of white-spotted bamboo sharks (Chiloscyllium plagiosum) hypothesized that motions of the hyoid, branchial arches and pectoral girdle, generate caudal motion of the food through the long oropharynx of modern sharks. To test these hypotheses, we measured food and cartilage motion with XROMM during intra-oropharyngeal transport and swallowing (N=3 individuals, 2-3 trials per individual). After entering the mouth, food does not move smoothly toward the esophagus, but rather moves in distinct steps with relatively little retrograde motion. Caudal food motion coincides with hyoid elevation and a closed mouth, supporting earlier studies showing that hyoid motion contributes to intra-oropharyngeal food transport by creating caudally directed water currents. Little correspondence between pectoral girdle and food motion was found, indicating minimal contribution of pectoral girdle motion. Transport speed was fast as food entered the mouth, slower and step-wise through the pharyngeal region and then fast again as it entered the esophagus. The food's static periods in the step-wise motion and its high velocity during swallowing could not be explained by hyoid or girdle motion, suggesting these sharks may also use the branchial arches for intra-oropharyngeal transport and swallowing.

Hydrodynamic function of dorsal fins in spiny dogfish and bamboo sharks during steady swimming.

A key feature of fish functional design is the presence of multiple fins that allow thrust vectoring and redirection of fluid momentum to contribute to both steady swimming and maneuvering. A number of previous studies have analyzed the function of dorsal fins in teleost fishes in this context, but the hydrodynamic function of dorsal fins in freely swimming sharks has not been analyzed, despite the potential for differential functional roles between the anterior and posterior dorsal fins. Previous anatomical research has suggested a primarily stabilizing role for shark dorsal fins. We evaluated the generality of this hypothesis by using time-resolved particle image velocimetry to record water flow patterns in the wake of both the anterior and posterior dorsal fins in two species of freely swimming sharks: bamboo sharks (Chiloscyllium plagiosum) and spiny dogfish (Squalus acanthias). Cross-correlation analysis of consecutive images was used to calculate stroke-averaged mean longitudinal and lateral velocity components, and vorticity. In spiny dogfish, we observed a velocity deficit in the wake of the first dorsal fin and flow acceleration behind the second dorsal fin, indicating that the first dorsal fin experiences net drag while the second dorsal fin can aid in propulsion. In contrast, the wake of both dorsal fins in bamboo sharks displayed increased net flow velocity in the majority of trials, reflecting a thrust contribution to steady swimming. In bamboo sharks, fluid flow in the wake of the second dorsal fin had higher absolute average velocity than that for first dorsal fin, and this may result from a positive vortex interaction between the first and second dorsal fins. These data suggest that the first dorsal fin in spiny dogfish has primarily a stabilizing function, while the second dorsal fin has a propulsive function. In bamboo sharks, both dorsal fins can contribute thrust and should be considered as propulsive adjuncts to the body during steady swimming. The function of shark dorsal fins can thus differ considerably among fins and species, and is not limited to a stabilizing role.

Function of the hypobranchial muscles and hyoidiomandibular ligament during suction capture and bite processing in white-spotted bamboo sharks, Chiloscyllium plagiosum.

Suction feeding in teleost fish is a power-dependent behavior, requiring rapid and forceful expansion of the orobranchial cavity by the hypobranchial and trunk muscles. To increase power production for expansion, many species employ in-series tendons and catch mechanisms to store and release elastic strain energy. Suction feeding sharks such as Chiloscyllium plagiosum lack large in-series tendons on the hypobranchials, yet two of the hypobranchials, the coracohyoideus and coracoarcualis (CH and CA; hyoid depressors), are arranged in-series, and run deep and parallel to a third muscle, the coracomandibularis (CM, jaw depressor). The arrangement of the CH and CA suggests that C. plagiosum is using the CH muscle rather than a tendon to store and release elastic strain energy. Here we describe the anatomy of the feeding apparatus, and present data on hyoid and jaw kinematics and fascicle shortening in the CM, CH and CA quantified using sonomicrometry, with muscle activity and buccal pressure recorded simultaneously. Results from prey capture show that prior to jaw and hyoid depression the CH is actively lengthened by shortening of the in-series CA. The active lengthening of the CH and pre-activation of the CH and CA suggest that the CH is functioning to store and release elastic energy during prey capture. Catch mechanisms are proposed involving a dynamic moment arm and four-bar linkage between the hyoidiomandibular ligament (LHML), jaws and ceratohyals that is influenced by the CM. Furthermore, the LHML may be temporarily disengaged during behaviors such as bite processing to release linkage constraints.

Dual function of the pectoral girdle for feeding and locomotion in white-spotted bamboo sharks.

Positioned at the intersection of the head, body and forelimb, the pectoral girdle has the potential to function in both feeding and locomotor behaviours-although the latter has been studied far more. In ray-finned fishes, the pectoral girdle attaches directly to the skull and is retracted during suction feeding, enabling the ventral body muscles to power rapid mouth expansion. However, in sharks, the pectoral girdle is displaced caudally and entirely separate from the skull (as in tetrapods), raising the question of whether it is mobile during suction feeding and contributing to suction expansion. We measured three-dimensional kinematics of the pectoral girdle in white-spotted bamboo sharks during suction feeding with X-ray reconstruction of moving morphology, and found the pectoral girdle consistently retracted about 11° by rotating caudoventrally about the dorsal scapular processes. This motion occurred mostly after peak gape, so it likely contributed more to accelerating captured prey through the oral cavity and pharynx, than to prey capture as in ray-finned fishes. Our results emphasize the multiple roles of the pectoral girdle in feeding and locomotion, both of which should be considered in studying the functional and evolutionary morphology of this structure.

Ontogeny of head and caudal fin shape of an apex marine predator: The tiger shark (Galeocerdo cuvier).

How morphology changes with size can have profound effects on the life history and ecology of an animal. For apex predators that can impact higher level ecosystem processes, such changes may have consequences for other species. Tiger sharks (Galeocerdo cuvier) are an apex predator in tropical seas, and, as adults, are highly migratory. However, little is known about ontogenetic changes in their body form, especially in relation to two aspects of shape that influence locomotion (caudal fin) and feeding (head shape). We captured digital images of the heads and caudal fins of live tiger sharks from Southern Florida and the Bahamas ranging in body size (hence age), and quantified shape of each using elliptical Fourier analysis. This revealed changes in the shape of the head and caudal fin of tiger sharks across ontogeny. Smaller juvenile tiger sharks show an asymmetrical tail with the dorsal (upper) lobe being substantially larger than the ventral (lower) lobe, and transition to more symmetrical tail in larger adults, although the upper lobe remains relatively larger in adults. The heads of juvenile tiger sharks are more conical, which transition to relatively broader heads over ontogeny. We interpret these changes as a result of two ecological transitions. First, adult tiger sharks can undertake extensive migrations and a more symmetrical tail could be more efficient for swimming longer distances, although we did not test this possibility. Second, adult tiger sharks expand their diet to consume larger and more diverse prey with age (turtles, mammals, and elasmobranchs), which requires substantially greater bite area and force to process. In contrast, juvenile tiger sharks consume smaller prey, such as fishes, crustaceans, and invertebrates. Our data reveal significant morphological shifts in an apex predator, which could have effects for other species that tiger sharks consume and interact with.

Performance of teeth of lingcod, Ophiodon elongatus, over ontogeny.

Fish teeth can play several roles during feeding; capture, retention, and processing. In many fish lineages teeth may be present on non-jaw cranial bones that lack opposing teeth, such as the vomer and palatine. We hypothesized that teeth on different bones have different functions, and that the function of a set of teeth may vary over ontogeny. In this study, puncture, and draw performance of in situ vomerine teeth are compared to premaxillary teeth of the piscivorous lingcod, Ophiodon elongatus. The force required to pierce prey and to draw prey out of the mouth once the teeth were embedded was measured in ten individuals ranging from 205 to 836 mm SL to test for ontogenetic effects. Vomerine teeth in juvenile lingcod required proportionally less force to puncture prey items than adult lingcod, while premaxillary teeth showed the opposite trend. Draw force required to remove prey from the grasp of both toothed bones show the same shift with ontogeny. These results suggest that there is a shift in tooth function from vomerine to premaxillary teeth over ontogeny of lingcods. In juvenile lingcod, vomerine teeth function more effectively during initial puncture. In contrast, the premaxillary teeth pierce more effectively in adults. Juvenile lingcod are expected to use the premaxillary teeth while adult lingcod are expected to use the vomerine teeth to retain prey due to the larger force required for the prey to escape. The curvature of vomerine teeth increases over ontogeny suggesting increasing functional performance in retaining prey.

Eating with a saw for a jaw: functional morphology of the jaws and tooth-whorl in Helicoprion davisii.

The recent reexamination of a tooth-whorl fossil of Helicoprion containing intact jaws shows that the symphyseal tooth-whorl occupies the entire length of Meckel's cartilage. Here, we use the morphology of the jaws and tooth-whorl to reconstruct the jaw musculature and develop a biomechanical model of the feeding mechanism in these early Permian predators. The jaw muscles may have generated large bite-forces; however, the mechanics of the jaws and whorl suggest that Helicoprion was better equipped for feeding on soft-bodied prey. Hard shelled prey would tend to slip anteriorly from the closing jaws due to the curvature of the tooth-whorl, lack of cuspate teeth on the palatoquadrate (PQ), and resistance of the prey. When feeding on soft-bodied prey, deformation of the prey traps prey tissue between the two halves of the PQ and the whorl. The curvature of the tooth-whorl and position of the exposed teeth relative to the jaw joint results in multiple tooth functions from anterior to posterior tooth that aid in feeding on soft-bodied prey. Posterior teeth cut and push prey deeper into the oral cavity, while middle teeth pierce and cut, and anterior teeth hook and drag more of the prey into the mouth. Furthermore, the anterior-posterior edges of the teeth facilitate prey cutting with jaw closure and jaw depression. The paths traveled by each tooth during jaw depression are reminiscent of curved pathways used with slashing weaponry such as swords and knifes. Thus, the jaws and tooth-whorl may have formed a multifunctional tool for capturing, processing, and transporting prey by cyclic opening and closing of the lower jaw in a sawing fashion.

Anatomy and muscle activity of the dorsal fins in bamboo sharks and spiny dogfish during turning maneuvers.

Stability and procured instability characterize two opposing types of swimming, steady and maneuvering, respectively. Fins can be used to manipulate flow to adjust stability during swimming maneuvers either actively using muscle control or passively by structural control. The function of the dorsal fins during turning maneuvering in two shark species with different swimming modes is investigated here using musculoskeletal anatomy and muscle function. White-spotted bamboo sharks are a benthic species that inhabits complex reef habitats and thus have high requirements for maneuverability. Spiny dogfish occupy a variety of coastal and continental shelf habitats and spend relatively more time cruising in open water. These species differ in dorsal fin morphology and fin position along the body. Bamboo sharks have a larger second dorsal fin area and proportionally more muscle insertion into both dorsal fins. The basal and radial pterygiophores are plate-like structures in spiny dogfish and are nearly indistinguishable from one another. In contrast, bamboo sharks lack basal pterygiophores, while the radial pterygiophores form two rows of elongated rectangular elements that articulate with one another. The dorsal fin muscles are composed of a large muscle mass that extends over the ceratotrichia overlying the radials in spiny dogfish. However, in bamboo sharks, the muscle mass is divided into multiple distinct muscles that insert onto the ceratotrichia. During turning maneuvers, the dorsal fin muscles are active in both species with no differences in onset between fin sides. Spiny dogfish have longer burst durations on the outer fin side, which is consistent with opposing resistance to the medium. In bamboo sharks, bilateral activation of the dorsal in muscles could also be stiffening the fin throughout the turn. Thus, dogfish sharks passively stiffen the dorsal fin structurally and functionally, while bamboo sharks have more flexible dorsal fins, which result from a steady swimming trade off.

Jaws for a spiral-tooth whorl: CT images reveal novel adaptation and phylogeny in fossil Helicoprion.

New CT scans of the spiral-tooth fossil, Helicoprion, resolve a longstanding mystery concerning the form and phylogeny of this ancient cartilaginous fish. We present the first three-dimensional images that show the tooth whorl occupying the entire mandibular arch, and which is supported along the midline of the lower jaw. Several characters of the upper jaw show that it articulated with the neurocranium in two places and that the hyomandibula was not part of the jaw suspension. These features identify Helicoprion as a member of the stem holocephalan group Euchondrocephali. Our reconstruction illustrates novel adaptations, such as lateral cartilage to buttress the tooth whorl, which accommodated the unusual trait of continuous addition and retention of teeth in a predatory chondrichthyan. Helicoprion exemplifies the climax of stem holocephalan diversification and body size in Late Palaeozoic seas, a role dominated today by sharks and rays.

Prey handling using whole-body fluid dynamics in batoids.

Fluid flow generated by body movements is a foraging tactic that has been exploited by many benthic species. In this study, the kinematics and hydrodynamics of prey handling behavior in little skates, Leucoraja erinacea, and round stingrays, Urobatis halleri, are compared using kinematics and particle image velocimetry. Both species use the body to form a tent to constrain the prey with the pectoral fin edges pressed against the substrate. Stingrays then elevate the head, which increases the volume between the body and the substrate to generate suction, while maintaining pectoral fin contact with the substrate. Meanwhile, the tip of the rostrum is curled upwards to create an opening where fluid is drawn under the body, functionally analogous to suction-feeding fishes. Skates also rotate the rostrum upwards although with the open rostral sides and the smaller fin area weaker fluid flow is generated. However, skates also use a rostral strike behavior in which the rostrum is rapidly rotated downwards pushing fluid towards the substrate to potentially stun or uncover prey. Thus, both species use the anterior portion of the body to direct fluid flow to handle prey albeit in different ways, which may be explained by differences in morphology. Rostral stiffness and pectoral fin insertion onto the rostrum differ between skates and rays and this corresponds to behavioral differences in prey handling resulting in distinct fluid flow patterns. The flexible muscular rostrum and greater fin area of stingrays allow more extensive use of suction to handle prey while the stiff cartilaginous rostrum of skates lacking extensive fin insertion is used as a paddle to strike prey as well as to clear away sand cover.

Fluid dynamics of feeding behaviour in white-spotted bamboo sharks.

Although the motor control of feeding is presumed to be generally conserved, some fishes are capable of modulating the feeding behaviour in response to prey type and or prey size. This led to the 'feeding modulation hypothesis', which states that rapid suction strikes are pre-programmed stereotyped events that proceed to completion once initiated regardless of sensory input. If this hypothesis holds true, successful strikes should be indistinguishable from unsuccessful strikes owing to a lack of feedback control in specialized suction feeding fishes. The hydrodynamics of suction feeding in white-spotted bamboo sharks (Chiloscyllium plagiosum) was studied in three behaviours: successful strikes, intraoral transports of prey and unsuccessful strikes. The area of the fluid velocity region around the head of feeding sharks was quantified using time-resolved digital particle image velocimetry (DPIV). The maximal size of the fluid velocity region is 56% larger in successful strikes than unsuccessful strikes (10.79 cm2 vs 6.90 cm2), but they do not differ in duration, indicating that strikes are modulated based on some aspect of the prey or simply as a result of decreased effort on the part of the predator. The hydrodynamic profiles of successful and unsuccessful strikes differ after 21 ms, a period probably too short to provide time to react through feedback control. The predator-to-prey distance is larger in missed strikes compared with successful strikes, indicating that insufficient suction is generated to compensate for the increased distance. An accuracy index distinguishes unsuccessful strikes (-0.26) from successful strikes (0.45 to 0.61). Successful strikes occur primarily between the horizontal axis of the mouth and the dorsal boundary of the ingested parcel of water, and missed prey are closer to the boundary or beyond. Suction transports are shorter in duration than suction strikes but have similar maximal fluid velocity areas to move the prey through the oropharyngeal cavity into the oesophagus (54 ms vs 67 ms).

Suction generation in white-spotted bamboo sharks Chiloscyllium plagiosum.

After the divergence of chondrichthyans and teleostomes, the structure of the feeding apparatus also diverged leading to alterations in the suction mechanism. In this study we investigated the mechanism for suction generation during feeding in white-spotted bamboo sharks, Chiloscyllium plagiosum and compared it with that in teleosts. The internal movement of cranial elements and pressure in the buccal, hyoid and pharyngeal cavities that are directly responsible for suction generation was quantified using sonomicrometry and pressure transducers. Backward stepwise multiple linear regressions were used to explore the relationship between expansion and pressure, accounting for 60-96% of the variation in pressure among capture events. The progression of anterior to posterior expansion in the buccal, hyoid and pharyngeal cavities is accompanied by the sequential onset of subambient pressure in these cavities as prey is drawn into the mouth. Gape opening triggers the onset of subambient pressure in the oropharyngeal cavities. Peak gape area coincides with peak subambient buccal pressure. Increased velocity of hyoid area expansion is primarily responsible for generating peak subambient pressure in the buccal and hyoid regions. Pharyngeal expansion appears to function as a sink to receive water influx from the mouth, much like that of compensatory suction in bidirectional aquatic feeders. Interestingly, C. plagiosum generates large suction pressures while paradoxically compressing the buccal cavity laterally, delaying the time to peak pressure. This represents a fundamental difference from the mechanism used to generate suction in teleost fishes. Interestingly, pressure in the three cavities peaks in the posterior to anterior direction. The complex shape changes that the buccal cavity undergoes indicate that, as in teleosts, unsteady flow predominates during suction feeding. Several kinematic variables function together, with great variation over long gape cycles to generate the low subambient pressures used by C. plagiosum to capture prey.

Evolution of asynchronous motor activity in paired muscles: effects of ecology, morphology, and phylogeny.

Many studies of feeding behavior have implanted electrodes unilaterally (in muscles on only one side of the head) to determine the basic motor patterns of muscles controlling the jaws. However, bilateral implantation has the potential to achieve a more comprehensive understanding of modification of the motor activity that may be occurring between the left and right sides of the head. In particular, complex processing of prey is often characterized by bilaterally asynchronous and even unilateral activation of the jaw musculature. In this study, we bilaterally implant feeding muscles in species from four orders of elasmobranchs (Squaliformes, Orectolobiformes, Carcharhiniformes, Rajoidea) in order to characterize the effects of type of prey, feeding behavior, and phylogeny on the degree of asynchronous muscle activation. Electrodes were implanted in three of the jaw adductors, two divisions of the quadratomandibularis and the preorbitalis, as well as in a cranial elevator in sharks, the epaxialis. The asynchrony of feeding events (measured as the degree to which activity of members of a muscle pair is out of phase) was compared across species for capture versus processing and simple versus complex prey, then interpreted in the contexts of phylogeny, morphology, and ecology to clarify determinants of asynchronous activity. Whereas capture and processing of prey were characterized by statistically similar degrees of asynchrony for data pooled across species, events involving complex prey were more asynchronous than were those involving simple prey. The two trophic generalists, Squalus acanthias and Leucoraja erinacea, modulated the degree of asynchrony according to type of prey, whereas the two behavioral specialists, Chiloscyllium plagiosum and Mustelus canis, activated the cranial muscles synchronously regardless of type of prey. These differences in jaw muscle activity would not have been detected with unilateral implantation. Therefore, we advocate bilateral implantation in studies of cranial muscle function in fishes, particularly when investigating behaviors associated with processing complex prey. Incorporating this methodology will provide a more detailed understanding of the coordination and evolution of paired-muscle function in the feeding apparatus relative to behavioral and ecological performance.

Morphology and mechanics of the teeth and jaws of white-spotted bamboo sharks (Chiloscyllium plagiosum).

The teeth of white-spotted bamboo sharks (Chiloscyllium plagiosum) are used to clutch soft-bodied prey and crush hard prey; however, the dual function is not evident from tooth morphology alone. Teeth exhibit characteristics that are in agreement with a clutching-type tooth morphology that is well suited for grasping and holding soft-bodied prey, but not for crushing hard prey. The dual role of this single tooth morphology is facilitated by features of the dental ligament and jaw joint. Tooth attachment is flexible and elastic, allowing movement in both sagittal and frontal planes. During prey capture spike-like tooth cusps pierce the flesh of soft prey, thereby preventing escape. When processing prey harder than the teeth can pierce the teeth passively depress, rotating inward towards the oral cavity such that the broader labial faces of the teeth are nearly parallel to the surface of the jaws and form a crushing surface. Movement into the depressed position increases the tooth surface area contacting prey and decreases the total stress applied to the tooth, thereby decreasing the risk of structural failure. This action is aided by a jaw joint that is ventrally offset from the occlusal planes of the jaws. The offset joint position allows many teeth to contact prey simultaneously and orients force vectors at contact points between the jaws and prey in a manner that shears or rolls prey between the jaws during a bite, thus, aiding in processing while reducing forward slip of hard prey from the mouth. Together the teeth, dental ligament, and jaws form an integrated system that may be beneficial to the feeding ecology of C. plagiosum, allowing for a diet that includes prey of varying hardness and elusiveness.

Evolution and ecology of feeding in elasmobranchs.

Paleozoic chondrichthyans had a large gape, numerous spike-like teeth, limited cranial kinesis, and a non-suspensory hyoid, suggesting a feeding mechanism dominated by bite and ram. Modern sharks are characterized by a mobile upper jaw braced by a suspensory hyoid arch that is highly kinetic. In batoids, the upper jaw is dissociated from the cranium permitting extensive protrusion of the jaws. Similar to actinopterygians, the evolution of highly mobile mandibular and hyoid elements has been correlated with extensive radiation of feeding modes in elasmobranchs, particularly that of suction. Modern elasmobranchs possess a remarkable variety of feeding modes for a group containing so few species. Biting, suction or filter-feeding may be used in conjunction with ram to capture prey, with most species able to use a combination of behaviors during a strike. Suction-feeding has repeatedly arisen within all recent major elasmobranch clades and is associated with a suite of morphological and behavioral specializations. Prey capture in a diverse assemblage of purported suction-feeding elasmobranchs is investigated in this study. Drop in water pressure measured in the mouth and at the location of the prey shows that suction inflow drops off rapidly with distance from the predator's mouth. Elasmobranchs specializing in suction-feeding may be limited to bottom associated prey and because of their small gape may have a diet restricted to relatively small prey. Behavior can affect performance and overcome constraints imposed by the fluid medium. Suction performance can be enhanced by proximity to a substrate or by decreasing distance from predator to prey using various morphological and/or behavioral characteristics. Benthic suction-feeders benefit by the increased strike radius due to deflection of water flow when feeding close to a substrate, and perhaps require less accuracy when capturing prey. Suction and ram-suction-feeding elasmobranchs can also use suction inflow to draw prey to them from a short distance, while ram-feeding sharks must accelerate and overtake the prey. The relationship between feeding strategy and ecology may depend in part on ecological, mechanistic or evolutionary specialization. Mechanistic suction-feeding specialist elasmobranchs are primarily benthic, while most epibenthic and pelagic elasmobranchs are generalists and use ram, suction, and biting to catch a diversity of prey in various habitats. Some shark species are considered to be ecological specialists in choosing certain kinds of prey over others. Batoids are evolutionary specialists in having a flattened morphology and most are generalist feeders. Filter-feeding elasmobranchs are ecological, mechanistic, and evolutionary specialists.

Eating without hands or tongue: specialization, elaboration and the evolution of prey processing mechanisms in cartilaginous fishes.

The ability to separate edible from inedible portions of prey is integral to feeding. However, this is typically overlooked in favour of prey capture as a driving force in the evolution of vertebrate feeding mechanisms. In processing prey, cartilaginous fishes appear handicapped because they lack the pharyngeal jaws of most bony fishes and the muscular tongue and forelimbs of most tetrapods. We argue that the elaborate cranial muscles of some cartilaginous fishes allow complex prey processing in addition to their usual roles in prey capture. The ability to manipulate prey has evolved twice along different mechanical pathways. Batoid chondrichthyans (rays and relatives) use elaborate lower jaw muscles to process armored benthic prey, separating out energetically useless material. In contrast, megacarnivorous carcharhiniform and lamniform sharks use a diversity of upper jaw muscles to control the jaws while gouging, allowing for reduction of prey much larger than the gape. We suggest experimental methods to test these hypotheses empirically.