The determinants of sexual segregation in the scalloped hammerhead shark,Sphyrna lewini

Synopsis Female scalloped hammerhead sharks move offshore at a smaller size than do males to form schools composed primarily of intermediate size female sharks. This movement results in smaller females feeding more on pelagic prey than do males and with greater predatory success. It is contended that this change in habitat causes females to grow more rapidly to reproductive size. Intermediate size females grow at a more rapid rate than males. Female scalloped hammerhead sharks mature at a size larger than males. For many elasmobranch species, females: (1) occupy a different habitat, (2) grow more rapidly prior to maturity and continue growth following maturation, (3) feed on different prey with increased feeding success, and (4) reproduce at a size larger than males. It is suggested that female segregation increases fitness, resulting in more rapid growth for the former sex. The females reach maturity at the larger size necessary to support embryonic young, yet similar age to males, matching the female reproductive lifetime to that of males.     

New microsatellite loci for the endangered scalloped hammerhead shark, Sphyrna lewini

We isolated 15 microsatellite markers for the scalloped hammerhead shark, Sphyrna lewini. Loci were tested on 80 specimens of S. lewini from four Eastern Pacific samples. The number of alleles per locus ranged from 6 to 31 (mean = 14). Observed and expected levels of heterozygosity per locus ranged from 0.39 to 0.91 (mean = 0.70) and from 0.54 to 0.90 (mean = 0.76), respectively. No pairs of loci were in gametic disequilibrium after Bonferroni correction of α. One locus showed significantly lower heterozygosity than expected under Hardy–Weinberg proportions in two populations, possibly caused by null alleles.     

Global phylogeography of the scalloped hammerhead shark (Sphyrna lewini)

Large marine fishes typically have little population genetic structure. The exceptions are associated with sedentary behaviour, disjunct distributions, or reproductive philopatry. Scalloped hammerhead sharks (Sphyrna lewini) incorporate the contrasting traits of oceanic habitat (usually associated with high dispersal) and possible fidelity to nursery grounds (for reproductive females). To evaluate the expectations of these contrasting behaviours, we examined the global genetic structure of S. lewini based on collections (n = 271 individuals) from 20 nursery areas. A 548-bp fragment of mitochondrial DNA control region revealed 22 polymorphic sites, 24 haplotypes, and three lineages distinguished by 2.56-3.77% sequence divergence. Coalescence analyses based on a provisional molecular clock indicate an origin in the Indo-West Pacific with late Pleistocene radiations into the central Pacific (Hawaii) and eastern Pacific (Central America), as well as recent interchange between oceans via southern Africa. Population subdivisions are strong (overall Phi(ST) = 0.749, P < 0.0001 and among oceans Phi(ST) = 0.598, P < 0.0098). Genetic discontinuity within oceans (Phi(ST) = 0.519, P < 0.0001) is primarily associated with oceanic barriers (migration across oceans M approximately 0), with much less structure along continental margins (M > 10). We conclude that nursery populations linked by continuous coastline have high connectivity, but that oceanic dispersal by females is rare. Although we cannot rule out philopatry to natal nurseries, oceanic barriers appear to have a much stronger influence on the genetic architecture of this species and may indicate a mechanism for recent evolutionary radiations in the genus Sphyrna.     

Report on fetal mummification in the scalloped hammerhead shark Sphyrna lewini

This study describes the fetal mummification process in two embryos of a 310 cm total length scalloped hammerhead shark Sphyrna lewini caught in southeastern Brazil, in December 2017. Fourteen embryos were observed in total, in which two males in the left uterus presented different stages of mummification. Both mummified embryos were covered by an exudate (i.e., a mucous substance), indicating a hematic mummification process. All embryos were at the placentotrophic stage of development, indicating that they were close to parturition. An intrinsic characteristic is suggested as possible etiology for this condition, such as umbilical torsion, because both embryos were at different sizes and, therefore, at different development stages. In addition, the sample size did not allow the authors to presume any pollution effect once only one female was observed. Finally, fetal mummification and other embryonic development disorders might have populational impacts due to reduction in embryo survival and, consequently, recruitment. For this reason and considering that S. lewini is categorized as a “critically endangered species,” this study's results have conservational relevance.     

Age,growth and reproductive biology of the silky shark,Carcharhinus falciformis,and the scalloped hammerhead,Sphyrna lewini,from the northwestern Gulf of Mexico

Synopsis The silky shark, Carcharhinus falciformis, and scalloped hammerhead, Sphyrna lewini, represent >80% of the shark by-catch of the winter swordfish/tuna longline fishery of the northwestern Gulf of Mexico. This catch represents a potential supplemental fishery, yet little is known of the life histories of the two species. This report relates reproductive biology data to age and growth estimates for 135 C. falciformis and 78 S. lewini. Unlike other regional populations, C. falciformis in the Gulf of Mexico may have a seasonal 12 month gestation period. Males mature at 210–220 cm TL (6–7 yr); females at >225 cm TL (7–9 yr). Application of age at length data for combined sexes produced von Bertalanffy growth model parameter estimates of L_∞ = 291 cm TL, K = 0.153, t₀ = −2.2 yr. Adult male S. lewini outnumbered adult females in catches because of differences in the distributions of the sexually segregated population. Males mature at 180 cm TL (10 yr); females at 250 cm TL (15 yr). von Bertalanffy parameter estimates for combined sexes of this species were L_∞ = 329 cm TL, K = 0.073, t_o = −2.2 yr.     

Demographic processes underlying subtle patterns of population structure in the scalloped hammerhead shark, Sphyrna lewini

Genetic diversity (θ), effective population size (N(e)), and contemporary levels of gene flow are important parameters to estimate for species of conservation concern, such as the globally endangered scalloped hammerhead shark, Sphyrna lewini. Therefore, we have reconstructed the demographic history of S. lewini across its Eastern Pacific (EP) range by applying classical and coalescent population genetic methods to a combination of 15 microsatellite loci and mtDNA control region sequences. In addition to significant population genetic structure and isolation-by-distance among seven coastal sites between central Mexico and Ecuador, the analyses revealed that all populations have experienced a bottleneck and that all current values of θ are at least an order of magnitude smaller than ancestral θ, indicating large decreases in N(e) (θ = 4N(e)μ), where μ is the mutation rate. Application of the isolation-with-migration (IM) model showed modest but significant genetic connectivity between most sampled sites (point estimates of Nm = 0.1-16.7), with divergence times (t) among all populations significantly greater than zero. Using a conservative (i.e., slow) fossil-based taxon-specific phylogenetic calibration for mtDNA mutation rates, posterior probability distributions (PPDs) for the onset of the decline in N(e) predate modern fishing in this region. The cause of decline over the last several thousand years is unknown but is highly atypical as a post-glacial demographic history. Regardless of the cause, our data and analyses suggest that S. lewini was far more abundant throughout the EP in the past than at present.     

Predominance of genetic monogamy by females in a hammerhead shark, Sphyrna tiburo: implications for shark conservation

There is growing interest in the mating systems of sharks and their relatives (Class Chondrichthyes) because these ancient fishes occupy a key position in vertebrate phylogeny and are increasingly in need of conservation due to widespread overexploitation. Based on precious few genetic and field observational studies, current speculation is that polyandrous mating strategies and multiple paternity may be common in sharks as they are in most other vertebrates. Here, we test this hypothesis by examining the genetic mating system of the bonnethead shark, Sphyrna tiburo, using microsatellite DNA profiling of 22 litters (22 mothers, 188 embryos genotyped at four polymorphic loci) obtained from multiple locations along the west coast of Florida. Contrary to expectations based on the ability of female S. tiburo to store sperm, the social nature of this species and the 100% multiple paternity observed in two other coastal shark species, over 81% of sampled bonnethead females produced litters sired by a single male (i.e. genetic monogamy). When multiple paternity occurred in S. tiburo, there was an indication of increased incidence in larger mothers with bigger litters. Our data suggest that sharks may exhibit complex genetic mating systems with a high degree of interspecific variability, and as a result some species may be more susceptible to loss of genetic variation in the face of escalating fishing pressure. Based on these findings, we suggest that knowledge of elasmobranch mating systems should be an important component of conservation and management programmes for these heavily exploited species.     

Notes on the long‐term transport of the scalloped hammerhead shark (Sphyrna lewini)

The capture and transport of scalloped hammerhead sharks (Sphyrna lewini Griffith and Smith, 1834) historically has represented a difficult, expensive, and uncertain undertaking for the public aquarium community. In this study, techniques were developed to improve the successful long‐term transport of S. lewini by mitigating some of the deleterious effects associated with hyperactivity and impaired swimming patterns. The relationship between the transport vessel size and shark sizes, numbers, and swimming behavior was considered when formulating the transport regime. By balancing these factors and adopting a comprehensive water treatment method, it was possible to extend the duration of a successful transport by up to 60 hr. Implications for the future transport of S. lewini and other free‐swimming sharks are discussed. Zoo Biol 21:243–251, 2002. © 2002 Wiley‐Liss, Inc.     

New insights into the trophic ecology of the scalloped hammerhead shark,Sphyrna lewini,in the eastern tropical Pacific Ocean

Scalloped hammerhead shark, Sphyrna lewini, is considered a generalist predator, showing individual specializations and different trophic levels. This species inhabits both oceanic islands and coastal nursery areas in the eastern tropical Pacific (ETP) Ocean. Existing trophic ecology studies on S. lewini have been focused on stomach contents. This study provides new insights into its foraging preferences, habitat use, and trophic position, using stable isotope analysis (SIA) of muscle tissue. Stable isotope signatures of δ¹³C and δ¹⁵N were determined in S. lewini muscles (n?=?29) from the Ecuadorian Pacific in 2013. Trophic position (TP), isotopic niches, and overlap of S. lewini were estimated by sex, age, and maturity stages to infer their dietary habits throughout life stages. SIA revealed complex movement patterns related to sex and age classes of S. lewini, highlighting high degrees of dietary plasticity and habitat use, with a stronger relation to coastal regions than previously reported. This study provides crucial information regarding essential areas and the related migration behavior of S. lewini, with important implications for their conservation and management in the ETP.

     

Correction to: New insights into the trophic ecology of the scalloped hammerhead shark,Sphyrna lewini,in the eastern tropical Pacific Ocean

Environmental Biology of Fishes -     

Periodicity of the growth-band formation in vertebrae of juvenile scalloped hammerhead shark Sphyrna lewini from the Mexican Pacific Ocean

The age of 296 juvenile scalloped hammerhead sharks Sphyrna lewini caught by several fisheries in the Mexican Pacific Ocean from March 2007 to September 2017 were estimated from growth band counts in thin-sectioned vertebrae. Marginal-increment analysis (MIA) and centrum-edge analysis (CEA) were used to verify the periodicity of formation of the growth bands, whereas elemental profiles obtained from LA-ICP-MS transect scans in vertebrae of 15 juveniles were used as an alternative approach to verify the age of the species for the first time. Age estimates ranged from 0 to 10+ years (42–158.7 cm total length; L_T). The index of average percentage error (I_APE 3.6%), CV (5.2%), bias plots and Bowker's tests of symmetry showed precise and low-biased age estimation. Both MIA and CEA indicated that in the vertebrae of juveniles of S. lewini a single translucent growth band was formed during winter (November–March) and an opaque band during summer (July–September), a period of faster growth, apparently correlated with a higher sea surface temperature. Peaks in vertebral P and Mn content spatially corresponded with the annual banding pattern in most of the samples, displaying 1.19 and 0.88 peaks per opaque band, respectively, which closely matched the annual deposition rate observed in this study. Although the periodicity of growth band formation needs to be verified for all sizes and ages representing the population of the species in the region, this demonstration of the annual formation of the growth bands in the vertebrae of juveniles should lead to a re-estimation of the growth parameters and productivity of the population to ensure that it is harvested at sustainable levels.     

Global phylogeography with mixed-marker analysis reveals male-mediated dispersal in the endangered scalloped hammerhead shark (Sphyrna lewini)

Background

The scalloped hammerhead shark, Sphyrna lewini, is a large endangered predator with a circumglobal distribution, observed in the open ocean but linked ontogenetically to coastal embayments for parturition and juvenile development. A previous survey of maternal (mtDNA) markers demonstrated strong genetic partitioning overall (global Φ_ST = 0.749) and significant population separations across oceans and between discontinuous continental coastlines.

Methodology/Principal Findings

We surveyed the same global range with increased sample coverage (N = 403) and 13 microsatellite loci to assess the male contribution to dispersal and population structure. Biparentally inherited microsatellites reveal low or absent genetic structure across ocean basins and global genetic differentiation (F _ST = 0.035) over an order of magnitude lower than the corresponding measures for maternal mtDNA lineages (Φ_ST = 0.749). Nuclear allelic richness and heterozygosity are high throughout the Indo-Pacific, while genetic structure is low. In contrast, allelic diversity is low while population structure is higher for populations at the ends of the range in the West Atlantic and East Pacific.

Conclusions/Significance

These data are consistent with the proposed Indo-Pacific center of origin for S. lewini, and indicate that females are philopatric or adhere to coastal habitats while males facilitate gene flow across oceanic expanses. This study includes the largest sampling effort and the most molecular loci ever used to survey the complete range of a large oceanic predator, and findings emphasize the importance of incorporating mixed-marker analysis into stock assessments of threatened and endangered shark species.     

A review of the biology, fisheries and conservation of the whale shark Rhincodon typus

Although the whale shark Rhincodon typus is the largest extant fish, it was not described until 1828 and by 1986 there were only 320 records of this species. Since then, growth in tourism and marine recreation globally has lead to a significant increase in the number of sightings and several areas with annual occurrences have been identified, spurring a surge of research on the species. Simultaneously, there was a great expansion in targeted R. typus fisheries to supply the Asian restaurant trade, as well as a largely un-quantified by-catch of the species in purse-seine tuna fisheries. Currently R. typus is listed by the IUCN as vulnerable, due mainly to the effects of targeted fishing in two areas. Photo-identification has shown that R. typus form seasonal size and sex segregated feeding aggregations and that a large proportion of fish in these aggregations are philopatric in the broadest sense, tending to return to, or remain near, a particular site. Somewhat conversely, satellite tracking studies have shown that fish from these aggregations can migrate at ocean-basin scales and genetic studies have, to date, found little graphic differentiation globally. Conservation approaches are now informed by observational and environmental studies that have provided insight into the feeding habits of the species and its preferred habitats. Notwithstanding these advances, there remain notable gaps in the knowledge of this species particularly with respect to the life history of neonates and adults who are not found in the feeding aggregations.     

Possible female philopatry of the smooth hammerhead shark Sphyrna zygaena revealed by genetic structure patterns

We assessed the spatial pattern of genetic structure of smooth hammerhead shark Sphyrna zygaena in 10 localities from the Northern Mexican Pacific. A total of 35 haplotypes were identified in 129 sequences of the mtDNA control region. The results showed slight but significant genetic structure among localities (Φ_ST = 0.044, P < 0.001). In addition, the localities with highest number of juveniles were genetically different (Φ_ST = 0.058, P < 0.024), which may be representative of nursery areas. The genetic differentiation pattern can be associated to female philopatry and preference for particular birthing sites. Finally, historical demography shows that S. zygaena populations present a recent demographic expansion that occurred during glacial events in the late Pleistocene to early Holocene.     

Population genetic divergence as consequence of past range expansion of the smooth hammerhead shark Sphyrna zygaena

Hydrobiologia - We investigated the roles of a benthivorous (Prochilodus brevis, Steindachner 1875) and a planktivorous (Oreochromis niloticus, Linnaeus, 1758) fish in translocating phosphorus from...     

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.     

The Mediterranean monk seal Monachus monachus: status,biology, threats,and conservation priorities

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