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.

A comprehensive petrochemical vulnerability index for marine fishes in the Gulf of Mexico.

Oil and gas extraction activities occur across the globe, yet species-specific toxicological information on the biological and ecological impacts of exposure to petrochemicals is lacking for the vast majority of marine species. To help prioritize species for recovery, mitigation, and conservation in light of significant toxicological data gaps, a trait-based petrochemical vulnerability index was developed and applied to the more than 1700 marine fishes present across the entire Gulf of Mexico, including all known bony fishes, sharks, rays and chimaeras. Using life history and other traits related to likelihood of exposure, physiological sensitivity to exposure, and population resiliency, final calculated petrochemical vulnerability scores can be used to provide information on the relative sensitivity, or resilience, of marine fish populations across the Gulf of Mexico to oil and gas activities. Based on current knowledge of traits, marine fishes with the highest vulnerability scores primarily occur in areas of high petrochemical activity, are found at or near the surface, and have low reproductive turnover rates and/or highly specialized diet and habitat requirements. Relative population vulnerability scores for marine fishes can be improved with additional toxicokinetic studies, including those that account for the synergistic or additive effect of multiple stressors, as well as increased research on ecological and life history traits, especially for deep living species.

Map-like use of Earth's magnetic field in sharks.

Migration is common in marine animals,1-5 and use of the map-like information of Earth's magnetic field appears to play an important role.2,6-9 While sharks are iconic migrants10-12 and well known for their sensitivity to electromagnetic fields,13-20 whether this ability is used for navigation is unresolved.14,17,21,22 We conducted magnetic displacement experiments on wild-caught bonnetheads (Sphyrna tiburo) and show that magnetic map cues can elicit homeward orientation. We further show that use of a magnetic map to derive positional information may help explain aspects of the genetic structure of bonnethead populations in the northwest Atlantic.23-26 These results offer a compelling explanation for the puzzle of how migratory routes and population structure are maintained in marine environments, where few physical barriers limit movements of vagile species. VIDEO ABSTRACT.

Blood biochemical status of deep-sea sharks following longline capture in the Gulf of Mexico.

Prior to the Deepwater Horizon (DWH) oil spill, little research effort was focused on studying deep-sea sharks in the Gulf of Mexico (GoM). While the biology of these fishes remains virtually unknown, they are routinely captured in commercial fisheries as bycatch. In the absence of basic biological data, and with the probability of post-release survival unknown for most species, effective management plans cannot be formulated, making populations highly susceptible to overfishing. Any potential detrimental effects of the DWH oil spill, which occurred at 1500 m deep, are also unknown. Following longline capture, we characterized the physiological blood biochemical parameters related to secondary stress and compared them among seven shark species occurring on the continental shelf edge and slope in the GoM at depths ranging from 200 to 2000 m. We also investigated the relationship between blood parameters and depth as well as proximity to the oil spill site. The deep-sea sharks examined here exhibited variability in blood chemistry associated with the secondary stress response, with values falling within published records for previously studied elasmobranchs. Results suggested that there is greater relative physiological stress in shallower-dwelling sharks as well as smaller-bodied sharks. Further, the rate of core temperature warming was fastest in smaller bodied sharks, which likely contributes to greater physiological stress. The core temperatures of the larger-bodied, deeper-dwelling species were not altered as drastically as the smaller-bodied sharks after being hauled to the surface. Any chronic physiological effects of the oil spill were not detectable as there were no relevant correlations between blood chemistry metrics and proximity to the DWH oil spill site.

A multi-taxonomic framework for assessing relative petrochemical vulnerability of marine biodiversity in the Gulf of Mexico.

A fundamental understanding of the impact of petrochemicals and other stressors on marine biodiversity is critical for effective management, restoration, recovery, and mitigation initiatives. As species-specific information on levels of petrochemical exposure and toxicological response are lacking for the majority of marine species, a trait-based assessment to rank species vulnerabilities to petrochemical activities in the Gulf of Mexico can provide a more comprehensive and effective means to prioritize species, habitats, and ecosystems for improved management, restoration and recovery. To initiate and standardize this process, we developed a trait-based framework, applicable to a wide range of vertebrate and invertebrate species, that can be used to rank relative population vulnerabilities of species to petrochemical activities in the Gulf of Mexico. Through expert consultation, 18 traits related to likelihood of exposure, individual sensitivity, and population resilience were identified and defined. The resulting multi-taxonomic petrochemical vulnerability framework can be adapted and applied to a wide variety of species groups and geographic regions. Additional recommendations and guidance on the application of the framework to rank species vulnerabilities under specific petrochemical exposure scenarios, management needs or data limitations are also discussed.

Mercury bioaccumulation in tilefish from the northeastern Gulf of Mexico 2 years after the Deepwater Horizon oil spill: Insights from Hg, C, N and S stable isotopes.

Mercury (Hg) concentration in fish of the Gulf of the Mexico (GoM) is a major concern due to the importance of the GoM for U.S. fisheries. The Deepwater Horizon (DWH) oil spill in April 2010 in the northern GoM resulted in large amounts of oil and dispersant released to the water column, which potentially modified Hg bioaccumulation patterns in affected areas. We measured Hg species (methylmercury (MMHg) and inorganic Hg (IHg)) concentrations, and light (C, N and S) and Hg stable isotopes in muscle and liver tissues from tilefish (Lopholatilus chamaleonticeps) sampled in 2012 and 2013 along the shelf break of the northeastern GoM. Fish located close to the mouth of the Mississippi River (MR) and northwest of the DWH well-head (47 km) showed significantly lower Hg levels in muscle and liver than fish located further northeast of the DWH (>109 km), where 98% of tilefish had Hg levels in the muscle above US consumption advisory thresholds (50% for tilefish close to the DWH). Differences in light and Hg stable isotopes signatures were observed between these two areas, showing higher δ15N, and lower δ202Hg, Δ199Hg and δ34S in fish close to the DWH/MR. This suggests that suspended particles from the MR reduces Hg bioavailability at the base of the GoM food chains. This phenomenon can be locally enhanced by the DWH that resulted in increased particles in the water column as evidenced by the marine snow layer in the sediments. On the other hand, freshly deposited Hg associated with organic matter in more oligotrophic marine waters enhanced Hg bioaccumulation in local food webs. Comparing Hg isotopic composition in liver and muscle of fish indicates specific metabolic response in fish having accumulated high levels of MMHg.

Squalus clarkae sp. nov., a new dogfish shark from the Northwest Atlantic and Gulf of Mexico, with comments on the Squalus mitsukurii species complex.

Sharks of the genus Squalus have slow reproductive rates coupled with low genetic diversity, as is typical of deep-water sharks, making this group slow to rebound from depletion due to overfishing. The number of species within Squalus has been expanding recently due to increased attention on taxonomic revision, and a growing research focus on little-known deep-water sharks in general. Here we use genetics and morphology to describe a new species of dogfish shark, Squalus clarkae sp. nov. from the Gulf of Mexico (GoM) which replaces Squalus mitsukurii in this region, and place it in the context of congeners from the Atlantic and elsewhere. Previously, S. clarkae sp. nov. was considered a part of the Squalus mitsukurii species complex, a group of closely related but distinct species. We sequenced the mitochondrial cytochrome oxidase I and the NADH Dehydrogenase II gene of S. mitsukurii from the type location in Japan, S. clarkae sp. nov. from the GoM, as well as three closely related species (S. cubensis, S. blainville, and S. megalops) and S. cf. mitsukurii from Brazil. Squalus clarkae sp. nov. is genetically distinct from other species with significant statistical support (>98.6% bootstrap support/posterior probability), and 2.8% divergent from S. mitsukurii in the type location of Japan. Morphological estimates also revealed differences between S. clarkae sp. nov., S. mitsukurii, and other Atlantic Squalus species, with S. clarkae sp. nov. exhibiting a longer body, smaller interorbital space, shorter caudal fin, and a differently-proportioned first dorsal fin. In general, dogfish sharks in the Atlantic and GoM are characterized by similar but distinct morphology, significant genetic variation, and small species ranges.

Origins of the Greenland shark (Somniosus microcephalus): Impacts of ice-olation and introgression.

Herein, we use genetic data from 277 sleeper sharks to perform coalescent-based modeling to test the hypothesis of early Quaternary emergence of the Greenland shark (Somniosus microcephalus) from ancestral sleeper sharks in the Canadian Arctic-Subarctic region. Our results show that morphologically cryptic somniosids S. microcephalus and Somniosus pacificus can be genetically distinguished using combined mitochondrial and nuclear DNA markers. Our data confirm the presence of genetically admixed individuals in the Canadian Arctic and sub-Arctic, and temperate Eastern Atlantic regions, suggesting introgressive hybridization upon secondary contact following the initial species divergence. Conservative substitution rates fitted to an Isolation with Migration (IM) model indicate a likely species divergence time of 2.34 Ma, using the mitochondrial sequence DNA, which in conjunction with the geographic distribution of admixtures and Pacific signatures likely indicates speciation associated with processes other than the closing of the Isthmus of Panama. This time span coincides with further planetary cooling in the early Quaternary period followed by the onset of oscillating glacial-interglacial cycles. We propose that the initial S. microcephalus-S. pacificus split, and subsequent hybridization events, were likely associated with the onset of Pleistocene glacial oscillations, whereby fluctuating sea levels constrained connectivity among Arctic oceanic basins, Arctic marginal seas, and the North Atlantic Ocean. Our data demonstrates support for the evolutionary consequences of oscillatory vicariance via transient oceanic isolation with subsequent secondary contact associated with fluctuating sea levels throughout the Quaternary period-which may serve as a model for the origins of Arctic marine fauna on a broad taxonomic scale.

Validation of a portable, waterproof blood pH analyser for elasmobranchs.

Quantifying changes in blood chemistry in elasmobranchs can provide insights into the physiological insults caused by anthropogenic stress, and can ultimately inform conservation and management strategies. Current methods for analysing elasmobranch blood chemistry in the field are often costly and logistically challenging. We compared blood pH values measured using a portable, waterproof pH meter (Hanna Instruments HI 99161) with blood pH values measured by an i-STAT system (CG4+ cartridges), which was previously validated for teleost and elasmobranch fishes, to gauge the accuracy of the pH meter in determining whole blood pH for the Cuban dogfish (Squalus cubensis) and lemon shark (Negaprion brevirostris). There was a significant linear relationship between values derived via the pH meter and the i-STAT for both species across a wide range of pH values and temperatures (Cuban dogfish: 6.8-7.1 pH 24-30°C; lemon sharks: 7.0-7.45 pH 25-31°C). The relative error in the pH meter's measurements was ~±2.7%. Using this device with appropriate correction factors and consideration of calibration temperatures can result in both a rapid and accurate assessment of whole blood pH, at least for the two elasmobranch species examined here. Additional species should be examined in the future across a wide range of temperatures to determine whether correction factors are universal.

Depth as a driver of evolution in the deep sea: Insights from grenadiers (Gadiformes: Macrouridae) of the genus Coryphaenoides.

Here we consider the role of depth as a driver of evolution in a genus of deep-sea fishes. We provide a phylogeny for the genus Coryphaenoides (Gadiformes: Macrouridae) that represents the breadth of habitat use and distributions for these species. In our consensus phylogeny species found at abyssal depths (>4000m) form a well-supported lineage, which interestingly also includes two non-abyssal species, C. striaturus and C. murrayi, diverging from the basal node of that lineage. Biogeographic analyses suggest the genus may have originated in the Southern and Pacific Oceans where contemporary species diversity is highest. The abyssal lineage seems to have arisen secondarily and likely originated in the Southern/Pacific Oceans but diversification of this lineage occurred in the Northern Atlantic Ocean. All abyssal species are found in the North Atlantic with the exception of C. yaquinae in the North Pacific and C. filicauda in the Southern Ocean. Abyssal species tend to have broad depth ranges and wide distributions, indicating that the stability of the deep oceans and the ability to live across wide depths may promote population connectivity and facilitate large ranges. We also confirm that morphologically defined subgenera do not agree with our phylogeny and that the Giant grenadier (formerly Albatrossia pectoralis) belongs to Coryphaenoides, indicating that a taxonomic revision of the genus is needed. We discuss the implications of our findings for understanding the radiation and diversification of this genus, and the likely role of adaptation to the abyss.

Critical assessment and ramifications of a purported marine trophic cascade.

When identifying potential trophic cascades, it is important to clearly establish the trophic linkages between predators and prey with respect to temporal abundance, demographics, distribution, and diet. In the northwest Atlantic Ocean, the depletion of large coastal sharks was thought to trigger a trophic cascade whereby predation release resulted in increased cownose ray abundance, which then caused increased predation on and subsequent collapse of commercial bivalve stocks. These claims were used to justify the development of a predator-control fishery for cownose rays, the "Save the Bay, Eat a Ray" fishery, to reduce predation on commercial bivalves. A reexamination of data suggests declines in large coastal sharks did not coincide with purported rapid increases in cownose ray abundance. Likewise, the increase in cownose ray abundance did not coincide with declines in commercial bivalves. The lack of temporal correlations coupled with published diet data suggest the purported trophic cascade is lacking the empirical linkages required of a trophic cascade. Furthermore, the life history parameters of cownose rays suggest they have low reproductive potential and their populations are incapable of rapid increases. Hypothesized trophic cascades should be closely scrutinized as spurious conclusions may negatively influence conservation and management decisions.

Feeding biomechanics of the cownose ray, Rhinoptera bonasus, over ontogeny.

Growth affects the performance of structure, so the pattern of growth must influence the role of a structure and an organism. Because animal performance is linked to morphological specialization, ontogenetic change in size may influence an organism's biological role. High bite force generation is presumably selected for in durophagous taxa. Therefore, these animals provide an excellent study system for investigating biomechanical consequences of growth on performance. An ontogenetic series of 27 cownose rays (Rhinoptera bonasus) were dissected in order to develop a biomechanical model of the feeding mechanism, which was then compared with bite forces measured from live rays. Mechanical advantage of the feeding apparatus was generally conserved throughout ontogeny, while an increase in the mass and cross-sectional area of the jaw adductors resulted in allometric gains in bite force generation. Of primary importance to forceful biting in this taxon is the use of a fibrocartilaginous tendon associated with the insertion of the primary jaw adductor division. This tendon may serve to redirect muscle forces anteriorly, transmitting them within the plane of biting. Measured bite forces obtained through electrostimulation of the jaw adductors in live rays were higher than predicted, possibly due to differences in specific tension of actual batoid muscle and that used in the model. Mass-specific bite forces in these rays are the highest recorded for elasmobranchs. Cownose rays exemplify a species that, through allometric growth of bite performance and morphological novelties, have expanded their ecological performance over ontogeny.

A re-evaluation of the size of the white shark (Carcharodon carcharias) population off California, USA.

White sharks are highly migratory and segregate by sex, age and size. Unlike marine mammals, they neither surface to breathe nor frequent haul-out sites, hindering generation of abundance data required to estimate population size. A recent tag-recapture study used photographic identifications of white sharks at two aggregation sites to estimate abundance in "central California" at 219 mature and sub-adult individuals. They concluded this represented approximately one-half of the total abundance of mature and sub-adult sharks in the entire eastern North Pacific Ocean (ENP). This low estimate generated great concern within the conservation community, prompting petitions for governmental endangered species designations. We critically examine that study and find violations of model assumptions that, when considered in total, lead to population underestimates. We also use a Bayesian mixture model to demonstrate that the inclusion of transient sharks, characteristic of white shark aggregation sites, would substantially increase abundance estimates for the adults and sub-adults in the surveyed sub-population. Using a dataset obtained from the same sampling locations and widely accepted demographic methodology, our analysis indicates a minimum all-life stages population size of >2000 individuals in the California subpopulation is required to account for the number and size range of individual sharks observed at the two sampled sites. Even accounting for methodological and conceptual biases, an extrapolation of these data to estimate the white shark population size throughout the ENP is inappropriate. The true ENP white shark population size is likely several-fold greater as both our study and the original published estimate exclude non-aggregating sharks and those that independently aggregate at other important ENP sites. Accurately estimating the central California and ENP white shark population size requires methodologies that account for biases introduced by sampling a limited number of sites and that account for all life history stages across the species' range of habitats.

Myological variability in a decoupled skeletal system: batoid cranial anatomy.

Chondrichthyans (sharks, batoids, and chimaeras) have simple feeding mechanisms owing to their relatively few cranial skeletal elements. However, the indirect association of the jaws to the cranium (euhyostylic jaw suspension) has resulted in myriad cranial muscle rearrangements of both the hyoid and mandibular elements. We examined the cranial musculature of an abbreviated phylogenetic representation of batoid fishes, including skates, guitarfishes and with a particular focus on stingrays. We identified homologous muscle groups across these taxa and describe changes in gross morphology across developmental and functional muscle groups, with the goal of exploring how decoupling of the jaws from the skull has effected muscular arrangement. In particular, we focus on the cranial anatomy of durophagous and nondurophagous batoids, as the former display marked differences in morphology compared to the latter. Durophagous stingrays are characterized by hypertrophied jaw adductors, reliance on pennate versus fusiform muscle fiber architecture, tendinous rather than aponeurotic muscle insertions, and an overall reduction in mandibular kinesis. Nondurophagous stingrays have muscles that rely on aponeurotic insertions onto the skeletal structure, and display musculoskeletal specialization for jaw protrusion and independent lower jaw kinesis, relative to durophagous stingrays. We find that among extant chondrichthyans, considerable variation exists in the hyoid and mandibular muscles, slightly less so in hypaxial muscles, whereas branchial muscles are overwhelmingly conserved. As chondrichthyans occupy a position sister to all other living gnathostomes, our understanding of the structure and function of early vertebrate feeding systems rests heavily on understanding chondrichthyan cranial anatomy. Our findings highlight the incredible variation in muscular complexity across chondrichthyans in general and batoids in particular.