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.     

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.     

Huffmanela selachii n. sp. (Nematoda: Trichosomoididae: Huffmanelinae): A new species infecting the skin of the great hammerhead shark (Sphyrna mokarran) and the blacktip reef shark (Carcharhinus melanopterus) in the Arabian Gulf,off-shore Saudi Arabia

From January 2017 - December 2019, 75 out of 850 (8.8 %) great hammerhead sharks from the Arabian Gulf had skin lesions of black irregular discolorations on the ventral surface of the head. The lesions consisted of pencil-like lineations often advancing forward by about 2 mm in back-and-forth looped scribbles often forming a relatively linear bands of about 5–7 cm wide. Similar lesions were also found in the blacktip reef shark from the same area within the same period, and consisted of straight to irregular black lines, extended indiscriminately across the skin of the sharks. Microscopic examination of the skin revealed the presence of dark-brown eggs exhibiting the spindle or ellipsoidal eggs characteristic of Huffmanela sp. The morphometrics of eggs from both hosts were similar (62.9–89.9 μm long and 29.3–56.1 μm wide). The eggshells were smooth with polar plugs protruding or not, with an abruptly truncated crown-like or shoulder-like collar surrounding the plug. The eggs were only found in the epidermal layer of the skin. Based on the unique morphometrics of the eggs, we report a new species, named: Huffmanela selachii n. sp.. This appears to be the first report of Huffmanela from either the great hammerhead shark or the blacktip reef shark, and the third reported Huffmanela in sharks from the Arabian Gulf. It is also one of few species reported from connecting waters of the greater Indian Ocean. This new finding contributes to our understanding of the diversity and ubiquity of Huffmanela sp. in marine creatures.     

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.     

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.     

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.     

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.     

Huffmanela mexicana n. sp. (Nematoda: Trichosomoididae) from the marine fish Sphoeroides annulatus in Mexico

A new species of trichosomoidid nematodes, Huffmanela mexicana n. sp., is established on the basis of its egg morphology and biological characters. The dark-shelled, embryonated eggs of this histozoic parasite occur in masses in the inner layer of the swimbladder wall of the bullseye puffer fish Sphoeroides annulatus Jenyns, 1843 (Tetraodontidae, Tetraodontiformes) off the Pacific coast of Mexico (Sinaloa State) and are characterized mainly by their shape and size (63-69 x 30-33 microm), an aspinose surface, relatively small polar plugs, and their thick egg wall (4-5 microm). A key to Huffmanela species based on egg morphology is presented.     

<Emphasis Type="Italic">Huffmanela lata</Emphasis> n. sp. (Nematoda: Trichosomoididae: Huffmanelinae) from the shark <Emphasis Type="Italic">Carcharhinus amblyrhynchos</Emphasis> (Elasmobranchii: Carcharhinidae) off New Caledonia

Huffmanela lata n. sp. is described from eggs only, which were found in a small black spot on the skin near the gill opening of a shark, Carcharhinus amblyrhynchos, caught off Nouméa, New Caledonia, South-West Pacific. The eggs are 77–88 (mean 84) μm in length and 52–63 (mean 57) μm in width, with a thick (6–8 μm) shell, apparently spinose. Mobile larvae, 200–250 μm in length, were visible in the eggs. The species is distinguished from other members of the genus by the dimensions and shape of its eggs, which are the broadest ever reported. This is the second species of Huffmanela described from a shark, after H. carcharhini (MacCallum, 1925), to which it appears closely related in terms of its site in the skin and the nature of the egg surface.     

Purification of a novel heterodimer from shark (Carcharhinus plumbeus) serum by gel-immobilized metal chromatography.

1. Resolution of the fraction of sandbar shark (Carcharhinus plumbeus) serum that was soluble in 50% saturated ammonium sulfate by gel-immobilized metal-affinity chromatography allowed the isolation of a novel disulfide-bonded heterodimer of intact mass 70 kDa. 2. Following reduction, the molecule could be resolved into two chains of apparent mass 36 and 24 kDa. 3. The molecules were glycoproteins as determined by an observed reduction in molecular weight following enzymatic glycosylation. 4. The two separate chains were related to one another on the basis of amino-acid composition analysis and by comparison of the N-terminal amino acids (seven out of 10 identities). 5. The exact relationship of this molecule to characterized heterodimers of higher vertebrates is unknown. 6. Cross-linked agarose-acetate was synthesized and proved to be an efficient concentrating agent and also a hydrophobic interaction adsorbant.     

A new Huffmanela species (Nematoda: Trichosomoididae) parasitizing atherinid fishes in north Patagonian Gulfs, Argentina

Huffmanela moraveci n. sp. (Nematode: Trichosomoididae) is described from the skin of fins and gills of the silversides Odontesthes smitti Lahille, 1949, and Odontesthes nigricans Richardson, 1848 (Pisces: Atheriniformes), from southwestern Atlantic coast, on the basis of the morphology of adults and eggs found in the host tissues. Huffmanela moraveci n. sp. can be distinguished from Huffmanela carcharini, H. banningi, H. japonica, H. mexicana, H. paronai, H. schouteni, H. shikokuensis, H. branchialis, H. filamentosa, H. ossicola, and H. lata by the size and shape of the eggs. The adult stage of H. moraveci n. sp. can be differentiated from H. hufmani and H. canadensis by the position of the nerve ring and by the total esophagus length/body length ratio. Huffmanela moraveci n. sp. also differs from H. huffmani by the presence of bacillary bands and by the body length. This is the first report of a species of Huffmanela in South America, the first report of a species of Huffmanela with Odontesthes spp. as host, and the third known adult form in the genus.     

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.     

First record of Huffmanela schouteni (Nematoda: Trichosomoididae), a histozoic parasite of flyingfishes, in Europe

Examination of 3 specimens of the Mediterranean flyingfish Cheilopogon heterurus (Rafinesque) from the Ligurian Sea, Italy, revealed the presence of numerous, dark-shelled trichinelloid eggs (69 to 75 x 30 to 33 microm) inside the innermost layer of the swimbladder, mostly located in clumps arranged in a distinct pattern. These were identified as Huffmanela schouteni Moravec et Campbell, 1991, a species known so far as eggs in flyingfishes and in the stools of humans in Cura?ao. This is the first finding of this parasite in Europe and C. heterurus represents a new host record.     

Histological and morphological aspects of reproduction in the sandbar shark Carcharhinus plumbeus in the U.S. south‐eastern Atlantic Ocean and Gulf of Mexico

The reproduction of the sandbar shark Carcharhinus plumbeus in the U.S. south‐eastern Atlantic Ocean including the Gulf of Mexico was examined using a combination of histological and morphological characteristics of C. plumbeus collected through fishery‐dependent and ‐independent sampling programmes (n = 1,567). Indices of maturity were constructed using measurements of gonads, reproductive tracts and claspers, and sandbar sharks exhibited 50% maturity sizes of 140 and 148 cm fork length for males and females respectively. Gonado‐somatic indices and variation in reproductive tract condition were used to determine seasonal trends in reproduction of mature C. plumbeus. Sandbar sharks have discrete seasonal reproductive cycles in which males produce sperm from January to May with a peak in May and females develop eggs from January to May with ovulation occurring in June. Females were shown to exhibit a >2 year reproductive cycle. Embryonic development was assessed through measurements of masses and lengths of uterine contents. Gestation was 12 months, from July to the following June, with parturition in late June. This research highlights a difference from previously reported data on the periodicity of female reproduction in C. plumbeus in the U.S. south‐eastern Atlantic Ocean and Gulf of Mexico, which may have major effects on future C. plumbeus stock management.     

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.     

Three new species of Huffmanela Moravec, 1987 (Nematoda: Trichosomoididae) from the gills of marine fish off New Caledonia

Three new species of Huffmanela, here described from eggs only, are reported from the gills of marine fish caught off Nouméa, New Caledonia. Eggs of Huffmanela branchialis n. sp., from Nemipterus furcosus (Nemipteridae), are 45-52 (mean 48) microm in length and 23-30 (mean 25) microm in width, with thin shells. Each egg is enclosed in a thin membrane forming a spindle-shaped envelope 53-85 (mean 63) microm in length. Eggs of H. filamentosa n. sp., from Gymnocranius grandoculis (Lethrinidae), are 48-53 (mean 50) microm in length and 25-30 (mean 27) microm in width, with thin shells. Each egg bears a few long (150 microm), thin filaments. Eggs of these two new species were compared to those of H. paronai Moravec & Garibaldi, 2000, which are redescribed. Eggs of H. ossicola n. sp. were found within the branchial arch bone of Bodianus loxozonus (Labridae) and also filled the spinal chord bone and other bones. This is the first species of Huffmanela reported from bone tissue. Eggs are large, 72-88 (mean 79) microm in length and 32-40 (mean 36) microm in width, with a very thick shell. Each egg is covered with numerous filaments enclosed in a thin envelope. Fresh eggs were unembryonated, but embryos were visible after incubation in seawater. The three new species can be distinguished from other species of Huffmanela by size and the nature of the egg covering. Egg morphology of and their location in the host suggest different life-cycles: those of the first two species (small eggs, thin shells, egg covering possibly favouring flotation) are released from the gill mucosa with the turnover of living tissues and immediately continue their life-cycle, but eggs of H. ossicola (large eggs, thick shell) are only available for the continuation of the life-cycle after the host's death.     

progressive ontogenetic changes in the soluble nuclear eye lens proteins of the sandbar shark,Carcharhinus milberti (Valenciennes)

• 1.1. Progressive ontogenetic changes in soluble nuclear eye lens proteins was demonstrated for the first time in a large species of shark.
• 2.2. Comparison of fetal and post-natal sharks showed the shift in proteins to be gradual rather than statically related to life history events such as birth and sexual maturation.
• 3.3. Studies considering the use of soluble nuclear eye lens proteins as a means of shark stock identification should at their onset consider ontogenetic changes as a possible source of protein variation.

     

Aggregation behavior of the grey reef shark, Carcharhinus amblyrhynchos, at Johnston Atoll, Central Pacific Ocean

Free-ranging female grey reef sharks, Carcharhinus amblyrhynchos, were observed forming daily aggregations offshore Sand Island, Johnston Atoll between March and late May from 1992 to 1995. Daily water temperatures were recorded at the aggregation area from 1993 to 1995. The annual aggregation cycle did not coincide with maximum or minimum annual water temperatures. During the 1994 aggregation, temperatures were recorded at least every hour at 7 sites in the Atoll. The sharks aggregated most frequently and in highest numbers at the largest and shallowest site, which also contained the fewest underwater structures. The water temperature at this site was 1–2 °C warmer than at neighboring sites and at a reef channel between the lagoon and the open ocean. The pattern of movement of sharks to and from the aggregation area was correlated with daily fluctuations of water temperature (r=0.38; p < 0.001), tide (r=0.30; p < 0.001) and light level (r=0.21; p < 0.001) after adjustment for autocorrelated errors. The daily maximum number of aggregating sharks coincided with the daily maximum water temperature. However, the number of aggregating sharks did not necessarily increase during days of higher water temperature. During the 1994 aggregation period, four sharks were fed ultrasonic telemetry tags, and telemetry stations were deployed at three sites within the aggregation area. Individual sharks were tracked returning to the aggregation area for durations of one to five days. The end of detection of tagged sharks is thought to be due to the regurgitation of the indigestible tags, rather than the sharks permanent departure from the aggregating area. Sharks aggregated only during the day; none of the tagged sharks were recorded and no sharks were ever seen at the aggregation area during night time. This revised version was published online in July 2006 with corrections to the Cover Date.