Analysis of prey capture and food transport kinematics in two Asian box turtles,Cuora amboinensis and Cuora flavomarginata (Chelonia,Geoemydidae), with emphasis on terrestrial feeding patterns

This study examines the kinematics and morphology of the feeding apparatus of two geoemydid chelonians, the Malayan (Amboina) box turtle (Cuora amboinensis) and the yellow-margined box turtle (Cuora flavomarginata). Both species are able to feed on land as well as in water. Feeding patterns were analysed by high-speed cinematography. The main focus of the present study is on the terrestrial feeding strategies in both Asian box turtles, because feeding on land has probably evolved de novo within the ancestrally aquatic genus Cuora. During terrestrial feeding (analysed for both species), the initial food prehension is always done by the jaws, whereas intraoral food transport and pharyngeal packing actions are tongue-based. The food uptake modes in Cuoras differ considerably from those described for purely terrestrial turtles. Lingual food prehension is typical of all tortoises (Testudinidae), but is absent in C. amboinensis and C. flavomarginata. A previous study on Terrapene carolina shows that this emydid turtle protrudes the tongue during ingestion on land, but that the first contact with the food item occurs by the jaws. Both Asian box turtles investigated here have highly movable, fleshy tongues; nonetheless, the hyolingual complex remains permanently retracted during initial prey capture. In aquatic feeding (analysed for C. amboinensis only), the prey is captured by a fast forward strike of the head (ram feeding). As opposed to ingestion on land, in the underwater grasp the hyoid protracts prior to jaw opening. The head morphology of the investigated species differs. In contrast to the Malayan box turtle, C. flavomarginata exhibits a more complexly structured dorsal lingual epithelium, a considerable palatal vault, weaker jaw adductor muscles and a simplified trochlear complex. The differences in the hyolingual morphology reflect the kinematic patterns of the terrestrial feeding transport.     

Modulation of prey capture kinematics and the role of lingual sensory feedback in the lizard Pogona vitticeps

Most organisms feed on a variety of prey that may differ dramatically in their physical and behavioural characteristics (e.g. mobility, mass, texture, etc.). Thus the ability to modulate prey capture behaviour in accordance with the characteristics of the food appears crucial. In animals that use rapid tongue movements to capture prey (frogs and chameleons), the coordination of jaws and tongue is based on visual cues gathered prior to the prey capture event. However, most iguanian lizards have much slower tongue-based prey capture systems suggesting that sensory feedback from the tongue may play an important role in coordinating jaw and tongue movements. We investigated the modulation of prey capture kinematics in the agamid lizard Pogona vitticeps when feeding on a range of food items differing in their physical characteristics. As the lizard is a dietary generalist, we expected it to be able to modulate its prey capture kinematics as a function of the (mechanical) demands imposed by the prey. Additionally, we investigated the role of lingual sensory feedback by transecting the trigeminal sensory afferents. Our findings demonstrated that P. vitticeps modulates its prey capture kinematics according to specific prey properties (e.g. size). In addition, transection of the trigeminal sensory nerves had a strong effect on prey capture kinematics. However, significant prey type effects and prey type by transection effects suggest that other sources of sensory information are also used to modulate the prey capture kinematics in P. vitticeps.     

Morphology and function of the tongue and hyoid apparatus in Varanus (Varanidae, Lacertilia)

The morphology and function of the tongue and hyoid apparatus in Varanus were examined by anatomical and experimental techniques. Morphological features unique to Varanus include a highly protrusible tongue that has lost a roughened dorsal surface, an exceptionally strong and mobile hyobranchial apparatus, a well-defined joint between the ceratohyal and anterior process, and a series of distinct muscles inserting at the anterior hyobranchial region. Varanus is also unusual among lizards in a number of feeding behaviors; it ingests prey entirely by inertial feeding, as the tongue does not participate in food transport. Further specializations include an increased reliance on hyobranchial movements in drinking and pharyngeal packing and compression. The long, narrow tongue is most likely related to the mechanics of tongue protrusion; the increased amount, strength, and complexity of hyobranchial movement is related to the fact that the hyobranchium in Varanus replaces the tongue in many functions. Previous hypotheses for the origin of these adaptations are discussed, and the difficulties of attributing these specializations to any specific scenario of adaptation or constraint are emphasized.     

Kinematics of the jaw and hyolingual apparatus during feeding in Caiman crocodilus

Movements of the neck, jaws, and hyolingual apparatus during inertial feeding in Caiman crocodilus were studied by cineradiography. Analysis reveals two kinds of cycles: inertial bites (reposition, kill/crush, and transport) and swallowing cycles. They differ in their gape profile and in displacement of the neck, cranium, and hyolingual apparatus. Inertial bites are initiated by an elevation of the neck and cranium; the head is then retracted backward, the prey simultaneously being lifted by the hyolingual apparatus. Next the lower jaw is depressed, and the prey is rapidly pushed further upward by the hyolingual apparatus. Thereafter fast mouth-closure occurs with the neck and cranium being abruptly depressed, the lower jaw elevated, and the hyolingual apparatus rapidly retracted ventrally. Depression of the neck and cranium thrusts the head forward and impacts the backward moving prey more posteriorly in the oral cavity. Swallowing cycles initially involve movement of the hyoid in front of the prey followed by rapid posteroventrad retraction of the hyoid, forcing the prey into the esophagus during opening and closing of the mouth. After mouth-closure, the hyoid apparatus is again protracted. Jaws, neck, tongue, and hyoid apparatus play an active role during intertial feeding sequences. At the beginning of a feeding sequence, the hyolingual apparatus mainly moves dorsoventrally, whereas toward the end of a sequence anteroposterior displacements of the hyoid are prominent. © 1992 Wiley-Liss, Inc.     

Modulatory complexity of the feeding repertoire in scincid lizards

The kinematics of jaws and tongue, and jaw muscle activity patterns were investigated in the omnivorous lizard Tiliqua rugosa, and the herbivorous Corucia zebrata (Scincidae) during feeding. Small metal markers were inserted into different parts of the skull, the jaws, and the tongue. Video and cineradiographic images were digitized and displacements of the head, jaws, and tongue were quantified. Additionally, muscle activity patterns were recorded, digitized and several variables were determined quantitatively. The effect of food type on the jaw and hyolingual movement patterns and the jaw muscle activity patterns was investigated for both species. The kinematic data indicate that distinct aspects of gape and tongue cycles are modulated in response to the food characteristics. Similarly, in both species, muscle activity patterns are altered in response to the type of food eaten. A comparison of kinematic and electromyographic patterns during intraoral transport cycles for both species shows that these can be related to food characteristics such as toughness and mobility. Differences between both species in the response to changes in food characteristics are minor. Clearly both species are able to fine tune the activation of the jaw muscles, resulting in the appropriate movement patterns for the type of food eaten. Accepted: 30 January 1999     

The scaling of tongue projection in the veiled chameleon,Chamaeleo calyptratus

Within a year of hatching, chameleons can grow by up to two orders of magnitude in body mass. Rapid growth of the feeding mechanism means that bones, muscles, and movements change as chameleons grow while needing to maintain function. A previous morphological study showed that the musculoskeletal components of the feeding apparatus grow with negative allometry relative to snout–vent length (SVL) in chameleons. Here, we investigate the scaling of prey capture kinematics and muscle physiological cross-sectional area in the veiled chameleon, Chamaeleo calyptratus. The chameleons used in this study varied in size from approximately 3 to 18 cm SVL (1–200 g). Feeding sequences of 12 chameleons of different sizes were filmed and the timing of movements and the displacements and velocities of the jaws, tongue, and the hyolingual apparatus were quantified. Our results show that most muscle cross-sectional areas as well as tongue and hyoid mass scaled with isometry relative to mandible length, yet with negative allometry relative to SVL. Durations of movement also scaled with negative allometry relative to SVL and mandible length. Distances and angles generally scaled as predicted under geometric similarity (slopes of 1 and 0, respectively), while velocities generally scaled with slopes greater than 0 relative to SVL and mandible length. These data indicate that the velocity of jaw and tongue movements is generally greater in adults compared to juveniles. The discrepancy between the scaling of cross-sectional areas versus movements suggests changes in the energy storage and release mechanisms implicated in tongue projection.     

Kinematics of prey capture and chewing in the lizard Agama agama (squamata: Agamidae)

High speed video recordings (200 fields per second) of prey capture and food processing in Agama agama permit the identification of strikes, chews and transport movements. Ten variables from strike movements and seven variables from chewing sequences are digitized; transport movements are inspected only. Univariate and multivariate statistical analyses disclose significant interindividual differences for three variables (maximum gape distance, maximum head angle, and maximum throat distance); but neither these nor principal components analysis show differences between strikes and chews for any of the gape change and hyoid depression variables. However, strikes and chews obviously differ in tongue protrusion and body movements. Chewing may be divided into four stages, comparable to those of transport cycles of other lizards and the generalized tetrapod model. Transport differs from chewing by having a shorter power stroke and relatively more cranial and less jaw movement. The kinematics of feeding in Agama agama are compared with those of other lizards studied previously.     

Separating the effects of prey size and speed on the kinematics of prey capture in the omnivorous lizard Gerrhosaurus major

Feeding behavior is known to be modulated as prey properties change. During prey capture, external prey properties, including size and mobility, are likely some of the most important components in predator–prey interactions. Whereas prey size has been demonstrated to elicit modulation of jaw movements during capture, how prey speed affects the approach and capture of prey remains unknown. We quantified the kinematics associated with movements of both the feeding and locomotor systems during prey capture in a lizard, Gerrhosaurus major, while facing prey differing in size and mobility (newborn mice, grasshoppers, and mealworms). Our data show that the feeding and locomotor systems were recruited differently in response to changes in the size or speed of the prey. The timing of jaw movements and of the positioning of the head are affected by changes in prey size—and speed, to a lesser extent. Changes in prey speed resulted in concomitant changes in the speed of strike and an early and greater elevation of the neck. External prey properties, and prey mobility in particular, are relevant in predator–prey interactions and elicit specific responses in different functional systems.     

Piscivorous cyprinid fish modulates suction feeding kinematics to capture elusive prey

Previous studies have shown that evasive prey generally elicit a different kinematical pattern of prey capture from suction feeding fish compared to non-evasive types of prey. However, no evidence exists that predatory fish can modulate their prey capture kinematics in response to whether or not an elusive prey performs an escape response. Here, we analyse prey capture kinematics of a specialist piscivore (asp, Aspius aspius) during feeding on untethered, live goldfish, which regularly displayed escape attempts when attacked by the asp. Significant modulation occurred in function of the escape attempts of prey: mouth opening was prolonged and increased in magnitude, and one individual also showed an increased hyoid depression when feeding on prey trying to escape. As the orientation of the prey with respect to the predator prior to the start of mouth opening was related to the probability of observing an escape attempt, asp could theoretically perform this type of modulation by a priori choosing a pre-programmed motor pattern. However, since contact between the prey and the asp's mouth appeared to be a factor improving the timing of mouth closing, this fine-tuning of prey capture kinematics is more likely to be caused by reflexive neural feedback control.     

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.     

The use of the tongue and hyoid apparatus during feeding in lizards (Ctenosaura similis and Tupinambis nigropunctatus)

The use of the tongue and hyoid is examined in cineradiographic and electromyographic investigations of feeding in two species of lizards, Ctenosaura similis (Iguanidae) and Tupinambis nigropunctatus (Teiidae). In both animals food is transported through the oral cavity by regular cycles of the tongue. Tongue movements correlate with jaw and hyoid movement. Similarities between the two animals in the use of the tongue in food transport, lapping, pharyngeal packing, and pharyngeal emptying are detailed. Mechanisms of tongue protrusion are examined and it is shown that the tongue in Tupinambis is relatively more protrusible than in Ctenosaura. This difference is complementary with data on the greater reliance of Tupinambis on the tongue as a sensory organ. Tupinambis further differs from Ctenosaura in possessing a greater mobility of the hyoid. In many features of tongue use in food transport, lizards resemble mammals, supporting postulations of a basic pattern of intra-oral food transport. However, whether this pattern can be attributed to convergence or a common, primitive neural pattern of control cannot be distinguished. Lizards lack two major characteristics of mammalian food transport: regular masticatory cycles and an internal swallowing mechanism.     

Morphology and histochemistry of the hyolingual apparatus in chameleons

We reexamined the morphological and functional properties of the hyoid, the tongue pad, and hyolingual musculature in chameleons. Dissections and histological sections indicated the presence of five distinctly individualized pairs of intrinsic tongue muscles. An analysis of the histochemical properties of the system revealed only two fiber types in the hyolingual muscles: fast glycolytic and fast oxidative glycolytic fibers. In accordance with this observation, motor-endplate staining showed that all endplates are of the en-plaque type. All muscles show relatively short fibers and large numbers of motor endplates, indicating a large potential for fine muscular control. The connective tissue sheet surrounding the entoglossal process contains elastin fibers at its periphery, allowing for elastic recoil of the hyolingual system after prey capture. The connective tissue sheets surrounding the m. accelerator and m. hyoglossus were examined under polarized light. The collagen fibers in the accelerator epimysium are configured in a crossed helical array that will facilitate limited muscle elongation. The microstructure of the tongue pad as revealed by SEM showed decreased adhesive properties, indicating a change in the prey prehension mechanics in chameleons compared to agamid or iguanid lizards. These findings provide the basis for further experimental analysis of the hyolingual system.     

A forceful upper jaw facilitates picking-based prey capture: biomechanics of feeding in a butterflyfish,Chaetodon trichrous

Biomechanical models of feeding mechanisms elucidate how animals capture food in the wild, which, in turn, expands our understanding of their fundamental trophic niche. However, little attention has been given to modeling the protrusible upper jaw apparatus that characterizes many teleost species. We expanded existing biomechanical models to include upper jaw forces using a generalist butterflyfish, Chaetodon trichrous (Chaetodontidae) that produces substantial upper jaw protrusion when feeding on midwater and benthic prey. Laboratory feeding trials for C. trichrous were recorded using high-speed digital imaging; from these sequences we quantified feeding performance parameters to use as inputs for the biomechanical model. According to the model outputs, the upper jaw makes a substantial contribution to the overall forces produced during mouth closing in C. trichrous. Thus, biomechanical models that only consider lower jaw closing forces will underestimate total bite force for this and likely other teleost species. We also quantified and subsequently modeled feeding events for C. trichrous consuming prey from the water column versus picking attached prey from the substrate to investigate whether there is a functional trade-off between prey capture modes. We found that individuals of C. trichrous alter their feeding behavior when consuming different prey types by changing the timing and magnitude of upper and lower jaw movements and that this behavioral modification will affect the forces produced by the jaws during prey capture by dynamically altering the lever mechanics of the jaws. In fact, the slower, lower magnitude movements produced during picking-based prey capture should produce a more forceful bite, which will facilitate feeding on benthic attached prey items, such as corals. Similarities between butterflyfishes and other teleost lineages that also employ picking-based prey capture suggest that a suite of key behavioral and morphological innovations enhances feeding success for benthic attached prey items.     

Comparative anatomy and evolutionary history of suction feeding in cetaceans

Some odontocetes possess unique features of the hyolingual apparatus that are involved in suction feeding. The hyoid bone and associated musculature generates rapid, piston‐like retraction, and depression of the hyoid and tongue. “Capture” suction feeders (e.g., Globicephala) use suction for capturing and swallowing prey. “Combination” feeders (i.e., Lagenorhynchus) use both raptorial feeding (to capture prey) and suction (to ingest prey). In “capture” suction feeders, features of the hyoid and skull have been attributed to creating suction (i.e., large surface area and mandibular bluntness). In addition to odontocetes, a mysticete, the gray whale (Eschrichtius robustus), is considered a benthic suction feeder. However, anatomical studies of purported suction‐feeding structures of the gray whale are lacking. In addition, few studies have utilized evolutionary approaches to understand the history of suction feeding in cetaceans. This study incorporates quantitative and qualitative hyoid and cranial data from 35 extant and 14 extinct cetacean species into a multivariate principal component analysis and comparative phylogenetic analyses. Conclusions from these analyses are that some commonly attributed features (i.e., ventral throat grooves and mandibular bluntness) and one principal component are significantly correlated with suction feeding. Finally, ancestral state reconstructions indicate that suction feeding likely evolved once, early in cetacean evolutionary history.     

Morphology and function of the feeding apparatus of the lungfish, Lepidosiren paradoxa (Dipnoi)

The feeding mechanism of the South American lungfish, Lepidosiren paradoxa retains many primitive teleostome characteristics. In particular, the process of initial prey capture shares four salient functional features with other primitive vertebrates: 1) prey capture by suction feeding, 2) cranial elevation at the cranio-vertebral joint during the mouth opening phase of the strike, 3) the hyoid apparatus plays a major role in mediating expansion of the oral cavity and is one biomechanical pathway involved in depressing the mandible, and 4) peak hyoid excursion occurs after maximum gape is achieved. Lepidosiren also possesses four key morphological and functional specializations of the feeding mechanism: 1) tooth plates, 2) an enlarged cranial rib serving as a site for the origin of muscles depressing the hyoid apparatus, 3) a depressor mandibulae muscle, apparently not homologous to that of amphibians, and 4) a complex sequence of manipulation and chewing of prey in the oral cavity prior to swallowing. The depressor mandibulae is always active during mouth opening, in contrast to some previous suggestions. Chewing cycles include alternating adduction and transport phases. Between each adduction, food may be transported in or out of the buccal cavity to position it between the tooth plates. The depressor mandibulae muscle is active in a double-burst pattern during chewing, with the larger second burst serving to open the mouth during prey transport. Swallowing is characterized by prolonged activity in the hyoid constrictor musculature and the geniothoracicus. Lepidosiren uses hydraulic transport achieved by movements of the hyoid apparatus to position prey within the oral cavity. This function is analogous to that of the tongue in many tetrapods.     

The Craniolingual Morphology of Waterfowl (Aves,Anseriformes) and Its Relationship with Feeding Mode Revealed Through Contrast-Enhanced X-Ray Computed Tomography and 2D Morphometrics

Within Anseriformes, waterfowl (ducks, geese, and swans) exhibit three specialized feeding modes that are distinctive among Aves: filter-feeding with fine and dense keratinous lamellae on a flat, mediolaterally expanded bill; cropping or grazing vegetation with large and robust lamellae with a dorsoventrally expanded bill; and sharp lamellae associated with a narrow bill used in acquiring mixed invertebrates and fish underwater mainly by grasping. Here we assess morphometric variation in cranial and hyolingual structures as well as hyolingual myology in a diverse sample of Anatidae to explore the relationship of tongue variation and feeding mode. Phylogenetically informed principal component analysis (phyl.PCA) of cranial-lingual measurements for 67 extant and two extinct anatids recovers grazers and filter-feeding taxa in largely distinct areas of morphospace, while underwater graspers and other mixed feeders show less distinct clustering. The relationship between morphometric differences in skeletal features and muscular variation was further explored through a reassessment of hyolingual musculature enabled by contrast-enhanced X-ray computed tomography (CT) imagery acquired from three exemplar species (Branta canadensis, Chen caerulescens, and Aythya americana) with distinctive ecologies and morphologies of the bony hyoid. Data for these duck and geese exemplars reveal further significant, and previously unstudied, morphological differences between filter-feeding and grazing species. Grazers have a larger hyolingual apparatus with highly-developed extrinsic hyoid muscles; while filter-feeding species are characterized by relatively more diminutive extrinsic hyoid muscles and larger intrinsic hyoid muscles. The feeding modes of two extinct taxa (i.e., Presbyornis and Thambetochen) were also estimated from morphometric data. The results indicate a derived terrestrial browsing or grazing ecology for Thambetochen but do not unequivocally support a specialized filter-feeding ecology for Presbyornis, which is recovered with mixed feeders including swans. The combination of detailed, CT-mediated acquisition of fine muscular anatomy with morphometric approaches shows promise for illuminating form–function relationships in extant taxa more generally.     

The functional morphology of lingual prey capture in a scincid lizard,Tiliqua scincoides (Reptilia: Squamata)

We investigated the functional morphology of lingual prey capture in the blue‐tongued skink, Tiliqua scincoides, a lingual‐feeding lizard nested deep within the family Scincidae, which is presumed to be dominated by jaw‐feeding. We used kinematic analysis of high‐speed video to characterize jaw and tongue movements during prey capture. Phylogenetically informed principal components analysis of tongue morphology showed that, compared to jaw‐feeding scincids and lacertids, T. scincoides and another tongue‐feeding scincid, Corucia zebrata, are distinct in ways suggesting an enhanced ability for hydrostatic shape change. Lingual feeding kinematics show substantial quantitative and qualitative variation among T. scincoides individuals. High‐speed video analysis showed that T. scincoides uses significant hydrostatic elongation and deformation during protrusion, tongue‐prey contact, and retraction. A key feature of lingual prey capture in T. scincoides is extensive hydrostatic deformation to increase the area of tongue‐prey contact, presumably to maximize wet adhesion of the prey item. Adhesion is mechanically reinforced during tongue retraction through formation of a distinctive “saddle” in the foretongue that supports the prey item, reducing the risk of prey loss during retraction.     

Horned lizards (Phrynosoma) incapacitate dangerous ant prey with mucus

Horned lizards (Iguanidae, Phrynosomatinae, Phrynosoma) are morphologically specialized reptiles characterized by squat, tank-like bodies, short limbs, blunt snouts, spines and cranial horns, among other traits. They are unusual among lizards in the degree to which they specialize on a diet of ants, but exceptional in the number of pugnacious, highly venomous, stinging ants they consume, especially harvester ants (genus Pogonomyrmex). Like other iguanian lizards, they capture insect prey on the tongue, but unlike other lizards, they neither bite nor chew dangerous prey before swallowing. Instead, they employ a unique kinematic pattern in which prey capture, transport and swallowing are combined into a single feeding stage, apparently leaving the mouth, pharynx, esophagus and stomach vulnerable to bites and stings. Nevertheless, horned lizards consume dozens of harvester ants without harm. We show that their derived feeding kinematics are associated with unique, mucus-secreting pharyngeal papillae that apparently serve to immobilize and incapacitate dangerous ants as they are swallowed by compacting them and binding them in mucus strands. Radially branched esophageal folds provide additional mucus-secreting surfaces the ants pass through as they are swallowed. Ants extracted from fresh-killed horned lizard stomachs are curled ventrally into balls and bound in mucus. We conclude that the pharyngeal papillae, in association with a unique form of hyolingual prey transport and swallowing, are horned lizard adaptations related to a diet of dangerous prey. Harvester ant defensive weapons, along with horned lizard adaptations against such weapons, suggest a long-term, predator-prey, co-evolutionary arms race between Phrynosoma and Pogonomyrmex. J. Exp. Zool. 309A:447-459, 2008. (c) 2008 Wiley-Liss, Inc.     

Structure and function of the hyolingual system in Hynobius and its bearing on the evolution of prey capture in terrestrial salamanders

The Hynobiidae is generally regarded as the most phylogenetically basal and least derived extant family of terrestrial salamanders. As in the other families of terrestrial salamanders, prey capture in the Hynobiidae is accomplished by lingual prehension. In Hynobius, the prey capture system appears to be a mosaic of derived and primitive features. This, in conjunction with previous studies, suggests that the hyolingual systems of all families of terrestrial salamanders have evolved various degrees of specialization since the appearance of the common ancestral condition. We propose that the generalized feeding system for the extant terrestrial salamanders includes a hyolingual skeleton comprised of one basibranchial, one pair of radial or radial-like structures, two pairs of ceratobranchials, two pairs of epibranchials, one pair of ceratohyals, and one urohyal arranged in a configuration similar to that of Hynobius; a simple, sac-like secondary tongue pad; a lift and thrust system of tongue projection; a four-part gape cycle; and a forward head and body surge. Modifications to this general plan, previously described for the disparate families, include various changes in the size, shape, and definition of the tongue pad, changes in the specific types of structures and configurations in the anterior hyolingual skeleton, secondary ossification in the posterior hyolingual skeleton, the appearance of various protrusion, projection, and flipping systems for tongue protraction, simplification of the kinematic gape profile, and loss of the forward head and body surge. The evolutionary trends in these modifications have provided a rich data set from which much phylogenetic information has been inferred. © 1996 Wiley-Liss, Inc.     

A KINEMATIC STUDY OF SUCTION FEEDING AND ASSOCIATED BEHAVIOR IN THE LONG-FINNED PILOT WHALE, GLOBICEPHALA MELAS (TRAILL)

Analysis of videotaped feeding sequences provides novel documentation of suction feeding in captive juvenile long-finned pilot whales ( Globicephala melas ). Swimming and stationary whales were videotaped while feeding at the surface, mid-water, and bottom. The ingestion sequence includes a preparatory phase with partial gape followed by jaw opening and rapid hyoid depression to suck in prey at a mean distance of 14 cm (duration 90 msec), although prey were taken from much greater distances. Depression and retraction of the large, piston-like tongue generate negative intraoral pressures for prey capture and ingestion. Food was normally ingested without grasping by teeth yet was manipulated with lingual, hyoid, and mandibular movement for realignment; suction was then used to transport prey into the oropharynx. Whales frequently rolled or inverted before taking prey, presumably to avoid grasping and repositioning. Prey were sucked off the bottom or sides of the pool without direct contact; lateral suction was used to ingest items from the sides of the mouth.