Philopatry influences the genetic population structure of the blacktip shark (Carcharhinus limbatus) at multiple spatial scales.

Understanding how interactions among microevolutionary forces generate genetic population structure of exploited species is vital to the implementation of management policies that facilitate persistence. Philopatry displayed by many coastal shark species can impact gene flow and facilitate selection, and has direct implications for the spatial scales of management. Here, genetic structure of the blacktip shark (Carcharhinus limbatus) was examined using a mixed-marker approach employing mitochondrial control region sequences and 4339 SNP-containing loci generated using ddRAD-Seq. Genetic variation was assessed among young-of-the-year sampled in 11 sites in waters of the United States in the western North Atlantic Ocean, including the Gulf of Mexico. Spatial and environmental analyses detected 68 nuclear loci putatively under selection, enabling separate assessments of neutral and adaptive genetic structure. Both mitochondrial and neutral SNP data indicated three genetically distinct units-the Atlantic, eastern Gulf, and western Gulf-that align with regional stocks and suggest regional philopatry by males and females. Heterogeneity at loci putatively under selection, associated with temperature and salinity, was observed among sites within Gulf units, suggesting local adaptation. Furthermore, five pairs of siblings were identified in the same site across timescales corresponding with female reproductive cycles. This indicates that females re-used a site for parturition, which has the potential to facilitate the sorting of adaptive variation among neighbouring sites. The results demonstrate differential impacts of microevolutionary forces at varying spatial scales and highlight the importance of conserving essential habitats to maintain sources of adaptive variation that may buffer species against environmental change.

Escaped bait: Bull shark Carcharhinus leucas with an intentionally attached harness rig.

A 193 cm total length female bull shark Carcharhinus leucas was captured in Florida bearing intentionally attached materials which resembled a harness. Harness-type live bait rigs are commonly used for small baitfish; some anglers use such devices with small sharks when targeting large sharks and bony fish. Biofouling on the apparatus and the extent of the injuries indicated the material had likely been on the shark for several years. This case highlights the dangers of using these types of devices on juveniles of long-lived species that attain a large body size.

Complex patterns of genetic population structure in the mouthbrooding marine catfish, Bagre marinus, in the Gulf of Mexico and U.S. Atlantic.

Patterns of genetic variation reflect interactions among microevolutionary forces that vary in strength with changing demography. Here, patterns of variation within and among samples of the mouthbrooding gafftopsail catfish (Bagre marinus, Family Ariidae) captured in the U.S. Atlantic and throughout the Gulf of Mexico were analyzed using genomics to generate neutral and non-neutral SNP data sets. Because genomic resources are lacking for ariids, linkage disequilibrium network analysis was used to examine patterns of putatively adaptive variation. Finally, historical demographic parameters were estimated from site frequency spectra. The results show four differentiated groups, corresponding to the (1) U.S. Atlantic, and the (2) northeastern, (3) northwestern, and (4) southern Gulf of Mexico. The non-neutral data presented two contrasting signals of structure, one due to increases in diversity moving west to east and north to south, and another to increased heterozygosity in the Atlantic. Demographic analysis suggested that recently reduced long-term effective population size in the Atlantic is likely an important driver of patterns of genetic variation and is consistent with a known reduction in population size potentially due to an epizootic. Overall, patterns of genetic variation resemble that of other fishes that use the same estuarine habitats as nurseries, regardless of the presence/absence of a larval phase, supporting the idea that adult/juvenile behavior and habitat are important predictors of contemporary patterns of genetic structure.

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.

Sexually Monomorphic Maps and Dimorphic Responses in Rat Genital Cortex.

Mammalian external genitals show sexual dimorphism [1, 2] and can change size and shape upon sexual arousal. Genitals feature prominently in the oldest pieces of figural art [3] and phallic depictions of penises informed psychoanalytic thought about sexuality [4, 5]. Despite this longstanding interest, the neural representations of genitals are still poorly understood [6]. In somatosensory cortex specifically, many studies did not detect any cortical representation of genitals [7-9]. Studies in humans debate whether genitals are represented displaced below the foot of the cortical body map [10-12] or whether they are represented somatotopically [13-15]. We wondered what a high-resolution mapping of genital representations might tell us about the sexual differentiation of the mammalian brain. We identified genital responses in rat somatosensory cortex in a region previously assigned as arm/leg cortex. Genital responses were more common in males than in females. Despite such response dimorphism, we observed a stunning anatomical monomorphism of cortical penis and clitoris input maps revealed by cytochrome-oxidase-staining of cortical layer 4. Genital representations were somatotopic and bilaterally symmetric, and their relative size increased markedly during puberty. Size, shape, and erect posture give the cortical penis representation a phallic appearance pointing to a role in sexually aroused states. Cortical genital neurons showed unusual multi-body-part responses and sexually dimorphic receptive fields. Specifically, genital neurons were co-activated by distant body regions, which are touched during mounting in the respective sex. Genital maps indicate a deep homology of penis and clitoris representations in line with a fundamentally bi-sexual layout [16] of the vertebrate brain.

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.

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.

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.

The function of bilateral odor arrival time differences in olfactory orientation of sharks.

The direction of an odor signal source can be estimated from bilateral differences in signal intensity and/or arrival time. The best-known examples of the use of arrival time differences are in acoustic orientation. For chemoreception, animals are believed to orient by comparing bilateral odor concentration differences, turning toward higher concentrations. However, time differences should not be ignored, because odor plumes show chaotic intermittency, with the concentration variance several orders of magnitude greater than the concentration mean. We presented a small shark species, Mustelus canis, with carefully timed and measured odor pulses directly into their nares. They turned toward the side stimulated first, even with delayed pulses of higher concentration. This is the first conclusive evidence that under seminatural conditions and without training, bilateral time differences trump odor concentration differences. This response would steer the shark into an odor patch each time and thereby enhance its contact with the plume, i.e., a stream of patches. Animals with more widely spaced nares would be able to resolve smaller angles of attack at higher swimming speeds, a feature that may have contributed to the evolution of hammerhead sharks. This constitutes a novel steering algorithm for tracking odor plumes.

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.

Sharks need the lateral line to locate odor sources: rheotaxis and eddy chemotaxis.

Odor plumes are complex, dynamic, three-dimensional structures used by many animals to locate food, mates, home sites, etc. Yet odor itself has no directional properties. Animals use a variety of different senses to obtain directional information. Since most odor plumes are composed of dispersing odor patches and dissipating vorticity eddies, aquatic animals may localize odor sources by simultaneous analysis of chemical and hydrodynamic dispersal fields, a process referred to as eddy chemotaxis. This study examines the contributions of olfaction, mechanoreception and vision to odor source localization in a shark, the smooth dogfish Mustelus canis. Two parallel, turbulent plumes were created in an 8 m flume: squid rinse odor and seawater control. Minimally turbulent ;oozing' sources of odor and seawater control were physically separated from sources of major turbulence by placing a brick downstream from each oozing source, creating two turbulent wakes, one or the other flavored with food odor. This created four separate targets for the sharks to locate. Animals were tested under two light conditions (fluorescent and infrared) and in two sensory conditions (lateral line intact and lateral line lesioned by streptomycin). Intact animals demonstrated a preference for the odor plume over the seawater plume and for the source of odor/turbulence (the brick on the odor side) over the source of the odor alone (the odor-oozing nozzle). Plume and target preference and search time were not significantly affected by light condition. In the light, lesioning the lateral line increased search time but did not affect success rate or plume preference. However, lesioned animals no longer discriminated between sources of turbulent and oozing odor. In the dark, search time of lesioned animals further increased, and the few animals that located any of the targets did not discriminate between odor and seawater plumes, let alone targets. These results demonstrate for the first time that sharks require both olfactory and lateral line input for efficient and precise tracking of odor-flavored wakes and that visual input can improve food-finding performance when lateral line information is not available. We distinguish between rheotaxis: orientation to the large-scale flow field (olfaction, vision and superficial lateral line), eddy chemotaxis: tracking the trail of small-scale, odor-flavored turbulence (olfaction and lateral line canals), and pinpointing the source of the plume (lateral line canals and olfaction).