Ocean ecosystems are under pressure from overfishing, climate change, habitat destruction and pollution. These pressures have led to documented declines of some fishes in some places, such as those living in coral reefs and on the high seas. However, it is not clear whether these population declines are isolated one-off examples or, instead, if they are sufficiently widespread to risk the extinction of large numbers of species.

Most fishes have a skeleton that is made of bone, but sharks and rays have a skeleton that is made of cartilage. A total of 1,041 species has such a skeleton and they are collectively known as the Chondrichthyes. To find out how well these fish are faring, Dulvy et al. worked with more than 300 scientists around the world to assess the conservation status of all 1,041 species.

Based on this, Dulvy et al. estimate that one in four of these species are threatened with extinction, mainly as a result of overfishing. Moreover, just 389 species (37.4% of the total) are considered to be safe, which is the lowest fraction of safe species among all vertebrate groups studied to date.

The largest sharks and rays are in the most peril, especially those living in shallow waters that are accessible to fisheries. A particular problem is the ‘fin trade’: the fins of sharks and shark-like rays are a delicacy in some Asian countries, and more than half of the chondrichthyans that enter the fin trade are under threat. Whether targeted or caught by boats fishing for other species, sharks and rays are used to supply a market that is largely unmonitored and unregulated. Habitat degradation and loss also pose considerable threats, particularly for freshwater sharks and rays.

Dulvy et al. identified three main hotspots where the biodiversity of sharks and rays was particularly seriously threatened—the Indo-Pacific Biodiversity Triangle, Red Sea, and the Mediterranean Sea—and argue that national and international action is needed to protect them from overfishing.

Populations and species are the building blocks of the communities and ecosystems that sustain humanity through a wide range of services (Mace et al., 2005; Díaz et al., 2006). There is increasing evidence that human impacts over the past 10 millennia have profoundly and permanently altered biodiversity on land, especially of vertebrates (Schipper et al., 2008; Hoffmann et al., 2010). The oceans encompass some of the earth’s largest habitats and longest evolutionary history, and there is mounting concern for the increasing human influence on marine biodiversity that has occurred over the past 500 years (Jackson, 2010). So far our knowledge of ocean biodiversity change is derived mainly from retrospective analyses usually limited to biased subsamples of diversity, such as: charismatic species, commercially-important fisheries, and coral reef ecosystems (Carpenter et al., 2008; Collette et al., 2011; McClenachan et al., 2012; Ricard et al., 2012). Notwithstanding the limitations of these biased snapshots, the rapid expansion of fisheries and globalized trade are emerging as the principal drivers of coastal and ocean threat (Polidoro et al., 2008; Anderson et al., 2011b; McClenachan et al., 2012). The extent and degree of the global impact of fisheries upon marine biodiversity, however, remains poorly understood and highly contentious. Recent insights from ecosystem models and fisheries stock assessments of mainly data-rich northern hemisphere seas, suggest that the status of a few of the best-studied exploited species and ecosystems may be improving (Worm et al., 2009). However, this view is based on only 295 populations of 147 fish species and hence is far from representative of the majority of the world’s fisheries and fished species, especially in the tropics for which there are few data and often less management (Sadovy, 2005; Newton et al., 2007; Branch et al., 2011; Costello et al., 2012; Ricard et al., 2012).

Overfishing and habitat degradation have profoundly altered populations of marine animals (Hutchings, 2000; Lotze et al., 2006; Polidoro et al., 2012), especially sharks and rays (Stevens et al., 2000; Simpfendorfer et al., 2002; Dudley and Simpfendorfer, 2006; Ferretti et al., 2010). It is not clear, however, whether the population declines of globally distributed species are locally reversible or symptomatic of an erosion of resilience and chronic accumulation of global marine extinction risk (Jackson, 2010; Neubauer et al., 2013). In response, we evaluate the scale and intensity of overfishing through a global systematic evaluation of the relative extinction risk for an entire lineage of exploited marine fishes—sharks, rays, and chimaeras (class Chondrichthyes)—using the Red List Categories and Criteria of the International Union for the Conservation of Nature (IUCN). We go on to identify, (i) the life history and ecological attributes of species (and taxonomic families) that render them prone to extinction, and (ii) the geographic locations with the greatest number of species of high conservation concern.

Chondrichthyans make up one of the oldest and most ecologically diverse vertebrate lineages: they arose at least 420 million years ago and rapidly radiated out to occupy the upper tiers of aquatic food webs (Compagno, 1990; Kriwet et al., 2008). Today, this group is one of the most speciose lineages of predators on earth that play important functional roles in the top-down control of coastal and oceanic ecosystem structure and function (Ferretti et al., 2010; Heithaus et al., 2012; Stevens et al., 2000). Sharks and their relatives include some of the latest maturing and slowest reproducing of all vertebrates, exhibiting the longest gestation periods and some of the highest levels of maternal investment in the animal kingdom (Cortés, 2000). The extreme life histories of many chondrichthyans result in very low population growth rates and weak density-dependent compensation in juvenile survival, rendering them intrinsically sensitive to elevated fishing mortality (Musick, 1999b; Cortés, 2002; García et al., 2008; Dulvy and Forrest, 2010).

Chondrichthyans are often caught as incidental, but are often retained as valuable bycatch of fisheries that focus on more productive teleost fish species, such as tunas or groundfishes (Stevens et al., 2005). In many cases, fishing pressure on chondrichthyans is increasing as teleost target species become less accessible (due to depletion or management restrictions) and because of the high, and in some cases rising, value of their meat, fins, livers, and/or gill rakers (Fowler et al., 2002; Clarke et al., 2006; Lack and Sant, 2009). Fins, in particular, have become one of the most valuable seafood commodities: it is estimated that the fins of between 26 and 73 million individuals, worth US$400-550 million, are traded each year (Clarke et al., 2007). The landings of sharks and rays, reported to the Food and Agriculture Organization of the United Nations (FAO), increased steadily to a peak in 2003 and have declined by 20% since (Figure 1A). True total catch, however, is likely to be 3–4 times greater than reported (Clarke et al., 2006; Worm et al., 2013). Most chondrichthyan catches are unregulated and often misidentified, unrecorded, aggregated, or discarded at sea, resulting in a lack of species-specific landings information (Barker and Schluessel, 2005; Clarke et al., 2006; Iglésias et al., 2010; Bornatowski et al., 2013). Consequently, FAO could only be ‘hopeful’ that the catch decline is due to improved management rather than being symptomatic of worldwide overfishing (FAO, 2010). The reported chondrichthyan catch has been increasingly dominated by rays, which have made up greater than half of reported taxonomically-differentiated landings for the past four decades (Figure 1B). Chondrichthyan landings were worth US$1 billion at the peak catch in 2003, since then the value has dropped to US$800 million as catch has declined (Musick and Musick, 2011). A main driver of shark fishing is the globalized trade to meet Asian demand for shark fin soup, a traditional and usually expensive Chinese dish. This particularly lucrative trade in fins (not only from sharks, but also of shark-like rays such as wedgefishes and sawfishes) remains largely unregulated across the 86 countries and territories that exported >9,500 mt of fins to Hong Kong (a major fin trade hub) in 2010 (Figure 1C).

Figure 1

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In a world of limited funding, conservation priorities are often based on immediacy of extinction, the value of biodiversity and conservation opportunity (Marris, 2007). In this study, we provide the first estimates of the threat status and hence risk of extinction of chondrichthyans. Our systematic global assessment of the status of this lineage that includes many iconic predators reveals a risky combination of high threat (17% observed and 23.9% estimated), low safety (Least Concern, 23% observed and >37% estimated), and high uncertainty in their threat status (Data Deficient, 46% observed and 8.7% estimated). Over half of species are predicted to be threatened or Near Threatened (n = 561, 53.9%, Table 1). While no species has been driven to global extinction—as far as we know—at least 28 populations of sawfishes, skates, and angel sharks are locally or regionally extinct (Dulvy et al., 2003; Dulvy and Forrest, 2010). Several shark species have not been seen for many decades. The Critically Endangered Pondicherry shark (Carcharhinus hemiodon) is known only from 20 museum specimens that were captured in the heavily-fished inshore waters of Southeast Asia: it has not been seen since 1979 (Cavanagh et al., 2003). The now ironically-named and Critically Endangered Common skate (Dipturus batis) and Common angel shark (Squatina squatina) are regionally extinct from much of their former geographic range in European waters (Cavanagh and Gibson, 2007; Gibson et al., 2008; Iglésias et al., 2010). The Largetooth sawfish (Pristis pristis) and Smalltooth sawfish (Pristis pectinata) are possibly extinct throughout much of the Eastern Atlantic, particularly in West Africa (Robillard and Séret, 2006; Harrison and Dulvy, 2014).

Our analysis provides an unprecedented understanding of how many chondrichthyan species are actually or likely to be threatened. A very high percentage of species are DD (46%, 487 species); that is one of the highest rates of Data Deficiency of any taxon to date (Hoffmann et al., 2010). This high level of uncertainty in status further elevates risk and presents a key challenge for future assessment efforts. We outline a first step through our estimation that 68 DD species are likely to be threatened based on their life histories and distribution. Numerous studies have retrospectively explained extinction risk, but few have made a priori predictions of risk (Dulvy and Reynolds, 2002; Davidson et al., 2012). Across many taxa, extinction risk has been shown to be a function of an extrinsic driver or threat (Jennings et al., 1998; Davies et al., 2006) and the corresponding life history and ecological traits: large body size (low intrinsic rate of population increase, high trophic level), small geographic range size, and ecological specialization. Maximum body size is an essential predictor of threat status, we presume because of the close relationship between body size and the intrinsic rate of population increase in sharks and rays (Smith et al., 1998; Frisk et al., 2001; Hutchings et al., 2012). Though we note that this proximate link may be mediated ultimately through the time-related traits of growth and mortality (Barnett et al., 2013; Juan-Jordá et al., 2013). Our novel contribution is to show that depth-related geographic traits are more important for explaining risk than geographic range per se. The shallowness of species (minimum depth limit) and the narrowness of their depth range are important risk factors (Figure 3). We hypothesize that this is so because shallower species are more accessible to fishing gears and those with narrower depth ranges have lower likelihood that a proportion of the species distribution remains beyond fishing activity. For example, the Endangered Barndoor skate (Dipturus laevis) was eliminated throughout much of its geographic range and depth distribution due to bycatch in trawl fisheries, yet may have rebounded because a previously unknown deepwater population component lay beyond the reach of most fisheries (Dulvy, 2000; Kulka et al., 2002; COSEWIC, 2010). We find that geographic range (measured as Extent of Occurrence) is largely unrelated to extinction risk. This is in marked contrast to extinction risk patterns on land (Jones et al., 2003; Cardillo et al., 2005; Anderson et al., 2011a) and in the marine fossil record (Harnik et al., 2012a, 2012b), where small geographic range size is the principal correlate of extinction risk. We suggest that this is because fishing activity is now widespread throughout the world’s oceans (Swartz et al., 2010), and even species with the largest ranges are exposed and often entirely encompassed by the footprint of fishing activity. By contrast, with a few exceptions (mainly eastern Atlantic slopes, Figure 7E), fishing has a narrow depth penetration and hence species found at greater depths can still find refuge from exploitation (Morato et al., 2006; Lam and Sadovy de Mitcheson, 2010).

The status of chondrichthyans is arguably among the worst reported for any major vertebrate lineage considered thus far, apart from amphibians (Stuart et al., 2004; Hoffmann et al., 2010). The percentage and absolute number of threatened amphibians is high (>30% are threatened), but a greater percentage are Least Concern (38%), and uncertainty of status is lower (32% DD) than for chondrichthyans. Our discovery of the high level of threat in freshwater chondrichthyans (36%) is consistent with the emerging picture of the intense and unmanaged extinction risk faced by many freshwater and estuarine species (Darwall et al., 2011).

Our threat estimate is comparable to other marine biodiversity status assessments, but our findings caution that ‘global’ fisheries assessments may be underestimating risk. The IUCN Global Marine Species Assessment is not yet complete, but reveals varying threat levels among taxa and regions (Polidoro et al., 2008, 2012). The only synoptic summary to-date focused on charismatic Indo-Pacific coral reef ecosystem species. Of the 1,568 IUCN-assessed marine vertebrates and invertebrates, 16% (range: 12–34% among families) were threatened (McClenachan et al., 2012). This is a conservative estimate of marine threat level because although they may be more intrinsically sensitive to extinction drivers, charismatic species are more likely to garner awareness of their status and support for monitoring and conservation (McClenachan et al., 2012). The predicted level of chondrichthyan threat (>24%) is distinctly greater than that provided by global fisheries risk assessments. These studies provide modeled estimates of the percentage of collapsed bony fish (teleost) stocks in both data-poor unassessed fisheries (18%, Costello et al., 2012) and data-rich fisheries (7–13%, Branch et al., 2011). This could be because teleosts are generally more resilient than elasmobranchs (Hutchings et al., 2012), but in addition we caution that analyses of biased geographic and taxonomic samples may be underestimating risk of collapse in global fisheries, particularly for species with less-resilient life histories.

Our work relies on consensus assessments by more than 300 scientists. However, given the uncertainty in some of the underlying data that inform our understanding of threat status, such as fisheries catch landings data, it is worth considering whether these uncertainties mean our assessments are downplaying the true risk. While there are methods of propagating uncertainty through the IUCN Red List Assessments (Akcakaya et al., 2000), in our experience this approach was uninformative for even the best-studied species, because it generated confidence intervals that spanned all IUCN Categories. Instead it is worth considering whether our estimates of threat are consistent with independent quantitative estimates of status. The Mediterranean Red List Assessment workshop in 2005 prompted subsequent quantitative analyses of catch landings, research trawl surveys, and sightings data. Quantitative trends could be estimated for five species suggesting they had declined by 96% to >99.9% relative to their former abundance suggesting they would meet the highest IUCN Threat category of Critically Endangered (Ferretti et al., 2008). By comparison the earlier IUCN regional assessment for these species, while suggesting they were all threatened, was more conservative for two of the five species: Hammerhead sharks (Sphyrna spp.)—Critically Endangered, Porbeagle shark (Lamna nasus)—Critically Endangered, Shortfin mako (Isurus oxyrinchus)—Critically Endangered, Blue shark (Prionace glauca)—Vulnerable, and Thresher shark (Alopias vulpinus)—Vulnerable.

We can also make a complementary comparison to a recent analysis of the status of 112 shark and ray fisheries (Costello et al., 2012). The median biomass relative to the biomass at Maximum Sustainable Yield (B/B_MSY) of these 112 shark and ray fisheries was 0.37, making them the most overfished groups of any of the world’s unassessed fisheries. Assuming B_MSY occurs at 0.3 to 0.5 of unexploited biomass then the median biomass of shark and ray fisheries had declined by between 81% and 89% by 2009. These biomass declines would be sufficient to qualify all of these 112 shark and ray fisheries for the Endangered IUCN category if they occurred within a three-generation time span. By comparison our results are considerably more conservative. Empirical analyses show that an IUCN threatened category listing is triggered only once teleost fishes (with far higher density-dependent compensation) have been fished down to below B_MSY (Dulvy et al., 2005; Porszt et al., 2012). Hence, our findings are consistent with only around one-quarter of chondrichthyan species having been fished down below the B_MSY target reference point. While there may be concern that expert assessments may overstate declines and threat, it is more likely that our conservative consensus-based approach has understated declines and risk in sharks and rays.

For marine species, predicting absolute risk of extinction remains highly uncertain because, even with adequate evidence of severe decline, in many instances the absolute population size remains large (Mace, 2004). There remains considerable uncertainty as to the relationship between census and effective population size (Reynolds et al., 2005). Therefore, Red List categorization of chondrichthyans should be interpreted as a comparative measure of relative extinction risk, in recognition that unmanaged steep declines, even of large populations, may ultimately lead to ecosystem perturbations and eventually biological extinction. The Red List serves to raise red flags calling for conservation action, sooner rather than later, while there is a still chance of recovery and of forestalling permanent biodiversity loss.

Despite more than two decades of rising awareness of chondrichthyan population declines and collapses, there is still no global mechanism to ensure financing, implementation and enforcement of chondrichthyan fishery management plans that is likely to rebuild populations to levels where they would no longer be threatened (Lack and Sant, 2009; Techera and Klein, 2011). This management shortfall is particularly problematic given the large geographic range of many species. Threat increased only slightly when geographic range is measured as the Extent of Occurrence; however, geographic range becomes increasingly important when it is measured as the number of countries (legal jurisdictions) spanned by each species. The proportion of species that are threatened increases markedly with geographic size measured by number of Exclusive Economic Zones (EEZs) spanned; one-quarter of threatened species span the EEZs of 18 or more countries (Figure 10). Hence, their large geographic ranges do not confer safety, but instead exacerbates risk because sharks and rays require coherent, effective international management.

Figure 10

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With a few exceptions (e.g., Australia and USA), many governments still lack the resources, expertise, and political will necessary to effectively conserve the vast majority of shark and rays, and indeed many other exploited organisms (Veitch et al., 2012). More than 50 sharks are included in Annex I (Highly Migratory Species) of the 1982 Law of the Sea Convention, implemented on the high seas under the 1992 Fish Stocks Agreement, but currently only a handful enjoy species-specific protections under the world’s Regional Fishery Management Organizations (Table 6), and many of these have yet to be implemented domestically. The Migratory Sharks Memorandum of Understanding (MoU) adopted by the Parties to the Convention on Migratory Species (CMS) so far only covers seven sharks, yet there may be more than 150 chondrichthyans that regularly migrate across national boundaries (Fowler, 2012). To date, only one of the United Nations Environment Programme’s Regional Seas Conventions, the Barcelona Convention for the Conservation of the Mediterranean Sea, includes chondrichthyan fishes and only a few of its Parties have taken concrete domestic action to implement these listings. Despite two decades of effort, only ten sharks and rays had been listed by the Convention on International Trade in Endangered Species (CITES) up to 2013 (Vincent et al., 2014). A further seven species of shark and ray were listed by CITES in 2013—the next challenge is to ensure effective implementation of these trade regulations (Mundy-Taylor and Crook, 2013). OSPAR (the Convention for the Protection of the marine Environment of the North-East Atlantic) lists many threatened shark and ray species, but its remit excludes fisheries issues. Many chondrichthyans qualify for listing under CITES, CMS, and various regional seas conventions, and should be formally considered for such action as a complement to action by Regional Fisheries Management Organizations (RFMOs) (Table 6).

Bans on ‘finning’ (slicing off a shark’s fins and discarding the body at sea) are the most widespread shark conservation measures. While these prohibitions, particularly those that require fins to remain attached through landing, can enhance monitoring and compliance, they have not significantly reduced shark mortality or risk to threatened species (Clarke et al., 2013). Steep declines and the high threat levels in migratory oceanic pelagic sharks suggest raising the priority of improved management of catch and trade through concerted actions by national governments working through RFMOs as well as CITES, and CMS (Table 7).

A high proportion of catch landings come from nations with a large number of threatened chondrichthyans and less-than-comprehensive chondrichthyan fishery management plans. Future research is required to down-scale these global Red List assessments and analyses to provide country-by-country diagnoses of the link between specific fisheries and specific threats to populations of more broadly distributed species (Wallace et al., 2010). Such information could be used to focus fisheries management and conservation interventions that are tailored to specific problems. There is no systematic global monitoring of shark and ray populations and the national fisheries catch landings statistics provide invaluable data for tracking fisheries trends in unmanaged fisheries (Newton et al., 2007; Worm et al., 2013). However, the surveillance power of such data could be greatly improved if collected at greater taxonomic resolution. While there have been continual improvements, catches are under-reported (Clarke et al., 2006), and for those that are reported only around one-third is reported at the species level (Fischer et al., 2012). To complement improved catch landings data, we recommend the development of repeat regional assessments of the Red List Status of chondrichthyans to provide an early warning of adverse changes in status and to detect and monitor the success of management initiatives and interventions. Aggregate Red List Threat indices for chondrichthyans, like those available for mammals, birds, amphibians, and hard corals (Carpenter et al., 2008) would provide one of the few global scale indicators of progress toward international biodiversity goals (Walpole et al., 2009; Butchart et al., 2010).

Our global status assessment of sharks and rays reveals the principal causes and severity of global marine biodiversity loss, and the threat level they face exposes a serious shortfall in the conservation management of commercially-exploited aquatic species (McClenachan et al., 2012). Chondrichthyans have slipped through the jurisdictional cracks of traditional national and international management authorities. Rather than accept that many chondrichthyans will inevitably be driven to economic, ecological, or biological extinction, we warn that dramatic changes in the enforcement and implementation of the conservation and management of threatened chondrichthyans are urgently needed to ensure a healthy future for these iconic fishes and the ecosystems they support.

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Author details

1. Nicholas K Dulvy

   1. IUCN Species Survival Commission Shark Specialist Group, Department of Biological Sciences, Simon Fraser University, Burnaby, Canada
   2. Earth to Ocean Research Group, Department of Biological Sciences, Simon Fraser University, Burnaby, Canada

   Contribution

   NKD, Conception and design, Acquisition of data, Analysis and interpretation of data, Drafting or revising the article, Contributed unpublished essential data or reagents

   For correspondence

   dulvy@sfu.ca

   Competing interests

   The authors declare that no competing interests exist.

2. Sarah L Fowler

   IUCN Species Survival Commission Shark Specialist Group, NatureBureau International, Newbury, United Kingdom

   Contribution

   SLF, Conception and design, Acquisition of data, Analysis and interpretation of data, Drafting or revising the article, Contributed unpublished essential data or reagents

   Competing interests

   The authors declare that no competing interests exist.

3. John A Musick

   Virginia Institute of Marine Science, College of William and Mary, Gloucester Point, United States

   Contribution

   JAM, Acquisition of data, Drafting or revising the article

   Competing interests

   The authors declare that no competing interests exist.

4. Rachel D Cavanagh

   British Antarctic Survey, Natural Environment Research Council, Cambridge, United Kingdom

   Contribution

   RDC, Conception and design, Acquisition of data, Analysis and interpretation of data, Drafting or revising the article

   Competing interests

   The authors declare that no competing interests exist.

5. Peter M Kyne

   Research Institute for the Environment and Livelihoods, Charles Darwin University, Darwin, Australia

   Contribution

   PMK, Acquisition of data, Analysis and interpretation of data, Drafting or revising the article

   Competing interests

   The authors declare that no competing interests exist.

6. Lucy R Harrison

   1. IUCN Species Survival Commission Shark Specialist Group, Department of Biological Sciences, Simon Fraser University, Burnaby, Canada
   2. Earth to Ocean Research Group, Department of Biological Sciences, Simon Fraser University, Burnaby, Canada

   Contribution

   LRH, Acquisition of data, Analysis and interpretation of data, Drafting or revising the article

   Competing interests

   The authors declare that no competing interests exist.

7. John K Carlson

   Southeast Fisheries Science Center, NOAA/National Marine Fisheries Service, Panama City, United States

   Contribution

   JKC, Analysis and interpretation of data, Drafting or revising the article

   Competing interests

   The authors declare that no competing interests exist.

8. Lindsay NK Davidson

   1. IUCN Species Survival Commission Shark Specialist Group, Department of Biological Sciences, Simon Fraser University, Burnaby, Canada
   2. Earth to Ocean Research Group, Department of Biological Sciences, Simon Fraser University, Burnaby, Canada

   Contribution

   LNKD, Analysis and interpretation of data, Drafting or revising the article

   Competing interests

   The authors declare that no competing interests exist.

9. Sonja V Fordham

   Shark Advocates International, The Ocean Foundation, Washington, DC, United States

   Contribution

   SVF, Analysis and interpretation of data, Drafting or revising the article, Contributed unpublished essential data or reagents

   Competing interests

   The authors declare that no competing interests exist.

10. Malcolm P Francis

    National Institute of Water and Atmospheric Research, Wellington, New Zealand

    Contribution

    MPF, Analysis and interpretation of data, Drafting or revising the article

    Competing interests

    The authors declare that no competing interests exist.

11. Caroline M Pollock

    Global Species Programme, International Union for the Conservation of Nature, Cambridge, United Kingdom

    Contribution

    CMP, Acquisition of data, Analysis and interpretation of data

    Competing interests

    The authors declare that no competing interests exist.

12. Colin A Simpfendorfer

    1. Centre for Sustainable Tropical Fisheries and Aquaculture, James Cook University, Townsville, Australia
    2. School of Earth and Environmental Sciences, James Cook University, Townsville, Australia

    Contribution

    CAS, Conception and design, Analysis and interpretation of data, Drafting or revising the article

    Competing interests

    The authors declare that no competing interests exist.

13. George H Burgess

    Florida Program for Shark Research, Florida Museum of Natural History, University of Florida, Gainsville, United States

    Contribution

    GHB, Analysis and interpretation of data, Drafting or revising the article

    Competing interests

    The authors declare that no competing interests exist.

14. Kent E Carpenter

    1. IUCN Species Programme Species Survival Commission, Old Dominion University, Norfolk, United States
    2. Conservation International Global Marine Species Assessment, Old Dominion University, Norfolk, United States

    Contribution

    KEC, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

15. Leonard JV Compagno

    Shark Research Center, Iziko, South African Museum, Cape Town, South Africa

    Contribution

    LJVC, Acquisition of data, Drafting or revising the article, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

16. David A Ebert

    Pacific Shark Research Center, Moss Landing Marine Laboratories, Moss Landing, United States

    Contribution

    DAE, Drafting or revising the article, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

17. Claudine Gibson

    IUCN Species Survival Commission Shark Specialist Group, NatureBureau International, Newbury, United Kingdom

    Contribution

    CG, Acquisition of data, Drafting or revising the article

    Competing interests

    The authors declare that no competing interests exist.

18. Michelle R Heupel

    School of Earth and Environmental Sciences, Australian Institute of Marine Science, Townsville, Australia

    Contribution

    MRH, Drafting or revising the article, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

19. Suzanne R Livingstone

    Global Marine Species Assessment, Biodiversity Assessment Unit, IUCN Species Programme, Conservation International, Arlington, United States

    Contribution

    SRL, Analysis and interpretation of data, Drafting or revising the article

    Competing interests

    The authors declare that no competing interests exist.

20. Jonnell C Sanciangco

    1. IUCN Species Programme Species Survival Commission, Old Dominion University, Norfolk, United States
    2. Conservation International Global Marine Species Assessment, Old Dominion University, Norfolk, United States

    Contribution

    JCS, Analysis and interpretation of data, Drafting or revising the article, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

21. John D Stevens

    Marine and Atmospheric Research, Commonwealth Scientific and Industrial Research Organisation, Hobart, Australia

    Contribution

    JDS, Analysis and interpretation of data, Drafting or revising the article, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

22. Sarah Valenti

    IUCN Species Survival Commission Shark Specialist Group, NatureBureau International, Newbury, United Kingdom

    Contribution

    SV, Acquisition of data, Analysis and interpretation of data, Drafting or revising the article

    Competing interests

    The authors declare that no competing interests exist.

23. William T White

    Marine and Atmospheric Research, Commonwealth Scientific and Industrial Research Organisation, Hobart, Australia

    Contribution

    WTW, Drafting or revising the article, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

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