Sharks occupy diverse ecological niches and play critical roles in marine ecosystems, often acting as apex predators. They are considered a slow-evolving lineage and have been suggested to exhibit exceptionally low cancer rates. These two features could be explained by a low nuclear mutation rate. Here, we provide a direct estimate of the nuclear mutation rate in the epaulette shark (Hemiscyllium ocellatum). We generate a high-quality reference genome, and resequence the whole genomes of parents and nine offspring to detect de novo mutations. Using stringent criteria, we estimate a mutation rate of 7×10-10 per base pair, per generation. This represents one of the lowest directly estimated mutation rates for any vertebrate clade, indicating that this basal vertebrate group is indeed a slowly evolving lineage whose ability to restore genetic diversity following a sustained population bottleneck may be hampered by a low mutation rate.

Interactions between pelagic thresher sharks (Alopias pelagicus) and cleaner wrasse were investigated at a seamount in the Philippines. Cleaning associations between sharks and teleosts are poorly understood, but the observable interactions seen at this site may explain why these mainly oceanic sharks regularly venture into shallow coastal waters where they are vulnerable to disturbance from human activity. From 1,230 hours of observations recorded by remote video camera between July 2005 and December 2009, 97 cleaner-thresher shark events were analyzed, 19 of which were interrupted. Observations of pelagic thresher sharks interacting with cleaners at the seamount were recorded at all times of day but their frequency declined gradually from morning until evening. Cleaners showed preferences for foraging on specific areas of a thresher shark's body. For all events combined, cleaners were observed to conduct 2,757 inspections, of which 33.9% took place on the shark's pelvis, 23.3% on the pectoral fins, 22.3% on the caudal fin, 8.6% on the body, 8.3% on the head, 2.1% on the dorsal fin, and 1.5% on the gills respectively. Cleaners did not preferentially inspect thresher sharks by time of day or by shark sex, but there was a direct correlation between the amount of time a thresher shark spent at a cleaning station and the number of inspections it received. Thresher shark clients modified their behavior by "circular-stance-swimming," presumably to facilitate cleaner inspections. The cleaner-thresher shark association reflected some of the known behavioral trends in the cleaner-reef teleost system since cleaners appeared to forage selectively on shark clients. Evidence is mounting that in addition to acting as social refuges and foraging grounds for large visiting marine predators, seamounts may also support pelagic ecology by functioning as cleaning stations for oceanic sharks and rays.

Sharks are one of the most threatened groups of marine animals, as high exploitation rates coupled with low resilience to fishing pressure have resulted in population declines worldwide. Designing conservation strategies for this group depends on basic knowledge of the geographic distribution and diversity of known species. So far, this information has been fragmented and incomplete. Here, we have synthesized the first global shark diversity pattern from a new database of published sources, including all 507 species described at present, and have identified hotspots of shark species richness, functional diversity and endemicity from these data. We have evaluated the congruence of these diversity measures and demonstrate their potential use in setting priority areas for shark conservation. Our results show that shark diversity across all species peaks on the continental shelves and at mid-latitudes (30-40 degrees N and S). Global hotspots of species richness, functional diversity and endemicity were found off Japan, Taiwan, the East and West coasts of Australia, Southeast Africa, Southeast Brazil and Southeast USA. Moreover, some areas with low to moderate species richness such as Southern Australia, Angola, North Chile and Western Continental Europe stood out as places of high functional diversity. Finally, species affected by shark finning showed different patterns of diversity, with peaks closer to the Equator and a more oceanic distribution overall. Our results show that the global pattern of shark diversity is uniquely different from land, and other well-studied marine taxa, and may provide guidance for spatial approaches to shark conservation. However, similar to terrestrial ecosystems, protected areas based on hotspots of diversity and endemism alone would provide insufficient means for safeguarding the diverse functional roles that sharks play in marine ecosystems.

Estuarine habitats are frequently used as nurseries by elasmobranch species for their protection and abundant resources; however, global climate change is increasing the frequency and severity of environmental challenges in these estuaries that may negatively affect elasmobranch physiology. Hyposmotic events are particularly challenging for marine sharks that osmoconform, and species-specific tolerances are not well known. Therefore, we sought to determine the effects of an acute (48 h) ecologically relevant hyposmotic event (25.8 ppt) on the physiology of two juvenile shark species, namely the school shark (Galeorhinus galeus), listed by the Australian Environmental Protection and Biodiversity Conservation Act as 'conservation dependent', and the gummy shark (Mustelus antarcticus), from the Pittwater Estuary (Australia). In both species, we observed a decrease in plasma osmolality brought about by selective losses of NaCl, urea and trimethylamine N-oxide, as well as decreases in haemoglobin, haematocrit and routine oxygen consumption. Heat-shock protein levels varied between species during the exposure, but we found no evidence of protein damage in any of the tissues tested. Although both species seemed to be able to cope with this level of osmotic challenge, overall the school sharks exhibited higher gill Na+/K+-ATPase activity and ubiquitin concentrations in routine and experimental conditions, a larger heat-shock protein response and a smaller decrease in routine oxygen consumption during the hyposmotic exposure, suggesting that there are species-specific responses that could potentially affect their ability to withstand longer or more severe changes in salinity. Emerging evidence from acoustic monitoring of sharks has indicated variability in the species found in the Pittwater Estuary during hyposmotic events, and together, our data may help to predict species abundance and distribution in the face of future global climate change.

In Southeast Asia, elasmobranchs are particularly threatened. We synthesized knowledge from the peer-reviewed and gray literature on elasmobranchs in the region, including their fisheries, status, trade, biology, and management. We found that 59% of assessed species are threatened with extinction and 72.5% are in decline; rays were more threatened than sharks. Research and conservation is complicated by the socioeconomic contexts of the countries, geopolitical issues in the South China Sea, and the overcapacity and multispecies nature of fisheries that incidentally capture elasmobranchs. The general paucity of data, funds, personnel, and enforcement hinders management. Reduced capacity in the general fishery sector and marine protected areas of sufficient size (for elasmobranchs and local enforcement capabilities) are among recommendations to strengthen conservation.

Ancient cartilaginous vertebrates, such as sharks, skates and rays, possess specialized electrosensory organs that detect weak electric fields and relay this information to the central nervous system1-4. Sharks exploit this sensory modality for predation, whereas skates may also use it to detect signals from conspecifics 5 . Here we analyse shark and skate electrosensory cells to determine whether discrete physiological properties could contribute to behaviourally relevant sensory tuning. We show that sharks and skates use a similar low threshold voltage-gated calcium channel to initiate cellular activity but use distinct potassium channels to modulate this activity. Electrosensory cells from sharks express specially adapted voltage-gated potassium channels that support large, repetitive membrane voltage spikes capable of driving near-maximal vesicular release from elaborate ribbon synapses. By contrast, skates use a calcium-activated potassium channel to produce small, tunable membrane voltage oscillations that elicit stimulus-dependent vesicular release. We propose that these sensory adaptations support amplified indiscriminate signal detection in sharks compared with selective frequency detection in skates, potentially reflecting the electroreceptive requirements of these elasmobranch species. Our findings demonstrate how sensory systems adapt to suit the lifestyle or environmental niche of an animal through discrete molecular and biophysical modifications.

Chiloscyllium, commonly called bamboo shark, can be found inhabiting the waters of the Indo-West Pacific around East Asian countries such as Malaysia, Myanmar, Thailand, Singapore, and Indonesia. The International Union for Conservation of Nature (IUCN) Red List has categorized them as nearly threatened sharks out of their declining population status due to overexploitation. A molecular study was carried out to portray the systematic relationships within Chiloscyllium species using 12S rRNA and cytochrome b gene sequences. Maximum parsimony and Bayesian were used to reconstruct their phylogeny trees. A total of 381 bp sequences' lengths were successfully aligned in the 12S rRNA region, with 41 bp sites being parsimony-informative. In the cytochrome b region, a total of 1120 bp sites were aligned, with 352 parsimony-informative characters. All analyses yield phylogeny trees on which C. indicum has close relationships with C. plagiosum. C. punctatum is sister taxon to both C. indicum and C. plagiosum while C. griseum and C. hasseltii formed their own clade as sister taxa. These Chiloscyllium classifications can be supported by some morphological characters (lateral dermal ridges on the body, coloring patterns, and appearance of hypobranchials and basibranchial plate) that can clearly be used to differentiate each species.

In the era of "Anthropocene defaunation," large species are often no longer detected in habitats where they formerly occurred. However, it is unclear whether this apparent missing, or "dark," diversity of megafauna results from local species extirpations or from failure to detect elusive remaining individuals. We find that despite two orders of magnitude less sampling effort, environmental DNA (eDNA) detects 44% more shark species than traditional underwater visual censuses and baited videos across the New Caledonian archipelago (south-western Pacific). Furthermore, eDNA analysis reveals the presence of previously unobserved shark species in human-impacted areas. Overall, our results highlight a greater prevalence of sharks than described by traditional survey methods in both impacted and wilderness areas. This indicates an urgent need for large-scale eDNA assessments to improve monitoring of threatened and elusive megafauna. Finally, our findings emphasize the need for conservation efforts specifically geared toward the protection of elusive, residual populations.

Multiple paternity appears to be a common trait of elasmobranch mating systems, with its occurrence likely driven by convenience, due to females seeking to minimize the stress of male harassment. Here we use molecular markers to analyse the frequency of multiple paternity in two related viviparous sharks, Mustelus mustelus and Mustelus punctulatus. We first applied molecular methods to assign pregnant females, embryos and additional reference adults (N = 792) to one of the two species. Paternity analysis was performed using a total of 9 polymorphic microsatellites on 19 females and 204 embryos of M. mustelus, and on 13 females and 303 embryos of M. punctulatus. Multiple paternity occurs in both species, with 47% of M. mustelus and 54% of M. punctulatus litters sired by at least two fathers. Female fecundity is not influenced by multiple mating and in 56% of polyandrous litters paternity is skewed, with one male siring most of the pups. Genetic analyses also revealed hybridization between the two species, with a M. punctulatus female bearing pups sired by a M. mustelus male. The frequency of polyandrous litters in these species is consistent with aspects of their reproductive biology, such as synchronous ovulation and possible occurrence of breeding aggregations.

Coastal and estuarine systems provide critical shark habitats due to their relatively high productivity and shallow, protected waters. The young (neonates, young-of-the-year, and juveniles) of many coastal shark species occupy a diverse range of habitats and areas where they experience environmental variability, including acute and seasonal shifts in local salinities and temperatures. Although the location and functioning of essential shark habitats has been a focus in recent shark research, there is a paucity of data from the South Pacific. In this study, we document the temporal and spatial distribution, age class composition, and environmental parameters of young bull sharks (Carcharhinus leucas) in the Rewa, Sigatoka, and Navua Rivers, Fiji's three largest riverine systems. One hundred and seventy-two young bull sharks were captured in fisheries-independent surveys from January 2016 to April 2018. The vast majority of the captures were neonates. Seasonality in patterns of occurrence of neonate individuals suggests a defined parturition period during summer. Environmental parameters between the Rewa and the Sigatoka River differed significantly, as did the recorded young bull sharks abundance. According to the surveys, young bull sharks occur in all three rivers with the Rewa River likely representing essential habitat for newly born bull sharks. These results enhance the understanding of bull shark ecology in Fiji and provide a scientific basis for the implementation of local conservation strategies that contribute to the protection of critical habitats.

No-take marine protected areas (MPAs) are a commonly applied tool to reduce human fishing impacts on marine and coastal ecosystems. However, conservation outcomes of MPAs for mobile and long-lived predators such as sharks are highly variable. Here, we use empirical animal tracking data from 459 individual sharks and baited remote underwater video surveys undertaken in 36 countries to construct an empirically supported individual-based model that estimates the conservation effectiveness of MPAs for five species of coral reef-associated sharks (Triaenodon obesus, Carcharhinus melanopterus, Carcharhinus amblyrhynchos, Carcharhinus perezi, and Ginglymostoma cirratum). We demonstrate how species-specific individual movement traits can contribute to fishing mortality of sharks found within MPAs as they move outside to adjacent fishing grounds. We discovered that the world's officially recorded coral reef-based managed areas (with a median width of 9.4 km) would need to be enforced as strict no-take MPAs and up to 5 times larger to expect protection of the majority of individuals of the five investigated reef shark species. The magnitude of this effect depended on local abundances and fishing pressure, with MPAs required to be 1.6-2.6 times larger to protect the same number of Atlantic and Caribbean species, which occur at lower abundances than similar species in the western Pacific. Furthermore, our model was used to quantify partially substantial reductions (>50%) in fishing mortality resulting from small increases in MPA size, allowing us to bridge a critical gap between traditional conservation planning and fisheries management. Overall, our results highlight the challenge of relying on abundance data alone to ensure that estimates of shark conservation impacts of MPAs follow the precautionary approach.

The evolution of gigantism in extinct otodontid sharks was paralleled by a series of drastic modifications in their dentition including widening of the crowns, loss of lateral cusplets, and acquisition of serrated cutting edges. These traits have generally been interpreted as key functional features that enabled the transition from piscivory to more energetic diets based on marine mammals, ultimately leading to the evolution of titanic body sizes in the most recent forms (including the emblematic Otodus megalodon). To investigate this hypothesis, we evaluate the biomechanics of the anterior, lateral, and posterior teeth of five otodontid species under different loading conditions by using two-dimensional finite element analysis. Stress distribution patterns are remarkably similar among all models under puncture and draw (i.e., when subjected to vertical and lateral forces, respectively). Contrary to expectation, higher average stress values are detected under both loading scenarios in more recent species. Altogether, this suggests little correlation between tooth morphology and key aspects of biomechanical behaviour in otodontids, making it difficult to frame the morphological trend of their dentitions within an adaptive scenario. We propose that this pattern most likely emerged as a non-functional by-product of heterochronic processes driven by selection towards larger body sizes.

In Australian and New Zealand waters, current knowledge on white shark (Carcharodon carcharias) movement ecology is based on individual tracking studies using relatively small numbers of tags. These studies describe a species that occupies highly variable and complex habitats. However, uncertainty remains as to whether the proposed movement patterns are representative of the wider population. Here, we tagged 103 immature Australasian white sharks (147-350 cm fork length) with both acoustic and satellite transmitters to expand our current knowledge of population linkages, spatiotemporal dynamics and coastal habitats. Eighty-three sharks provided useable data. Based on individual tracking periods of up to 5 years and a total of 2,865 days of tracking data, we were able to characterise complex movement patterns over ~45° of latitude and ~72° of longitude and distinguish regular/recurrent patterns from occasional/exceptional migration events. Shark movements ranged from Papua New Guinea to sub-Antarctic waters and to Western Australia, highlighting connectivity across their entire Australasian range. Results over the 12-year study period yielded a comprehensive characterisation of the movement ecology of immature Australasian white sharks across multiple spatial scales and substantially expanded the body of knowledge available for population assessment and management.

Sharks play essential roles in ocean food webs and human culture, but also face population declines worldwide due to human activity. The relationship between sharks and the microbes on and in the shark body is unclear, despite research on other animals showing the microbiome as intertwined with host physiology, immunity, and ecology. Research on shark-microbe interactions faces the significant challenge of sampling the largest and most elusive shark species. We leveraged a unique sampling infrastructure to compare the microbiomes of two apex predators, the white (Carcharodon carcharias) and tiger shark (Galeocerdo cuvier), to those of the filter-feeding whale shark (Rhincodon typus), allowing us to explore the effects of feeding mode on intestinal microbiome diversity and metabolic function, and environmental exposure on the diversity of microbes external to the body (on the skin, gill).

Overfishing is the most significant threat facing sharks and rays. Given the growth in consumption of seafood, combined with the compounding effects of habitat loss, climate change, and pollution, there is a need to identify recovery paths, particularly in poorly managed and poorly monitored fisheries. Here, we document conservation through fisheries management success for 11 coastal sharks in US waters by comparing population trends through a Bayesian state-space model before and after the implementation of the 1993 Fisheries Management Plan for Sharks. We took advantage of the spatial and temporal gradients in fishing exposure and fisheries management in the Western Atlantic to analyze the effect on the Red List status of all 26 wide-ranging coastal sharks and rays. We show that extinction risk was greater where fishing pressure was higher, but this was offset by the strength of management engagement (indicated by strength of National and Regional Plan of Action for sharks and rays). The regional Red List Index (which tracks changes in extinction risk through time) declined in all regions until the 1980s but then improved in the North and Central Atlantic such that the average extinction risk is currently half that in the Southwest. Many sharks and rays are wide ranging, and successful fisheries management in one country can be undone by poorly regulated or unregulated fishing elsewhere. Our study underscores that well-enforced, science-based management of carefully monitored fisheries can achieve conservation success, even for slow-growing species.

Spatial structuring and segregation by sex and size is considered to be an intrinsic attribute of shark populations. These spatial patterns remain poorly understood, particularly for oceanic species such as blue shark (Prionace glauca), despite its importance for the management and conservation of this highly migratory species. This study presents the results of a long-term electronic tagging experiment to investigate the migratory patterns of blue shark, to elucidate how these patterns change across its life history and to assess the existence of a nursery area in the central North Atlantic. Blue sharks belonging to different life stages (n = 34) were tracked for periods up to 952 days during which they moved extensively (up to an estimated 28.139 km), occupying large parts of the oceanic basin. Notwithstanding a large individual variability, there were pronounced differences in movements and space use across the species' life history. The study provides strong evidence for the existence of a discrete central North Atlantic nursery, where juveniles can reside for up to at least 2 years. In contrast with previously described nurseries of coastal and semi-pelagic sharks, this oceanic nursery is comparatively vast and open suggesting that shelter from predators is not its main function. Subsequently, male and female blue sharks spatially segregate. Females engage in seasonal latitudinal migrations until approaching maturity, when they undergo an ontogenic habitat shift towards tropical latitudes. In contrast, juvenile males generally expanded their range southward and apparently displayed a higher degree of behavioural polymorphism. These results provide important insights into the spatial ecology of pelagic sharks, with implications for the sustainable management of this heavily exploited shark, especially in the central North Atlantic where the presence of a nursery and the seasonal overlap and alternation of different life stages coincides with a high fishing mortality.

Squaliform sharks represent approximately 27 % of extant shark diversity, comprising more than 130 species with a predominantly deep-dwelling lifestyle. Many Squaliform species are highly specialized, including some that are bioluminescent, a character that is reported exclusively from Squaliform sharks within Chondrichthyes. The interfamiliar relationships within the order are still not satisfactorily resolved. Herein we estimate the phylogenetic interrelationships of a generic level sampling of "squaloid" sharks and closely related taxa using aligned sequences derived from a targeted gene capture approach. The resulting phylogenetic estimate is further used to evaluate the age of first occurrence of bioluminescence in Squaliformes.

All vertebrates initially feed their offspring using yolk reserves. In some live-bearing species these yolk reserves may be supplemented with extra nutrition via a placenta. Sharks belonging to the Carcharhinidae family are all live-bearing, and with the exception of the tiger shark (Galeocerdo cuvier), develop placental connections after exhausting yolk reserves. Phylogenetic relationships suggest the lack of placenta in tiger sharks is due to secondary loss. This represents a dramatic shift in reproductive strategy, and is likely to have left a molecular footprint of positive selection within the genome.

Teeth are an integral component of feeding ecology, with a clear link between tooth morphology and diet, as without suitable dentition prey cannot be captured nor broken down for consumption. Bull sharks, Carcharhinus leucas, undergo an ontogenetic niche shift from freshwater to marine habitats, which raises the question: does tooth morphology change with ontogeny? Tooth shape, surface area and thickness were measured using both morphometrics and elliptic Fourier analysis to determine if morphology varied with position in the jaw and if there was an ontogenetic change concordant with this niche shift. Significant ontogenetic differences in tooth morphology as a function of position in the jaw and shark total length were found, with upper and lower jaws of bull sharks presenting two different tooth morphologies. Tooth shape and thickness fell into two groupings, anterior and posterior, in both the upper and lower jaws. Tooth surface area, however, indicated three groupings, mesial, intermediate and distal, in both the upper and lower jaws. While tooth morphology changed significantly with size, showing an inflection at sharks of 135 cm total length, each morphological aspect retained the same tooth groupings throughout. These ontogenetic differences in tooth morphologies reflect tooth strength, prey handling and heterodonty.

The rapid expansion of human activities threatens ocean-wide biodiversity. Numerous marine animal populations have declined, yet it remains unclear whether these trends are symptomatic of a chronic accumulation of global marine extinction risk. We present the first systematic analysis of threat for a globally distributed lineage of 1,041 chondrichthyan fishes-sharks, rays, and chimaeras. We estimate that one-quarter are threatened according to IUCN Red List criteria due to overfishing (targeted and incidental). Large-bodied, shallow-water species are at greatest risk and five out of the seven most threatened families are rays. Overall chondrichthyan extinction risk is substantially higher than for most other vertebrates, and only one-third of species are considered safe. Population depletion has occurred throughout the world's ice-free waters, but is particularly prevalent in the Indo-Pacific Biodiversity Triangle and Mediterranean Sea. Improved management of fisheries and trade is urgently needed to avoid extinctions and promote population recovery. DOI: http://dx.doi.org/10.7554/eLife.00590.001.