Food habits and ontogenetic changes in the diet of the sandbar shark, Carcharhinus plumbeus, in Hawaii

The sandbar shark, Carcharhinus plumbeus, is a wide-ranging coastal species in tropical and temperate regions, and it is the most common species of shark in Hawaii, as in many locations where it occurs. Information on the diet and feeding habits of this species in the Pacific Ocean are extremely limited. For this study we quantified the diet of sandbar sharks in Hawaii based on records collected during the Hawaii Cooperative Shark Research and Control Program from 1967 to 1969. During this program a total of 565 stomachs were examined, of which 265 contained food. Sharks ranged in size from 59 to 190 cm total length. Teleosts were the most common prey group, but both cephalopods and crustaceans also occurred frequently. Ontogenetic changes in diet of sandbar sharks were apparent, with crustaceans forming a greater proportion of the diet of smaller sharks. Both cephalopods and elasmobranchs increased in importance with increasing shark size. Prey diversity also increased with size, with large, mobile, and reef prey species found more commonly in the diet of larger sharks. Mature male and female sharks appeared to segregate by depth, though major differences in the diet between the sexes were not apparent. However, there was some evidence of dietary differences between sharks caught in different depths and seasons. The results of this study suggest that sandbar sharks in Hawaii and throughout the world, are primarily piscivores, but also consume a variety of invertebrate prey, and that their diet varies with geographical location and stage of development.     

Gastric evacuation in the sandbar shark, Carcharhinus plumbeus

Sandbar sharks maintained in an enclosure in the natural environment were fed meals of soft blue crab, Callinectes sapidus , or menhaden, Brevoortia lyrannus , and were sacrificed at intervals after feeding to measure rates of gastric emptying. Regression analysis was used to evaluate the adequacy of various models in describing the decrease in stomach contents with time after feeding. A Gompertz growth curve provided the best fit to the data for both food types. This model suggests that gastric emptying is characterized by an initial lag phase during which evacuation rate increases, a period of maximal evacuation, and a decreasing evacuation rate during the later stages of digestion. Passage of food from the stomach was slow relative to other species of fish studied. The time required to evacuate 98% of a meal was 92–3 and 70–7 h for sharks fed menhaden and blue crab respectively. Evacuation rates did not differ between day and night periods.     

Teeth penetration force of the tiger shark Galeocerdo cuvier and sandbar shark Carcharhinus plumbeus

This study examined the minimum force required of functional teeth and replacement teeth in the tiger shark Galeocerdo cuvier and the sandbar shark Carcharhinus plumbeus to penetrate the scales and muscle of sheepshead Archosargus probatocephalus and pigfish Orthopristis chrysoptera. Penetration force ranged from 7·7–41·9 and 3·2–26·3 N to penetrate A. probatocephalus and O. chrysoptera, respectively. Replacement teeth required significantly less force to penetrate O. chrysoptera for both shark species, most probably due to microscopic wear of the tooth surfaces supporting the theory shark teeth are replaced regularly to ensure sharp teeth that are efficient for prey capture.     

Isolation and characterization of five dinucleotide microsatellite loci in the sandbar shark,Carcharhinus plumbeus

Five dinucleotide markers were isolated and optimized from a microsatellite‐enriched genomic library obtained from the sandbar shark, Carcharhinus plumbeus. Genotypic distributions of all markers were found to be in conformance with the expectations of Hardy–Weinberg equilibrium with four to 39 alleles present per locus. We amplified these loci in two female sharks and their litters. A maternal allele was recovered at each locus in all progeny indicating reliable amplification. More than two paternal alleles were recovered across both litters indicating genetic polyandry. Additionally, these markers were amplified across 10 carcharhiniform species to examine their utility in other studies.     

Clearance of a dermal Huffmanela sp. in a sandbar shark (Carcharhinus plumbeus) using levamisole

A wild-caught captive sandbar shark Carcharhinus plumbeus developed a contiguous network of darkly pigmented linear tracks that progressed from the snout to the ventral cervical region. Microscopic examination of a skin scraping revealed nematode eggs of the genus Huffmanela, a group of histozoic nematodes that is known to parasitize requiem sharks and marine and freshwater teleosts. The fresh eggs were darkly pigmented with bipolar plugs, contained a larva, and measured 73.3 to 86.4 by 39.0 to 47.4 microm (n = 10). Formalin-fixed and paraffin-embedded eggs were significantly smaller (Wilcoxon rank sums test, p < 0.005), measuring 70.5 to 78.9 by 33.6 to 41.3 microm (n = 13). These measurements do not correlate with previously reported species of Huffmanela. Serial treatment with levamisole (10 mg kg(-1), intramuscular [i.m.]) cleared the egg tracks within 21 d, with no recurrence or apparent complications.     

Age and growth of the sandbar shark, Carcharhinus plumbeus, in Hawaiian waters through vertebral analysis

Age and growth estimates were determined for the sandbar shark, Carcharhinus plumbeus, from Oahu, Hawaii in the central Pacific Ocean. Age estimates were obtained through vertebral centra analysis of 187 sharks. We verified our age estimates through marginal increment analysis of centra and oxytetracycline marking methods of at liberty sandbar sharks. Sizes of sampled sharks ranged from 46 to 147 cm pre-caudal length. Four growth models were fitted to length-at-age data; two forms of the von Bertalanffy growth model, the Gompertz growth model, and a logistic growth model. Males and females exhibited statistically significant differences in growth, indicating that females grow slower and attain larger sizes than males. Growth parameter estimates revealed slower growth rates than previously estimated (based on captive specimens) for Hawaiian sandbar sharks. The von Bertalanffy growth model using empirical length-at-birth provided the best biological and statistical fit to the data. This model gave parameter estimates of L _∞ = 138.5 cm PCL and k = 0.12 year⁻¹ for males and L _∞ = 152.8 cm PCL, k = 0.10 year⁻¹ for females. Male and female sandbar sharks mature at approximately 8 and 10 years of age, respectively.     

World phylogeography and male‐mediated gene flow in the sandbar shark,Carcharhinus plumbeus

The sandbar shark, Carcharhinus plumbeus, is a large, cosmopolitan, coastal species. Females are thought to show philopatry to nursery grounds while males potentially migrate long distances, creating an opportunity for male‐mediated gene flow that may lead to discordance in patterns revealed by mitochondrial DNA (mtDNA) and nuclear markers. While this dynamic has been investigated in elasmobranchs over small spatial scales, it has not been examined at a global level. We examined patterns of historical phylogeography and contemporary gene flow by genotyping 329 individuals from nine locations throughout the species’ range at eight nuclear microsatellite markers and sequencing the complete mtDNA control region. Pairwise comparisons often resulted in fixation indices and divergence estimates of greater magnitude using mtDNA sequence data than microsatellite data. In addition, multiple methods of estimation suggested fewer populations based on microsatellite loci than on mtDNA sequence data. Coalescent analyses suggest divergence and restricted migration among Hawaii, Taiwan, eastern and western Australia using mtDNA sequence data and no divergence and high migration rates, between Taiwan and both Australian sites using microsatellite data. Evidence of secondary contact was detected between several localities and appears to be discreet in time rather than continuous. Collectively, these data suggest complex spatial/temporal relationships between shark populations that may feature pulses of female dispersal and more continuous male‐mediated gene flow.     

Huffmanela cf. carcharhini (Nematoda: Trichosomoididae: Huffmanelinae) from skin of a sandbar shark, Carcharhinus plumbeus, in the Pacific Ocean

Eggs of Huffmanela cf. carcharhini from the skin of an aquarium-held, juvenile sandbar shark, Carcharhinus plumbeus , from the Pacific Ocean were studied using light and scanning electron microscopy. Grossly, eggs imparted a scribble-like skin marking approximately 130 × 60 mm on the right side of the shark's snout adjacent to its eye and nostril. Fresh (unfixed) eggs were elliptical, 75-95 μm long (xˉ = 85 μm, SD = ±4.5; n = 75), 48-63 μm wide (53 ± 3.4; 75), 8-10 μm in shell thickness (9 ± 1.3; 27), 45-68 μm in vitelline mass length (52 ± 6.9; 8); had a smooth shell surface and nonprotruding polar plugs 8-13 μm wide (10 ± 1.5; 73); lacked thin filaments, superficial envelope, and shell spines; sank in 35 ppt artificial seawater; and did not spontaneously hatch after 12 hr in 35 ppt artificial seawater. Formalin-fixed eggs measured 193 days postfixation were 75-95 μm long (84 ± 3.9; 150), 45-60 μm wide (50 ± 2.2; 150), 5-10 μm in shell thickness (8 ± 1.2; 87), 45-60 μm in vitelline mass length (51 ± 3.0; 92), and 30-40 μm in vitelline mass width (33 ± 2.0; 84), and had nonprotruding polar plugs that were 10-15 μm long (11 ± 1.4; 93) and 8-10 μm wide (9 ± 1.1; 108). Forcibly hatched first-stage larvae (unfixed) were filiform, 188-273 μm long (212 ± 25.5; 13), 8-13 μm wide (10 ± 1.2; 13), and had fine transverse striations. Eggs infected the epidermis only. Histology revealed intra-epithelial inflammation with eosinophilic granulocytes and hyperplasia, plus dermal lymphofollicular hyperplasia associated with the infection. The eggs of H. cf. carcharhini likely undergo considerable ex utero development before being sloughed (unhatched) from the host, along with epidermal cells.     

Genetic polyandry and sexual conflict in the sandbar shark, Carcharhinus plumbeus, in the western North Atlantic and Gulf of Mexico

To investigate patterns of polyandry in the sandbar shark (Carcharhinus plumbeus), 20 pregnant females were sampled from the western North Atlantic and Gulf of Mexico. Five species-specific microsatellite markers were used to genotype each shark and its litter. Of 20 litters, 17 (85%) were shown to have multiple sires. In multiply sired litters, the estimated minimum number of sires ranged from two to five with an average of 2.3 males per litter. Regression analysis did not demonstrate a significant relationship between female reproductive success and female body size or sire number and female body size. There was a high incidence of reproductive skew noted in litters, and two groups of males with significantly different mean reproductive success were observed. Analyses using Bateman's principles suggest that there is less direct benefit for females that acquire multiple mates than for males who bias paternity within litters. In light of past morphological and behavioural studies, these data suggest that patterns of polyandry in elasmobranchs may be determined by coercive mating, and that breeding behaviour has likely evolved in the context of sexual conflict.     

Effective size closely approximates the census size in the heavily exploited western Atlantic population of the sandbar shark, Carcharhinus plumbeus

The sandbar shark, Carcharhinus plumbeus, is a long-lived species with low lifetime fecundity that is heavily fished in the western North Atlantic. Inshore nursery grounds increase survivorship of sandbar shark pups and the principal nurseries are in the mid-Atlantic region. We calculated effective number of breeders (N_b) and effective population size (N_e) for adults utilizing the nursery grounds of the Delaware Bay and the Eastern Shore of Virginia by genotyping 902 animals across five cohorts at eight microsatellite loci. Estimates of N_b and N_e were compared to estimates of census size (N_c) of cohorts obtained from Delaware Bay. The estimated N_e/N_c and N_b/N_c ratios were 0.45 or higher whether the Delaware Bay cohorts were considered as distinct year classes or combined. This is in contrast to estimated N_e/N_c ratios in other exploited marine fishes, which are several orders of magnitude smaller. Instead, the N_e/N_c ratio of sandbar sharks is similar to that found in marine and terrestrial mammals.     

Ontogenetic changes in the diet of the sandbar shark, <Emphasis Type="Italic">Carcharhinus plumbeus</Emphasis>, in lower Chesapeake Bay and Virginia (USA) coastal waters

This study describes the diet of the sandbar shark, Carcharhinus plumbeus, highlighting differences in diet within various regions of the Virginia (USA) nursery area, as well as ontogenetic changes in diet. Stomach samples were obtained in 2001 and 2002 from 232 sharks caught by gillnets or longlines. Historical data from the Virginia Institute of Marine Science (VIMS) Shark Ecology Program were also analyzed. Ontogenetic changes in diet were evident, with crustacean prey decreasing in frequency with increasing shark size, and elasmobranch prey importance increasing with increasing shark size. Whereas previous research in Chincoteague Bay, VA showed the blue crab, Callinectes sapidus, was the dominant crustacean in sandbar shark diet, the mantis shrimp, Squilla empusa, dominated the crustacean portion of the diet in this study. Differences in diet of sharks were observed among locations within the study area. Small juveniles (≤80 cm precaudal length) in the lower Chesapeake Bay ate more fishes, whereas Eastern Shore juveniles ate more crustaceans. Crustacean prey items varied among locations along the Eastern Shore, with more portunid crabs consumed in waters near Wachapreague and more mantis shrimp consumed near Sand Shoal Inlet. Our study showed that Carcharhinus plumbeus is a generalist predator and is thus unlikely to strongly impact the population of any particular prey species, and in turn is not likely to be strongly affected by fluctuations in abundance of a single prey species.     

Isolation,purification and characterization of pepsin soluble collagen isolated from silvertip shark (Carcharhinus albimarginatus) skeletal and head bone

Type II pepsin soluble collagens (PSC) were isolated from skeletal and head bone of silvertip shark; and examined for their biochemical and structural properties. Among the raw materials, the protein content (8.99%) was high in skeletal bone and the ash content (28%) was high in head bone. After the collagen extraction, the raw materials contained higher amount of ash content ranging from 82 to 88%. The hydroxyproline content of skeletal and skeletal PSC (30 and 113 mg/g) was higher than those head and head PSC. Both collagens were composed of two different α-chains (α₁- and α₂-chains) and were characterized as type II collagen. Amino acid analysis of skeletal and head PSC indicated imino acid contents of 156 and 175 amino acid residues per 1000 residues, respectively. Similar, Fourier transform infrared spectra of SCII and HCII were observed, which suggested that the isolation process did not affect the secondary structure and molecular order of collagen, particularly the triple–helical structure. Denaturation temperature of skeletal PSC (31 °C) was higher than that of head PSC. SEM microstructure of the collagens depicted a porous, fibrillary and multi-layered structure. These results suggested that the PSC isolated from skeletal and head bone of silvertip shark were found to be suitable biomaterial in commercial applications as alternatives to mammalian collagen.     

Biology of the Galapagos shark,Carcharhinus galapagensis, in Hawai'i

Synopsis Catch records from the Hawai'i Cooperative Shark Research and Control Program, which operated in Hawai'i from 1967–1969, were examined and data on the Galapagos shark,Carcharhinus galapagensis were analyzed. A total of 304 Galapagos sharks was caught, predominantly with longlines. More female sharks were caught than males, and the catch was skewed geographically. On the island of O'ahu the highest catch rates occurred along the north and south coasts. High catch rates also occurred near points of land, where longshore currents converge. Average depth of capture was greater for juveniles (45.1 m) and mature males (60.2 m), than for subadults (38.8 m) and mature female sharks (34.2 m). Males appear to reach maturity between 205 and 239 cm total length, and females between 215 and 245 cm. Litter size ranged from 4 to 16 pups, with an average of 8.7. In Hawaiian waters Galapagos sharks are born at just over 80 cm total length. Mating and parturition apparently occur early in the year, and gestation is estimated to be about 12 months. Stomach contents consisted mainly of teleosts and benthic prey, and ontogenetic changes in diet occurred as sharks increased in size. Sharks consumed a smaller proportion of teleosts and more elasmobranchs with increasing size. Dietary diversity also increased with increasing size of shark.     

The biology of the finetooth shark,Carcharhinus isodon

Synopsis The finetooth shark inhabits shallow coastal waters of the western Atlantic from North Carolina to Brazil. It is common off the southeastern United States, where it spends the summer off Georgia and the Carolinas and winters off Florida. The species appears in the nursery and mating areas of South Carolina when the surface water temperature rises above 20° C in late April and early May. Both adults and juveniles are common in the shallow coastal waters of South Carolina through the summer, where they feed primarily on menhaden. The finetooth shark leaves the Carolinas in early fall and migrates southward as the surface water temperature decreases below 20° C. Females reach maturity at about 1350 mm TL. Males mature at about 1300 mm TL. The finetooth shark has consecutive, year-long ovarian and gestation cycles, like most carcharhinid sharks. Mating occurs from early May to early June. Freshly mated females bear a large spermozeugma at the base of each uterus. The spermozeugmata are large almond shaped masses of individual spermatozoa embedded in a supporting matrix. Embryos are lecithotrophic during their first fifteen weeks of development. Subsequently, the embryos establish a placental connection to the mother. Implantation occurs when the embryos measure about 130 mm or at about the fifteenth week of gestation. Gravid females carrying young 480–550 mm TL enter the shallow water nurseries off South Carolina in late May. Parturition occurs from late May to mid-June, after a gestation period of about twelve months, plus or minus two weeks. The young measure 480–580 mm TL at birth. Oocytes grow little during the gestation cycle. After parturition, a cohort of oocytes begins to develop, that will be ovulated the following May. Thus, the ovarian cycle lasts about a year, although most of the oocyte growth occurs in the months just prior to ovulation.     

Bioenergetics of the bull shark,Carcharhinus leucas,maintained in captivity

Understanding the energy requirements for captive sharks is important for their successful long-term maintenance. This information is critical in assessing the health of the animals and the suitability of their environment. We studied five bull sharks (Carcharhinus leucas) for up to 7 years in a 2.5 × 10⁶ liter oceanarium. Individual animal feedings provided information for food intake analysis. During the first 3 years, fork length increase was estimated to have averaged 1.9 cm/month (s.d. = 0.1), or 23.0 cm/year. Biannual measurements, begun in the fourth year, showed that growth rates decreased during the next 4 years to a mean rate of 0.6 cm/month (s.d. = 0.2), or about 7.0 cm/year. Mean food consumption from June 1988 to December 1992 was 3.4% body weight per week. Caloric conversion of weights incorporated into a simple bioenergetics model providing mean metabolic expenditures per animal was 5.7 (s.d. = 0.3) and 4 (s.d. = 0.5) kcal/kg/day for 1991 and 1992, respectively. © 1994 Wiley-Liss, Inc.     

New medium for solubilizing proteins of the eye lens core (nucleus)

The eye lens core (nucleus) is a practical source of proteins for use as genetic markers. A new, simple extraction solution for these proteins is described. The solution was tested on fish (mullet, rockfish, and barracuda) lens nuclei, which produced weak extracts with other extraction media. A new solution, in addition to producing the strongest extracts, demonstrated desirable qualities relating to electrophoresis and storage. This solution may serve as a general extraction solution for nuclear lens proteins.