Global collision-risk hotspots of marine traffic and the world's largest fish, the whale shark.

Marine traffic is increasing globally yet collisions with endangered megafauna such as whales, sea turtles, and planktivorous sharks go largely undetected or unreported. Collisions leading to mortality can have population-level consequences for endangered species. Hence, identifying simultaneous space use of megafauna and shipping throughout ranges may reveal as-yet-unknown spatial targets requiring conservation. However, global studies tracking megafauna and shipping occurrences are lacking. Here we combine satellite-tracked movements of the whale shark, Rhincodon typus, and vessel activity to show that 92% of sharks' horizontal space use and nearly 50% of vertical space use overlap with persistent large vessel (>300 gross tons) traffic. Collision-risk estimates correlated with reported whale shark mortality from ship strikes, indicating higher mortality in areas with greatest overlap. Hotspots of potential collision risk were evident in all major oceans, predominantly from overlap with cargo and tanker vessels, and were concentrated in gulf regions, where dense traffic co-occurred with seasonal shark movements. Nearly a third of whale shark hotspots overlapped with the highest collision-risk areas, with the last known locations of tracked sharks coinciding with busier shipping routes more often than expected. Depth-recording tags provided evidence for sinking, likely dead, whale sharks, suggesting substantial "cryptic" lethal ship strikes are possible, which could explain why whale shark population declines continue despite international protection and low fishing-induced mortality. Mitigation measures to reduce ship-strike risk should be considered to conserve this species and other ocean giants that are likely experiencing similar impacts from growing global vessel traffic.

Microbiome structure in large pelagic sharks with distinct feeding ecologies.

BACKGROUND: Sharks play essential roles in ocean food webs and human culture, but also face population declines worldwide due to human activity. The relationship between sharks and the microbes on and in the shark body is unclear, despite research on other animals showing the microbiome as intertwined with host physiology, immunity, and ecology. Research on shark-microbe interactions faces the significant challenge of sampling the largest and most elusive shark species. We leveraged a unique sampling infrastructure to compare the microbiomes of two apex predators, the white (Carcharodon carcharias) and tiger shark (Galeocerdo cuvier), to those of the filter-feeding whale shark (Rhincodon typus), allowing us to explore the effects of feeding mode on intestinal microbiome diversity and metabolic function, and environmental exposure on the diversity of microbes external to the body (on the skin, gill). RESULTS: The fecal microbiomes of white and whale sharks were highly similar in taxonomic and gene category composition despite differences in host feeding mode and diet. Fecal microbiomes from these species were also taxon-poor compared to those of many other vertebrates and were more similar to those of predatory teleost fishes and toothed whales than to those of filter-feeding baleen whales. In contrast, microbiomes of external body niches were taxon-rich and significantly influenced by diversity in the water column microbiome. CONCLUSIONS: These results suggest complex roles for host identity, diet, and environmental exposure in structuring the shark microbiome and identify a small, but conserved, number of intestinal microbial taxa as potential contributors to shark physiology.

Elasmobranch microbiomes: emerging patterns and implications for host health and ecology.

Elasmobranchs (sharks, skates and rays) are of broad ecological, economic, and societal value. These globally important fishes are experiencing sharp population declines as a result of human activity in the oceans. Research to understand elasmobranch ecology and conservation is critical and has now begun to explore the role of body-associated microbiomes in shaping elasmobranch health. Here, we review the burgeoning efforts to understand elasmobranch microbiomes, highlighting microbiome variation among gastrointestinal, oral, skin, and blood-associated niches. We identify major bacterial lineages in the microbiome, challenges to the field, key unanswered questions, and avenues for future work. We argue for prioritizing research to determine how microbiomes interact mechanistically with the unique physiology of elasmobranchs, potentially identifying roles in host immunity, disease, nutrition, and waste processing. Understanding elasmobranch-microbiome interactions is critical for predicting how sharks and rays respond to a changing ocean and for managing healthy populations in managed care.

Connecting post-release mortality to the physiological stress response of large coastal sharks in a commercial longline fishery.

Bycatch mortality is a major factor contributing to shark population declines. Post-release mortality (PRM) is particularly difficult to quantify, limiting the accuracy of stock assessments. We paired blood-stress physiology with animal-borne accelerometers to quantify PRM rates of sharks caught in a commercial bottom longline fishery. Blood was sampled from the same individuals that were tagged, providing direct correlation between stress physiology and animal fate for sandbar (Carcharhinus plumbeus, N = 130), blacktip (C. limbatus, N = 105), tiger (Galeocerdo cuvier, N = 52), spinner (C. brevipinna, N = 14), and bull sharks (C. leucas, N = 14). PRM rates ranged from 2% and 3% PRM in tiger and sandbar sharks to 42% and 71% PRM in blacktip and spinner sharks, respectively. Decision trees based on blood values predicted mortality with >67% accuracy in blacktip and spinner sharks, and >99% accuracy in sandbar sharks. Ninety percent of PRM occurred within 5 h after release and 59% within 2 h. Blood physiology indicated that PRM was primarily associated with acidosis and increases in plasma potassium levels. Total fishing mortality reached 62% for blacktip and 89% for spinner sharks, which may be under-estimates given that some soak times were shortened to focus on PRM. Our findings suggest that no-take regulations may be beneficial for sandbar, tiger, and bull sharks, but less effective for more susceptible species such as blacktip and spinner sharks.

First insights into the vertical habitat use of the whitespotted eagle ray Aetobatus narinari revealed by pop-up satellite archival tags.

The whitespotted eagle ray Aetobatus narinari is a tropical to warm-temperate benthopelagic batoid that ranges widely throughout the western Atlantic Ocean. Despite conservation concerns for the species, its vertical habitat use and diving behaviour remain unknown. Patterns and drivers in the depth distribution of A. narinari were investigated at two separate locations, the western North Atlantic (Islands of Bermuda) and the eastern Gulf of Mexico (Sarasota, Florida, U.S.A.). Between 2010 and 2014, seven pop-up satellite archival tags were attached to A. narinari using three methods: a through-tail suture, an external tail-band and through-wing attachment. Retention time ranged from 0 to 180 days, with tags attached via the through-tail method retained longest. Tagged rays spent the majority of time (82.85 ± 12.17% S.D.) within the upper 10 m of the water column and, with one exception, no rays travelled deeper than ~26 m. One Bermuda ray recorded a maximum depth of 50.5 m, suggesting that these animals make excursions off the fore-reef slope of the Bermuda Platform. Individuals occupied deeper depths (7.42 ± 3.99 m S.D.) during the day versus night (4.90 ± 2.89 m S.D.), which may be explained by foraging and/or predator avoidance. Each individual experienced a significant difference in depth and temperature distributions over the diel cycle. There was evidence that mean hourly depth was best described by location and individual variation using a generalized additive mixed model approach. This is the first study to compare depth distributions of A. narinari from different locations and describe the thermal habitat for this species. Our study highlights the importance of region in describing A. narinari depth use, which may be relevant when developing management plans, whilst demonstrating that diel patterns appear to hold across individuals.

Correlations of metabolic rate and body acceleration in three species of coastal sharks under contrasting temperature regimes.

The ability to produce estimates of the metabolic rate of free-ranging animals is fundamental to the study of their ecology. However, measuring the energy expenditure of animals in the field has proved difficult, especially for aquatic taxa. Accelerometry presents a means of translating metabolic rates measured in the laboratory to individuals studied in the field, pending appropriate laboratory calibrations. Such calibrations have only been performed on a few fish species to date, and only one where the effects of temperature were accounted for. Here, we present calibrations between activity, measured as overall dynamic body acceleration (ODBA), and metabolic rate, measured through respirometry, for nurse sharks (Ginglymostoma cirratum), lemon sharks (Negaprion brevirostris) and blacktip sharks (Carcharhinus limbatus). Calibrations were made at a range of volitional swimming speeds and experimental temperatures. Linear mixed models were used to determine a predictive equation for metabolic rate based on measured ODBA values, with the optimal model using ODBA in combination with activity state and temperature to predict metabolic rate in lemon and nurse sharks, and ODBA and temperature to predict metabolic rate in blacktip sharks. This study lays the groundwork for calculating the metabolic rate of these species in the wild using acceleration data.

Modulation of shark prey capture kinematics in response to sensory deprivation.

The ability of predators to modulate prey capture in response to the size, location, and behavior of prey is critical to successful feeding on a variety of prey types. Modulating in response to changes in sensory information may be critical to successful foraging in a variety of environments. Three shark species with different feeding morphologies and behaviors were filmed using high-speed videography while capturing live prey: the ram-feeding blacktip shark, the ram-biting bonnethead, and the suction-feeding nurse shark. Sharks were examined intact and after sensory information was blocked (olfaction, vision, mechanoreception, and electroreception, alone and in combination), to elucidate the contribution of the senses to the kinematics of prey capture. In response to sensory deprivation, the blacktip shark demonstrated the greatest amount of modulation, followed by the nurse shark. In the absence of olfaction, blacktip sharks open the jaws slowly, suggestive of less motivation. Without lateral line cues, blacktip sharks capture prey from greater horizontal angles using increased ram. When visual cues are absent, blacktip sharks elevate the head earlier and to a greater degree, allowing them to overcome imprecise position of the prey relative to the mouth, and capture prey using decreased ram, while suction remains unchanged. When visual cues are absent, nurse sharks open the mouth wider, extend the labial cartilages further, and increase suction while simultaneously decreasing ram. Unlike some bony fish, neither species switches feeding modalities (i.e. from ram to suction or vice versa). Bonnetheads failed to open the mouth when electrosensory cues were blocked, but otherwise little to no modulation was found in this species. These results suggest that prey capture may be less plastic in elasmobranchs than in bony fishes, possibly due to anatomical differences, and that the ability to modulate feeding kinematics in response to available sensory information varies by species, rather than by feeding modality.

Vertical Movements and Patterns in Diving Behavior of Whale Sharks as Revealed by Pop-Up Satellite Tags in the Eastern Gulf of Mexico.

The whale shark (Rhincodon typus) is a wide-ranging, filter-feeding species typically observed at or near the surface. This shark's sub-surface habits and behaviors have only begun to be revealed in recent years through the use of archival and satellite tagging technology. We attached pop-up satellite archival transmitting tags to 35 whale sharks in the southeastern Gulf of Mexico off the Yucatan Peninsula from 2003-2012 and three tags to whale sharks in the northeastern Gulf off Florida in 2010, to examine these sharks' long-term movement patterns and gain insight into the underlying factors influencing their vertical habitat selection. Archived data were received from 31 tags deployed on sharks of both sexes with total lengths of 5.5-9 m. Nine of these tags were physically recovered facilitating a detailed long-term view into the sharks' vertical movements. Whale sharks feeding inshore on fish eggs off the northeast Yucatan Peninsula demonstrated reverse diel vertical migration, with extended periods of surface swimming beginning at sunrise followed by an abrupt change in the mid-afternoon to regular vertical oscillations, a pattern that continued overnight. When in oceanic waters, sharks spent about 95% of their time within epipelagic depths (<200 m) but regularly undertook very deep ("extreme") dives (>500 m) that largely occurred during daytime or twilight hours (max. depth recorded 1,928 m), had V-shaped depth-time profiles, and comprised more rapid descents (0.68 m sec-1) than ascents (0.50 m sec-1). Nearly half of these extreme dives had descent profiles with brief but conspicuous changes in vertical direction at a mean depth of 475 m. We hypothesize these stutter steps represent foraging events within the deep scattering layer, however, the extreme dives may have additional functions. Overall, our results demonstrate complex and dynamic patterns of habitat utilization for R. typus that appear to be in response to changing biotic and abiotic conditions influencing the distribution and abundance of their prey.

Smells Like Home: The Role of Olfactory Cues in the Homing Behavior of Blacktip Sharks, Carcharhinus limbatus.

Animal navigation in the marine environment is believed to be guided by different sensory cues over different spatial scales. Geomagnetic cues are thought to guide long-range navigation, while visual or olfactory cues allow animals to pinpoint precise locations, but the complete behavioral sequence is not yet understood. Terra Ceia Bay is a primary nursery area for blacktip sharks, Carcharhinus limbatus, on southwestern Florida's Gulf of Mexico coast. Young-of-the-year animals show strong fidelity to a specific home range in the northeastern end of the bay and rapidly return when displaced. Older juveniles demonstrate annual philopatry for the first few years, migrating as far south as the Florida Keys each fall, then returning to Terra Ceia Bay each spring. To examine the sensory cues used in homing, we captured neonate (<3 weeks old) blacktip sharks from within their home range, fitted them with acoustic tags, and translocated them to sites 8 km away in adjacent Tampa Bay and released them. Intact animals returned to their home range, within 34 h on average, and remained there. With olfaction blocked, fewer animals returned to their home range and they took longer to do so, 130 h on average. However, they did not remain there but instead moved throughout Terra Ceia Bay and in and out of Tampa Bay. Since sharks from both treatments returned at night in tannic and turbid water, vision is likely not playing a major role in navigation by these animals. The animals in this study also returned on incoming or slack tides, suggesting that sharks, like many other fish, may use selective tidal stream transport to conserve energy and aid navigation during migration. Collectively, these results suggest that while other cues, possibly geomagnetic and/or tidal information, might guide sharks over long distances, olfactory cues are required for recognizing their specific home range.

Population Structure and Seasonal Migration of the Spotted Eagle Ray, Aetobatus narinari.

Few studies have reported on the fine-scale population genetics of batoid species in the Atlantic basin. Here, we investigate the genetic diversity and population structure of the spotted eagle ray, Aetobatus narinari, sampled in the northeastern and southwestern parts of the Gulf of Mexico and in the northwestern Caribbean Sea. Samples were collected from 286 individuals sampled across 3 geographic localities. Estimates of divergence based on the mitochondrial cytochrome b gene and 10 nuclear microsatellite loci reveal weak but significant genetic structure among A. narinari populations in this region. Analysis of molecular variance estimates based on both marker types indicate significant differentiation between Florida and Mexico populations, while comparisons with Cuba suggest high levels of gene flow with rays from both Mexico and Florida. Conflicting results were found from the different marker types when sexes were analyzed separately underscoring the importance of applying multiple marker types when making inferences about population structure and sex-biased dispersal. Results from Bayesian clustering analyses suggest rays may be migrating south out of the Gulf of Mexico and into the northwestern Caribbean Sea. Given the impacts of fisheries on this species, coupled with the lack of population genetic data available, these findings offer valuable information to aid with conservation management strategies.

Constructional morphology within the head of hammerhead sharks (sphyrnidae).

The study of functional trade-offs is important if a structure, such as the cranium, serves multiple biological roles, and is, therefore, shaped by multiple selective pressures. The sphyrnid cephalofoil presents an excellent model for investigating potential trade-offs among sensory, neural, and feeding structures. In this study, hammerhead shark species were chosen to represent differences in head form through phylogeny. A combination of surface-based geometric morphometrics, computed tomography (CT) volumetric analysis, and phylogenetic analyses were utilized to investigate potential trade-offs within the head. Hammerhead sharks display a diversity of cranial morphologies where the position of the eyes and nares vary among species, with only minor changes in shape, position, and volume of the feeding apparatus through phylogeny. The basal winghead shark, Eusphyra blochii, has small anteriorly positioned eyes. Through phylogeny, the relative size and position of the eyes change, such that derived species have larger, more medially positioned eyes. The lateral position of the external nares is highly variable, showing no phylogenetic trend. Mouth size and position are conserved, remaining relatively unchanged. Volumetric CT analyses reveal no trade-offs between the feeding apparatus and the remaining cranial structures. The few trade-offs were isolated to the nasal capsule volume's inverse correlation with braincase, chondrocranial, and total cephalofoil volume. Eye volume also decreased as cephalofoil width increased. These data indicate that despite considerable changes in head shape, much of the head is morphologically conserved through sphyrnid phylogeny, particularly the jaw cartilages and their associated feeding muscles, with shape change and morphological trade-offs being primarily confined to the lateral wings of the cephalofoil and their associated sensory structures.

There and back again: a review of residency and return migrations in sharks, with implications for population structure and management.

The overexploitation of sharks has become a global environmental issue in need of a comprehensive and multifaceted management response. Tracking studies are beginning to elucidate how shark movements shape the internal dynamics and structure of populations, which determine the most appropriate scale of these management efforts. Tracked sharks frequently either remain in a restricted geographic area for an extended period of time (residency) or return to a previously resided-in area after making long-distance movements (site fidelity). Genetic studies have shown that some individuals of certain species preferentially return to their exact birthplaces (natal philopatry) or birth regions (regional philopatry) for either parturition or mating, even though they make long-distance movements that would allow them to breed elsewhere. More than 80 peer-reviewed articles, constituting the majority of published shark tracking and population genetic studies, provide evidence of at least one of these behaviors in a combined 31 shark species from six of the eight extant orders. Residency, site fidelity, and philopatry can alone or in combination structure many coastal shark populations on finer geographic scales than expected based on their potential for dispersal. This information should therefore be used to scale and inform assessment, management, and conservation activities intended to restore depleted shark populations.

Movable shark scales act as a passive dynamic micro-roughness to control flow separation.

Shark scales on fast-swimming sharks have been shown to be movable to angles in excess of 50°, and we hypothesize that this characteristic gives this shark skin a preferred flow direction. During the onset of separation, flow reversal is initiated close to the surface. However, the movable scales would be actuated by the reversed flow thereby causing a greater resistance to any further flow reversal and this mechanism would disrupt the process leading to eventual flow separation. Here we report for the first time experimental evidence of the separation control capability of real shark skin through water tunnel testing. Using skin samples from a shortfin mako Isurus oxyrinchus, we tested a pectoral fin and flank skin attached to a NACA 4412 hydrofoil and separation control was observed in the presence of movable shark scales under certain conditions in both cases. We hypothesize that the scales provide a passive, flow-actuated mechanism acting as a dynamic micro-roughness to control flow separation.

Multisensory integration and behavioral plasticity in sharks from different ecological niches.

The underwater sensory world and the sensory systems of aquatic animals have become better understood in recent decades, but typically have been studied one sense at a time. A comprehensive analysis of multisensory interactions during complex behavioral tasks has remained a subject of discussion without experimental evidence. We set out to generate a general model of multisensory information extraction by aquatic animals. For our model we chose to analyze the hierarchical, integrative, and sometimes alternate use of various sensory systems during the feeding sequence in three species of sharks that differ in sensory anatomy and behavioral ecology. By blocking senses in different combinations, we show that when some of their normal sensory cues were unavailable, sharks were often still capable of successfully detecting, tracking and capturing prey by switching to alternate sensory modalities. While there were significant species differences, odor was generally the first signal detected, leading to upstream swimming and wake tracking. Closer to the prey, as more sensory cues became available, the preferred sensory modalities varied among species, with vision, hydrodynamic imaging, electroreception, and touch being important for orienting to, striking at, and capturing the prey. Experimental deprivation of senses showed how sharks exploit the many signals that comprise their sensory world, each sense coming into play as they provide more accurate information during the behavioral sequence of hunting. The results may be applicable to aquatic hunting in general and, with appropriate modification, to other types of animal behavior.

A physiological analysis of color vision in batoid elasmobranchs.

The potential for color vision in elasmobranchs has been studied in detail; however, a high degree of variation exists among the group. Evidence for ultraviolet (UV) vision is lacking, despite the presence of UV vision in every other vertebrate class. An integrative physiological approach was used to investigate color and ultraviolet vision in cownose rays and yellow stingrays, two batoids that inhabit different spectral environments. Both species had peaks in UV, short, medium, and long wavelength spectral regions in dark-, light-, and chromatic-adapted electroretinograms. Although no UV cones were found with microspectrophotometric analysis, both rays had multiple cone visual pigments with λ max at 470 and 551 nm in cownose rays (Rhinoptera bonasus) and 475, 533, and 562 nm in yellow stingrays (Urobatis jamaicensis). The same analysis demonstrated that both species had rod λ max at 500 and 499 nm, respectively. The lens and cornea of cownose rays maximally transmitted wavelengths greater than 350 nm and greater than 376 nm in yellow stingrays. These results support the potential for color vision in these species and future investigations should reveal the extent to which color discrimination is significant in a behavioral context.

Horizontal movements, migration patterns, and population structure of whale sharks in the Gulf of Mexico and northwestern Caribbean sea.

Whale sharks, Rhincodon typus, aggregate by the hundreds in a summer feeding area off the northeastern Yucatan Peninsula, Mexico, where the Gulf of Mexico meets the Caribbean Sea. The aggregation remains in the nutrient-rich waters off Isla Holbox, Isla Contoy and Isla Mujeres, Quintana Roo for several months in the summer and then dissipates between August and October. Little has been known about where these sharks come from or migrate to after they disperse. From 2003-2012, we used conventional visual tags, photo-identification, and satellite tags to characterize the basic population structure and large-scale horizontal movements of whale sharks that come to this feeding area off Mexico. The aggregation comprised sharks ranging 2.5-10.0 m in total length and included juveniles, subadults, and adults of both sexes, with a male-biased sex ratio (72%). Individual sharks remained in the area for an estimated mean duration of 24-33 days with maximum residency up to about 6 months as determined by photo-identification. After leaving the feeding area the sharks showed horizontal movements in multiple directions throughout the Gulf of Mexico basin, the northwestern Caribbean Sea, and the Straits of Florida. Returns of individual sharks to the Quintana Roo feeding area in subsequent years were common, with some animals returning for six consecutive years. One female shark with an estimated total length of 7.5 m moved at least 7,213 km in 150 days, traveling through the northern Caribbean Sea and across the equator to the South Atlantic Ocean where her satellite tag popped up near the Mid-Atlantic Ridge. We hypothesize this journey to the open waters of the Mid-Atlantic was for reproductive purposes but alternative explanations are considered. The broad movements of whale sharks across multiple political boundaries corroborates genetics data supporting gene flow between geographically distinct areas and underscores the need for management and conservation strategies for this species on a global scale.

Feeding anatomy, filter-feeding rate, and diet of whale sharks Rhincodon typus during surface ram filter feeding off the Yucatan Peninsula, Mexico.

The feeding anatomy, behavior and diet of the whale shark Rhincodon typus were studied off Cabo Catoche, Yucatan Peninsula, Mexico. The filtering apparatus is composed of 20 unique filtering pads that completely occlude the pharyngeal cavity. A reticulated mesh lies on the proximal surface of the pads, with openings averaging 1.2mm in diameter. Superficial to this, a series of primary and secondary cartilaginous vanes support the pads and direct the water across the primary gill filaments. During surface ram filter feeding, sharks swam at an average velocity of 1.1m/s with 85% of the open mouth below the water's surface. Sharks on average spent approximately 7.5h/day feeding at the surface on dense plankton dominated by sergestids, calanoid copepods, chaetognaths and fish larvae. Based on calculated flow speed and underwater mouth area, it was estimated that a whale shark of 443 cm total length (TL) filters 326 m(3)/h, and a 622 cm TL shark 614 m(3)/h. With an average plankton biomass of 4.5 g/m(3) at the feeding site, the two sizes of sharks on average would ingest 1467 and 2763 g of plankton per hour, and their daily ration would be approximately 14,931 and 28,121 kJ, respectively. These values are consistent with independently derived feeding rations of captive, growing whale sharks in an aquarium. A feeding mechanism utilizing cross-flow filtration of plankton is described, allowing the sharks to ingest plankton that is smaller than the mesh while reducing clogging of the filtering apparatus.

Temporal resolution and spectral sensitivity of the visual system of three coastal shark species from different light environments.

Visual temporal resolution and scotopic spectral sensitivity of three coastal shark species (bonnethead Sphyrna tiburo, scalloped hammerhead Sphyrna lewini, and blacknose shark Carcharhinus acronotus) were investigated by electroretinogram. Temporal resolution was quantified under photopic and scotopic conditions using response waveform dynamics and maximum critical flicker-fusion frequency (CFF). Photopic CFF(max) was significantly higher than scotopic CFF(max) in all species. The bonnethead had the shortest photoreceptor response latency time (23.5 ms) and the highest CFF(max) (31 Hz), suggesting that its eyes are adapted for a bright photic environment. In contrast, the blacknose had the longest response latency time (34.8 ms) and lowest CFF(max) (16 Hz), indicating its eyes are adapted for a dimmer environment or nocturnal lifestyle. Scotopic spectral sensitivity revealed maximum peaks (480 nm) in the bonnethead and blacknose sharks that correlated with environmental spectra measured during twilight, which is a biologically relevant period of heightened predation.

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.

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.