Selachohemecus benzi n. sp. (Digenea: Sanguinicolidae) from the blacktip shark Carcharhinus limbatus (Carcharhinidae) in the northern Gulf of Mexico

Selachohemecus benzi Bullard & Overstreet n. sp. infects the heart and kidney of the blacktip shark Carcharhinus limbatus in the northern Gulf of Mexico off Florida and Mississippi, USA. Specimens of S. olsoni Short, 1954, the only congener and only other named blood fluke reported from a chondrichthyan in the Gulf of Mexico, were collected from the heart of the Atlantic sharpnose shark Rhizoprionodon terraenovae from two new localities, Apalachicola Bay, Florida, and Mississippi Sound, Mississippi, USA. The new species differs from S. olsoni by having a larger body (1.4-3.8 mm long), robust tegumental body spines numbering 51-63 along each lateral body margin, a testis extending from the posterior caeca to the ovary, and a medial ovary with lobes. We amend the diagnosis of Selachohemecus Short, 1954 to accommodate it and provide a diagnostic key for all named chondrichthyan blood flukes.     

Scaling of bite force in the blacktip shark Carcharhinus limbatus

Although bite force is a frequently studied performance measure of feeding ecology, changes in bite force over ontogeny have rarely been investigated. Biting by the blacktip shark Carcharhinus limbatus was theoretically modeled over ontogeny to investigate the scaling of bite force, the morphological basis of the observed scaling relationship, the ecological consequences of ontogenetic changes in performance, and whether cranial morphometrics can be used as an accurate proxy for bite force. Theoretical bite force, which was positively allometric with respect to total length (TL), ranged from 32 N (61 cm TL) to 423 N (152 cm TL) at the anterior tips of the jaws and from 107 (61 cm TL) to 1083 N (152 cm TL) at the posterior teeth. This observation is attributed to positive allometry in the mechanical advantage of the jaw-adducting mechanism and the cross-sectional area of all four jaw-adducting muscles. Theoretical bite force was accurately predicted by cranial morphometrics including prebranchial length and head width as well. Although positive allometry of bite force in C. limbatus would seem to indicate an ecological necessity for this phenomenon, dietary analyses do not necessarily indicate any ontogenetic shift in prey types requiring larger bite forces. The positively allometric increase in theoretical bite force may be associated with numerous other selective pressures including maintenance of an apical position within the ecosystem.     

Dionchus postoncomiracidia (Monogenea: Dionchidae) from the skin of blacktip sharks, Carcharhinus limbatus (Carcharhinidae)

Postoncomiracidia of Dionchus sp. are described from specimens collected from the skin of 2 blacktip sharks Carcharhinus limbatus, captured in the northern Gulf of Mexico. The parasites resemble nonciliated oncomiracidia hatched from eggs laid by Dionchus sp. on gills of a cobia Rachycentron canadum and adults of Dionchus remorae that were collected from gills of a common sharksucker Echeneis naucrates, captured in association with a third blacktip shark. The hamuli of the postoncomiracidia were morphologically similar to those of adult D. remorae. This is the first report of dionchids from an elasmobranch and from a location other than the gills. These findings support the idea that some dionchid oncomiracidia colonize the skin of sharks or other aquatic vertebrates that sponsor remoras, prior to transferring to other remoras and maturing.     

New insights into the trophic ecology of blacktip sharks (Carcharhinus limbatus) from a subtropical estuary in the western Gulf of Mexico

As environmental change persists, understanding resource use patterns is of value to predict the consequences of shifting trophic structures. While many sharks are opportunistic predators, some exhibit prey selectivity, putting them at higher risk compared to species with greater trophic plasticity. In the Gulf of Mexico (GOM), Clupeids and Sciaenids comprise 69% of blacktip shark (Carcharhinus limbatus) diets, which is consequential considering potential responses of these prey groups to disturbance and over harvesting. We assessed if blacktips exhibit selectivity for Clupeids and Sciaenids in the western GOM based on stomach contents from sharks in coastal Texas. Clupeids comprised <2% of diets, while striped mullet (Mugil cephalus) and red drum (Sciaenops ocellatus) comprised >70% of identifiable prey. Ontogenetic shifts from smaller (Clupeids, small Sciaenids) to larger, higher trophic level (Ariidae, Elasmobranchii) prey fits our understanding of foraging among coastal sharks, and suggests our regional understanding of blacktip trophic ecology may be limited by the sizes of sampled sharks. Observed increases in blacktip densities coupled with declines in prey (Mugilids, Sciaenids) is concerning if blacktips have limited diet plasticity. Yet GOM blacktips may be more generalized than previously thought, which is promising for conservation and management.     

Parthenogenesis in a large‐bodied requiem shark,the blacktip Carcharhinus limbatus

Genetic evidence is provided for parthenogenesis in a large‐bodied shark, the blacktip Carcharhinus limbatus, from the speciose and commercially important family Carcharhinidae, the first verified case of asexual development in this lineage and only the second for any chondrichthyan. The parthenogenetic embryo exhibited elevated homozygosity relative to its mother, indicating that automictic parthenogenesis is the most likely mechanism. Although this finding shows that parthenogenesis is more common and widespread in sharks than previously realized and supports the early existence of parthenogenetic abilities in vertebrates, the adaptive significance of automixis in these ancient fishes remains unclear.     

Microsatellite and mitochondrial DNA analyses of the genetic structure of blacktip shark (Carcharhinus limbatus) nurseries in the northwestern Atlantic, Gulf of Mexico, and Caribbean Sea

Abstract We investigated the genetic structure of blacktip shark (Carcharhinus limbatus) continental nurseries in the northwestern Atlantic Ocean, Gulf of Mexico, and Caribbean Sea using mitochondrial DNA control region sequences and eight nuclear microsatellite loci scored in neonate and young-of-the-year sharks. Significant structure was detected with both markers among nine nurseries (mitochondrial PhiST = 0.350, P < 0.001; nuclear PhiST = 0.007, P < 0.001) and sharks from the northwestern Atlantic, eastern Gulf of Mexico, western Gulf of Mexico, northern Yucatan, and Belize possessed significantly different mitochondrial DNA haplotype frequencies. Microsatellite differentiation was limited to comparisons involving northern Yucatan and Belize sharks with nuclear genetic homogeneity throughout the eastern Gulf of Mexico, western Gulf of Mexico, and northwestern Atlantic. Differences in the magnitude of maternal vs. biparental genetic differentiation support female philopatry to northwestern Atlantic, Gulf of Mexico, and Caribbean Sea natal nursery regions with higher levels of male-mediated gene flow. Philopatry has produced multiple reproductive stocks of this commercially important shark species throughout the range of this study.     

First record of dicephalia in a bull shark Carcharhinus leucas (Chondrichthyes: Carcharhinidae) foetus from the Gulf of Mexico,U.S.A.

The first recorded incidence of dicephalia in a bull shark Carcharhinus leucas is reported from a foetus collected by a fisherman in the Gulf of Mexico near Florida, U.S.A. External examination, Radiography and magnetic resonance imaging revealed a case of monosomic dicephalia where the axial skeleton and internal organs were found to divide into parallel systems anterior to the pectoral girdle resulting in two well‐developed heads.     

Scale morphology and flexibility in the shortfin mako Isurus oxyrinchus and the blacktip shark Carcharhinus limbatus

We quantified placoid scale morphology and flexibility in the shortfin mako Isurus oxyrinchus and the blacktip shark Carcharhinus limbatus. The shortfin mako shark has shorter scales than the blacktip shark. The majority of the shortfin mako shark scales have three longitudinal riblets with narrow spacing and shallow grooves. In comparison, the blacktip shark scales have five to seven longitudinal riblets with wider spacing and deeper grooves. Manual manipulation of the scales at 16 regions on the body and fins revealed a range of scale flexibility, from regions of nonerectable scales such as on the leading edge of the fins to highly erectable scales along the flank of the shortfin mako shark body. The flank scales of the shortfin mako shark can be erected to a greater angle than the flank scales of the blacktip shark. The shortfin mako shark has a region of highly flexible scales on the lateral flank that can be erected to at least 50°. The scales of the two species are anchored in the stratum laxum of the dermis. The attachment fibers of the scales in both species appear to be almost exclusively collagen, with elastin fibers visible in the stratum laxum of both species. The most erectable scales of the shortfin mako shark have long crowns and relatively short bases that are wider than long. The combination of a long crown length to short base length facilitates pivoting of the scales. Erection of flank scales and resulting drag reduction is hypothesized to be passively driven by localized flow patterns over the skin. J. Morphol. 2012. © 2012 Wiley Periodicals, Inc.     

Inclusion of South American samples reveals new population structuring of the blacktip shark (Carcharhinus limbatus) in the western Atlantic

Carcharhinus limbatus has a cosmopolitan distribution and marked genetic structuring, mainly because of its philopatric behavior. However, analysis of this structuring has not previously included South American populations. In the present study, we analyzed a sample of adult individuals collected on the northern coast of Brazil and compared the sequences of the mitochondrial control region with those of populations already genotyped. Relatively high haplotype diversity (12 haplotypes, genetic diversity of 0.796) was observed, similar to that in other populations but with a much larger number of private alleles. In contrast to populations studied previously, which were represented by neonates, the pronounced allelic variability found in the South American individuals may have resulted from migrations from other populations in the region that have yet to be genotyped. This population was also genetically distinct from the other Atlantic populations (F_st > 0.8), probably because of female philopatry, and apparently separated from the northwestern Atlantic group 1.39 million years ago. These findings indicate that the C. limbatus population from northern Brazil is genetically distinct from all other populations and should be considered as a different management unit for the protection of stocks.     

Megenteron manteri n. sp. and Steringophorus sp. (Digenea: Fellodistomidae) from Monomitopus agassizzi (Ophidiidae) in the Gulf of Mexico

During a study of digenean parasites of deep-sea fishes, 81% (17 of 21) of Monomitopus agassizzi (Ophidiidae) from the Gulf of Mexico were infected with Megenteron manteri n. sp. and 10% (2 of 21) were infected with what appears to be a new species of Steringophorus. M. manteri n. sp. differs from M. crassum in having a longer body (3,733–9,394 m), shorter caeca and a uterus that extends posteriorly beyond the ends of the caeca. The species of Steringophorus differs from other species in the genus in having smaller eggs (19–23 m long) and caeca that extend into the posterior half of the hindbody.     

Gill surface area provides a clue for the respiratory basis of brain size in the blacktip shark (Carcharhinus limbatus)

Brain size varies dramatically, both within and across species, and this variation is often believed to be the result of trade-offs between the cognitive benefits of having a large brain for a given body size and the energetic cost of sustaining neural tissue. One potential consequence of having a large brain is that organisms must also meet the associated high energetic demands. Thus, a key question is whether metabolic rate correlates with brain size. However, using metabolic rate to measure energetic demand yields a relatively instantaneous and dynamic measure of energy turnover, which is incompatible with the longer evolutionary timescale of changes in brain size within and across species. Morphological traits associated with oxygen consumption, specifically gill surface area, have been shown to be correlates of oxygen demand and energy use, and thus may serve as integrated correlates of these processes, allowing us to assess whether evolutionary changes in brain size correlate with changes in longer-term oxygen demand and energy use. We tested how brain size relates to gill surface area in the blacktip shark Carcharhinus limbatus. First, we examined whether the allometric slope of brain mass (i.e., the rate that brain mass changes with body mass) is lower than the allometric slope of gill surface area across ontogeny. Second, we tested whether gill surface area explains variation in brain mass, after accounting for the effects of body mass on brain mass. We found that brain mass and gill surface area both had positive allometric slopes, with larger individuals having both larger brains and larger gill surface areas compared to smaller individuals. However, the allometric slope of brain mass was lower than the allometric slope of gill surface area, consistent with our prediction that the allometric slope of gill surface area could pose an upper limit to the allometric slope of brain mass. Finally, after accounting for body mass, individuals with larger brains tended to have larger gill surface areas. Together, our results provide clues as to how fishes may evolve and maintain large brains despite their high energetic cost, suggesting that C. limbatus individuals with a large gill surface area for their body mass may be able to support a higher energetic turnover, and, in turn, a larger brain for their body mass.     

Spirocamallanus cricotus sp. n. and S. halitrophus sp. n. (Nematoda: Camallanidea) from fishes in the northern Gulf of Mexico

Spirocamallanus cricotus sp. n. (= S. pereirai, in part) and S. halitrophus sp. n. are described from marine fishes of the northern Gulf of Mexico. Spirocamallanus cricotus has a ledge anterior to the basal ring in the buccal capsule, similar spicules with a ratio of 1:1.4 to 2.1, 3 pre- and 5 postcloacal papillae, and 8 rectal glands in the female; S. halitrophus lacks the ledge and possesses dissimilar spicules with a ratio of 1:1.3 to 1.8, 3 pre- and 6 postcloacal papillae, and 4 rectal glands in the female.     

Natural variability and effects of cleaning and storage procedures on vertebral chemistry of the blacktip shark Carcharhinus limbatus

Key methodological assumptions regarding the degree of natural variability and influence of sample handling and storage of elasmobranch vertebral chemistry were assessed using laser‐ablation inductively coupled plasma mass spectrometry. Vertebral chemistry of juvenile blacktip sharks Carcharhinus limbatus was examined to identify whether differences existed among different regions of the vertebral column, between thoracic vertebrae of individual fish or within individual vertebrae. Additionally, the effects of bleach exposure and storage in ethanol on vertebral chemistry were compared. No significant variation in vertebral chemistry was found among different regions of the vertebral column or between thoracic vertebrae, but significant differences among transect locations within individual vertebrae were observed. The variation at all three levels appears comparable with published data on sagittal otoliths of bilaterally symmetrical teleost fishes. The experimental assessment of potential treatment effects indicated vertebral chemistry was not significantly affected by bleach or exposure to ethanol. Taken together, these results support the idea that vertebrae taken from the same region of the vertebral column can be treated as equivalent and at least certain elements remain robust to exposure to bleach and ethanol.     

Age and growth estimates of the bull shark,Carcharhinus leucas,from the northern Gulf of Mexico

Synopsis Length at age and growth rates for 59 bull sharks, Carcharhinus leucas, collected from the northern Gulf of Mexico were estimated from the band patterns formed seasonally in the vertebral centra. The combined age at length data for both sexes were applied to a von Bertalanffy growth model producing parameter estimates of L = 285 cm TL, K = .076, t₀ = –3.0 yr. Lengths at age for males and females were similar except that males did not attain as great a length as females. Growth was apparently slow and varied among individuals, but in general, was estimated to be 15–20 cm yr^–1 for the first five years, 10 cm yr^–1 for years 6–10, 5–7 cm yr^–1 for years 11–16, and less than 4–5 cm yr^–1 thereafter. Males mature at 210–220 cm TL or 14–15 yr of age; females mature at>225 cm TL or 18+ yr of age. The largest male (245 cm TL) was 21.3 yr old; the largest female (268 cm TL) was 24.2 yr old.     

A novel field method to distinguish between cryptic carcharhinid sharks,Australian blacktip shark Carcharhinus tilstoni and common blacktip shark C. limbatus,despite the presence of hybrids

Multivariate and machine‐learning methods were used to develop field identification techniques for two species of cryptic blacktip shark. From 112 specimens, precaudal vertebrae (PCV) counts and molecular analysis identified 95 Australian blacktip sharks Carcharhinus tilstoni and 17 common blacktip sharks Carcharhinus limbatus. Molecular analysis also revealed 27 of the 112 were C. tilstoni × C. limbatus hybrids, of which 23 had C. tilstoni PCV counts and four had C. limbatus PCV counts. In the absence of further information about hybrid phenotypes, hybrids were assigned as either C. limbatus or C. tilstoni based on PCV counts. Discriminant analysis achieved 80% successful identification, but machine‐learning models were better, achieving 100% successful identification, using six key measurements (fork length, caudal‐fin peduncle height, interdorsal space, second dorsal‐fin height, pelvic‐fin length and pelvic‐fin midpoint to first dorsal‐fin insertion). Furthermore, pelvic‐fin markings could be used for identification: C. limbatus has a distinct black mark >3% of the total pelvic‐fin area, while C. tilstoni has markings with diffuse edges, or has smaller or no markings. Machine learning and pelvic‐fin marking identification methods were field tested achieving 87 and 90% successful identification, respectively. With further refinement, the techniques developed here will form an important part of a multi‐faceted approach to identification of C. tilstoni and C. limbatus and have a clear management and conservation application to these commercially important sharks. The methods developed here are broadly applicable and can be used to resolve species identities in many fisheries where cryptic species exist.     

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.     

Worldwide phylogeography of the blacktip shark (Carcharhinus limbatus) inferred from mitochondrial DNA reveals isolation of western Atlantic populations coupled with recent Pacific dispersal

Although many coastal shark species have widespread distributions, the genetic relatedness of worldwide populations has been examined for few species. The blacktip shark, (Carcharhinus limbatus), inhabits tropical and subtropical coastal waters throughout the world. In this study, we examined the genetic relationships of blacktip shark populations (n = 364 sharks) throughout the majority of the species' range using the entire mitochondrial control region (1067-1070 nucleotides). Two geographically distinct maternal lineages (western Atlantic, Gulf of Mexico, and Caribbean Sea clades, and eastern Atlantic, Indian, and Pacific Ocean clades) were identified and shallow population structure was detected throughout their geographic ranges. These findings indicate that a major population subdivision exists across the Atlantic Ocean, but not the Pacific Ocean. The historical dispersal of this widespread, coastal species may have been interrupted by the rise of the Isthmus of Panama. This scenario implies historical dispersal across the Pacific Ocean (supported by the recovery of the same common haplotype from the Philippines, Hawaii, and the Gulf of California reflecting recent/contemporary dispersal abilities) and an oceanic barrier to recent migration across the Atlantic. Genetic structure within the eastern Atlantic/Indo-Pacific (Phi(ST) = 0.612, P < 0.001) supports maternal philopatry throughout this area, expanding previous western Atlantic findings. Eastern Atlantic/Indo-Pacific C. limbatus control region haplotypes were paraphyletic to Carcharhinus tilstoni haplotypes in our maximum-parsimony analysis. The greater divergence of western Atlantic C. limbatus than C. tilstoni from eastern Atlantic/Indo-Pacific C. limbatus reflects the taxonomic uncertainty of western Atlantic C. limbatus.     

Chaulioleptos haywardi n. gen., n. sp. (Digenea: Sanguinicolidae) from Filimanus heptadactyla (Perciformes: Polynemidae) of Moreton Bay, Australia

Here we describe the first species of sanguinicolid blood fluke (Trematoda: Digenea) from a polynemid fish. Chaulioleptos haywardi n. gen., n. sp. is described from Filimanus heptadactyla Cuvier, 1829 (Perciformes: Polynemidae), the sevenfinger threadfin from Sandgate, Moreton Bay (southeast Queensland, Australia). Chaulioleptos haywardi differs from existing sanguinicolid genera in the combined possession of the following 7 characters: 2 testes, an entirely postovarian uterus, a uterine chamber, separate genital pores, an H-shaped intestine with abbreviated anterior caeca, tegumental spines in incomplete ventro-marginal transverse rows that are continuous along the length of the body, and vitelline follicles that are tightly compacted and subsequently appear to form a solid branching mass occupying the area anterior to intestinal bifurcation and extending posteriorly to the level of the posterior margin of the anterior testis. Chaulioleptos haywardi is most closely related to Paracardicola Martin, 1960 and Adelomyllos Nolan and Cribb, 2004.