Seasonally mediated niche partitioning in a vertically compressed pelagic predator guild.

Niche partitioning among closely related, sympatric species is a fundamental concept in ecology, and its mechanisms are of broad interest for understanding ecosystem functioning and predicting the impacts of human-driven environmental change. However, identifying mechanisms by which top marine predators partition available resources has been especially challenging given the difficulty of quantifying resource use of large pelagic animals. In the eastern tropical Pacific (ETP), three large, highly mobile and ecologically similar pelagic predators (blue marlin (Makaira nigricans), black marlin (Istiompax indica) and sailfish (Istiophorus platypterus)) coexist in a vertically compressed habitat. To evaluate each species' ecological niche, we leveraged a decade of recreational fisheries data, multi-year satellite tracking with high-resolution dive data, and stable isotope analysis. Fishery interaction and telemetry-based three-dimensional seasonal utilization distributions suggested high spatial and temporal overlap among species; however, seasonal and diel variability in diving behaviour produced spatial partitioning, leading to low trophic overlap among species. Expanding oxygen minimum zones will reduce the available vertical habitat within predator guilds, likely leading to increases in interspecific competition. Thus, understanding the mechanisms of habitat partitioning among predators in the vertically compressed ETP can provide insight into how predators in other ocean regions may respond to vertically limited habitats.

Patrolling the border: Billfish exploit the hypoxic boundary created by the world's largest oxygen minimum zone.

Pelagic predators must contend with low prey densities that are irregularly distributed and dynamic in space and time. Based on satellite imagery and telemetry data, many pelagic predators will concentrate horizontal movements on ephemeral surface fronts-gradients between water masses-because of enhanced local productivity and increased forage fish densities. Vertical fronts (e.g. thermoclines, oxyclines) can be spatially and temporally persistent, and aggregate lower trophic level and diel vertically migrating organisms due to sharp changes in temperature, water density or available oxygen. Thus, vertical fronts represent a stable and potentially energy rich habitat feature for diving pelagic predators but remain little explored in their capacity to enhance foraging opportunities. Here, we use a novel suite of high-resolution biologging data, including in situ derived oxygen saturation and video, to document how two top predators in the pelagic ecosystem exploit the vertical fronts created by the oxygen minimum zone of the eastern tropical Pacific. Prey search behaviour was dependent on dive shape, and significantly increased near the thermocline and hypoxic boundary for blue marlin Makaira nigricans and sailfish Istiophorus platypterus, respectively. Further, we identify a behaviour not yet reported for pelagic predators, whereby the predator repeatedly dives below the thermocline and hypoxic boundary (and by extension, below the prey). We hypothesize this behaviour is used to ambush prey concentrated at the boundaries from below. We describe how habitat fronts created by low oxygen environments can influence pelagic ecosystems, which will become increasingly important to understand in the context of global change and expanding oxygen minimum zones. We anticipate that our findings are shared among many pelagic predators where strong vertical fronts occur, and additional high-resolution tagging is warranted to confirm this.

Elucidating the role of competition in driving spatial and trophic niche patterns in sympatric juvenile sharks.

The coexistence of ecologically and morphologically similar species is often facilitated by the partitioning of ecological niches. While subordinate species can reduce competition with dominant competitors through spatial and/or trophic segregation, empirical support from wild settings, particularly those involving large-bodied taxa in marine ecosystems, are rare. Shark nursery areas provide an opportunity to investigate the mechanisms of coexistence. We used experimental and field studies of sympatric juvenile sharks (blacktip reef shark, Carcharhinus melanopterus; sicklefin lemon shark, Negaprion acutidens) to investigate how competitive ability influenced realized niches at St. Joseph Atoll, Seychelles. Captive trials revealed that sicklefin lemon sharks were dominant over blacktip reef sharks, consistently taking food rewards. In the field, blacktip reef sharks were captured over a broader area than sicklefin lemon sharks, but daily space use of actively tracked sharks showed a high degree of overlap across microhabitats. While stomach contents analysis revealed that blacktip reef shark diets included a broader range of prey items, stable isotope analysis demonstrated significantly higher mean δ13C values for sicklefin lemon sharks, suggesting diverging dietary preferences. Overall, our results matched theoretical predictions of subordinate competitors using a greater range of habitats and displaying broader feeding niches than competitively dominant species. While separating the realized and fundamental niche of marine predators is complicated, we provide evidence that resource partitioning is at least partially driven by interspecific competition.

Genomes of endangered great hammerhead and shortfin mako sharks reveal historic population declines and high levels of inbreeding in great hammerhead.

Despite increasing threats of extinction to Elasmobranchii (sharks and rays), whole genome-based conservation insights are lacking. Here, we present chromosome-level genome assemblies for the Critically Endangered great hammerhead (Sphyrna mokarran) and the Endangered shortfin mako (Isurus oxyrinchus) sharks, with genetic diversity and historical demographic comparisons to other shark species. The great hammerhead exhibited low genetic variation, with 8.7% of the 2.77 Gbp genome in runs of homozygosity (ROH) > 1 Mbp and 74.4% in ROH >100 kbp. The 4.98 Gbp shortfin mako genome had considerably greater diversity and <1% in ROH > 1 Mbp. Both these sharks experienced precipitous declines in effective population size (Ne) over the last 250 thousand years. While shortfin mako exhibited a large historical Ne that may have enabled the retention of higher genetic variation, the genomic data suggest a possibly more concerning picture for the great hammerhead, and a need for evaluation with additional individuals.

Hunting behavior of a solitary sailfish Istiophorus platypterus and estimated energy gain after prey capture.

Foraging behavior and interaction with prey is an integral component of the ecological niche of predators but is inherently difficult to observe for highly mobile animals in the marine environment. Billfishes have been described as energy speculators, expending a large amount of energy foraging, expecting to offset high costs with periodic high energetic gain. Surface-based group feeding of sailfish, Istiophorus platypterus, is commonly observed, yet sailfish are believed to be largely solitary roaming predators with high metabolic requirements, suggesting that individual foraging also represents a major component of predator-prey interactions. Here, we use biologging data and video to examine daily activity levels and foraging behavior, estimate metabolic costs, and document a solitary predation event for a 40 kg sailfish. We estimate a median active metabolic rate of 218.9 ± 70.5 mgO2 kg-1 h-1 which increased to 518.8 ± 586.3 mgO2 kg-1 h-1 during prey pursuit. Assuming a successful predation, we estimate a daily net energy gain of 2.4 MJ (5.1 MJ acquired, 2.7 MJ expended), supporting the energy speculator model. While group hunting may be a common activity used by sailfish to acquire energy, our calculations indicate that opportunistic individual foraging events offer a net energy return that contributes to the fitness of these highly mobile predators.

Global collision-risk hotspots of marine traffic and the world's largest fish, the whale shark.

Marine traffic is increasing globally yet collisions with endangered megafauna such as whales, sea turtles, and planktivorous sharks go largely undetected or unreported. Collisions leading to mortality can have population-level consequences for endangered species. Hence, identifying simultaneous space use of megafauna and shipping throughout ranges may reveal as-yet-unknown spatial targets requiring conservation. However, global studies tracking megafauna and shipping occurrences are lacking. Here we combine satellite-tracked movements of the whale shark, Rhincodon typus, and vessel activity to show that 92% of sharks' horizontal space use and nearly 50% of vertical space use overlap with persistent large vessel (>300 gross tons) traffic. Collision-risk estimates correlated with reported whale shark mortality from ship strikes, indicating higher mortality in areas with greatest overlap. Hotspots of potential collision risk were evident in all major oceans, predominantly from overlap with cargo and tanker vessels, and were concentrated in gulf regions, where dense traffic co-occurred with seasonal shark movements. Nearly a third of whale shark hotspots overlapped with the highest collision-risk areas, with the last known locations of tracked sharks coinciding with busier shipping routes more often than expected. Depth-recording tags provided evidence for sinking, likely dead, whale sharks, suggesting substantial "cryptic" lethal ship strikes are possible, which could explain why whale shark population declines continue despite international protection and low fishing-induced mortality. Mitigation measures to reduce ship-strike risk should be considered to conserve this species and other ocean giants that are likely experiencing similar impacts from growing global vessel traffic.

Three complete mitochondrial genomes of shortfin mako sharks, Isurus oxyrinchus, from the Atlantic and Pacific Oceans.

We present complete mitogenome sequences of three shortfin mako sharks (Isurus oxyrinchus) sampled from the western Pacific, and eastern and western Atlantic oceans. Mitogenome sequence lengths ranged between 16,699 bp and 16,702 bp, and all three mitogenomes contained one non-coding control region, two rRNA genes, 22 tRNA genes, and 13 protein-coding genes. Comparative assessment of five mitogenomes from globally distributed shortfin makos (the current three and two previously published mitogenomes) yielded 98.4% identity, with the protein-coding genes ATP8, ATP6, and ND5 as the most variable regions (sequence identities of 96.4%, 96.5%, and 97.6%, respectively). These mitogenome sequences contribute resources for assessing the genetic population dynamics of this endangered oceanic apex predator.

Genetic population dynamics of the critically endangered scalloped hammerhead shark (Sphyrna lewini) in the Eastern Tropical Pacific.

The scalloped hammerhead shark, Sphyrna lewini, is a Critically Endangered, migratory species known for its tendency to form iconic and visually spectacular large aggregations. Herein, we investigated the population genetic dynamics of the scalloped hammerhead across much of its distribution in the Eastern Tropical Pacific (ETP), ranging from Costa Rica to Ecuador, focusing on young-of-year animals from putative coastal nursery areas and adult females from seasonal aggregations that form in the northern Galápagos Islands. Nuclear microsatellites and partial mitochondrial control region sequences showed little evidence of population structure suggesting that scalloped hammerheads in this ETP region comprise a single genetic stock. Galápagos aggregations of adults were not comprised of related individuals, suggesting that kinship does not play a role in the formation of the repeated, annual gatherings at these remote offshore locations. Despite high levels of fisheries exploitation of this species in the ETP, the adult scalloped hammerheads here showed greater genetic diversity compared with adult conspecifics from other parts of the species' global distribution. A phylogeographic analysis of available, globally sourced, mitochondrial control region sequence data (n = 1818 sequences) revealed that scalloped hammerheads comprise three distinct matrilines corresponding to the three major world ocean basins, highlighting the need for conservation of these evolutionarily unique lineages. This study provides the first view of the genetic properties of a scalloped hammerhead aggregation, and the largest sample size-based investigation of population structure and phylogeography of this species in the ETP to date.

Genomic Assessment of Global Population Structure in a Highly Migratory and Habitat Versatile Apex Predator, the Tiger Shark (Galeocerdo cuvier).

Understanding the population dynamics of highly mobile, widely distributed, oceanic sharks, many of which are overexploited, is necessary to aid their conservation management. We investigated the global population genomics of tiger sharks (Galeocerdo cuvier), a circumglobally distributed, apex predator displaying remarkable behavioral versatility in its diet, habitat use (near coastal, coral reef, pelagic), and individual movement patterns (spatially resident to long-distance migrations). We genotyped 242 tiger sharks from 10 globally distributed locations at more than 2000 single nucleotide polymorphisms. Although this species often conducts massive distance migrations, the data show strong genetic differentiation at both neutral (FST = 0.125-0.144) and candidate outlier loci (FST = 0.570-0.761) between western Atlantic and Indo-Pacific sharks, suggesting the potential for adaptation to the environments specific to these oceanic regions. Within these regions, there was mixed support for population differentiation between northern and southern hemispheres in the western Atlantic, and none for structure within the Indian Ocean. Notably, the results demonstrate a low level of population differentiation of tiger sharks from the remote Hawaiian archipelago compared with sharks from the Indian Ocean (FST = 0.003-0.005, P < 0.01). Given concerns about biodiversity loss and marine ecosystem impacts caused by overfishing of oceanic sharks in the midst of rapid environmental change, our results suggest it imperative that international fishery management prioritize conservation of the evolutionary potential of the highly genetically differentiated Atlantic and Indo-Pacific populations of this unique apex predator. Furthermore, we suggest targeted management attention to tiger sharks in the Hawaiian archipelago based on a precautionary biodiversity conservation perspective.

Spatio-temporal genetic tagging of a cosmopolitan planktivorous shark provides insight to gene flow, temporal variation and site-specific re-encounters.

Migratory movements in response to seasonal resources often influence population structure and dynamics. Yet in mobile marine predators, population genetic consequences of such repetitious behaviour remain inaccessible without comprehensive sampling strategies. Temporal genetic sampling of seasonally recurring aggregations of planktivorous basking sharks, Cetorhinus maximus, in the Northeast Atlantic (NEA) affords an opportunity to resolve individual re-encounters at key sites with population connectivity and patterns of relatedness. Genetic tagging (19 microsatellites) revealed 18% of re-sampled individuals in the NEA demonstrated inter/multi-annual site-specific re-encounters. High genetic connectivity and migration between aggregation sites indicate the Irish Sea as an important movement corridor, with a contemporary effective population estimate (Ne) of 382 (CI = 241-830). We contrast the prevailing view of high gene flow across oceanic regions with evidence of population structure within the NEA, with early-season sharks off southwest Ireland possibly representing genetically distinct migrants. Finally, we found basking sharks surfacing together in the NEA are on average more related than expected by chance, suggesting a genetic consequence of, or a potential mechanism maintaining, site-specific re-encounters. Long-term temporal genetic monitoring is paramount in determining future viability of cosmopolitan marine species, identifying genetic units for conservation management, and for understanding aggregation structure and dynamics.

The complete mitochondrial genome of a gray reef shark, Carcharhinus amblyrhynchos (Carcharhiniformes: Carcharhinidae), from the Western Indian Ocean.

We present the mitochondrial genome sequence of a gray reef shark, Carcharhinus amblyrhynchos (Bleeker 1856), a coral reef associated species. This is the first mitogenome for this species from the western Indian Ocean. The mitogenome is 16,705 bp in length, has 13 protein-coding genes, 22 tRNA genes, 2 rRNA genes and a non-coding control region, and demonstrates a gene arrangement congruent with other shark and most vertebrate species. This mitogenome provides a genomic resource for assisting with population, evolutionary and conservation studies for the gray reef shark, which is increasingly under threat from fisheries.

White shark genome reveals ancient elasmobranch adaptations associated with wound healing and the maintenance of genome stability.

The white shark (Carcharodon carcharias; Chondrichthyes, Elasmobranchii) is one of the most publicly recognized marine animals. Here we report the genome sequence of the white shark and comparative evolutionary genomic analyses to the chondrichthyans, whale shark (Elasmobranchii) and elephant shark (Holocephali), as well as various vertebrates. The 4.63-Gbp white shark genome contains 24,520 predicted genes, and has a repeat content of 58.5%. We provide evidence for a history of positive selection and gene-content enrichments regarding important genome stability-related genes and functional categories, particularly so for the two elasmobranchs. We hypothesize that the molecular adaptive emphasis on genome stability in white and whale sharks may reflect the combined selective pressure of large genome sizes, high repeat content, high long-interspersed element retrotransposon representation, large body size, and long lifespans, represented across these two species. Molecular adaptation for wound healing was also evident, with positive selection in key genes involved in the wound-healing process, as well as Gene Ontology enrichments in fundamental wound-healing pathways. Sharks, particularly apex predators such as the white shark, are believed to have an acute sense of smell. However, we found very few olfactory receptor genes, very few trace amine-associated receptors, and extremely low numbers of G protein-coupled receptors. We did however, identify 13 copies of vomeronasal type 2 (V2R) genes in white shark and 10 in whale shark; this, combined with the over 30 V2Rs reported previously for elephant shark, suggests this gene family may underlie the keen odorant reception of chondrichthyans.

The complete mitochondrial genome of an Atlantic Ocean Shortfin Mako Shark, Isurus oxyrinchus.

We report the first complete mitochondrial genome of a shortfin mako shark from the Atlantic Ocean. The genome had 16,700 base pairs and contained 13 protein-coding genes, 2 rRNA genes, 22 tRNA genes, and a non-coding D-loop. There were 81 individual differences compared to the published mitochondrial genome of a shortfin mako from the Pacific Ocean, with most variability found in protein coding genes, especially ND5, ND3, and ND1. These highly variable genes may be useful population markers in future studies, and availability of a second mitogenome will assist with future, genome-scale studies of this IUCN Endangered species.

Transcriptome-Derived Microsatellites Demonstrate Strong Genetic Differentiation in Pacific White Sharks.

Recent advances in genome-scale sequencing technology have allowed the development of high resolution genetic markers for the study of nonmodel taxa. In particular, transcriptome sequencing has proven to be highly useful in generating genomic markers for use in population genetic studies, allowing for insight into species connectivity, as well as local adaptive processes as many transcriptome-derived markers are found within or associated with functional genes. Herein, we developed a set of 30 microsatellite markers from a heart transcriptome for the white shark (Carcharodon carcharias), a widely distributed and globally vulnerable marine predator. Using these markers as well as 10 published anonymous genomic microsatellite loci, we provide 1) the first nuclear genetic assessment of the cross-Pacific connectivity of white sharks, and 2) a comparison of the levels of inferred differentiation across microsatellite marker sets (i.e., transcriptome vs. anonymous) to assess their respective utility to elucidate the population genetic dynamics of white sharks. Significant (FST = 0.083, P = 0.05; G″ST = 0.200; P = 0.001) genetic differentiation was found between Southwestern Pacific (n = 19) and Northeastern Pacific (n = 20) white sharks, indicating restricted, cross Pacific gene flow in this species. Transcriptome-derived microsatellite marker sets identified much higher (up to 2×) levels of genetic differentiation than anonymous genomic markers, underscoring potential utility of transcriptome markers in identifying subtle population genetic differences within highly vagile, globally distributed marine species.Subject areas: Population structure and phylogeography; Conservation genetics and biodiversity.

DNA analysis of traded shark fins and mobulid gill plates reveals a high proportion of species of conservation concern.

Continuously increasing demand for plant and animal products causes unsustainable depletion of biological resources. It is estimated that one-quarter of sharks and rays are threatened worldwide and although the global fin trade is widely recognized as a major driver, demand for meat, liver oil, and gill plates also represents a significant threat. This study used DNA barcoding and 16 S rRNA sequencing as a method to identify shark and ray species from dried fins and gill plates, obtained in Canada, China, and Sri Lanka. 129 fins and gill plates were analysed and searches on BOLD produced matches to 20 species of sharks and five species of rays or - in two cases - to a species pair. Twelve of the species found are listed or have been approved for listing in 2017 in the appendices of the Convention on International Trade in Endangered Species of Fauna and Flora (CITES), including the whale shark (Rhincodon typus), which was surprisingly found among both shark fin and gill plate samples. More than half of identified species fall under the IUCN Red List categories 'Endangered' and 'Vulnerable', raising further concerns about the impacts of this trade on the sustainability of these low productivity species.

Biogeophysical and physiological processes drive movement patterns in a marine predator.

BACKGROUND: Blue sharks (Prionace glauca) are among the most abundant and widely distributed of oceanic elasmobranchs. Millions are taken annually in pelagic longline fisheries and comprise the highest component of auctioned fin weight in the international shark fin trade. Though studies of blue sharks outnumber those of other large pelagic sharks, the species' complicated and sexually segregated life history still confound current understanding of Atlantic movement patterns. Lack of detailed information regarding movement and vertical behavior continues to limit management efforts that require such data for stock assessment and sustainable catch modeling. Therefore, this study aims to describe behavioral and ecological patterns distinct to aggregating and migrating blue sharks, and compare the findings to existing Atlantic movement models. RESULTS: Data collected from 23 blue sharks instrumented with pop-up satellite archival tags were used in statistical predictive regression models to investigate habitat use during a localized aggregation in the northwest Atlantic, while undergoing seasonal migrations, and with respect to environmental variables. Deployment durations ranged from 4 to 273 days, with sharks inhabiting both productive coastal waters and the open ocean, and exhibiting long-distance seasonal movements exceeding 3700 km. While aggregating on the continental shelf of the northwest Atlantic, blue sharks displayed consistent depth use independent of sex and life stage, and exhibited varied response to environmental (temperature and chlorophyll a) factors. As sharks dispersed from the aggregation site, depth use was influenced by bathymetry, latitude, demography, and presence in the Gulf Stream. Mature females were not observed at the New England tagging site, however, two mature females with recent mating wounds were captured and tagged opportunistically in The Bahamas, one of which migrated to the Mid-Atlantic Ridge. CONCLUSIONS: Vertical behaviors displayed by blue sharks varied greatly among locales; depth use off the continental shelf was significantly greater, and individuals exhibited a greater frequency of deep-diving behavior, compared to periods of aggregation on the continental shelf. Sexual segregation was evident, suggesting mature and immature males, and immature females may be subjected to high levels of anthropogenic exploitation in this region during periods of aggregation. Analysis of the spatio-temporal tracks revealed that nine individuals traveled beyond the United States EEZ, including a mature female captured in The Bahamas that migrated to the Mid-Atlantic Ridge. These results reflect and augment existing Atlantic migration models, and highlight the complex, synergistic nature of factors affecting blue shark ecology and the need for a cooperative management approach in the North Atlantic.

Comparative transcriptomics of elasmobranchs and teleosts highlight important processes in adaptive immunity and regional endothermy.

BACKGROUND: Comparative genomic and/or transcriptomic analyses involving elasmobranchs remain limited, with genome level comparisons of the elasmobranch immune system to that of higher vertebrates, non-existent. This paper reports a comparative RNA-seq analysis of heart tissue from seven species, including four elasmobranchs and three teleosts, focusing on immunity, but concomitantly seeking to identify genetic similarities shared by the two lamnid sharks and the single billfish in our study, which could be linked to convergent evolution of regional endothermy. RESULTS: Across seven species, we identified an average of 10,877 Swiss-Prot annotated genes from an average of 32,474 open reading frames within each species' heart transcriptome. About half of these genes were shared between all species while the remainder included functional differences between our groups of interest (elasmobranch vs. teleost and endotherms vs. ectotherms) as revealed by Gene Ontology (GO) and selection analyses. A repeatedly represented functional category, in both the uniquely expressed elasmobranch genes (total of 259) and the elasmobranch GO enrichment results, involved antibody-mediated immunity, either in the recruitment of immune cells (Fc receptors) or in antigen presentation, including such terms as "antigen processing and presentation of exogenous peptide antigen via MHC class II", and such genes as MHC class II, HLA-DPB1. Molecular adaptation analyses identified three genes in elasmobranchs with a history of positive selection, including legumain (LGMN), a gene with roles in both innate and adaptive immunity including producing antigens for presentation by MHC class II. Comparisons between the endothermic and ectothermic species revealed an enrichment of GO terms associated with cardiac muscle contraction in endotherms, with 19 genes expressed solely in endotherms, several of which have significant roles in lipid and fat metabolism. CONCLUSIONS: This collective comparative evidence provides the first multi-taxa transcriptomic-based perspective on differences between elasmobranchs and teleosts, and suggests various unique features associated with the adaptive immune system of elasmobranchs, pointing in particular to the potential importance of MHC Class II. This in turn suggests that expanded comparative work involving additional tissues, as well as genome sequencing of multiple elasmobranch species would be productive in elucidating the regulatory and genome architectural hallmarks of elasmobranchs.

The complete mitochondrial genome of the Critically Endangered Angelshark, Squatina squatina.

Here, we describe the first mitochondrial genome of the angelshark, Squatina squatina. The genome is 16,689 bp in length with 13 protein-coding genes, 22 tRNA genes, 2 rRNA genes, and a non-coding control region. Base composition of the mitogenome has an A + T bias (62.9%), seen commonly in other elasmobranchs. This genome provides a key resource for future investigations of the population genetic dynamics and evolution of this Critically Endangered shark.

The complete mitochondrial genome of the endangered great hammerhead shark, Sphyrna mokarran.

We present the first mitochondrial genome sequence of the great hammerhead shark, Sphyrna mokarran. This species is of considerable conservation concern throughout its global distribution, and currently listed as Endangered on the IUCN Red List. The mitochondrial genome is 16,719 bp in length with 13 protein-coding genes, 22 tRNA genes, 2 rRNA genes and a non-coding control region. The gene arrangement is congruent with other shark and most vertebrate species. This S. mokarran mitogenome provides a genomic resource for assisting with population studies and conservation efforts for this highly depleted species.

Complete mitogenome sequences of smooth hammerhead sharks, Sphyrna zygaena, from the eastern and western Atlantic.

We report the first mitogenome sequences of the circumglobally distributed, highly mobile, smooth hammerhead shark, Sphyrna zygaena, from the eastern and western Atlantic. Both genomes were 16,729 bp long with 13 protein-coding genes, two rRNAs, 22 tRNAs and a non-coding control region. The two Atlantic shark sequences differ from each other by 13 SNPs, and by 43 and 44 SNPs from the published mitogenome of an S. zygaena specimen from the eastern Pacific Ocean. The cross-Atlantic mitogenome sequences reported here provide a resource to assist with population genetics studies of this widely exploited species of conservation concern.