Oceanic sharks clean at coastal seamount

Interactions between pelagic thresher sharks (Alopias pelagicus) and cleaner wrasse were investigated at a seamount in the Philippines. Cleaning associations between sharks and teleosts are poorly understood, but the observable interactions seen at this site may explain why these mainly oceanic sharks regularly venture into shallow coastal waters where they are vulnerable to disturbance from human activity. From 1,230 hours of observations recorded by remote video camera between July 2005 and December 2009, 97 cleaner-thresher shark events were analyzed, 19 of which were interrupted. Observations of pelagic thresher sharks interacting with cleaners at the seamount were recorded at all times of day but their frequency declined gradually from morning until evening. Cleaners showed preferences for foraging on specific areas of a thresher shark's body. For all events combined, cleaners were observed to conduct 2,757 inspections, of which 33.9% took place on the shark's pelvis, 23.3% on the pectoral fins, 22.3% on the caudal fin, 8.6% on the body, 8.3% on the head, 2.1% on the dorsal fin, and 1.5% on the gills respectively. Cleaners did not preferentially inspect thresher sharks by time of day or by shark sex, but there was a direct correlation between the amount of time a thresher shark spent at a cleaning station and the number of inspections it received. Thresher shark clients modified their behavior by "circular-stance-swimming," presumably to facilitate cleaner inspections. The cleaner-thresher shark association reflected some of the known behavioral trends in the cleaner-reef teleost system since cleaners appeared to forage selectively on shark clients. Evidence is mounting that in addition to acting as social refuges and foraging grounds for large visiting marine predators, seamounts may also support pelagic ecology by functioning as cleaning stations for oceanic sharks and rays.     

The denticle surface of thresher shark tails: Three-dimensional structure and comparison to other pelagic species

Shark skin denticles (scales) are diverse in morphology both among species and across the body of single individuals, although the function of this diversity is poorly understood. The extremely elongate and highly flexible tail of thresher sharks provides an opportunity to characterize gradients in denticle surface characteristics along the length of the tail and assess correlations between denticle morphology and tail kinematics. We measured denticle morphology on the caudal fin of three mature and two embryo common thresher sharks (Alopias vulpinus), and we compared thresher tail denticles to those of eleven other shark species. Using surface profilometry, we quantified 3D-denticle patterning and texture along the tail of threshers (27 regions in adults, and 16 regions in embryos). We report that tails of thresher embryos have a membrane that covers the denticles and reduces surface roughness. In mature thresher tails, surfaces have an average roughness of 5.6 μm which is smoother than some other pelagic shark species, but similar in roughness to blacktip, porbeagle, and bonnethead shark tails. There is no gradient down the tail in roughness for the middle or trailing edge regions and hence no correlation with kinematic amplitude or inferred magnitude of flow separation along the tail during locomotion. Along the length of the tail there is a leading-to-trailing-edge gradient with larger leading edge denticles that lack ridges (average roughness = 9.6 μm), and smaller trailing edge denticles with 5 ridges (average roughness = 5.7 μm). Thresher shark tails have many missing denticles visible as gaps in the surface, and we present evidence that these denticles are being replaced by new denticles that emerge from the skin below.     

Application of multiplex PCR approaches for shark molecular identification: feasibility and applications for fisheries management and conservation in the Eastern Tropical Pacific

Here we describe the application of new and existing multiplex PCR methodologies for shark species molecular identification. Four multiplex systems (group ID, thresher sharks, hammerhead sharks and miscellaneous shark) were employed with primers previously described and some designed in this study, which allow for species identification after running PCR products through an agarose gel. This system was implemented for samples (bodies and fins) collected from unidentified sharks landed in the port of Buenaventura and from confiscated tissues obtained from illegal fishing around the Malpelo Island Marine Protected Area, Pacific Coast of Colombia. This method has allowed reliable identification, to date, of 407 samples to the genus and/or species levels, most of them (380) identified as the pelagic thresher shark (Alopias pelagicus). Another seven samples were identified as scalloped hammerhead sharks (Sphyrna lewini). This is an easy-to-implement and reliable identification method that could even be used locally to monitor shark captures in the main fishing ports of developed and developing countries.     

Lionfish predators use flared fin displays to initiate cooperative hunting

Despite considerable study, mystery surrounds the use of signals that initiate cooperative hunting in animals. Using a labyrinth test chamber, we examined whether a lionfish, Dendrochirus zebra, would initiate cooperative hunts with piscine partners. We found that D. zebra uses a stereotyped flared fin display to alert conspecific and heterospecific lionfish species Pterois antennata to the presence of prey. Per capita success rate was significantly higher for cooperative hunters when compared with solitary ones, with hunt responders assisting hunt initiators in cornering the prey using their large extended pectoral fins. The initiators would most often take the first strike at the group of prey, but both hunters would then alternate striking at the remaining prey. Results suggest that the cooperative communication signal may be characteristic to the lionfish family, as interspecific hunters were equally coordinated and successful as intraspecific hunters. Our findings emphasize the complexity of collaborative foraging behaviours in lionfish; the turn-taking in strikes suggests that individuals do not solely try to maximize their own hunting success: instead they equally share the resources between themselves. Communicative group hunting has enabled Pteroine fish to function as highly efficient predators.     

Comparative feeding ecology of shortfin mako, blue and thresher sharks in the California Current

This study describes the feeding ecology of three pelagic shark species in the California Current: shortfin mako (Isurus oxyrinchus); blue (Prionace glauca); and thresher (Alopias vulpinus) sharks. Stomach contents of sharks collected from 2002 to 2008 were identified to the lowest taxonomic level and analyzed using univariate and multivariate methods. Of 330 mako sharks sampled (53 to 248?cm fork length [FL]), 238 stomachs contained 42 prey taxa, with jumbo squid (Dosidicus gigas) and Pacific saury (Cololabis saira) representing the most important prey based on the geometric index of importance (GII). In addition, 158 blue sharks were sampled (76 to 248?cm FL) and 114 stomachs contained 38 prey taxa, with jumbo and Gonatus spp. squids representing the most important prey. Lastly, 225 thresher sharks were sampled (108 to 228?cm FL) and 157 stomachs contained 18 prey taxa with northern anchovy (Engraulis mordax) and Pacific sardine (Sardinops sagax) identified as the most important prey. Overall, mako sharks had the most diverse diet based upon Simpson??s diversity index (1/D) (8.43?±?1.16), feeding on many species of teleosts and cephalopods, followed by blue sharks (6.20?±?2.11) which consumed a wide range of prey (primarily cephalopods), while thresher sharks were most specialized (2.62?±?0.34), feeding primarily on coastal pelagic teleosts. Dietary overlap was lowest between blue and thresher sharks (S?rensen similarity index?=?0.321 and Simplified Morisita Horn index?=?0.006), and seasonal variability in diet was greatest for blue sharks (Simplified Morisita Horn index?=?0.260, Analysis of Similarity (ANOSIM) p?<?0.001). In addition, size class, and subregion were significant factors that affected diet of each species differently (ANOSIM p?<?0.001). Despite similarities in life history characteristics and spatial and temporal overlap in habitat, diets of these three common shark species are distinct in the California Current.     

The Species and Origin of Shark Fins in Taiwan’s Fishing Ports,Markets, and Customs Detention: A DNA Barcoding Analysis

The increasing consumption of shark products, along with the shark’s fishing vulnerabilities, has led to the decrease in certain shark populations. In this study we used a DNA barcoding method to identify the species of shark landings at fishing ports, shark fin products in retail stores, and shark fins detained by Taiwan customs. In total we identified 23, 24, and 14 species from 231 fishing landings, 316 fin products, and 113 detained shark fins, respectively. All the three sample sources were dominated by Prionace glauca, which accounted for more than 30% of the collected samples. Over 60% of the species identified in the fin products also appeared in the port landings, suggesting the domestic-dominance of shark fin products in Taiwan. However, international trade also contributes a certain proportion of the fin product markets, as four species identified from the shark fin products are not found in Taiwan’s waters, and some domestic-available species were also found in the customs-detained sample. In addition to the species identification, we also found geographical differentiation in the cox1 gene of the common thresher sharks (Alopias vulpinus), the pelagic thresher shark (A. pelagicus), the smooth hammerhead shark (Sphyrna zygaena), and the scalloped hammerhead shark (S. lewini). This result might allow fishing authorities to more effectively trace the origins as well as enforce the management and conservation of these sharks.     

Gill morphometrics of the thresher sharks (Genus Alopias): Correlation of gill dimensions with aerobic demand and environmental oxygen

Gill morphometrics of the three thresher shark species (genus Alopias) were determined to examine how metabolism and habitat correlate with respiratory specialization for increased gas exchange. Thresher sharks have large gill surface areas, short water–blood barrier distances, and thin lamellae. Their large gill areas are derived from long total filament lengths and large lamellae, a morphometric configuration documented for other active elasmobranchs (i.e., lamnid sharks, Lamnidae) that augments respiratory surface area while limiting increases in branchial resistance to ventilatory flow. The bigeye thresher, Alopias superciliosus, which can experience prolonged exposure to hypoxia during diel vertical migrations, has the largest gill surface area documented for any elasmobranch species studied to date. The pelagic thresher shark, A. pelagicus, a warm‐water epi‐pelagic species, has a gill surface area comparable to that of the common thresher shark, A. vulpinus, despite the latter's expected higher aerobic requirements associated with regional endothermy. In addition, A. vulpinus has a significantly longer water–blood barrier distance than A. pelagicus and A. superciliosus, which likely reflects its cold, well‐oxygenated habitat relative to the two other Alopias species. In fast‐swimming fishes (such as A. vulpinus and A. pelagicus) cranial streamlining may impose morphological constraints on gill size. However, such constraints may be relaxed in hypoxia‐dwelling species (such as A. superciliosus) that are likely less dependent on streamlining and can therefore accommodate larger branchial chambers and gills. J. Morphol. 276:589–600, 2015. © 2015 Wiley Periodicals, Inc.     

Ontogenetic dietary shifts and feeding behavior of the tiger shark, Galeocerdo cuvier, in Hawaiian waters

Synopsis Stomach content data from 281 tiger sharks caught during shark control programs in Hawaii between 1967 and 1969, and during 1976 were analyzed to examine feeding habits and ontogenetic shifts in diet. As sharks increased in size, prey diversity and frequency of occurrence of large prey items increased. The percent occurrence of teleosts and cephalopods in stomachs decreased as sharks increased in length, while occurrence of elasmobranchs, turtles, land mammals, crustaceans, and undigestible items increased. Comparisons between the diets of tiger sharks from Hawaii and other locations indicate that ontogenetic shifts are universal in this species and that tiger sharks may be opportunistic feeders that prey heavily on abundant, easy to capture prey. Small tiger sharks may be spatially segregated from medium and large sharks and appear to be primarily nocturnal, bottom feeders. Large tiger sharks feed near the bottom at night, but also feed at the surface during the day. Prey, similar in size to humans, begin to occur in the diet of tiger sharks approximately 230 cm TL, and therefore sharks of this size and larger may pose the greatest threat to humans. Ontogenetic shifts in diet may be attributed to increased size of sharks, expanded range and exploitation of habitats of larger sharks, and/or improved hunting skill of larger sharks.Deceased 1974     

Ventricle morphology in pelagic elasmobranch fishes

Ventricle weights of the warm-bodied great white shark, Atlantic shortfin mako, and the common thresher shark (the latter presumed to be warm-bodied) are similar to those of ectothermic blue sharks, sandbar sharks, dusky sharks, tiger sharks and scalloped hammerhead sharks. Ventricle muscularity, as estimated by the ratio of cortical to spongy layer thickness, is almost twice as great in the former three species than in the latter elasmobranchs. Measurements of ventricular volumes suggest that the ventricles of the great white, Atlantic shortfin mako and common thresher sharks are better adapted to respond to demands for increases in cardiac output via increased heartbeat frequency in comparison with ectothermic species of shark.     

Size and distribution of Pandarus satyrus (Copepoda: Pandaridae) on the blue shark Prionace glauca (Carcharhiniformes: Carcharhinidae) in Costa Rica.

A total of 80 specimens of Pandarus satyrus, a cosmopolitan ectoparasitic copepod, were taken from fishery catches of blue sharks (Prionace glauca) in the Eastern Pacific coast of Costa Rica. All specimens were found in the dorsal surface of pectoral fins (8-30 per shark). Longer specimens were most abundant.     

Body Form and Locomotion in Sharks

A revised interpretation of the mode of action of the heterocercaltail in sharks shows that the upturned tail axis tends to producea thrust directed downwards behind the centre of balance ofthe fish and thus gives a moment turning the head upwards. Thisis countered in two ways—by the rotation of the tail alongits longitudinal axis during each lateral beat, and throughthe action of the ventral hypochordal lobe. The shape of thetail and the mode of action of the tail in all sharks so farconsidered reflects a balance between these three factors, inall of them the net effect being the production of a forwardthrust from the tail that passes directly through the centreof balance of the fiish. There is normally therefore no tendencyfor the fish to turn around the centre of balance in a sagittalplane but there is a net sinking effect that is countered bythe planning effect of the pectoral fins and the ventral surfaceof the head. A study of 56 species of sharks shows that the tail is constructedaccording to a remarkably consistent common plan, the extremesbeing the high angled rather symmetrical tail of pelagic sharkssuch as hums, Lamna and Rhincodon and the straight tails ofbenthic sharks such as Ginglymostoma in which a ventral hypochordallobe is absent. When the general body shape of sharks, includingthe position of insertion of the median and paired fins andthe pattern of growth of fin surface areas is considered, theuniformity of the shark body plan and locomolor function isfurther emphasised. Four patterns of body form in sharks are recognised: 1) Thefast swimming pelagic sharks and the whale sharks have a tailwith a high aspect ratio, a conical head, a lateral fluke onthe caudal peduncle. 2) The generalised sharks typified by theCarcharhinidae, have lower heterocercal angles, a flattenedventral surface on the head and lack the caudal fluke. 3) Thedemersal sharks typified by the catsharks (Scyliorhinidae) havea very low, almost straight tail. The ventral hypochordal lobeis absent and the first dorsal fin is posterior in position.4) The squalomorph sharks are distinct in the absence of theanal fin, presence of a marked epicaudal lobe in the tail andoften an elevated insertion of the pectorals. The anal and second dorsal fins are always the smallest finsand the pectorals grow at the fastest rate. In general thereis an inverse relationship between size and rale of growth ofall fins and the ventral surface of the head. In hammerheadsthe growth data confirms that the head has a significant planingaction in swimming. The pectoral, second dorsal and anal finsshow an extreme constancy of position of insertion in all sharksstudied. The locomotor mechanism of sharks is adapted for anefficient cruising swimming but at the same time, the potentialinstability in the sagittal plan allows for the production ofturning moments that are used in attack and feeding.     

Spatial distribution of sharks captures in the Pacific of Nicaragua and its relationship with several oceanographic variables

Between August 1995 and August 1997 long line fishing techniques and a bathythermograph were used to correlate some physical variables with the spatial distribution of four shark species in 26 fishing cruises off the Nicaraguan Pacific Coast. They were the thresher (Alopias vulpinus), blue (Prionace glaucea), gray (Carcharhinus falciformis) and hammer (Sphyra lewini). All species concentrated in the southeastern Nicaraguan Pacific, at the seasonal upwelling area of Papagayo Gulf. The range of sea surface temperatures in which the sharks were captured was 25-28 degrees C. We could clearly associate this physical parameter with shark availability. The vertical distribution of the captured sharks suggests that they occupy termocline levels above the 15 degrees C isotherm. Although these species are oceanic, the blue shark was captured in ocean waters over the 1800 m isobar, while the grey and thresher sharks where close to the continental shelf. Body length in decreasing order are: thresher (210-290 cm, SN = 21), blue (60-240 cm, SN = 13) and gray (80-200 cm, SN = 17).     

Ontogenetic diet shifts and prey selection in nursery bound lemon sharks, Negaprion brevirostris, indicate a flexible foraging tactic

Ontogenetic variations in shark diet are often qualitatively inferred from dietary analysis and hindered by high levels of unidentified prey or small sample sizes. This study focused on nursery bound lemon sharks (Negaprion brevirostris, n?=?396), enabling some control over the confounding variables of prey choice associated with ontogeny. Nursery bound lemon sharks exhibited weak ontogenetic variation in dietary composition with high levels of dietary overlap. Variation in prey preference of lemon sharks with ontogeny was complex, but revealed a continuous shift from predominantly opportunistic benthic foraging as neonates to more selective piscivory with increasing shark size while in the nursery. Lemon sharks demonstrated a discrete ontogenetic shift in the number of prey consumed and stomach content weight (Kruskal-Wallis tests p?<?0.01), as well as prey size (ANOVA, p?<?0.001). All sizes of sharks exhibited positive size selection of prey (Mann?CWhitney U tests, p?<?0.01). However, the lack of size preference by all but the largest lemon sharks for their major prey (yellowfin mojarra, Gerres cinereus), suggests neonate sharks, while capable of occasionally foraging on large prey, are relatively inept opportunistic foragers. This was evident in high diet breadth, low diversity of consumed prey and lower trophic level than larger sharks. This study represents the first quantitative analysis of ontogenetic variation in prey preference and size selection in sharks, indicating a flexible foraging tactic in lemon sharks and the importance of hunting ability and predator size in prey choice.     

The functional role of the caudal fin in the feeding ecology of the common thresher shark Alopias vulpinus

This study tests the hypothesis that the common thresher shark Alopias vulpinus uses its elongate caudal fin to both produce thrust and immobilize prey during feeding. Underwater video recorded in southern California from 2007 to 2009 revealed 34 feeding events, all of which were initiated with the upper lobe of the caudal fin.     

Functional morphology of the pectoral fins in bamboo sharks, Chiloscyllium plagiosum: benthic vs. pelagic station-holding

Bamboo sharks (Chiloscyllium plagiosum) are primarily benthic and use their relatively flexible pectoral and pelvic fins to rest on and move about the substrate. We examined the morphology of the pectoral fins and investigated their locomotory function to determine if pectoral fin function during both benthic station-holding and pelagic swimming differs from fin function described previously in leopard sharks, Triakis semifasciata. We used three-dimensional kinematics and digital particle image velocimetry (DPIV) to quantify pectoral fin function in five white-spotted bamboo sharks, C. plagiosum, during four behaviors: holding station on the substrate, steady horizontal swimming, and rising and sinking during swimming. During benthic station-holding in current flow, bamboo sharks decrease body angle and adjust pectoral fin angle to shed a clockwise fluid vortex. This vortex generates negative lift more than eight times that produced during open water vertical maneuvering and also results in an upstream flow that pushes against the posterior surface of the pectoral fin to oppose drag. In contrast, there is no evidence of significant lift force in the wake of the pectoral fin during steady horizontal swimming. The pectoral fin is held concave downward and at a negative dihedral angle during steady horizontal swimming, promoting maneuverability rather than stability, although this negative dihedral angle is much less than that observed previously in sturgeon and leopard sharks. During sinking, the pectoral fins are held concave upward and shed a clockwise vortex with a negative lift force, while in rising the pectoral fin is held concave downward and sheds a counterclockwise vortex with a positive lift force. Bamboo sharks appear to sacrifice maneuverability for stability when locomoting in the water column and use their relatively flexible fins to generate strong negative lift forces when holding position on the substrate and to enhance stability when swimming in the water column.     

Spiral valve parasites of blue and common thresher sharks as indicators of shark feeding behaviour and ecology

This study documented the parasite faunas of the spiral valves of blue sharks Prionace glauca (L. 1758) and common thresher sharks Alopias vulpinus (Bonnaterre, 1788) caught in the California Current Large Marine Ecosystem (CCLME) north of the Mexican border. The spiral valves of 18 blue and 19 thresher sharks caught in the CCLME from 2009 to 2013 were examined for parasites. Seven parasite taxa were found in blue sharks and nine in threshers. The tetraphyllidean cestode Anthobothrium sp. (78% prevalence) was the most common parasite in blue sharks, and the phyllobothriid cestode Paraorygmatobothrium sp. (90% prevalence) was the most common in threshers. An adult nematode of the genus Piscicapillaria was found in threshers for the first time and may be a new species. Adult individuals of Hysterothylacium sp. were found in both shark species. The adult acanthocephalan Rhadinorhynchus cololabis and remains of the parasitic copepod Pennella sp. – both parasites of Pacific saury, Cololabis saira – were found in the intestines of threshers, indicating recent feeding on saury. This study paves the way for a more comprehensive examination, including more samples and a wider variety of shark species, to provide a greater understanding of shark feeding behaviour and possibly provide information on shark population biology.     

Internal morphology and function of paired fins in the epaulette shark,hemiscyllium ocellatum

Paired fins and associated internal structures of the epauletic sharkHemiscyllium ocellatum, were described on the basis of three specimens. A comparison with other genera showed the epaulette shark to be, characterized by two elongated basal cartilages articulating with a distally projecting articular condyle on the coracoid, a loosely separated radial series with an intermediate series, a levator pectoralis inferior muscle and an anterolaterally developed depressor pectoralis muscle in the pectoral fin, and an elongated anterior pelvic basal cartilage articulating with a distally projecting articular condyle and an anterolaterally developed depressor pelvicus muscle in the pelvic fin. In captivity, the sharks exhibited both upright and crawling behavior on the bottom by using the pectoral and pelvic fins and bending the body. The distinctive morphological characters are shared by otherHemiscyllium species and are suggested as important factors enabling their unique behavior associated with a complex coral reef habitat.     

Genetic profiling reveals illegal international trade in fins of the great white shark, <Emphasis Type="Italic">Carcharodon carcharias</Emphasis>

Great white sharks are protected by national legislation in several countries, making this species the most widely protected elasmobranch in the world. Although the market demand for shark fins in general has continued to grow, the value and extent of utilization of white shark fins in trade has been controversial. We combine law enforcement with genetic profiling to demonstrate that illegal trade in fins of this species is occurring in the contemporary international market. Furthermore, we document the presence of fins from very young white sharks in the trade, suggesting a multiple-use market (food to trophies) exists for fins of this species. The presence of small fins in the trade contradicts the view that white shark fins have market value only as large display trophies, and not as food. Our findings indicate that effective conservation of protected shark species will require international management regimes that include monitoring of the shark fishery and trade on a species-specific basis.     

Morphological scaling of body form in four shark species differing in ecology and life history

Body form can change across ontogeny, and can influence how animals of different sizes move and feed. Scaling data on live apex predatory sharks are rare and, therefore, we examined patterns of scaling in ontogenetic series of four sympatric shark species exhibiting a range of sizes, ecologies and life histories (tiger, bull, blacktip, and nurse shark). We evaluated 13 linear morphological variables and two areas (caudal and dorsal) that could influence both animal condition and locomotor performance. These measurements included dimensions of the dorsal, pectoral, and caudal fins, as well as several dimensions of body circumference, and of the head. For all four species, the body axis (eye‐to‐eye, lateral span, frontal span, proximal span) scaled close to isometry (expected slope of 1.0). The two largest sharks (tiger and bull sharks) also showed significant negative allometry for elements of the caudal fin. We found significant negative allometry in the lengths of the upper lobe of the caudal fin (caudal fin 1) and the overall height of the caudal fin (caudal fin 2) in tiger and bull sharks, with slopes ranging from about 0.60 to 0.73. Further, tiger sharks showed negative allometry in caudal fin area. These results suggest that in terms of overall body dimensions, small sharks are roughly geometrically similar to large sharks, at least within the species we examined. However, juvenile tiger (and to a lesser extent bull sharks) are notable in having proportionately larger caudal fins compared to adult sharks. As the caudal fin contributes to generating thrust during forward locomotion, this scaling implies differences among adult and juvenile sharks in locomotor ability. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 114 , 126–135.