Functional morphology of extreme jaw protrusion in Neotropical cichlids

The New World cichlids Petenia splendida and Caquetaia spp. possess extraordinarily protrusible jaws. We investigated the feeding behavior of extreme (here defined as greater than 30% head length) and modest jaw-protruding Neotropical cichlids by comparing feeding kinematics, cranial morphology, and feeding performance. Digital high-speed video (500 fps) of P. splendida, C. spectabile, and Astronotus ocellatus feeding on live guppy prey was analyzed to generate kinematic and performance variables. All three cichlid taxa utilized cranial elevation, lower jaw depression, and rotation of the suspensorium to protrude the jaws during feeding experiments. Extreme anterior jaw protrusion in P. splendida and C. spectabile resulted from augmented lower jaw depression and anterior rotation of the suspensorium. Morphological comparisons among eight cichlid species revealed novel anterior and posterior points of flexion within the suspensorium of P. splendida and Caquetaia spp. The combination of anterior and posterior loosening within the suspensorium in P. splendida and Caquetaia spp. permitted considerable anterior rotation of the suspensorium and contributed to protrusion of the jaws. Petenia splendida and C. spectabile exhibited greater ram distance and higher ram velocities than did A. ocellatus, resulting primarily from increased jaw protrusion. Petenia splendida and C. spectabile exhibited lower suction feeding performance than A. ocellatus, as indicated by lower suction-induced prey movements and velocities. Thus, extreme jaw protrusion in these cichlids may represent an adaptation for capturing elusive prey by enhancing the ram velocity of the predator but does not enhance suction feeding performance.     

Evolution and ecology of feeding in elasmobranchs

Paleozoic chondrichthyans had a large gape, numerous spike-liketeeth, limited cranial kinesis, and a non-suspensory hyoid,suggesting a feeding mechanism dominated by bite and ram. Modernsharks are characterized by a mobile upper jaw braced by a suspensoryhyoid arch that is highly kinetic. In batoids, the upper jawis dissociated from the cranium permitting extensive protrusionof the jaws. Similar to actinopterygians, the evolution of highlymobile mandibular and hyoid elements has been correlated withextensive radiation of feeding modes in elasmobranchs, particularlythat of suction. Modern elasmobranchs possess a remarkable varietyof feeding modes for a group containing so few species. Biting,suction or filter-feeding may be used in conjunction with ramto capture prey, with most species able to use a combinationof behaviors during a strike. Suction-feeding has repeatedlyarisen within all recent major elasmobranch clades and is associatedwith a suite of morphological and behavioral specializations.Prey capture in a diverse assemblage of purported suction-feedingelasmobranchs is investigated in this study. Drop in water pressuremeasured in the mouth and at the location of the prey showsthat suction inflow drops off rapidly with distance from thepredator's mouth. Elasmobranchs specializing in suction-feedingmay be limited to bottom associated prey and because of theirsmall gape may have a diet restricted to relatively small prey.Behavior can affect performance and overcome constraints imposedby the fluid medium. Suction performance can be enhanced byproximity to a substrate or by decreasing distance from predatorto prey using various morphological and/or behavioral characteristics.Benthic suction-feeders benefit by the increased strike radiusdue to deflection of water flow when feeding close to a substrate,and perhaps require less accuracy when capturing prey. Suctionand ram-suction-feeding elasmobranchs can also use suction inflowto draw prey to them from a short distance, while ram-feedingsharks must accelerate and overtake the prey. The relationshipbetween feeding strategy and ecology may depend in part on ecological,mechanistic or evolutionary specialization. Mechanistic suction-feedingspecialist elasmobranchs are primarily benthic, while most epibenthicand pelagic elasmobranchs are generalists and use ram, suction,and biting to catch a diversity of prey in various habitats.Some shark species are considered to be ecological specialistsin choosing certain kinds of prey over others. Batoids are evolutionaryspecialists in having a flattened morphology and most are generalistfeeders. Filter-feeding elasmobranchs are ecological, mechanistic,and evolutionary specialists.     

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 degrees, 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 degrees. 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.     

Evolution and mechanics of long jaws in butterflyfishes (family Chaetodontidae)

We analyzed the functional morphology and evolution of the long jaws found in several butterflyfishes. We used a conservative reanalysis of an existing morphological dataset to generate a phylogeny that guided our selection of seven short- and long-jawed taxa in which to investigate the functional anatomy of the head and jaws: Chaetodon xanthurus, Prognathodes falcifer (formerly Chaetodon falcifer), Chelmon rostratus, Heniochus acuminatus, Johnrandallia nigrirostris, Forcipiger flavissimus, and F. longirostris. We used manipulations of fresh, preserved, and cleared and stained specimens to develop mechanical diagrams of how the jaws might be protruded or depressed. Species differed based on the number of joints within the suspensorium. We used high-speed video analysis of five of the seven species (C. xanthurus, Chel. rostratus, H. acuminatus, F. flavissimus, and F. longirostris) to test our predictions based on the mechanical diagrams: two suspensorial joints should facilitate purely anteriorly directed protrusion of the lower jaw, one joint should allow less anterior protrusion and result in more depression of the lower jaw, and no joints in the suspensorium should constrain the lower jaw to simple ventral rotation around the jaw joint, as seen in generalized perciform fishes. We found that the longest-jawed species, F. longirostris, was able to protrude its jaws in a predominantly anterior direction and further than any other species. This was achieved with little input from cranial elevation, the principal input for other known lower jaw protruders, and is hypothesized to be facilitated by separate modifications to the sternohyoideus mechanism and to the adductor arcus palatini muscle. In F. longirostris the adductor arcus palatini muscle has fibers oriented anteroposteriorly rather than medial-laterally, as seen in most other perciforms and in the other butterflyfish studied. These fibers are oriented such that they could rotate the ventral portion of the quadrate anteriorly, thus projecting the lower jaw anteriorly. The intermediate species lack modification of the adductor arcus palatini and do not protrude their jaws as far (in the case of F. flavissimus) or in a purely anterior fashion (in the case of Chel. rostratus). The short-jawed species both exhibit only ventral rotation of the lower jaw, despite the fact that H. acuminatus is closely related to Forcipiger.     

Feeding behavior and kinematics of the lesser electric ray, Narcine brasiliensis (Elasmobranchii: Batoidea)

Jaw protrusion is a major functional motif in fish feeding and can occur during mouth opening or closing. This temporal variation impacts the role that jaw protrusion plays in prey apprehension and processing. The lesser electric ray Narcine brasiliensis is a benthic elasmobranch (Batoidea: Torpediniformes) with an extreme and unique method of prey capture. The feeding kinematics of this species were investigated using high-speed videography and pressure transduction. The ray captures its food by protruding its jaws up to 100% of head length (approximately 20% of disc width) beneath the substrate and generating negative oral pressures (< or = 31 kPa) to suck worms into its mouth. Food is further winnowed from ingested sediment by repeated, often asymmetrical protrusions of the jaws (> 70 degrees deviation from the midline) while sand is expelled from the spiracles, gills and mouth. The pronounced ram contribution of capture (jaw protrusion) brings the mouth close enough to the food to allow suction feeding. Due to the anatomical coupling of the jaws, upper jaw protrusion occurs in the expansive phase (unlike most elasmobranchs and similar to bony fishes), and also exhibits a biphasic (slow-open, fast-open) movement similar to tetrapod feeding. The morphological restrictions that permit this unique protrusion mechanism, including coupled jaws and a narrow gape, may increase suction performance, but also likely strongly constrain dietary breadth.     

Functional morphology of bite mechanics in the great barracuda (Sphyraena barracuda)

The great barracuda, Sphyraena barracuda, is a voracious marine predator that captures fish with a swift ram feeding strike. While aspects of its ram feeding kinematics have been examined, an unexamined aspect of their feeding strategy is the bite mechanism used to process prey. Barracuda can attack fish larger than the gape of their jaws, and in order to swallow large prey, can sever their prey into pieces with powerful jaws replete with sharp cutting teeth. Our study examines the functional morphology and biomechanics of 'ram-biting' behavior in great barracuda where the posterior portions of the oral jaws are used to slice through prey. Using fresh fish and preserved museum specimens, we examined the jaw mechanism of an ontogenetic series of barracuda ranging from 20 g to 8.2 kg. Jaw functional morphology was described from dissections of fresh specimens and bite mechanics were determined from jaw morphometrics using the software MandibLever (v3.2). High-speed video of barracuda biting (1500 framess(-1)) revealed that prey are impacted at the corner of the mouth during capture in an orthogonal position where rapid repeated bites and short lateral headshakes result in cutting the prey in two. Predicted dynamic force output of the lower jaw nearly doubles from the tip to the corner of the mouth reaching as high as 58 N in large individuals. A robust palatine bone embedded with large dagger-like teeth opposes the mandible at the rear of the jaws providing for a scissor-like bite capable of shearing through the flesh and bone of its prey.     

PLASTICITY OF JAW AND SKULL MORPHOLOGY IN THE NEOTROPICAL CICHLIDS GEOPHAGUS BRASILIENSIS AND G. STEINDACHNERI

I examined plasticity of jaw and skull morphology induced by feeding different diets in two species of the neotropical cichlid genus Geophagus. The two species possess different modes of development, which affect the size at which young begin feeding. I hypothesized that the difference in size at first feeding could lead to a difference in the amount of change inducible in the two species. The young of the substrate-spawning species, G. brasiliensis, which begin feeding at a smaller size, were predicted to be more plastic than those of the mouthbrooding species, G. steindachneri. The two diets used to induce differences were brine shrimp nauplii and chironomid larvae. Numerous measures of the jaw and skull differed significantly between groups fed the two diets but the amount of plasticity induced was small and would not present a problem for taxonomists. Contrary to my prediction, both the magnitude and pattern of plasticity induced in the two species was similar. Thus, mode of parental care and the size at which young begin feeding do not affect the degree of plasticity. Fish fed brine shrimp nauplii were longer in oral jaw region, but were shorter and shallower in the area behind the oral jaws. An additional group of G. brasiliensis was fed flake food to compare the results of this study to other studies. The differences in measures between fish fed brine shrimp diets and flake food diets were greater than those between fish fed brine shrimp and chironomid larvae. A possible role of plasticity for enhancing rather than retarding morphological evolution is discussed.     

Comparative kinematics of cypriniform premaxillary protrusion

Premaxillary protrusion has evolved multiple times within teleosts, and has been implicated as contributing to the evolutionary success of clades bearing this adaptation. Cypriniform fishes protrude the jaws via the kinethmoid, a median sesamoid bone that is a synapomorphy for the order. Using five cypriniform species, we provide the first comparative kinematic study of jaw protrusion in this speciose order. Our goals were to compare jaw protrusion in cypriniforms to that in other clades that independently evolved upper jaw protrusion, assess the variation in feeding kinematics among members of the order, and test if variation in the shape of the kinethmoid has an effect on either jaw kinematics or the degree of suction or ram used during a feeding event. We also examined the coordination in the relative timings of upper and lower jaw movements to gain insight on the cypriniform protrusile mechanism. Overall, speed of protrusion in cypriniforms is slower than in other teleosts. Protrusion speed differed significantly among cypriniforms but this is likely not due to kinethmoid shape alone; rather, it may be a result of both kinethmoid shape and branching patterns of the A1 division of the adductor mandibulae. In the benthic cypriniforms investigated here, upper jaw protrusion contributed up to 60% of overall ram of the strikes and interestingly, these species also produced the most suction. There is relatively little coordination of upper and lower jaw movements in cypriniforms, suggesting that previous hypotheses of premaxillary protrusion via lower jaw depression are not supported within Cypriniformes. Significant variation in kinematics suggests that cypriniforms may have the ability to modulate feeding, which could be an advantage if presented with the challenge of feeding on different types of prey.     

Prey capture behavior and kinematics of the Atlantic cownose ray, Rhinoptera bonasus

The structurally reinforced jaws of the cownose ray, Rhinoptera bonasus testify to this species' durophagous diet of mollusks, but seem ill-suited to the behaviors necessary for excavating such prey. This study explores this discordance by investigating the prey excavation and capture kinematics of R. bonasus. Based on the basal suction feeding mechanism in this group of fishes, we hypothesized a hydraulic method of excavation. As expected, prey capture kinematics of R. bonasus show marked differences relative to other elasmobranchs, relating to prey excavation and use of the cephalic lobes (modified anterior pectoral fin extensions unique to derived myliobatiform rays). Prey are excavated by repeated opening and closing of the jaws to fluidize surrounding sand. The food item is then enclosed laterally by the depressed cephalic lobes, which transport it toward the mouth for ingestion by inertial suction. Unlike in most sharks, upper jaw protrusion and mandibular depression are simultaneous. During food capture, the ray's spiracle, mouth, and gill slit movements are timed such that water enters only the mouth (e.g., the spiracle closes prior to prey capture and reopens immediately following). Indigestible parts are then hydraulically winnowed from edible prey portions, by mouth movements similar to those used in excavation, and ejected through the mouth. The unique sensory/manipulatory capabilities of the cephalic lobes, as well as the cownose ray's hydraulic excavation/winnowing behaviors and suction feeding, make this species an effective benthic predator, despite its epibenthic lifestyle.     

Zooplankton capture by a coral reef fish: an adaptive response to evasive prey

Synopsis High-speed cinematography and video using modified Schlieren optics and laser illumination helped elicit details of prey capture mechanisms used by Chromis viridis while feeding on calanoid copepods and Artemia. Chromis viridis is capable of a ram-jaw, low-suction feeding, as well as a typical suction feeding behavior described for other species of planktivores. By adjusting the degree of jaw protrusion and amount of suction used during a feeding strike, this fish can modulate its feeding strikes according to the prey type being encountered. The ram-jaw feeding mode enables C. viridis to capture highly evasive calanoid copepods within 6 to 10 msec. The use of specialized feeding behavior for evasive prey and the ability to vary feeding behavior are adaptations for feeding on evasive prey.     

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

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

Advances in the Study of Feeding Behaviors, Mechanisms, and Mechanics of Sharks

Sharks as a group have a long history as highly successful predatory fishes. Although, the number of recent studies on their diet, feeding behavior, feeding mechanism, and mechanics have increased, many areas still require additional investigation. Dietary studies of sharks are generally more abundant than those on feeding activity patterns, and most of the studies are confined to relatively few species, many being carcharhiniform sharks. These studies reveal that sharks are generally asynchronous opportunistic feeders on the most abundant prey item, which are primarily other fishes. Studies of natural feeding behavior are few and many observations of feeding behavior are based on anecdotal reports. To capture their prey sharks either ram, suction, bite, filter, or use a combination of these behaviors. Foraging may be solitary or aggregate, and while cooperative foraging has been hypothesized it has not been conclusively demonstrated. Studies on the anatomy of the feeding mechanism are abundant and thorough, and far exceed the number of functional studies. Many of these studies have investigated the functional role of morphological features such as the protrusible upper jaw, but only recently have we begun to interpret the mechanics of the feeding apparatus and how it affects feeding behavior. Teeth are represented in the fossil record and are readily available in extant sharks. Therefore much is known about their morphology but again functional studies are primarily theoretical and await experimental analysis. Recent mechanistic approaches to the study of prey capture have revealed that kinematic and motor patterns are conserved in many species and that the ability to modulate feeding behavior varies greatly among taxa. In addition, the relationship of jaw suspension to feeding behavior is not as clear as was once believed, and contrary to previous interpretations upper jaw protrusibility appears to be related to the morphology of the upper jaw-chondrocranial articulation rather than the type of jaw suspension. Finally, we propose a set of specific hypotheses including: (1) The functional specialization for suction feeding hypothesis that morphological and functional specialization for suction feeding has repeatedly arisen in numerous elasmobranch lineages, (2) The aquatic suction feeding functional convergence hypothesis that similar hydrodynamic constraints in bony fishes and sharks result in convergent morphological and functional specializations for suction feeding in both groups, (3) The feeding modulation hypothesis that suction capture events in sharks are more stereotyped and therefore less modulated compared to ram and bite capture events, and (4) The independence of jaw suspension and feeding behavior hypothesis whereby the traditional categorization of jaw suspension types in sharks is not a good predictor of jaw mobility and prey capture behavior. Together with a set of questions these hypotheses help to guide future research on the feeding biology of sharks.     

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.     

Laboratory constraints on feeding behaviours in polymorphic bluegill sunfish (Lepomis macrochirus)

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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.     

Mechanics of biting in great white and sandtiger sharks

Although a strong correlation between jaw mechanics and prey selection has been demonstrated in bony fishes (Osteichthyes), how jaw mechanics influence feeding performance in cartilaginous fishes (Chondrichthyes) remains unknown. Hence, tooth shape has been regarded as a primary predictor of feeding behavior in sharks. Here we apply Finite Element Analysis (FEA) to examine form and function in the jaws of two threatened shark species, the great white (Carcharodon carcharias) and the sandtiger (Carcharias taurus). These species possess characteristic tooth shapes believed to reflect dietary preferences. We show that the jaws of sandtigers and great whites are adapted for rapid closure and generation of maximum bite force, respectively, and that these functional differences are consistent with diet and dentition. Our results suggest that in both taxa, insertion of jaw adductor muscles on a central tendon functions to straighten and sustain muscle fibers to nearly orthogonal insertion angles as the mouth opens. We argue that this jaw muscle arrangement allows high bite forces to be maintained across a wider range of gape angles than observed in mammalian models. Finally, our data suggest that the jaws of sub-adult great whites are mechanically vulnerable when handling large prey. In addition to ontogenetic changes in dentition, further mineralization of the jaws may be required to effectively feed on marine mammals. Our study is the first comparative FEA of the jaws for any fish species. Results highlight the potential of FEA for testing previously intractable questions regarding feeding mechanisms in sharks and other vertebrates.     

Largemouth bass (Micropterus salmoides) switch feeding modalities in response to sensory deprivation

Animals use a suite of sensory modalities to precisely locate and capture prey. While numerous studies have examined the effects of sensory deprivation on the behaviors leading to prey capture and while it is generally believed that information in the pre-strike period determines the way fish capture prey, this study is the first to examine the contribution of sensory information to jaw kinematics during capture. Largemouth bass were filmed using high-speed videography while capturing live mosquitofish. Bass were examined intact, with visual deprivation under infrared light, and with lateral line deprivation following treatment with cobalt chloride. Deprived of visual cues, this visual ram-feeding predator switches towards suction-based feeding to successfully capture prey. They approach prey slowly but open their mouths more rapidly, which has been shown to result in greater buccal pressure, causing their prey to move a greater distance at a more rapid velocity as they are being drawn into the predators' mouths. Deprived of lateral line cues, bass have higher forward velocities during capture and capture prey earlier in the gape cycle. This study demonstrates that sensory pre-strike information directly affects the capture modality employed by fishes and that fish can modulate between ram and suction not only by adjusting the amount of ram by increasing or decreasing their movements, but also by actively increasing the amount of suction used. These results suggest that the ability to modulate feeding behavior may allow animals to not only exploit a broader breadth of prey items, but also to be capable of doing so in a wider variety of environments.     

Effects of ram speed on prey capture kinematics of juvenile Indo-Pacific tarpon,Megalops cyprinoides

We examined the effects of variation in swimming speed, or ‘ram speed’, on the feeding kinematics of juvenile Indo-Pacific tarpon, Megalops cyprinoides. Tarpon were filmed feeding on non-elusive prey at 500 images s^?1. Prey items were offered at one end of the filming tank, the opposite end where tarpon grouped, to encourage them to use a ram strategy to capture their prey. We describe tarpon as ram-suction feeders. Ram speed varied among strikes from 0.19 to 1.38 m/s and each individual produced speeds that spanned at least 0.9 m/s across trials. Although suction distances were much less variable, prey movement towards the predator was present in all feeding trials. There was a strong positive relationship between initial predator – prey distance and ram speed (r²=0.72, P<0.001). When tarpon initiated their strike from further away, they achieved higher ram speeds, but also took longer to capture their prey. All other timing variables were unaffected by ram speed whereas at higher ram speeds tarpon exhibited greater expansion of the mouth and buccal cavity. Greater buccal expansion accomplished in the same period of time implies that both the total volume of water captured and the water flow rate entering the mouth was greater in strikes at higher ram speeds. Our results demonstrate how feeding kinematics may vary as a function of ram speed, and how fish predators that lack jaw protrusion and have a large gape capacity can maximize their feeding success by altering their swimming speed.     

Feeding of the megamouth shark (Pisces: Lamniformes: Megachasmidae) predicted by its hyoid arch: A biomechanical approach

Studies of the megamouth shark, one of three planktivorous sharks, can provide information about their evolutionary history. Megamouth shark feeding has never been observed in life animals, but two alternative hypotheses on biomechanics suggest either feeding, i.e., ram feeding or suction feeding. In this study, the second moment of area of the ceratohyal cartilages, which is an indicator of the flexural stiffness of the cartilages, is calculated for 21 species of ram‐ and suction‐feeding sharks using computed tomography. The results indicate that suction‐feeding sharks have ceratohyal cartilages with a larger second moment of area than ram‐feeding sharks. The result also indicates that the ram–suction index, which is an indicator of relative contribution of ram and suction behavior, is also correlated with the second moment of area of the ceratohyal. Considering that large bending stresses are expected to be applied to the ceratohyal cartilage during suction, the larger second moment of area of the ceratohyal of suction‐feeding sharks can be interpreted as an adaptation for suction feeding. Based on the small second moment of area of the ceratohyal cartilage of the megamouth shark, the feeding mode of the megamouth shark is considered to be ram feeding, similar to the planktivorous basking shark. From these results, an evolutionary scenario of feeding mechanics of three species of planktivorous sharks can be suggested. In this scenario, the planktivorous whale shark evolved ram feeding from a benthic suction‐feeding ancestor. Ram feeding in the planktivorous megamouth shark and the basking shark evolved from ram feeding swimming‐type ancestors and that both developed their unique filtering system to capture small‐sized prey. J. Morphol., 2011. © 2011 Wiley‐Liss, Inc.     

Trophic apparatus in cyprinodontiform fishes: Functional specializations for picking and scraping behaviors

Cyprinodontiforms are a diverse and speciose order that includes topminnows, pupfishes, swordtails, mosquitofishes, guppies, and mollies. Sister group to the Beloniformes and Atheriniformes, Cyprinodontiformes contains approximately twice the number of species of these other two orders combined. Recent studies suggest that this group is well suited to capturing prey by “picking” small items from the water surface, water column, and the substrate. Because picking places unusual performance demands on the feeding apparatus, this mode of prey capture may rely upon novel morphological modifications not found in more widespread ram‐ or suction‐based feeding mechanisms. To assess this evolutionary hypothesis, we describe the trophic anatomy of 16 cyprinodontiform species, selected to broadly represent the order as well as capture intrageneric variation. The group appears to have undergone gradual morphological changes to become increasingly specialized for picking and scraping behaviors. We also identify a suite of functional characters related to the acquisition of a novel and previously undescribed mechanism of premaxillary protrusion and retraction, including: modification of the “premaxillomandibular” ligament (which connects each side of the premaxilla to the ipsilateral mandible, or lower jaw), a novel architecture of the ligaments and bony elements that unite the premaxillae, maxillae and palatine bones, and novel insertions of the adductor muscles onto the jaws. These morphological changes to both the upper and lower jaws suggest an evolutionary trend within this group toward increased reliance on picking individual prey from the water column/substrate or for scraping encrusting material from the substrate. We propose that the suite of morphological characters described here enable a functional innovation, “picking,” which leads to novel trophic habits. J. Morphol., 2009. © 2008 Wiley‐Liss, Inc.