Cranial Elevation Research Articles

Epaxial and hypaxial co-contraction: a mechanism for modulating strike pressure and accuracy during suction feeding in channel catfish.

Most fish species use concentric epaxial and hypaxial contractions to suction feed, whereby both muscle groups produce cranial expansion and negative intraoral pressures. In contrast, channel catfish (Ictalurus punctatus) suction feed with little to no cranial elevation and epaxial shortening, generating suction power primarily with hypaxial shortening and pectoral girdle retraction. We hypothesized that channel catfish (1) actively anchor the head via isometric contraction of the epaxials and (2) vary feeding performance by modulating the absolute and relative outputs of the co-contracting muscles. We used a combination of electromyography, intraoral pressure recordings and specimen manipulation, and developed a new dual-lever model to explore this idea. We detected epaxial and hypaxial co-contraction prior to suction force development in all strikes. Our model revealed that the differential between the co-contracting muscles may be used to modulate suction pressure and strike accuracy.

Morphological comparison of the cranial movement apparatus in mudskippers (Gobiidae: Oxudercinae)

Possession of the neck allows vertebrates to move the head independently from the trunk. Fish do not have the neck and thus the cranial mobility could be limited. Oxudercine gobies show full range of habitat transition from aquatic to terrestrial environments and exhibit flexible cranial movement, yet the cranium-movement apparatus is little known. In this study, we investigated the anatomy of the structure of the eight oxudercine gobies, Oxuderces nexipinnis, Parapocryptes serperaster, Pseudapocryptes elongatus, Scartelaos histophorus, Boleophthalmus boddarti, Periophthalmus chrysospilos, Periophthalmodon schlosseri, and Periophthalmodon septemradiatus. These species share similarities in the specialized features of the craniovertebral joint and the epaxials attaching onto different locations of the neurocranium. On the other hand, large space between the ventral portions of the craniovertebral joint only occurs in O. nexipinnis, Pd. elongatus, Pn. schlosseri and Pn. septemradiatus. Hypaxials are hypertrophied at the insertion point and attach more anteriorly onto the ventral side of the neurocranium in B. boddarti, O. nexipinnis, Pa. serperaster, Pd. elongatus, and S. histophorus, whereas the muscles are small and attach posteriorly in the remaining species. There were significant differences in the area occupancy ratio of the post-cranial neural spines, the lever arm ratio of the cranial rotation, and the angle between the horizontal plane and the plane through the craniovertebral joint among the species. The cranial depression presumably facilitates grazing of oxudercine gobies in their early stage of terrestrial transition, whereas the cranial elevation parameters are contradictory to the terrestrial gradient. The cranium-movement morphometrics partially agree with the phylogeny.

A neck-like vertebral motion in fish.

Tetrapods use their neck to move the head three-dimensionally, relative to the body and limbs. Fish lack this anatomical neck, yet during feeding many species elevate (dorsally rotate) the head relative to the body. Cranial elevation is hypothesized to result from the craniovertebral and cranial-most intervertebral joints acting as a neck, by dorsally rotating (extending). However, this has never been tested due to the difficulty of visualizing and measuring vertebral motion in vivo. I used X-ray reconstruction of moving morphology to measure three-dimensional vertebral kinematics in rainbow trout (Oncorhynchus mykiss) and Commerson's frogfish (Antennarius commerson) during feeding. Despite dramatically different morphologies, in both species dorsoventral rotations extended far beyond the craniovertebral and cranial intervertebral joints. Trout combine small (most less than 3°) dorsal rotations over up to a third of their intervertebral joints to elevate the neurocranium. Frogfish use extremely large (often 20–30°) rotations of the craniovertebral and first intervertebral joint, but smaller rotations occurred across two-thirds of the vertebral column during cranial elevation. Unlike tetrapods, fish rotate large regions of the vertebral column to rotate the head. This suggests both cranial and more caudal vertebrae should be considered to understand how non-tetrapods control motion at the head–body interface.

Fishes can use axial muscles as anchors or motors for powerful suction feeding.

ABSTRACTSome fishes rely on large regions of the dorsal (epaxial) and ventral (hypaxial) body muscles to power suction feeding. Epaxial and hypaxial muscles are known to act as motors, powering rapid mouth expansion by shortening to elevate the neurocranium and retract the pectoral girdle, respectively. However, some species, like catfishes, use little cranial elevation. Are these fishes instead using the epaxial muscles to forcefully anchor the head, and if so, are they limited to lower-power strikes? We used X-ray imaging to measure epaxial and hypaxial length dynamics (fluoromicrometry) and associated skeletal motions (XROMM) during 24 suction feeding strikes from three channel catfish (Ictalurus punctatus). We also estimated the power required for suction feeding from oral pressure and dynamic endocast volume measurements. Cranial elevation relative to the body was small (<5 deg) and the epaxial muscles did not shorten during peak expansion power. In contrast, the hypaxial muscles consistently shortened by 4–8% to rotate the pectoral girdle 6–11 deg relative to the body. Despite only the hypaxial muscles generating power, catfish strikes were similar in power to those of other species, such as largemouth bass (Micropterus salmoides), that use epaxial and hypaxial muscles to power mouth expansion. These results show that the epaxial muscles are not used as motors in catfish, but suggest they position and stabilize the cranium while the hypaxial muscles power mouth expansion ventrally. Thus, axial muscles can serve fundamentally different mechanical roles in generating and controlling cranial motion during suction feeding in fishes.

Spatial and temporal changes in buccal pressure during prey-capture in the trumpetfish (Aulostomus maculatus)

Suction feeding involves the coordinated contraction of numerous muscles to rapidly expand the buccal cavity to create hydrodynamic loading. The expanding buccal cavity generates forces that draw water and the prey item in as the pressure is equalized between the buccal cavity and the external environment. Here we quantified the spatial and temporal aspects of subambient pressure generation in the elongated buccal cavity of trumpetfish (Aulostomus maculatus) during feeding strikes. Pressure was measured with two Millar micro-tip catheter pressure transducers (SPR-407) and was synchronized with high-speed videography. There was a statistically significant difference (p = 0.03) between the mean pressure generated in the caudal buccal cavity (− 24.67 ± 0.72 kPa) and the rostral buccal cavity (− 18.69 ± 0.66 kPa). There was also a statistically significant difference (p = 0.01) in the time between rostral and caudal peak pressure generation. The caudal peak pressure is generated 5.56 ± 0.38 ms before the rostral peak pressure and is nearly synchronous with maximum hyoid depression. Kinematic data reveal that, after subambient pressure is generated by hyoid depression, the buccal cavity first expands laterally at the mouth opening and proceeds in a caudal direction. Morphological data suggest that trumpetfish evolved lengthened sesamoid tendons that translate whole-body contractile forces to cranial elevation and buccal cavity expansion during feeding. These tendons may have evolved to compensate for the lengthy moment arm created by the elongated skull of trumpetfish in order to overcome the resultant increased water resistance.

New views of humpback whale flow dynamics and oral morphology during prey engulfment.

The rise of inexpensive, user-friendly cameras and editing software promises to revolutionize data collection with minimal disturbance to marine mammals. Video sequences recorded by aerial drones and GoPro cameras provided close-up views and unique perspectives of humpback whales engulfing juvenile salmon at or just below the water surface in Southeast Alaska and Prince William Sound. Although humpback feeding is famous for its flexibility, several stereotyped events were noted in the 47 lunges we analyzed. Engulfment was rapid (mean 2.07 s), and the entrance through which the tongue inverts into the ventral pouch was seen as water rushes in. Cranial elevation was a major contributor to gape, and pouch contraction sometimes began before full gape closure, with reverberating waves indicating rebounding flow of water within the expanded pouch. Expulsion of filtered water began with a small splash at the anterior of the mouth, followed by sustained excurrent flow in the mouth's central or posterior regions. Apart from a splash of rebounding water, water within the mouth was surprisingly turbulence-free during engulfment, but submersion of the whale's head created visible surface whirlpools and vortices which may aggregate prey for subsequent engulfment.

Bite force and feeding kinematics in the eastern North Pacific kyphosidae

Some fishes that feed on attached food items possess an intramandibular joint (IMJ), which is thought to increase maximum gape and facilitate contact between the tooth-bearing surface and the substrate. However, the mechanical consequences of using an IMJ to remove attached food items from the substrate are still poorly understood. We examined the most prominent eastern North Pacific kyphosid, the scraper: Girella nigricans and two other kyphosids, Medialuna californiensis and Hermosilla azurea, which occupy similar habitats. Of the three species, G. nigricans had the highest theoretical bite force per unit length. We examined the feeding mechanics of G. nigricans in two different feeding scenarios: a scraping behavior elicited on a block of brine shrimp gelatin and a picking behavior elicited on Ulva sp. We measured cranial elevation, lower jaw rotation, premaxillary protrusion, premaxillary rotation, gape maximum, and intramandibular rotation. Ulva treatments produced significantly greater cranial rotation, when compared to gelatin treatments. Gelatin treatments were associated with greater lower jaw rotation and larger gape. Premaxillary rotation and premaxillary protrusion did not differ between treatments. Intramandibular rotation occurred only when G. nigricans physically contacted the gelatin, suggesting the IMJ is a passive joint with no associated musculature. We also noted that G. nigricans do not appear to use suction to draw food into the mouth. The lack of suction and the presence of the IMJ suggest that the jaws of G. nigricans are specialized for maximizing jaw force when scraping.

Modeling the Relationship between Environmental Temperature and Feeding Performance in Florida (USA) Nonnative Fishes, with Implications for Invasive-Species Response to Climate Change

Aims: To determine the effect of environmental temperature on the feeding kinematics of two Florida (USA) invasive-fish species, pike killifish Belonesox belizanus and Mayan cichlid Cichlasoma urophthalmus, in an attempt to explore the adaptive response of wholeorganism performance to climate change. Study Design: Model I (Fixed-Effects Model) Linear Regression Analysis, y-Dependent Variable = Kinematics; x-Independent Variable = Environmental Temperature. Place and Duration of Study: Fish Ecophysiology Laboratory, Department of Biological Sciences, Florida Institute of Technology in May-December, 2011. Methodology: Four each of B. belizanus and C. urophthalmus, collected from invasive populations in Florida were acclimated in 38 liter experimental aquarium, trained, and filmed using high-speed video while eating fish-prey at 20°C, 25°C and 30°C. Four films per invasive-fish species at each temperature were analyzed using MaxTRAQ digitizing software. In each film, three kinematic-excursion (peak gape, peak hyoid depression and cranial elevation at peak gape) and three kinematic-timing (time to peak gape, hyoid depression, and cranial elevation) variables were measured. Each kinematic variable was regressed against temperature to model the relationship between feeding performance and Research Article Annual Research & Review in Biology, 4(1): 121-132, 2014 122 environmental temperature. Results: All experimental fishes behaved normally and fed aggressively in each of the three experimental temperature regimes. It is evident in the feeding kinematics-temperature plots that fish-feeding at each temperature was variable and unpredictable. Out of the 12 regression equations generated to model the relationship between feeding kinematics and environmental temperature, only one (peak gape in C. urophthalmus) showed a significant slope (Peak Gape = 1.42 + 0.01 Temperature; R = 0.22; P< 0.01). Conclusion: The models defining the relationship between feeding kinematics and environmental temperature are weak, as indicated by the extremely low values of the coefficient of determination (R). Empirical evidence indicates that the feeding performance of invasive B. belizanus and C. urophthalmus is not affected by temperature.

Prey processing in the Siamese fighting fish (Betta splendens)

We studied prey processing in the Siamese fighting fish (Betta splendens), involving slow, easily observed head-bobbing movements, which were compared with prey processing in other aquatic feeding vertebrates. We hypothesized that head-bobbing is a unique prey-processing behaviour, which alternatively could be structurally and functionally analogous with raking in basal teleosts, or with pharyngognathy in neoteleosts. Modulation of head-bobbing was elicited by prey with different motility and toughness. Head-bobbing involved sustained mouth occlusion and pronounced cranial elevation, similar to raking. However, the hyoid and pectoral girdle were protracted, and not retracted as in both raking and pharyngognathy. High-speed videofluoroscopy of hyoid movements confirmed that head-bobbing differs from other known aquatic prey-processing behaviours. Nevertheless, head-bobbing and other prey-processing behaviours converge on a recurrent functional theme in the trophic ecology of aquatic feeding vertebrates; the use of intraoral and oropharyngeal dentition surfaces to immobilize, reduce and process relatively large, tough or motile prey. Prey processing outside the pharyngeal region has not been described for neoteleosts previously, but morphological evidence suggests that relatives of Betta might use similar processing behaviours. Thus, our results suggest that pharyngognathy did not out-compete ancestral prey-processing mechanisms completely during the evolution of neoteleosts.

Sound production in the longnose butterflyfishes (genusForcipiger): cranial kinematics, muscle activity and honest signals

Many teleost fishes produce sounds for social communication with mechanisms that do not involve swim bladder musculature. Such sounds may reflect physical attributes of the sound-production mechanism, be constrained by body size and therefore control signal reliability during agonistic behaviors. We examined kinematics of the cranium, median fins and caudal peduncle during sound production in two territorial chaetodontid butterflyfish sister species: forcepsfish (Forcipiger flavissimus) and longnose butterflyfish (F. longirostris). During intraspecific agonistic encounters, both species emit a single pulse sound that precedes rapid cranial rotation at velocities and accelerations that exceed those of prey strikes by many ram- and suction-feeding fishes. Electromyography showed that onsets of activity for anterior epaxialis, sternohyoideus, A1 and A2 adductor mandibulae muscles and sound emission are coincident but precede cranial elevation. Observations indicate that sound production is driven by epaxial muscle contraction whereas a ventral linkage between the head and pectoral girdle is maintained by simultaneous activity from the adductor mandibulae and sternohyoideus. Thus, the girdle, ribs and rostral swim bladder are pulled anteriorly before the head is released and rotated dorsally. Predictions of the hypothesis that acoustic signals are indicators of body size and kinematic performance were confirmed. Variation in forcepsfish sound duration and sound pressure level is explained partly by cranial elevation velocity and epaxial electromyogram duration. Body size, however, explains most variation in duration and sound pressure level. These observed associations indicate that forcepsfish sounds may be accurate indicators of size and condition that are related to resource holding potential during social encounters.

The integration of locomotion and prey capture in divergent cottid fishes: functional disparity despite morphological similarity

Many mobile animals rely on the integration of locomotion and feeding to capture prey. Fishes commonly swim up to a prey item and utilize a combination of ram and suction feeding for prey capture. Marine cottids represent a diverse and abundant lineage of fishes that exhibit variation in feeding mode that is related to their mouth morphology. However, little is known regarding the integration of the locomotor and feeding systems during prey capture. We quantified the feeding kinematics, feeding performance and integration of locomotion and feeding in two species of divergent cottids: Blepsias cirrhosus (silver-spotted sculpin) and Oligocottus maculosus (tidepool sculpin). Individuals were caught from sympatric habitats near the Bamfield Marine Sciences Centre on Vancouver Island and filmed with a high-speed video camera (500 Hz) while feeding on amphipod prey. Two principal component axes summarize differences in integration and feeding mode despite similarity in attack velocity and feeding morphology (peak gape, peak cranial elevation and peak jaw protrusion). A greater number of correlations between locomotor and feeding variables in B. cirrhosus, compared with O. maculosus, indicate greater integration. We conclude that traditional measures of attack kinematics do not capture functionally and ecologically relevant differences between species. The mechanisms underlying differences in locomotor strategy likely result from unexplored morphological or ecological differences between species. In cottids, integration is apparent in more basal, subtidal species such as B. cirrhosus, and the need for integration may be superceded by demands from the habitat in more derived, shallow-water species such as O. maculosus.

Head and body kinematics of pulse sound generation and feeding in longnose butterflyfishes (Genus Forcipiger).

Both species of butterflyfish (Chaetododntidae) in the genus Forcipiger produce pulsatile sounds during territorial interactions. Sounds of one species, the forcepsfish (F. flavissimus), are approximately 20 ms and occur with rapid dorsoventral head motion (headbob). We performed laboratory sound production and feeding experiments with high‐speed video (600 fps) to test predictions of the hypothesis that sound production in this genus results from evolutionary modification of stereotyped feeding behavior. Cranial elevation is an important component of prey capture for a variety of teleost fishes and facilitates opening of the buccal cavity and oral jaws. Studies on kinematics of feeding in Forcipiger by other authors revealed a minor contribution of cranial elevation compared to other fishes. Preliminary results indicate cranial elevation during headbobs involves a comparable range of motion relative to prey capture (approximately 3°–7°), but maximal elevation occurs within 12 ms during the headbob compar...

A biomechanical model of feeding kinematics forDunkleosteus terrelli(Arthrodira, Placodermi)

Biomechanical models illustrate how the principles of physics and physiology determine function in organisms, allowing ecological inferences and functional predictions to be based on morphology. Dynamic lever and linkage models of the mechanisms of the jaw and skull during feeding in fishes predict function from morphology and have been used to compare the feeding biomechanics of diverse fish groups, including fossil taxa, and to test ideas in ecological morphology. Here we perform detailed computational modeling of the four-bar linkage mechanism in the skull and jaw systems ofDunkleosteus terrelli, using software that accepts landmark morphological data to simulate the movements and mechanics of the skull and jaws during prey capture. The linkage system is based on the quadrate and cranio-thoracic joints: Cranial elevation around the cranio-thoracic joint forces the quadrate joint forward, which, coupled with a jaw depressor muscle connecting the jaw to the thoracic shield, causes the jaw to rotate downward during skull expansion. Results show a high speed transmission for jaw opening, producing a rapid expansion phase similar to that in modern fishes that use suction during prey capture. During jaw closing, the model computes jaw and skull rotation and a series of mechanical metrics including effective mechanical advantage of the jaw lever and kinematic transmission of the skull linkage system. Estimates of muscle cross-sectional area based on the largest of five specimens analyzed allow the bite force and strike speed to be estimated. Jaw-closing muscles ofDunkleosteuspowered an extraordinarily strong bite, with an estimated maximal bite force of over 6000 N at the jaw tip and more than 7400 N at the rear dental plates, for a large individual (10 m total length). This bite force capability is among the most powerful bites in animals. The combination of rapid gape expansion and powerful bite meant thatDunkleosteus terrellicould both catch elusive prey and penetrate protective armor, allowing this apex predator to potentially eat anything in its ecosystem, including other placoderms.

Biomechanics of a convergently derived prey-processing mechanism in fishes: evidence from comparative tongue bite apparatus morphology and raking kinematics

A tongue-bite apparatus (TBA) governs raking behaviors in two major and unrelated teleost lineages, the osteoglossomorph and salmoniform fishes. We present data on comparative morphology and kinematics from two representative species, the rainbow trout (Oncorhynchus mykiss) and the Australian arowana (Scleropages jardinii), which suggest that both the TBA and raking are convergently derived in these lineages. Similar TBA morphologies were present, except for differences in TBA dentition and shape of the novel cleithrobranchial ligament (CBL), which is arc-shaped in O. mykiss and straight in S. jardinii. Eight kinematic variables were used to quantify motion magnitude and maximum-timing in the kinematic input mechanisms of the TBA. Five variables differed inter-specifically (pectoral girdle retraction magnitude and timing, cranial and hyoid elevation and gape-distance timing), yet an incomplete taxon separation across multivariate kinematic space demonstrated an overall similarity in raking behavior. An outgroup analysis using bowfin (Amia calva) and pickerel (Esox americanus) to compare kinematics of raking with chewing and prey-capture provided robust quantitative evidence of raking being a convergently derived behavior. Support was also found for the notion that raking more likely evolved from the strike, a functionally distinct behavior, than from chewing, an alternative prey-processing behavior. Based on raking kinematic and muscle-activity data, we propose biomechanical models of the three input mechanisms that govern kinematics of the basihyal output mechanism during the raking power stroke: (1) cranial elevation protracts the upper TBA jaw from the lower (basihyal) TBA jaw; (2) basihyal retraction is caused directly by contraction of the sternohyoideus (SH); (3) hypaxial shortening, relayed via the pectoral girdle and SH-CBL complex, is an indirect basihyal retraction mechanism modeled as a four-bar linkage. These models will aid future analyses mapping structural and functional traits to the evolution of behaviors.

Prey capture kinematics and four-bar linkages in the bay pipefish, Syngnathus leptorhynchus

Because of their modified cranial morphology, syngnathid pipefishes have been described as extreme suction feeders. The presumption is that these fishes use their elongate snout much like a pipette in capturing planktonic prey. In this study, we quantify the contribution of suction to the feeding strike and quantitatively describe the prey capture mechanics of the bay pipefish Syngnathus leptorhynchus, focusing specifically on the role of both cranial elevation and snout movement. We used high-speed video to capture feeding sequences from nine individuals feeding on live brine shrimp. Sequences were digitized in order to calculate kinematic variables that could be used to describe prey capture. Prey capture was very rapid, from 2 to 6 ms from the onset of cranial rotation. We found that suction contributed at most about one-eighth as much as ram to the reduction of the distance between predator and prey. This movement of the predator was due almost exclusively to movement of the snout and neurocranium rather than movement of the whole body. The body was positioned ventral and posterior to the prey and the snout was rotated dorsally by as much as 21°, thereby placing the mouth immediately behind the prey for capture. The snout did not follow the identical trajectory as the neurocranium, however, and reached a maximum angle of only about 10°. The snout consists, in part, of elongate suspensorial elements and the linkages among these elements are retained despite changes in shape. Thus, when the neurocranium is rotated, the four-bar linkage that connects this action with hyoid depression simultaneously acts to expand and straighten the snout relative to the neurocranium. We confirm the presence of a four-bar linkage that facilitates these kinematics by couplings between the pectoral girdle, urohyal, hyoid complex, and the neurocranium–suspensorium complex.

Congruence between muscle activity and kinematics in a convergently derived prey-processing behavior

Quantification of anatomical and physiological characteristics of the function of a musculoskeletal system may yield a detailed understanding of how the organizational levels of morphology, biomechanics, kinematics, and muscle activity patterns (MAPs) influence behavioral diversity. Using separate analyses of these organizational levels in representative study taxa, we sought patterns of congruence in how organizational levels drive behavioral modulation in a novel raking prey-processing behavior found in teleosts belonging to two evolutionarily distinct lineages. Biomechanically divergent prey (elusive, robust goldfish and sedentary, malleable earthworms) were fed to knifefish, Chitala ornata (Osteoglossomorpha) and brook trout, Salvelinus fontinalis (Salmoniformes). Electromyography recorded MAPs from the hyoid protractor, jaw adductor, sternohyoideus, epaxialis, and hypaxialis musculature, while sonomicrometry sampled deep basihyal kinesis and contractile length dynamics in the basihyal protractor and retractor muscles. Syntheses of our results with recent analyses of cranial morphology and raking kinematics showed that raking in Salvelinus relies on an elongated cranial out lever, extensive cranial elevation and a curved cleithrobranchial ligament (CBL), and that both raking MAPs and kinematics remain entirely unmodulated-a highly unusual trait, particularly among feeding generalists. Chitala had a shorter CBL and a raking power stroke involving increased retraction of the elongated pectoral girdle during raking on goldfish. The raking MAP was also modulated in Chitala, involving an extensive overlap between muscle activity of the preparatory and power stroke phases, driven by shifts in hypaxial timing and recruitment of the hyoid protractor muscle. Sonomicrometry revealed that the protractor hyoideus muscle stored energy from retraction of the pectoral girdle for ca. 5-20 ms after onset of the power stroke and then hyper-extended. This mechanism of elastic recoil in Chitala, which amplifies retraction of the basihyal during raking on goldfish without a significant increase in recruitment of the hypaxialis, suggests a unique mechanism of modulation based on performance-enhancing changes in the design and function of the musculoskeletal system.

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.

Coordinated mechanics of feeding, swimming, and eye movements in Tautoga onitis, and implications for the evolution of trophic strategies in fishes

Fish feeding behavior results from successful coordination of the fins, jaws, and sensory systems, and the organization of this behavior may affect the fish’s foraging abilities and trophic ecology. Using quantitative kinematic methods, movements of the jaws, fins and eyes of Tautoga onitis (Teleostei: Labridae) were analyzed during feeding events. Tautog feeding events consisted of three phases: approach, strike, and recovery, each defined by a combination of kinematic events. The approach was characterized by changes in fin movements and in body position, with the eyes directed forward at the food item. The strike began with the onset of jaw opening and protrusion, then cranial elevation, with the eyes no longer looking at the food item. The end of the strike and the beginning of the recovery involved a braking maneuver with the pectoral fins; the fish turns and swims away from the original food location item after prey capture. The coordination performance variables of tautog were quantitatively compared to published data from closely related cheiline wrasses and parrotfishes, which represent different feeding ecologies within a monophyletic assemblage. Fishes feeding on molluscs and benthic invertebrates (Cheilinus fasciatus and Tautoga onitis) represented an intermediate coordination condition, with herbivores (the parrotfishes, Scarus quoyi and Sparisoma radians) at one extreme, and fishes feeding on elusive prey (Epibulus insidiator and Oxycheilinus digrammus) at the other extreme. The analysis suggests that the biomechanical demands of coordination for feeding on benthic invertebrates may represent a generalized, and perhaps ancestral behavior in the wrasses, whereas more specialized trophic niches have evolved divergent, more specialized demands. Examining the movement and coordination of the jaws, fins, and eyes during fish feeding provides a detailed mechanistic basis for behavior, and comparison of coordination patterns during feeding among different taxa can measure how these trophic strategies differ. Understanding the evolution of feeding ecologies in these demersal fishes may have implications for understanding their role within their shallow water reef community.

Sound communication by the forceps fish, Forcipiger flavissimus (Chaetodontidae)

The forceps fish, Forcipiger flavissimus (family Chaetodontidae), lacks the anterior swim bladder horns and laterophysic connection that are proposed to improve processing of acoustic signals in the genus Chaetodon. Despite the lack of hearing specializations, laboratory experiments using hydrophones synchronized to video revealed the forceps fish produce at least three sounds: (1) a pulse (21‐ms duration, peak frequency=318 Hz) associated with rapid cranial elevation, (2) a low‐frequency pulse (9‐ms duration, peak frequency=41 Hz) associated with anal fin retraction and elevation, and (3) a high‐frequency broadband click (3‐ms duration, peak frequency=7924 Hz) associated with tail slap behavior. Auditory evoked potential experiments demonstrate that hearing thresholds for this species are 10–21 dB higher than Chaetodon in the 200–600‐Hz band. High‐speed kinematic analyses show that motor movements associated with some acoustic behaviors are different than those in Chaetodon, thus the presence of sound production may be the ancestral state for the family but the mechanisms are divergent. Field observations on Hawaiian reefs reveal that forceps fish occur in pairs and trios that may be haremic, and use acoustic communication among group members.[Work supported by NSF IBN0137883 to TT.]

Comparative and developmental functional morphology of the jaws of living and fossil gars (Actinopterygii: Lepisosteidae)

The feeding mechanism of gars (Ginglymodi : Lepisosteidae) is characterized by cranial elevation and lower jaw rotation but minimal cranial kinesis. Gar jaws have numerous, sharply pointed, elongate teeth for capture of evasive prey. Their mandibles range from relatively short to extremely long depending on the species. Jaw length and lever dimensions were hypothesized to affect the biomechanics of force and motion during feeding, according to simple mechanical models of muscles exerting force through first- or third-order levers. A morphometric protocol was used to measure the jaw structure of seven living and five fossil species of gar and these data were used to calculate the mechanical advantage (a measure of force transmission) for both opening and closing of the mandible. Gars were found to possess low mechanical advantage (MA) and high transmission of motion, although gars occupy a range of biomechanical states across the continuum of force vs. velocity transmission. The long-nose gar, Lepisosteus osseus, has one of the lowest jaw closing MAs (0.05) ever measured in fishes. Intraspecific lever mechanics were also calculated for a developmental series (from feeding larvae to adults) of L. osseus and Atractosteus spatula. A characteristic ontogenetic curve in MA of the lower jaw was obtained, with a large decrease in MA between larva and juvenile, followed by a steady increase during adult growth. This curve correlates with a change in prey type, with the small, robust-jawed individuals feeding mainly on crustaceans and insects and the large, long-jawed individuals of all species becoming mainly piscivorous. Principal components analysis of functionally important morphometrics shows that several gar species occupy different regions of functional morphospace. Some fossil gar species are also placed within functional morphospace using this approach.