Functional morphology of the feeding apparatus,feeding constraints,and suction performance in the nurse shark Ginglymostoma cirratum

The nurse shark, Ginglymostoma cirratum, is an obligate suction feeder that preys on benthic invertebrates and fish. Its cranial morphology exhibits a suite of structural and functional modifications that facilitate this mode of prey capture. During suction‐feeding, subambient pressure is generated by the ventral expansion of the hyoid apparatus and the floor of its buccopharyngeal cavity. As in suction‐feeding bony fishes, the nurse shark exhibits expansive, compressive, and recovery kinematic phases that produce posterior‐directed water flow through the buccopharyngeal cavity. However, there is generally neither a preparatory phase nor cranial elevation. Suction is generated by the rapid depression of the buccopharyngeal floor by the coracoarcualis, coracohyoideus, and coracobranchiales muscles. Because the hyoid arch of G. cirratum is loosely connected to the mandible, contraction of the rectus cervicis muscle group can greatly depress the floor of the buccopharyngeal cavity below the depressed mandible, resulting in large volumetric expansion. Suction pressures in the nurse shark vary greatly, but include the greatest subambient pressures reported for an aquatic‐feeding vertebrate. Maximum suction pressure does not appear to be related to shark size, but is correlated with the rate of buccopharyngeal expansion. As in suction‐feeding bony fishes, suction in the nurse shark is only effective within approximately 3 cm in front of the mouth. The foraging behavior of this shark is most likely constrained to ambushing or stalking due to the exponential decay of effective suction in front of the mouth. Prey capture may be facilitated by foraging within reef confines and close to the substrate, which can enhance the effective suction distance, or by foraging at night when it can more closely approach prey. J. Morphol., 2008. © 2008 Wiley‐Liss, Inc.     

Linking cranial kinematics,buccal pressure,and suction feeding performance in largemouth bass

The rate and magnitude of buccal expansion are thought to determine the pattern of water flow and the change in buccal pressure during suction feeding. Feeding events that generate higher flow rates should induce stronger suction pressure and allow predators to draw prey from further away. We tested these expectations by measuring the effects of prey capture kinematics on suction pressure and the effects of the latter on the distance from which prey were drawn-termed suction distance. We simultaneously, but not synchronously, recorded 500-Hz video and buccal pressure from 199 sequences of four largemouth bass, Micropterus salmoides, feeding on goldfish. From the video, we quantified several kinematic variables associated with the head and jaws of the feeding bass that were hypothesized to affect pressure. In a multiple regression, kinematic data accounted for 79.7% of the variation among strikes in minimum pressure. Faster mouth opening and hyoid depression were correlated with lower pressures, a larger area under the pressure curve, and a faster rate of pressure reduction. In contrast, buccal pressure variables explained only 16.5% of the variation in suction distance, and no single pressure variable had a significant relationship with suction distance. Thus, although expected relationships between head kinematics and buccal pressure were confirmed, suction distance was only weakly related to buccal pressure. Three explanations are considered. First, bass may not attempt to maximize the distance from which prey are drawn. Second, the response of prey items to suction-induced flow depends on prey behavior and orientation and is, therefore, subject to considerable variation. Third, previous theoretical work indicates that water velocity decays exponentially with distance from the predator's mouth, indicating that variation among strikes in flow at the mouth opening is compressed away from the mouth. These findings are consistent with other recent data and suggest that suction distance is a poor metric of suction feeding performance.     

Reconstruction of cranial and hyobranchial muscles in the triassic temnospondyl Gerrothorax provides evidence for akinetic suction feeding

The cranial and hyobranchial muscles of the Triassic temnospondyl Gerrothorax have been reconstructed based on direct evidence (spatial limitations, ossified muscle insertion sites on skull, mandible, and hyobranchium) and on phylogenetic reasoning (with extant basal actinopterygians and caudates as bracketing taxa). The skeletal and soft‐anatomical data allow the reconstruction of the feeding strike of this bottom‐dwelling, aquatic temnospondyl. The orientation of the muscle scars on the postglenoid area of the mandible indicates that the depressor mandibulae was indeed used for lowering the mandible and not to raise the skull as supposed previously and implies that the skull including the mandible must have been lifted off the ground during prey capture. It can thus be assumed that Gerrothorax raised the head toward the prey with the jaws still closed. Analogous to the bracketing taxa, subsequent mouth opening was caused by action of the strong epaxial muscles (further elevation of the head) and the depressor mandibulae and rectus cervicis (lowering of the mandible). During mouth opening, the action of the rectus cervicis muscle also rotated the hyobranchial apparatus ventrally and caudally, thus expanding the buccal cavity and causing the inflow of water with the prey through the mouth opening. The strongly developed depressor mandibulae and rectus cervicis, and the well ossified, large quadrate‐articular joint suggest that this action occurred rapidly and that powerful suction was generated. Also, the jaw adductors were well developed and enabled a rapid mouth closure. In contrast to extant caudate larvae and most extant actinopterygians (teleosts), no cranial kinesis was possible in the Gerrothorax skull, and therefore suction feeding was not as elaborate as in these extant forms. This reconstruction may guide future studies of feeding in extinct aquatic tetrapods with ossified hyobranchial apparatus. J. Morphol., 2013. © 2012 Wiley Periodicals, Inc.     

Prey capture in the pike-perch,Stizostedion lucioperca (teleostei,percidae): A structural and functional analysis

Summary The pike-perch,Stizostedion lucioperca, uses both suction and grasping during feeding. Type, size, and position of prey and predator determine the movement of catching. This is concluded from simultaneous motion analysis, electromyography, and the record of pressures inside the buccopharyngeal cavity during feeding. The EMG incorporates 24 muscles of the head, including the branchial basket and the anterior body musculature. When the pike-perch begins to feed acceleration and expansion of the head parts determine the negative buccopharyngeal pressure and therefore the suction force applied to different preys. Not the head muscles, but the epaxial and hypaxial body musculatures provide the main force for the rapid expansion of the head through movements of the neurocranium, pectoral girdle, and hyoid arch. Despite the lack of a true neck, the pike-perch is able to move its neurocranium in all directions to aim the suction force. The experiments revealed that ventral and lateral movements aid in the ingestion of a big prey after it has been grasped with the teeth. The head muscles are active as regulators of the opening movements and in the closing movements. Variable overlaps of ab- and adductor activity show that their contraction patterns are interdependent. Variations in the recorded pressures can be related largely to a series of EMGs showing different starting moments of adductor contraction. In this progressive series two patterns were distinguished, and their accompanying movements were compared and related to the type of prey. According to the feeding behavior and morphology, the pike-perch is classified as a rapacious predator. Comparison with some other voracious fishes shows that besides the total length of the lower jaw and the dentigerous area, the construction and dentition of the upper jaws and the anterior suspensorial and neurocranial parts are also important features of this ecological type. However it appears that the fishes selected for this comparison use suction rather than the teeth as the main means of catching the smaller, but commonly eaten prey. The teeth prevent escape after capture by sucking and they increase the maximum prey size that can be caught. In this group, the head ofStizostedion appears to be comparatively well adapted to sucking with grasping adaptations.     

THE EVOLUTION OF TETRAPOD FEEDING BEHAVIOR: KINEMATIC HOMOLOGIES IN PREY TRANSPORT

One of the major features of the aquatic-to-terrestrial transition in vertebrate evolution was the change in the mechanism used to transport prey from the jaws to the throat. Primarily, vertebrates use hydraulic transport, but the transition to terrestrial life was accompanied by modifications of the hyobranchial apparatus that permit tongue-based transport. Despite an extensive data base on amniote feeding systems and mechanisms of intraoral prey transport, few data are available on the mechanism of prey transport in anamniote tetrapods. Transport cycles of four Ambystoma tigrinum (Amphibia) feeding on worms and crickets were filmed at 150 flames per second to produce quantitative profiles of the intraoral transport cycles for the two prey types. During the transport cycle the head and body remain stationary relative to the background: transport in Ambystoma tigrinum thus does not involve inertial movements of the head or body. Prey type had little effect on the kinematics of prey transport. The process of prey transport may be divided into four phases: preparatory, fast opening, closing, and recovery. The preparatory phase itself is divided into two parts: an extended segment that may include slight slow opening and a static phase prior to mouth opening where no change in gape occurs. The kinematic profile of transport in terrestrial salamanders is extremely similar to that used by fishes during hydraulic (aquatic) prey transport. We hypothesize that the distinct recovery and preparatory phases in the transport cycle of anamniote tetrapods are together homologous to the slow opening phases of the amniote cycle, and that during the evolution of terrestrial prey processing systems the primitive extended preparatory phase has become greatly compressed and incorporated into the amniote gape cycle.     

Functional morphology of the feeding system in the ruff — Gymnocephalus cernua (L. 1758) — (Teleostei,Percidae)

The ruff, Gymnocephalus cernua, is a European freshwater fish that feeds by sucking up small invertebrates from the bottom of ponds and slow flowing rivers. The feeding movements have been studied by simultaneous electromyography of seventeen muscles of the head and cinematographic techniques. A theoretical model of movements imposes the functional demands of suction upon an abstraction of the form of a teleost head. Three phases in the feeding act, a preparatory phase, a suction phase and a transport phase, could be correlated with the observed movements and EMGs. Differences between the predicted and the actual movement are discussed. Two different types of feeding occur. The direction, magnitude and duration of the suction forces during feeding are modified, according to the position of the prey. A mechanism preventing early mandibular depression allows sudden and strong suction. Retardation of the suspensorial abduction during the overall expansion of the buccal cavity is ascribed to kinetic interrelations with the hyoid arch. Protrusion of the upper jaws also permits an earlier closure of the mouth and directs the food-containing waterflow posteriorly. When the fish is feeding on sinking prey, protrusion occurs later in the sequence of movements than when it is feeding from the bottom. As the protruded jaws produce a downwardly pointed mouth this retardation aims the suction force.     

The Effects of Satiation on Strike Mode and Prey Capture Kinematics in the Largemouth bass, Micropterus salmoides

The feeding mechanism and kinematics of prey capture have been studied in many fishes. However, the effects of satiation on the strike mode and prey capture kinematics have never been directly measured. We analyze 12 kinematic variables to determine the effects of satiation on prey capture in five largemouth bass, Micropterus salmoides, by using high speed videography. We also present the first experimental test for modulatory capabilities in response to satiation, by using the ram-suction index. Significant changes in the kinematic variables of maximum lower jaw depression, maximum gape distance, maximum hyoid depression, time to maximum hyoid depression, and time from maximum hyoid depression to recovery were seen with the effects of satiation. Change in the kinematic variables imply a decrease in jaw opening velocity and the magnitude of suction velocity created during repetitive strikes by M. salmoides with increasing satiation. The bass primarily uses a ram strike mode, with some suction bites occasionally. Ram-suction index analyses suggests that M. salmoides does not modulate strike mode in response to satiation. However, the bass modulate prey capture kinematics without altering strike mode with the effects of satiation. Prey capture success decreases in each bass, as the probability of a successful prey capture event becomes lower, with increasing satiation. These findings demonstrate that satiation can have major effects on prey capture kinematics and future studies of feeding kinematics should account for satiation in their analyses.     

Modulation of prey capture kinematics in the cheeklined wrasse Oxycheilinus digrammus (Teleostei: Labridae)

The ability to modulate prey capture behaviors is of interest to organismal biologists as it suggests that predators can perceive features of the prey and select suitable behaviors from an available repertoire to successfully capture the item. Thus, behavior may be as important a trait as morphology in determining an organism's diet. Using high-speed video, we measured prey capture kinematics in three cheeklined wrasse, Oxycheilinus digrammus. We studied the effects of three experimental prey treatments: live fish, dead prawn suspended in the water column, and dead prawn pieces anchored to the substrate in a clip. Live prey elicited significantly more rapid strikes than dead prey suspended in the water column, and the head of the predator was expanded to significantly larger maxima. These changes in prey capture kinematics suggest the generation of more inertial suction. With greater expansion of the head, more water can be accelerated into the buccal cavity. The attached prey treatment elicited strikes as rapid as those on live prey. We suggest that the kinematics of rapid strikes on attached prey are indicative of attempts to use suction to detach the prey item. More rapid expansion of the buccal or mouth cavity should lead to higher velocities of water entering the mouth and therefore to enhanced suction. Further modulation in response to the attached prey item, such as clipping or wrenching behaviors, was not observed. J. Exp. Zool. 290:88-100, 2001.     

The effects of intraspecific competition on the prey capture behavior and kinematics of the bluegill sunfish, Lepomis macrochirus

Competition has broad effects on fish and specifically the effects of competition on the prey capture kinematics and behavior are important for the assessment of future prey capture studies in bony fishes. Prey capture kinematics and behavior in bony fishes have been shown to be affected by temperature and satiation. The densities at which bony fish are kept have also been shown to affect their growth, behavior, prey selection, feeding and physiology. We investigated how density induced intraspecific competition for food affects the prey capture kinematics of juvenile bluegill sunfish, Lepomis macrochirus. High speed video was utilized to film five bold individuals feeding at three different densities representing different levels of intraspecific competition. We hypothesized that: (1) the feeding kinematics will be faster at higher levels of competition compared to lower levels of competition, and (2) bluegill should shift from more suction-based feeding towards more ram-based feeding with increasing levels of competition in order to outcompete conspecifics for a prey item. We found that, with increased intraspecific competition, prey capture became faster, involving more rapid jaw opening and therefore greater inertial suction, shorter mouth closing times, and shorter gape cycles. Furthermore, the attack velocity of the fish increased with increasing competition, however a shift towards primarily ram based feeding was not confirmed. Our study demonstrates that prey capture kinematics are affected by the presence of conspecifics and future studies need to consider the effects of competition on prey capture kinematics.     

Comparisons of aquatic versus terrestrial predatory strikes in the pitviper, Agkistrodon piscivorus

Recent studies comparing terrestrial versus aquatic locomotion in animals have shown that changes in kinematics, muscular activation patterns, and performance across media are often dramatic. Surprisingly, however, despite the importance of feeding to the survival of most animals, few studies have compared differences in feeding behaviour between media. The present study compares prey-capture behaviour, strike success, strike velocity and acceleration, and prey-capture kinematics in a semi-aquatic pitviper (Agkistrodon piscivorus) when capturing both terrestrial (mice) and aquatic (fish) prey in a standardized laboratory setting. Strike velocity and acceleration did not differ significantly between media, but instead were positively correlated with initial prey distance. By contrast, the kinematics of terrestrial and aquatic strikes differed significantly in several aspects: max gape angle during the retraction phase, angular velocity of mouth closing during the strike, and the initial head angle before the strike. Terrestrial strikes were associated with higher gape angles during the retraction phase, higher angular velocities of mouth closure, and a more inclined head angle at the onset of the strike. Finally, strike success differed significantly between strike types, with terrestrial strikes being considerably more successful than aquatic strikes. Strike success likely differed due to the relatively slow mouth-closing velocity of aquatic strikes.     

Feeding kinematics and morphology of the alligator gar (Atractosteus spatula,Lacépède, 1803)

Living gars are a small clade of seven species that occupy an important position on the actinopterygian phylogenetic tree as members of Holostei, sister-group to teleosts, and exhibit many plesiomorphic traits used to interpret and reconstruct early osteichthyan feeding mechanisms. Previous studies of gar feeding kinematics have focused on the ram-based, lateral-snapping mode of prey capture found in the narrow-snouted Lepisosteus genus, whereas this study focuses on a member of the broad-snouted Atractosteus sister-genus, the alligator gar (Atractosteus spatula, Lacépède, 1803). High-speed videography reveals that the feeding system of alligator gars is capable of rapid expansion from anterior to posterior, timed in a way to generate suction, counteract the effects of a bow-wave during ram-feeding, and direct a unidirectional flow of water through the feeding system. Reconstructed contrast-enhanced μCT-based cranial anatomy and three-dimensional modeling of linkage mechanics show that a lateral-sliding palatoquadrate, flexible intrasuspensorial joint, pivoting interhyal, and retractable pectoral girdle increase the range of motion and expansive capabilities of the alligator gar feeding mechanism. Reconstructions of muscular anatomy, inferences from in vivo kinematics, and in situ manipulations show that input from the hyoid constrictors and hypaxials play an important role in decoupling and modulating the dual roles of the sternohyoideus during feeding: hyoid retraction (jaw opening) and hyoid rotation (pharyngeal expansion). The alligator gar possesses an intricate feeding mechanism, capable of precise control with plesiomorphic muscles that represent one of the many ways the ancestral osteichthyan feeding mechanism has been modified for prey capture.     

Scaling of suction feeding performance in the catfish Clarias gariepinus

Ontogenetic changes in the absolute dimensions of the cranial system together with changes in kinematics during prey capture can cause differences in the spatiotemporal patterns of water flow generated during suction feeding. Because the velocity of this water flow determines the force that pulls prey toward and into the mouth cavity, this can affect suction feeding performance. In this study, size-related changes in the suction-induced flow patterns are determined. To do so, a mathematical suction model is applied to video recordings of prey capturing Clarias gariepinus ranging in total length from 111 to 923 mm. Although large C. gariepinus could be expected to have increasing peak velocities of water flow compared with small individuals, the results from the hydrodynamic model show that this is not the case. Yet, when C. gariepinus becomes larger, the expansive phase is prolonged, resulting in a longer sustained flow. This flow also reaches farther in front of the mouth almost proportionally with head size. Forward dynamical simulations with spherical prey that are subjected to the calculated water flows indicate that the absolute distance from which a given prey can be sucked into the mouth as well as the maximal prey diameter increase substantially with increasing head size. Consequently, the range of potential prey that can be captured through suction feeding will become broader during growth of C. gariepinus. This appears to be reflected in the natural diet of this species, where both the size and the number of evasive prey increase with increasing predator size.     

Intrinsic Mechanics and Control of Fast Cranio-Cervical Movements in Aquatic Feeding Turtles

Aquatic feeding strikes on agile prey in snake-necked turtlesinvolve fast neck extension, bucco-pharyngo-oesophageal expansion,and head retraction. The ultimate, rectilinear accelerationof the head towards the prey requires complex vertebral rotations,that vary widely from strike to strike. This poses complex motorcontrol issues for the numerous intrinsic neck-muscles, whichare the sole neck extensors. Mathematical modelling revealsthat extensor activity might be superfluous for this phase ofthe strike. The ultimate acceleration of the head at the endof the strike always coincides with forceful oropharyngeal expansion.The momentum of the induced flow of water is sufficient to pullthe head (and the neck) straight towards the prey. This buccalexpansion proceeds identically to that observed in primary aquaticfeeders: a rostro-caudal expansion sequence characterized byan optimal timing of the functional components supporting theexpansion wave. Yet distinct structural solutions, both at theskeletal, and muscular level, are involved. This points towardsprominent hydrodynamic constraints. Head and neck are retractedby extrinsic neck muscles. Given the high number of degreesof freedom, this musculo-skeletal system is obviously under-determined,which compromises control. We propose that erroneous foldingof the neck (i.e., diverging from the highly persistent retractedconfiguration) might be avoided through the presence of a subtleclick system at the level of the joint between cervical vertebrae5 and 6.     

Prey capture and processing behaviors vary with prey size and shape in Australian and subantarctic fur seals

When hunting at sea, pinnipeds should adapt their foraging behaviors to suit the prey they are targeting. We performed captive feeding trials with two species of otariid seal, Australian fur seals (Arctocephalus pusillus doriferus) and subantarctic fur seals (Arctocephalus tropicalis). This allowed us to record detailed observations of how their foraging behaviors vary when presented with prey items that cover the full range of body shapes and sizes encountered in the wild. Small prey were captured using suction alone, while larger prey items were caught in the teeth using raptorial biting. Small fish and long skinny prey items could then be swallowed whole or processed by shaking, while all prey items with body depths greater than 7.5 cm were processed by shaking at the water's surface. This matched opportunistic observations of feeding in wild Australian fur seals. Use of “shake feeding” as the main prey processing tactic also matches predictions that this method would be one of the only tactics available to aquatic tetrapods that are unable to secure prey using their forelimbs.     

Metamorphosis and evolution of feeding behaviour in salamanders of the family Plethodontidae

Plethodontid salamanders capture prey with enhanced tongue protraction relative to other salamander taxa, yet metamorphosing plethodontids are hypothesized to be constrained relative to direct-developing plethodontids in their degree of tongue evolution (protraction length and velocity) by the presence of a larval stage in development. In this biphasic life history the hyobranchial apparatus serves the conflicting functions of larval suction feeding and adult tongue protraction. The deletion of the larval stage removes one of the conflicting functions and has thus permitted direct-developing plethodontids to circumvent this constraint and evolve extremely long tongues, which in some species can be projected to 80% of body length. To evaluate this constraint hypothesis and explore taxonomic diversity of feeding behaviours, we studied feeding in larvae, adults and metamorphosing individuals of seven species of metamorphosing plethodontids from the basal taxa Desmognathinae and Hemidactyliini using direct observations, high-speed videography and kinematic analysis. We found that larval plethodontids suction feed, but feeding is suspended entirely during metamorphosis, and aquatic adults do not suction feed. Adults have exapted the terrestrial modes of tongue and jaw prehension for aquatic prey capture. These findings substantiate the premise that suction feeding and tongue protraction are conflicting functions, and thus our results support the constraint hypothesis. Plethodontid adults have evolved their extreme tongue protraction ability at the expense of adult suction feeding. The rapid metamorphosis that characterizes plethodontids may be an adaptation that minimizes the non-feeding period imposed by the evolution of derived tongue protraction in adults. © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society , 2002, 134 , 375–400.     

Prey capture by Luciocephalus pulcher: implications for models of jaw protrusion in teleost fishes

Synopsis Luciocephalus pulcher possesses one of the most protrusible jaws known among teleosts, the premaxillae extending anteriorly a distance of 33% of the head length during feeding. Jaw bone movement during feeding proceeds according to a stereotypical pattern and resembles that of other teleosts except for extreme cranial elevation and premaxillary protrusion. Anatomical specializations associated with cranial elevation include: a highly modified first vertebra with a separate neural spine, articular fossae on the posterior aspect, greatly enlarged zygapophyses on the second vertebra with complex articular condyles, and highly pinnate multi-layered epaxial musculature with multiple tendinous insertions on the skull. Luciocephalus, despite the extreme jaw protrusion, does not use suction during prey capture: rather, the prey is captured by a rapid lunge (peak velocity of about 150 cm per sec) and is surrounded by the open mouth. Previous hypotheses of the function of upper jaw protrusion are reviewed in relation to jaw movements inLuciocephalus. Protrusion is not obligatorily linked with suction feeding; behavioral aspects of the feeding process limit the possible range of biological roles of a given morphological specialization, and make prediction of role from structure risky.     

The ontogeny of functional design: metamorphosis of feeding behaviour in the tiger salamander (Ambystoma tigrinum)

The ontogeny of feeding behaviour was studied quantitatively in the tiger salamander, Ambystoma tigrinum , to elucidate the relative importance of morphological and environmental changes on form and function. High-speed films of prey capture provided data for a frame-by-frame analysis of seven kinematic parameters of feeding behaviour. By comparing underwater feeding of larval and metamorphosed individuals, the effect of morphological changes occurring at metamorphosis on the feeding kinematic pattern was determined. By comparing metamorphosed animals feeding in the water and on land, changes in feeding kinematics associated with the environmental transition (and thus the differing physical properties of water and air) were determined. Both univariate and multivariate analyses failed to demonstrate any differences between larval and metamorphosed aquatic feedings for seven kinematic variables. However, when individuals feed on land, a radical shift in hyoid kinematics was observed. In addition, multivariate analysis showed that terrestrial feedings differed from aquatic feedings in having longer duration head movements. The lack of a kinematic difference between larval and metamorphosed individuals feeding in the water indicates that the morphological changes occurring at metamorphosis do not impose any obligatory kinematic consequences. Rather, metamorphosed Ambystoma tigrinum acquire the ability to modulate their kinematic pattern depending on the environment.     

Aquatic Versus Terrestrial Feeding Modes: Possible Impacts on the Trophic Ecology of Vertebrates

On the basis of experimental work, a clear dichotomy in designof the feeding mechanism between aquatic and terrestrial vertebrateshas been found. The aquatic medium combined with suction andthe hydrodynamic tongue offer an unparalleled array of preycapture opportunities for aquatic vertebrates. In the terrestrialfeeding model, prey capture and prey processing require a precisefunctional design to match the nature of the prey. It is proposedthat this dichotomy in basic design would result in fundamentaldifferences in the aquatic and terrestrial paradigms in vertebrateecology. Four hypotheses and their respective supporting evidenceare presented: (1) Increased opportunism, more extensive preyswitching and food overlap are prevalent in aquatic systems;(2) Convergent evolution which is so pervasive in terrestrialfeeding systems is uncommon among aquatic vertebrate feedingsystems; (3) Competition is relaxed and character displacementis absent in aquatic vertebrate feeding systems; and (4) Patternsof diversity in aquatic vertebrate feeding systems defy theterrestrial paradigm.     

Morphological convergence as a consequence of extreme functional demands: examples from the feeding system of natricine snakes

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.     

Aquatic feeding in a terrestrial turtle: a functional-morphological study of the feeding apparatus in the Indochinese box turtle Cuora galbinifrons (Testudines, Geoemydidae)

The Indochinese box turtle Cuora galbinifrons is regarded as a purely terrestrial species, but our results demonstrate that it can feed both on land and in water. The inverse relationship between the relative development of the hyoid apparatus and the tongue found in the most investigated chelonians is not valid in the Indochinese box turtle. Our morphological analysis of the feeding apparatus reveals that the palate shape and the design of the tongue are consistent with terrestrial feeders, but the construction of the hyoid complex is more characteristic of aquatic feeders. Previous studies have demonstrated that tongue enlargement negatively impacts the capacity of the turtles to suction feed. The present study focuses on the aquatic intraoral prey transport kinematic patterns. Our analysis is based on high-speed films with 250 fr/s and high-speed cineradiography with 50 fr/s. The aquatic intraoral food transport mechanisms differ depending on prey size: small items are transported predominantly by “inertial suction”, whereas larger items are moved by the tongue—normally a clear terrestrial strategy. As the genus Cuora is ancestrally aquatic, the use of lingual food transport in the aquatic environment is presumably an aberrant modus typical only for the most terrestrial among the Asian box turtles.