Species-specific profiles of per- and polyfluoroalkyl substances (PFAS) in small coastal sharks along the South Atlantic Bight of the United States.

Per- and polyfluoroalkyl substances (PFAS) have gained widespread commercial use across the globe in various industrial and consumer products, such as textiles, firefighting foams, and surface coating materials. Studies have shown that PFAS exhibit a strong tendency to accumulate within aquatic food webs, primarily due to their high bioaccumulation potential and resistance to degradation. Despite such concerns, their impact on marine predators like sharks remains underexplored. This study aimed to investigate the presence of 34 PFAS in the plasma (n = 315) of four small coastal sharks inhabiting the South Atlantic Bight of the United States (U.S). Among the sharks studied, bonnetheads (Sphyrna tiburo) had the highest ∑PFAS concentration (3031 ± 1674 pg g - 1 plasma, n = 103), followed by the Atlantic sharpnose shark (Rhizoprionodon terraenovae, 2407 ± 969 pg g - 1, n = 101), blacknose shark (Carcharhinus acronotus, 1713 ± 662 pg g - 1, n = 83) and finetooth shark (Carcharhinus isodon, 1431 ± 891 pg g - 1, n = 28). Despite declines in the manufacturing of perfluorooctane sulfonate (PFOS) and perfluorooctanoic acid (PFOA), the long-chain (C8 - C13) perfluoroalkyl acids (PFAAs) were frequently detected, with PFOS, perfluorodecanoic acid (PFDA), and perfluorotridecanoic acid (PFTrDA) present as the most dominant PFAS. Furthermore, males exhibited significantly higher ∑PFAS concentrations than females in bonnetheads (p < 0.01), suggesting possible sex-specific PFAS accumulation or maternal offloading in some species. The results of this study underscore the urgency for more extensive biomonitoring of PFAS in aquatic/marine environments to obtain a comprehensive understanding of the impact and fate of these emerging pollutants on marine fauna.

Embryonic development in the bonnethead (Sphyrna tiburo), a viviparous hammerhead shark.

BACKGROUND: The hammerhead sharks (family Sphyrnidae) are an immediately recognizable group of sharks due to their unique head shape. Though there has long been an interest in hammerhead development, there are currently no explicit staging tables published for any members of the group. The bonnethead Sphyrna tiburo is the smallest member of Sphyrnidae and is abundant in estuarine and nearshore waters in the Gulf of Mexico and Western North Atlantic Ocean. Due to their relative abundance, close proximity to shore, and brief gestation period, it has been possible to collect and document multiple embryonic specimens at progressive stages of development. RESULTS: We present the first comprehensive embryonic staging series for the Bonnethead, a viviparous hammerhead shark. Our stage series covers a period of development from stages that match the vertebrate phylotypic period, from Stage 23, through stages of morphological divergence to complete development at birth-Stage 35). Notably, we use a variety of techniques to document crucial stages that lead to their extreme craniofacial diversity, resulting in the formation of one of the most distinctive characters of any shark species, the cephalofoil or hammer-like head. CONCLUSION: Documenting the development of hard-to-access vertebrates, like this viviparous shark species, offers important information about how new and diverse morphologies arise that otherwise may remain poorly studied. This work will serve as a platform for future comparative developmental research both within sharks and across the phylogeny of vertebrates, underpinning the extreme potential of craniofacial development and morphological diversity in vertebrate animals.

Philopatry influences the genetic population structure of the blacktip shark (Carcharhinus limbatus) at multiple spatial scales.

Understanding how interactions among microevolutionary forces generate genetic population structure of exploited species is vital to the implementation of management policies that facilitate persistence. Philopatry displayed by many coastal shark species can impact gene flow and facilitate selection, and has direct implications for the spatial scales of management. Here, genetic structure of the blacktip shark (Carcharhinus limbatus) was examined using a mixed-marker approach employing mitochondrial control region sequences and 4339 SNP-containing loci generated using ddRAD-Seq. Genetic variation was assessed among young-of-the-year sampled in 11 sites in waters of the United States in the western North Atlantic Ocean, including the Gulf of Mexico. Spatial and environmental analyses detected 68 nuclear loci putatively under selection, enabling separate assessments of neutral and adaptive genetic structure. Both mitochondrial and neutral SNP data indicated three genetically distinct units-the Atlantic, eastern Gulf, and western Gulf-that align with regional stocks and suggest regional philopatry by males and females. Heterogeneity at loci putatively under selection, associated with temperature and salinity, was observed among sites within Gulf units, suggesting local adaptation. Furthermore, five pairs of siblings were identified in the same site across timescales corresponding with female reproductive cycles. This indicates that females re-used a site for parturition, which has the potential to facilitate the sorting of adaptive variation among neighbouring sites. The results demonstrate differential impacts of microevolutionary forces at varying spatial scales and highlight the importance of conserving essential habitats to maintain sources of adaptive variation that may buffer species against environmental change.

Observations of fin injury closure in Great Hammerheads and implications for the use of fin-mounted geolocators.

OBJECTIVE: Sharks face a high risk of injuries throughout all life stages and are therefore expected to show a good wound closure capacity. METHODS: Here, the wound closure of one major injury and one minor injury to the first dorsal fins of two free-ranging, mature female Great Hammerheads Sphyrna mokarran is described macroscopically. RESULT: The sharks showed complete wound closure of single, clean-cut lacerations measuring 24.2 and 11.6 cm in length after an estimated 323 and 138 days. These estimates were based on the observed closure rate and visual confirmation of a complete wound closure upon multiple resightings of the same individuals. Additionally, the posterior lateral displacement of fin-mounted geolocators within the fin and outside of the fin without causing external damage was documented in three additional Great Hammerheads. CONCLUSION: These observations supplement findings about wound closure capabilities in elasmobranchs. The documented geolocator displacement furthers the discussion about the safe use of these geolocators to track shark movements but also has implications for future tagging studies.

Vertical space use and thermal range of the great hammerhead (Sphyrna mokarran), (Rüppell, 1837) in the western North Atlantic.

The great hammerhead (Sphyrna mokarran) is a highly mobile, large-bodied shark primarily found in coastal-pelagic and semi-oceanic waters across a circumtropical range. It is a target or by-catch species in multiple fisheries, and as a result, rapid population declines have occurred in many regions. These declines have contributed to the species being assessed as globally critically endangered on the IUCN Red List. Although conservation and management measures have yielded promising results in some regions, such as the United States, high levels of at-vessel and post-release mortality remain a major concern to the species population recovery. This examined the vertical space use and thermal range of pop-off archival satellite-tagged S. mokarran in the western North Atlantic Ocean, expanding the understanding of the ecological niche of this species and providing insight into by-catch mitigation strategies for fisheries managers. The results showed that S. mokarran predominantly used shallow depths (75% of records <30 m) and had a narrow temperature range (89% of records between 23 and 28°C). Individual differences in depth use were apparent, and a strong diel cycle was observed, with sharks occupying significantly deeper depths during the daytime. Furthermore, two individuals were confirmed pregnant with one migrating from the Bahamas to South Carolina, U.S.A., providing further evidence of regional connectivity and parturition off the U.S. East Coast. The findings suggest that S. mokarran may be vulnerable to incidental capture in the western North Atlantic commercial longline fisheries due to substantial vertical overlap between the species and the gear. The results can be incorporated into conservation and management efforts to develop and/or refine mitigation measures focused on reducing the by-catch and associated mortality of this species, which can ultimately aide S. mokarran population recovery in areas with poor conservation status.

Hexabothriidae and Monocotylidae (Monogenoidea) from the gills of neonate hammerhead sharks (Sphyrnidae) Sphyrna gilberti, Sphyrna lewini and their hybrids from the western North Atlantic Ocean.

Neonates of hammerhead sharks (Sphyrnidae), Sphyrna lewini (Griffith and Smith, 1834), the sympatric cryptic species, Sphyrna gilberti Quattro et al., 2013, and their hybrids were captured in the western North Atlantic, along the coast of South Carolina, USA, between 2018 and 2019 and examined for gill monogenoids. Parasites were identified and redescribed from the gills of 79 neonates, and DNA sequences from partial fragments of the nuclear 28S ribosomal RNA (rDNA) and cytochrome c oxidase I mitochondrial DNA (COI) genes were generated to confirm species identifications. Three species of monogenoids from Hexabothriidae Price, 1942 and Monocotylidae Taschenberg, 1879 were determined and redescribed. Two species of Hexabothriidae, Erpocotyle microstoma (Brooks, 1934) and Erpocotyle sphyrnae (MacCallum, 1931), infecting both species of Sphyrna and hybrids; and 1 species of Monocotylidae, Loimosina wilsoni Manter, 1944, infecting only S. lewini and hybrids. Loimosina wilsoni 28S rDNA sequences matched those of Loimosina sp. from the southern coast of Brazil. Based on limited morphological analysis, Loimosina parawilsoni is likely a junior synonym of L. wilsoni. This is the first taxonomic study of monogenoids infecting S. gilberti and hybrids of S. gilberti and S. lewini.

Multiple paternity in two populations of finetooth sharks (Carcharhinus isodon) with varying reproductive periodicity.

The mechanisms underlying polyandry and female mate choice in certain taxonomic groups remain widely debated. In elasmobranchs, several species have shown varying rates of polyandry based on genetic studies of multiple paternity (MP). We investigated MP in the finetooth shark, Carcharhinus isodon, in order to directly test the encounter rate hypothesis (ERH), which predicts that MP is a result of the frequency of encounters between mature conspecifics during the breeding season, and should therefore increase when more time is available for copulation and sperm storage. Female finetooth sharks in the northern Gulf of Mexico (GoM) have been found to reproduce with both annual periodicity and biennial periodicity, while finetooth sharks from the northwestern Atlantic Ocean have only been found to reproduce biennially, allowing us to compare mating opportunity to frequency of MP. Our results show high rates of MP with no significant difference in frequency between females in the GoM (83.0%) and Atlantic (88.2%, p = .8718) and varying but nonsignificant rates of MP between females in the GoM reproducing annually (93.0%) and biennially (76.6%, p = .2760). While the ERH is not supported by this study, it remains possible that reproductive periodicity and other physiological factors play a role in determining rates of MP in elasmobranchs, with potential benefits to individuals and populations.

Reproductive cycle and fecundity of the bonnethead Sphyrna tiburo L. from the northwest Atlantic Ocean.

The present study examined temporal changes in plasma sex hormone concentrations and the morphology and histology of reproductive organs in mature northwest Atlantic (NWA) bonnetheads Sphyrna tiburo L. to characterize reproductive cycle, breeding periodicity and fertility in this still poorly studied population. Progressive increases in testis width, epididymis head width, plasma testosterone (T) concentrations, and occurrence of mature spermatozoa were observed in male S. tiburo from June to September, demonstrating that spermatogenesis occurs during the summer. Nonetheless, increases in maximum follicle diameter, oviducal gland width, plasma 17ß-estradiol and T concentrations, and occurrence of vitellogenic follicles were not observed in mature females until between October and April, demonstrating non-synchronous patterns of gametogenesis in males and females. Fresh copulatory wounds were observed in females collected during late September along with histological evidence for sperm presence in the oviducal gland between September and April, confirming a 6- to 7 month period of female sperm storage. Ovulation occurred between mid-April and early May in concert with increases in female plasma progesterone concentrations. Gestation occurred during a 4.5- to 5 month period between May and early September, and 97% of mature females collected during this period were gravid, indicating a highly synchronized, annual reproductive periodicity. Brood size was significantly correlated with maternal size and ranged from 1 to 13 pups with a mean ± S.D. of 8.1 ± 2.2, which was significantly lower than reported in Gulf of Mexico (GOM) populations. The occurrence of non-fertile offspring was observed in 17% of broods with a range of 1-7 non-fertile eggs present in individual females. Thus, as previously reported in GOM S. tiburo, this unusual form of infertility also appears to be prevalent in the NWA population and requires further study. This study has demonstrated meaningful differences in reproductive biology of these populations, emphasizing the need for region-specific approaches for population management.

Histologic Observations of Dermal Wound Healing in a Free-Ranging Blacktip Shark from the Southeastern U.S. Atlantic Coast: A Case Report.

Despite indications that sharks have an exceptional capacity to heal from traumatic injuries, no detailed microscopic observations of integumental wounds have been reported for sharks. This study details the histopathological features of such wounds in a free-ranging shark. An adult male Blacktip Shark Carcharhinus limbatus was collected in 2017 during fisheries-independent sampling efforts in the coastal southeastern U.S. Atlantic. The shark had numerous lesions on his head, torso, and left pectoral fin that were compatible with shark bites. Representative samples from two wounds on the head were examined by light microscopy. The epidermal changes included hyperplasia and hypertrophy; intracellular edema; and the absence of goblet cells, denticles, and chromatophores. In the dermis, fibrinocellular exudation, granulation tissue, and marked skeletal muscle necrosis and regeneration were observed. The above features were comparable to wound healing in bony fish, albeit minor differences were found. Although this case documents exceptionally good regeneration of skeletal muscle in the shark, we found no evidence of unique morphological healing patterns. Further studies on wound healing are needed because recent molecular and genetic findings do suggest evolutionary adaptations enhancing healing in sharks.

Re-evaluation of reproductive cycle and fecundity of finetooth sharks Carcharhinus isodon (Valenciennes 1839) from the Northwest Atlantic Ocean, with new observations on ovarian cycle and reproductive endocrinology of biennially reproducing sharks.

To provide updated information on life history for improved fishery management, the reproductive cycle of the finetooth shark Carcharhinus isodon from Northwest Atlantic (NWA) populations was investigated by examining temporal changes in morphology and histology of reproductive organs. Changes in plasma concentrations of gonadal sex hormones in relation to reproductive stage were also examined. Increases in testis width, epididymis head width, plasma testosterone concentrations and occurrence of mature spermatozoa were observed in male sharks between December and April, suggesting a seasonal pattern in reproduction that culminates with copulatory activity in May. Increases in maximum follicular diameter, oviducal gland width, plasma 17ß-estradiol concentrations and occurrence of vitellogenic follicles were observed in non-pregnant female sharks during the same time period along with the occurrence of newly pregnant females in May, demonstrating strong synchronicity between male and female reproductive cycles. Pregnant females bearing full-term embryos were also observed in May, indicating that parturition occurs between mid-May and early June and gestation requires 12 months. Only transient temporal changes in follicle size and oviducal gland width were observed in pregnant females, indicating that reproductive periodicity is biennial; nonetheless, a single female exhibiting signs of concurrent vitellogenesis and pregnancy was observed. Mean brood size ± S.D. was 3.9 ± 0.9 offspring/female. Fecundity was not significantly correlated with female size, in part due to an unexpectedly high rate of early embryo mortality, which occurred in 11% of pregnant females, and was more common in larger individuals. Changes in ovarian activity during mid-pregnancy were observed, suggesting possible roles for the ovary in regulating some aspects of early to mid-gestation. This study confirms that earlier characterizations of the reproductive cycle and fecundity in NWA finetooth sharks remain valid for use in fishery management. This study also highlights unusual features of finetooth shark pregnancy (e.g., early embryo death, mid-pregnancy changes in ovarian function) that may have broader relevance to understanding elasmobranch reproduction.

Hybridization between sympatric hammerhead sharks in the western North Atlantic Ocean.

Hybridization between closely related species has been documented across a wide range of taxa but has not been well studied in elasmobranchs. Hammerhead sharks have drawn global conservation concern because they experience some of the highest mortality rates among sharks when interacting with fisheries. Here we report on the detection of hybrids between the globally distributed scalloped hammerhead ( Sphyrna lewini) and recently described Carolina hammerhead ( S. gilberti) which are only known from the western Atlantic Ocean. Using a genomics approach, 10 first-generation hybrids and 15-17 backcrosses were detected from 554 individuals. The identification of backcrosses demonstrates hybrids are viable, and all backcrosses but one involved a scalloped hammerhead. All hybrids but one possessed Carolina hammerhead mtDNA, indicating sex-biased gene flow between species. Repeated hybridization and backcrossing with scalloped hammerheads could lead to the loss of endemic Carolina hammerheads.

Longevity of Atlantic Sharpnose Sharks Rhizoprionodon terraenovae and Blacknose Sharks Carcharhinus acronotus in the western North Atlantic Ocean based on tag-recapture data and direct age estimates.

Longevity of Rhizoprionodon terraenovae and Carcharhinus acronotus in the western North Atlantic Ocean was examined using direct age estimates from vertebral sections and tag-recapture data. Time-at-liberty ranged from 7.7-14.0 years (mean =10.1) for R. terraenovae and 10.9-12.8 years (mean =11.9) for C. acronotus. Maximum estimated longevity was determined to be 19.8 years through tag-recapture data and 18.5 years from direct age estimates for R. terraenovae and 22.8 years through tag-recapture data and 20.5 years through direct age estimates for C. acronotus. These longevity estimates represent a large increase over previous estimates and may have significant effects on analyses that depend on longevity including lifetime fecundity, mortality rates, demographic analyses and stock assessments.