Effects of organism and substrate size on burial mechanics of English sole, Parophrys vetulus.

Flatfishes use cyclic body undulations to force water into the sediment and fluidize substrate particles, displacing them into the water column. When water velocity decreases, suspended particles settle back onto the fish, hiding it from view. Burial may become more challenging as flatfishes grow because the area to be covered increases exponentially with the second power of length. In addition, particle size is not uniform in naturally occurring substrates, and larger particles require higher water velocities for fluidization. We quantified the effects of organism and particle-size scaling on burial behavior of English sole, Parophrys vetulus We recorded burial events from a size range of individuals (5-32 cm total length, TL), while maintaining constant substrate grain size. Larger fish used lower cycle frequencies and took longer to bury, but overall burial performance was maintained (∼100% coverage). To test the effect of particle size on burial performance, individuals of similar lengths (5.7-8.1 cm TL) were presented with different substrate sizes (0.125-0.710 mm). Particle size did not affect cycle frequency or time to burial, but fish did not achieve 100% coverage with the largest particles because they could not fluidize this substrate. Taken together, these results suggest that both body size and substrate grain size can potentially limit the ability of flatfishes to bury: a very large fish (>150 cm) may move too slowly to fluidize all but the smallest substrate particles and some particles are simply too large for smaller individuals to fluidize.

Behavioral and physiological adaptations to high-flow velocities in chubs (Gila spp.) native to Southwestern USA.

Morphological streamlining is often associated with physiological advantages for steady swimming in fishes. Though most commonly studied in pelagic fishes, streamlining also occurs in fishes that occupy high-flow environments. Before the installation of dams and water diversions, bonytail (Cyprinidae, Gila elegans), a fish endemic to the Colorado River (USA), regularly experienced massive, seasonal flooding events. Individuals of G. elegans display morphological characteristics that may facilitate swimming in high-flow conditions, including a narrow caudal peduncle and a high aspect ratio caudal fin. We tested the hypothesis that these features improve sustained swimming performance in bonytail by comparing locomotor performance in G. elegans with that of the closely related roundtail chub (Gila robusta) and two non-native species, rainbow trout (Oncorhynchus mykiss) and smallmouth bass (Micropterus dolomieu), using a Brett-style respirometer and locomotor step-tests. Gila elegans had the lowest estimated drag coefficient and the highest sustained swimming speeds relative to the other three species. There were no detectible differences in locomotor energetics during steady swimming among the four species. When challenged by high-velocity water flows, the second native species examined in this study, G. robusta, exploited the boundary effects in the flow tank by pitching forward and bracing the pelvic and pectoral fins against the acrylic tank bottom to 'hold station'. Because G. robusta can station hold to prevent being swept downstream during high flows and G. elegans can maintain swimming speeds greater than those of smallmouth bass and rainbow trout with comparable metabolic costs, we suggest that management agencies could use artificial flooding events to wash non-native competitors downstream and out of the Colorado River habitat.

Functional morphology and kinematics of terrestrial feeding in the largescale foureyes (Anableps anableps).

A major challenge for aquatic vertebrates that invade land is feeding in the terrestrial realm. The capacity of the gape to become parallel with the ground has been shown to be a key factor to allow fishes to feed on prey lying on a terrestrial surface. To do so, two strategies have been identified that involve a nose-down tilting of the head: (1) by pivoting on the pectoral fins as observed in mudskippers, and (2) curling of the anterior part of the body supported by a long and flexible eel-like body as shown in eel-catfish. Although Anableps anableps successfully feeds on land, it does not possess an eel-like body or pectoral fins to support or lift the anterior part of the body. We identified the mechanism of terrestrial prey capture in A. anableps by studying kinematics and functional morphology of the cranial structures, using high-speed video and graphical 3D reconstructions from computed tomography scans. In contrast to the previously described mechanisms, A. anableps relies solely on upper and lower jaw movement for re-orientation of the gape towards the ground. The premaxilla is protruded anteroventrally, and the lower jaw is depressed to a right angle with the substrate. Both the lower and upper jaws are selectively positioned onto the prey. Anableps anableps thereby uses the jaw protrusion mechanism previously described for other cyprinodontiforms to allow a continued protrusion of the premaxilla even while closing the jaws. Several structural adaptations appear to allow more controlled movements and increased amplitude of anterior and ventral protrusion of the upper jaw compared with other cyprinodontiforms.

A Step Forward: Functional Diversity and Emerging Themes of Slow-Speed Locomotion in Vertebrates.

Walking can be defined broadly as a slow-speed movement produced when appendages interact with the ground to generate forward propulsion. Until recently, most studies of walking have focused on humans and a handful of domesticated vertebrates moving at a steady rate over highly simplified, static surfaces, which may bias our understanding of the unifying principles that underlie vertebrate locomotion. In the last few decades, studies have expanded to include a range of environmental contexts (e.g., uneven terrain, perturbations, deformable substrates) and greater phylogenetic breadth (e.g., non-domesticated species, small and/or ectothermic tetrapods and fishes); these studies have revealed that even a gait as superficially simple as walking is far more complex than previously thought. In addition, technological advances and accessibility of imaging systems and computational power have recently expanded our capabilities to test hypotheses about the locomotor movements of extant and extinct organisms in silico. In this symposium, scientists showcased diverse taxa (from extant fishes to extinct dinosaurs) moving through a range of variable conditions (speed perturbations, inclines, and deformable substrates) to address the causes and consequences of functional diversity in locomotor systems and discuss nascent research areas and techniques. From the symposium contributions, several themes emerged: (1) slow-speed, appendage-based movements in fishes are best described as walking-like movements rather than true walking gaits, (2) environmental variation (e.g., deformable substrates) and dynamic stimuli (e.g., perturbations) trigger kinematic and neuromuscular changes in animals that make defining a single gait or the transition between gaits more complicated than originally thought, and (3) computational advances have increased the ability to process large data sets, emulate the 3D motions of extant and extinct taxa, and even model species interactions in ancient ecosystems. Although this symposium allowed us to make great strides forward in our understanding of vertebrate walking, much ground remains to be covered. First, there is a much greater range of vertebrate appendage-based locomotor behaviors than has been previously recognized and existing terminology fails to accurately capture and describe this diversity. Second, despite recent efforts, the mechanisms that vertebrates use modify locomotor behaviors in response to predictable and unpredictable locomotor challenges are still poorly understood. Third, while computer-based models and simulations facilitate a greater understanding of the kinetics and kinematics of movement in both extant and extinct animals, a universal, one-size-fits-all, predictive model of appendage-based movement in vertebrates remains elusive.

Locomotor behavior across an environmental transition in the ropefish, Erpetoichthys calabaricus.

Many amphibious organisms undergo repeated aquatic to terrestrial transitions during their lifetime; limbless, elongate organisms that make such transitions must rely on axial-based locomotion in both habitats. How is the same anatomical structure employed to produce an effective behavior across such disparate habitats? Here, we examine an elongate amphibious fish, the ropefish (Erpetoichthys calabaricus), and ask: (1) how do locomotor movements change during the transition between aquatic and terrestrial environments and (2) do distantly related amphibious fishes demonstrate similar modes of terrestrial locomotion? Ropefish were examined moving in four experimental treatments (in which the water level was to lowered mimic the transition between environments) that varied from fully aquatic to fully terrestrial. Kinematic parameters (lateral excursion, wavelength, amplitude and frequency) were calculated for points along the midline of the body and compared across treatments. Terrestrial locomotion in the ropefish is characterized by long, slow, large-amplitude undulations down the length of the body; in contrast, aquatic locomotion is characterized by short-wavelength, small-amplitude, high-frequency undulations that gradually increase in an anterior to posterior direction. Experimental treatments with intermediate water levels were more similar to aquatic locomotion in that they demonstrated an anterior to posterior pattern of increasing lateral excursion and wave amplitude, but were more similar to terrestrial locomotion with regard to wavelength, which did not change in an anterior to posterior direction. Finally, the ropefish and another elongate amphibious fish, the eel, consistently exhibit movements characterized by 'path following' when moving on land, which suggests that elongate fishes exhibit functional convergence during terrestrial locomotion.

The fast and the tendinous? Locomotor modifications of the caudal peduncle in Gila spp. from the American Southwest.

In the American Southwest, the fishes within the genus Gila evolved in an environment with seasonal rainstorms that caused stochastic flooding. Some species within this genus, such as bonytail (Gila elegans), possess locomotor morphologies that are similar to those seen in high-performance swimmers such as tuna and lamnid sharks. These shared features include a shallow caudal peduncle, lunate tail, and mechanisms to transmit force from the anterior musculature to the tail fin. We compared the skeletal anatomy of the caudal region of bonytail to roundtail chub (Gila robusta) and humpback chub (Gila cypha) to determine which vertebral elements have been modified to create a shallow peduncle. We also tested the tensile strength of the red (slow oxidative) axial muscle by performing a standard stress test. If the muscle can withstand a large load, this suggests it may play a tendon-like role in transmitting force from the anterior muscle to the hypural plate of the tail. Lastly, we measured the collagen content of the red axial muscle (visualized using serial sections and Masson's trichrome stain) to determine if increased tensile strength is associated with increased collagen content. We found bonytail caudal peduncles are characterized by acute vertebral spines and have red axial muscle that can resist tearing under tension. Roundtail chub peduncles are characterized by relatively more obtuse angles and the red muscle tears easily under tension. Humpback chub possess an intermediate morphology, with relatively obtuse vertebral spine angles and the red muscle can resist tearing under tension. Bonytail have increased collagen content in posterior red axial muscle compared to the anterior musculature also suggesting a tendon-like role of the posterior red muscle. In combination with previous studies of swimming performance, our findings suggest that the axial musculature of bonytail may play a role in transmitting force directly to the shallow peduncle in a manner similar to that of the great lateral tendon of scombrids.

Integrating Studies of Anatomy, Physiology, and Behavior into Conservation Strategies for the Imperiled Cyprinid Fishes of the Southwestern United States.

Over the last 100 years, fishes native to the Southwestern United States have faced a myriad of biotic and abiotic pressures which has resulted in most being federally listed as endangered or threatened. Most notably, water diversions and the introduction of non-native fishes have been the primary culprits in causing the downfall of native fish populations. We describe how recent studies of morphology, physiology, and behavior yield insights into the failed (occasionally successful) management of this vanishing biota. We describe how understanding locomotor morphologies, physiologies, and behaviors unique to Southwestern native fishes can be used to create habitats that favor native fishes. Additionally, through realizing differences in morphologies and behaviors between native and non-native fishes, we describe how understanding predator-prey interactions might render greater survivorship of native fishes when stocked into the wild from repatriation programs. Understanding fundamental form-function relationships is imperative for managers to make educated decisions on how to best recover species of concern in the Southwestern United States and worldwide.

Are Extreme Anatomical Modifications Required for Fish to Move Effectively on Land? Comparative Anatomy of the Posterior Axial Skeleton in the Cyprinodontiformes.

Many teleost fishes with no apparent modifications for life on land are able to produce effective terrestrial locomotor behaviors, including a ballistic behavior called the "tail-flip" jump. Cyprinodontiformes (killifishes, Teleostei: Atherinomorpha) that live at the water's edge vary in morphology and inclination to emerge onto land. Do fish with an amphibious predisposition have extensive modification of the propulsive region of the body when compared to fully aquatic relatives? We quantified body shape and anatomy of the caudal peduncle and tail (the propulsive organ on land and in water) in 11 cyprinodontiform species and two outgroup taxa (Atherinomorpha). We hypothesized that amphibious species would have longer, "shallower" bodies (larger body fineness ratios), deeper (proportionally larger) caudal peduncles, and more robust bones in the tail fin (larger ossified area of the hypural/epural bones) to facilitate locomotor movements on land. We found no evidence of convergence in body shape or skeletal anatomy among species known to make voluntary sojourns onto land. In fact, deep-bodied species, shallow-bodied species, and species with intermediate morphologies all are able to emerge from the water and move on land. It is possible that there are as-yet-undocumented subtle soft-tissue (muscle, tendon, and ligament) modifications that enhance terrestrial locomotor performance in species known to spend large periods of time on land. However, it is also possible that extreme anatomical changes are not required for aquatic cyprinodontiform species to produce effective locomotor movements when they emerge out of the water and move across the land. Anat Rec, 2019. © 2019 American Association for Anatomy.

Morphology of a picky eater: a novel mechanism underlies premaxillary protrusion and retraction within cyprinodontiforms.

Upper jaw protrusion is hypothesized to improve feeding performance in teleost fishes by enhancing suction production and stealth of the feeding event. However, many cyprinodontiform fishes (mid-water feeders, such as mosquitofish, killifish, swordtails, mollies and pupfish) use upper jaw protrusion for "picking" prey out of the water column or off the substrate; this feeding mode may require improved jaw dexterity, but does not necessarily require increased stealth and/or suction production. We describe functional aspects of the bones, muscles and ligaments of the anterior jaws in three cyprinodontiform genera: Fundulus (Fundulidae), Gambusia and Poecilia (Poeciliidae). All three genera possess a premaxillomandibular ligament that connects the premaxilla of the upper jaw to the mandible. The architecture of this ligament is markedly different from the upper-lower jaw connections previously described for basal atherinomorphs or other teleosts, and this loose ligamentous connection allows for more pronounced premaxillary protrusion in this group relative to closely related outgroup taxa. Within poeciliids, a novel insertion of the second division of the adductor mandibulae (A2) onto the premaxilla has also evolved, which allows this jaw adductor to actively retract the premaxilla during mouth closing. This movement is in contrast with most other teleosts, where the upper jaw is retracted passively via pressure applied by the adduction of the lower jaw. We postulate that this mechanism of premaxillary protrusion mediates the cyprinodontiforms' ability to selectively pick specific food items from the water column, surface or bottom, as a picking-based feeding mechanism requires controlled and coordinated "forceps-like" movements of the upper and lower jaws. This mechanism is further refined in some poeciliids, where direct muscular control of the premaxillae may facilitate picking and/or scraping material from the substrate.

Premaxillary movements in cyprinodontiform fishes: an unusual protrusion mechanism facilitates "picking" prey capture.

Premaxillary protrusion is hypothesized to confer a number of feeding advantages to teleost fishes; however, most proposed advantages relate to enhanced stealth or suction production during prey capture. Cyprinodontiformes exhibit an unusual form of premaxillary protrusion where the descending process of the premaxilla does not rotate anteriorly to occlude the sides of the open mouth during prey capture. Instead, the premaxilla is protruded such that it gives the impression of a beak during prey capture. We quantified premaxillary kinematics during feeding in four cyprinodontiform taxa and compared them with three percomorph taxa to identify any performance consequences of this protrusion mechanism. Individual prey capture events were recorded using digital high-speed video at 250-500 frames per second (n >or= 4 individuals, >or= 4 strikes per individual). Species differed in the timing of movement and the maximum displacement of the premaxilla during the gape cycle and in the contribution of the premaxilla to jaw closing. Cyprinodontiform taxa produced less premaxillary protrusion than the percomorph taxa, and were consistently slower in the time to maximum gape. Further, it appears cyprinodontiforms can alter the contribution of the premaxilla to mouth closure on an event-specific basis. We were able to demonstrate that, within at least one species, this variability is associated with the location of the prey (bottom vs. water column). Cyprinodontiform upper jaw movements likely reflect increased dexterity associated with a foraging ecology where prey items are "picked" from a variety of locations: the bottom, water column, or surface. We postulate that dexterity requires slow, precisely controlled jaw movements; thus, may be traded off for some aspects of suction-feeding performance, such as protrusion distance and speed.

A walking behavior generates functional overland movements in the tidepool sculpin, Oligocottus maculosus.

Tidepool sculpins (Oligocottus maculosus) have been observed moving overland in the rocky intertidal, and we documented the terrestrial walking behavior that they use to accomplish this. We quantified the terrestrial movements of O. maculosus and compared them to (1) their aquatic locomotion, (2) terrestrial locomotion of closely-related subtidal species (Leptocottus armatus and Icelinus borealis), and (3) terrestrial movements of walking catfishes (Clarias spp.). We recorded sculpin movements (210 fps) on a terrestrial platform and in a water tank and tracked body landmarks for kinematic analysis. The axial-appendage-based terrestrial locomotion of O. maculosus is driven by cyclic lateral oscillations of the tail, synchronized with alternating rotations about the base of the pectoral fins, a behavior that appears similar to a military "army crawl." The pectoral fins do not provide propulsion, but act as stable points for the body to rotate around. In contrast, individuals of O. maculosus use primarily axial undulation during slow-speed swimming. The army crawl is a more effective terrestrial behavior (greater distance ratio) than the movements produced by L. armatus and I. borealis, which use rapid, cyclic oscillations of the tail, without coordinated pectoral fin movements. Relative to Clarias spp., O. maculosus rotated the body about the base of the pectoral fin, rather than the tip of the fin, which may cause O. maculosus to have a lower distance ratio. Since O. maculosus lack major morphological adaptations for terrestrial locomotion, instead relying on behavioral adaptations, we propose behavioral adaptations may evolutionarily predate morphological adaptations for terrestrial locomotion in vertebrates.

Botulinum toxin injections as a method for chemically denervating skeletal muscle to test functional hypotheses: a pilot study in Lepomis cyanellus.

In this study, we demonstrate that botulinum toxin can be used to chemically denervate muscles to test functional hypotheses. We injected research-grade type A botulinum toxin complex into pectoral fin abductors (abductor superficialis) of green sunfish (Lepomis cyanellus) to determine whether chemical denervation would eliminate the ability of a particular muscle to contribute to overall pectoral fin movements. Reduction of target muscle activity occurred within 8 d of the injection, and paralysis was confirmed using electromyography. No paralysis was seen in the adjacent muscles (abductor profundus) or in positive controls (saline injections). Paralysis occurred more slowly and at lower doses than previously documented for mammals. However, botulinum toxin complex (500 kDa) was used here, whereas previous studies have used purified toxin (150 kDa). Therefore, differences in physiological responses between fish and mammals cannot yet be distinguished from differences caused by the toxin type. However, we note that the toxin complex is less likely to diffuse across muscle fascia (because it is large), which should minimize paralytic effects on adjacent muscles. We suggest that botulinum toxin holds great promise as a chemical denervation agent in functional studies of animal locomotion and feeding behaviors.

Ontogeny of performance in vertebrates.

When competing for food or other resources, or when confronted with predators, young animals may be at a disadvantage relative to adults because of their smaller size. Additionally, the ongoing differentiation and growth of tissues and the development of sensory-motor integration during early ontogeny may constrain performance. Because ectothermic vertebrates show different growth regimes and energetic requirements when compared to endothermic vertebrates, differences in the ontogenetic trajectories of performance traits in these two groups might be expected. However, both groups of vertebrates show similar patterns of changes in performance with ontogeny. Evidence for compensation, resulting in relatively high levels of performance in juveniles relative to adults, appears common for traits related to locomotor and defensive behaviors. However, there is little evidence for compensation in traits associated with feeding and foraging. We suggest that this difference may be due to different selective regimes operating on locomotor versus feeding traits. As a result, relatively high levels of locomotor performance in juveniles and relatively high levels of feeding performance in adults are observed across a wide range of vertebrate groups.

Development of the escape response in teleost fishes: do ontogenetic changes enable improved performance?

Teleost fishes typically first encounter the environment as free-swimming embryos or larvae. Larvae are morphologically distinct from adults, and major anatomical structures are unformed. Thus, larvae undergo a series of dramatic morphological changes until they reach adult morphology (but are reproductively immature) and are considered juveniles. Free-swimming embryos and larvae are able to perform a C-start, an effective escape response that is used evade predators. However, escape response performance improves during early development: as young fish grow, they swim faster (length-specific maximum velocity increases) and perform the escape more rapidly (time to complete the behavior decreases). These improvements cease when fish become juveniles, although absolute swimming velocity (m s(-1)) continues to increase. We use studies of escape behavior and ontogeny in California halibut (Paralichthys californicus), rainbow trout (Oncorhynchus mykiss), and razorback suckers (Xyrauchen texanus) to test the hypothesis that specific morphological changes improve escape performance. We suggest that formation of the caudal fin improves energy transfer to the water and therefore increases thrust production and swimming velocity. In addition, changes to the axial skeleton during the larval period produce increased axial stiffness, which in turn allows the production of a more rapid and effective escape response. Because escape performance improves as adult morphology develops, fish that enter the environment in an advanced stage of development (i.e., those with direct development) should have a greater ability to evade predators than do fish that enter the environment at an early stage of development (i.e., those with indirect development).

Why does Gila elegans have a bony tail? A study of swimming morphology convergence.

Caudal-fin-based swimming is the primary form of locomotion in most fishes. As a result, many species have developed specializations to enhance performance during steady swimming. Specializations that enable high swimming speeds to be maintained for long periods of time include: a streamlined body, high-aspect-ratio (winglike) caudal fin, a shallow caudal peduncle, and high proportions of slow-twitch ("red") axial muscle. We described the locomotor specializations of a fish species native to the Colorado River and compared those specializations to other fish species from this habitat, as well as to a high-performance marine swimmer. The focal species for this study was the bonytail (Gila elegans), which has a distinct morphology when compared with closely related species from the Southwestern United States. Comparative species used in this study were the roundtail chub (Gila robusta), a closely related species from low-flow habitats; the common carp (Cyprinus carpio), an invasive cyprinid also found in low-flow habitats; and the chub mackerel (Scomber japonicus), a model high-performance swimmer from the marine environment. The bonytail had a shallow caudal peduncle and a high-aspect-ratio tail that were similar to those of the chub mackerel. The bonytail also had a more streamlined body than the roundtail chub and the common carp, although not as streamlined as the chub mackerel. The chub mackerel had a significantly higher proportion of red muscle than the other three species, which did not differ from one another. Taken together, the streamlined body, narrow caudal peduncle, and high-aspect-ratio tail of the bonytail suggest that this species has responded to the selection pressures of the historically fast-flowing Colorado River, where flooding events and base flows may have required native species to produce and sustain very high swimming speeds to prevent being washed downstream.

Look before you leap: Visual navigation and terrestrial locomotion of the intertidal killifish Fundulus heteroclitus.

Mummichogs (Fundulus heteroclitus; Cyprinodontiformes) are intertidal killifish that can breathe air and locomote on land. Our goals were to characterize the terrestrial locomotion of mummichogs and determine their method of navigation towards water in a terrestrial environment. We used high-speed video to record behavior during stranding experiments and found that mummichogs use a tail-flip jump to move overland, similarly to other Cyprinodontiformes. However, mummichogs also prop themselves upright into a prone position between each jump, a previously undescribed behavior. After becoming prone, mummichogs rotate about their vertical axis, directing the caudal fin towards the water. Then, they roll back onto their lateral aspect and use a tail-flip behavior to leap into a caudally-directed, ballistic flight path. We conducted experiments to determine the sensory stimulus used to locate a body of water by placing mummichogs on a square platform with one side adjacent to a sea table. Under artificial light, mummichogs moved towards the sea table with a higher frequency than towards the other sides. Under dark conditions, mummichogs did not show a preference for moving towards the sea table. When the surface of the water was covered with reflective foil, mummichogs moved towards it as if it were a body of water. These results suggest that mummichogs primarily use visual cues, specifically reflected light, to orient towards the water. The uprighting behavior that mummichogs perform between terrestrial jumps may provide an opportunity for these fish to receive visual information that allows them to safely return to the water. J. Exp. Zool. 325A:57-64, 2016. © 2015 Wiley Periodicals, Inc.

Cranial movements during suction feeding in teleost fishes: Are they modified to enhance suction production?

Suction is produced during prey capture by most teleost fishes. Here, we ask two questions about the functional basis of suction feeding. First, is there variation in the kinematic pattern produced by different species while suction feeding? Second, do species termed 'suction specialists' demonstrate similar modifications to their feeding behavior? We used 10 kinematic variables in a principal component analysis to identify axes of variation among 14 suction feeding teleost species (representing nine families and five orders within the Euteleostei) that demonstrate different feeding habits and habitats. MANOVA and Tukey post hoc tests were used to assess differences among species. Most species clustered together on the principal component axes, suggesting a generalized mechanism that facilitates unidirectional flow. Typically, only one species stood out as 'extreme' on each functional axis, and a species that stood out on one axis did not stand out on others. Only one species, the flatfish Pleuronichthys verticalis, an obligate benthic feeder, demonstrated modifications consistent with enhanced suction production. This species displayed a suite of changes that should enhance suction production, including large hyoid depression, large cranial rotation, and small gape. We suggest that suction performance may be greatest in such obligate benthic feeders because cranial morphology is highly modified and prey are captured from the substrate.

Suction, Ram, and Biting: Deviations and Limitations to the Capture of Aquatic Prey.

When feeding, most aquatic organisms generate suction that draws prey into the mouth. The papers in this volume are a demonstration of this fact. However, under what circumstances is suction ineffective as a feeding mechanism? Here we consider the interplay between suction, ram, and biting, and analyze the contribution of each to the capture of prey by a wide variety of species of fish. We find, not surprisingly, that ram is the dominant contributor to feeding because suction, and biting, are only effective when very close to the prey. As species utilize more strongly ram-dominated modes of feeding, they may be released from the morphological and behavioral constraints associated with the need to direct a current of water into the head. Morphological and behavioral changes that facilitate larger gapes and stronger jaws are explored here, including predators that lack a protrusile upper jaw, predators with elongate jaws, predators that rely on suspension feeding, and predators that bite. Interestingly, while the mobility of the jaws and the shape of the opening of the mouth are modified in species that have departed from a primary reliance on suction feeding, the anterior-to-posterior wave of expansion persists. This wave may be greatly slowed in ram and biting species, but its retention suggests a fundamental importance to aquatic feeding.

The Teleost Intramandibular Joint: A mechanism That Allows Fish to Obtain Prey Unavailable to Suction Feeders.

Although the majority of teleost fishes possess a fused lower jaw (or mandible), some lineages have acquired a secondary joint in the lower jaw, termed the intramandibular joint (IMJ). The IMJ is a new module that formed within the already exceptionally complex teleost head, and disarticulation of two bony elements of the mandible potentially creates a "double-jointed" jaw. The apparent independent acquisition of this new functional module in divergent lineages raises a suite of questions. (1) How many teleostean lineages contain IMJ-bearing species? (2) Does the IMJ serve the same purpose in all teleosts? (3) Is the IMJ associated with altered feeding kinematics? (4) Do IMJ-bearing fishes experience trade-offs in other aspects of feeding performance? (5) Is the IMJ used to procure prey that are otherwise unavailable? The IMJ is probably under-reported, but has been documented in at least 10 lineages within the Teleostei. Across diverse IMJ-bearing lineages, this secondary joint in the lower jaw serves a variety of functions, including: generating dynamic out-levers that allow fish to apply additional force to a food item during jaw closing; allowing fish to "pick" individual prey items with pincer-like jaws; and facilitating contact with the substrate by altering the size and orientation of the gape. There are no consistent changes in feeding kinematics in IMJ-bearing species relative to their sister taxa; however, some IMJ-bearing taxa produce very slow movements during the capture of food, which may compromise their ability to move prey into the mouth via suction. Despite diversity in behavior, all IMJ-bearing lineages have the ability to remove foods that are physically attached to the substrate or to bite off pieces from sessile organisms. Because such prey cannot be drawn into the mouth by suction, the IMJ provides a new mechanism that enables fish to obtain food that otherwise would be unavailable.