Author Correction: Global status and conservation potential of reef sharks.

An Amendment to this paper has been published and can be accessed via a link at the top of the paper.

Global status and conservation potential of reef sharks.

Decades of overexploitation have devastated shark populations, leaving considerable doubt as to their ecological status1,2. Yet much of what is known about sharks has been inferred from catch records in industrial fisheries, whereas far less information is available about sharks that live in coastal habitats3. Here we address this knowledge gap using data from more than 15,000 standardized baited remote underwater video stations that were deployed on 371 reefs in 58 nations to estimate the conservation status of reef sharks globally. Our results reveal the profound impact that fishing has had on reef shark populations: we observed no sharks on almost 20% of the surveyed reefs. Reef sharks were almost completely absent from reefs in several nations, and shark depletion was strongly related to socio-economic conditions such as the size and proximity of the nearest market, poor governance and the density of the human population. However, opportunities for the conservation of reef sharks remain: shark sanctuaries, closed areas, catch limits and an absence of gillnets and longlines were associated with a substantially higher relative abundance of reef sharks. These results reveal several policy pathways for the restoration and management of reef shark populations, from direct top-down management of fishing to indirect improvement of governance conditions. Reef shark populations will only have a high chance of recovery by engaging key socio-economic aspects of tropical fisheries.

Global spatial risk assessment of sharks under the footprint of fisheries.

Effective ocean management and the conservation of highly migratory species depend on resolving the overlap between animal movements and distributions, and fishing effort. However, this information is lacking at a global scale. Here we show, using a big-data approach that combines satellite-tracked movements of pelagic sharks and global fishing fleets, that 24% of the mean monthly space used by sharks falls under the footprint of pelagic longline fisheries. Space-use hotspots of commercially valuable sharks and of internationally protected species had the highest overlap with longlines (up to 76% and 64%, respectively), and were also associated with significant increases in fishing effort. We conclude that pelagic sharks have limited spatial refuge from current levels of fishing effort in marine areas beyond national jurisdictions (the high seas). Our results demonstrate an urgent need for conservation and management measures at high-seas hotspots of shark space use, and highlight the potential of simultaneous satellite surveillance of megafauna and fishers as a tool for near-real-time, dynamic management.

Risk profiles of elite breath-hold divers.

This study aimed to determine a typical profile of elite breath-hold divers (BHDs), in relation to loss of consciousness (LOC) and episodic memory. Forty-four BHDs were evaluated during a world championship with anthropometric and physiological measurements, psychosociological factors and memory assessment. Seventy-five percent of the BHDs had at least one LOC with the predominance being men (p < 0.05). Thirty six percent of BHDs presented a low-risk profile and 64% a high-risk profile with no particular psychological pattern. Stepwise multiple linear regression showed that body fat, years of BH practice, age and forced vital capacity explained a significant amount of the variance of LOC for all BHDs (F(4,39) = 16.03, p < 0.001, R2 = 0.622, R2Adjusted = 0.583). No correlation was found between resting physiological parameters and their training or depth performances. In conclusion, anthropometric data, pulmonary factors and breath-holding experience were predictive of LOC in elite BHDs, with men taking more risks. BHDs episodic memory was not impaired.

Coral reef restoration for coastal protection: Crafting technical and financial solutions.

Coastal erosion, aggravated by coral reef mortality is a major issue for Small Island Developing States. Traditionally gray infrastructure, financed by public budgets has been used to combat beach loss. We examined if three Nature-based Solutions (NbS): (i) coral restoration (green) (ii) restoration + limestone (hybrid) and (iii) restoration + 3D printed concrete (hybrid) could deliver positive outcomes for coastal protection and further incentivize cost sharing for reef conservation, with private beneficiaries. We modelled the impact of restoration on wave attenuation at two reefs off Barbados and simulated up-front and maintenance costs over a 25-year period. All solutions provide additionality when compared to gray infrastructure, especially in mitigating against Sea Level Rise. Restoration was the least costly with the highest risk of failure. The hybrid solutions, were less risky than the green as they provided immediate wave attenuation, alongside complementary services such as increased attractiveness due to the presence of reef fish. Their costs were however between +80% and +450% higher than gray solutions. While this might initially deter the use of NbS, blended finance and in some cases, Payments for Ecosystem Services, could provide options for governments and private beneficiaries to share costs, with ultimately greater benefits for themselves and coral reefs.

Life-history characteristics of an exploited bonefish Albula glossodonta population in a remote South Pacific atoll.

Bonefishes Albula spp. are important components of subsistence fisheries and lucrative sport fishing industries throughout their circumtropical distribution. In Oceania, however, Albula spp. have historically been overexploited and there is a growing need to balance the demands of competing fishing sectors, making the description of their life history a regional priority. To this aim, we collected biological samples from Albula spp. of Anaa atoll, French Polynesia, to identify the species that compose the stock and estimate their life-history parameters including age, growth, reproduction and natural mortality. Our results indicate that Albula glossodonta is the species of bonefish present, with a maximum age that is below the, 20 year longevity of the genus (8 years in males and 10 years in females). Differential growth patterns existed between the two sexes (L∞ = 58, 78 cm fork length (LF ) and K = 0.38, 0.21 for males and females, respectively). Males attained sexual maturity at 43 cm LF (c. 3 years) whereas females matured at 48 cm LF (c. 4 years) and oocyte production was significantly related to body mass, with a maximum batch fecundity of 1,133,767 oocytes in a 4406 g (70 cm LF ) female. The gonado-somatic index of harvested fishes indicated that the spawning season extends from March through September. Based on the observation of a, 20 year bonefish at the proximate Tetiaroa Atoll and several empirical models, estimates of natural mortality ranged from 0.21 to 0.68; however, an estimate of 0.21 was deemed most appropriate. This information facilitated the resurgence of a Rahui (temporary fishing closure) and community-based management to protect A. glossodonta during a critical portion of their spawning season and in this context our results provide an important demographic baseline in evaluating the recovery of this fishery.

Concede or clash? Solitary sharks competing for food assess rivals to decide.

To adapt to their environment, organisms can either directly interact with their surroundings or use social information (i.e. information provided by neighbouring individuals). Social information relates to the external features of surrounding peers, and little is known about its use by solitary species. Here, we investigated the use of social cues in a solitary marine predator by creating artificial aggregations of free-ranging sicklefin lemon sharks (Negaprion acutidens). Using a novel monitoring protocol, we analysed both dominance interactions and tolerance associations between sharks competing for food in relation with the number, the morphology and the behaviour of rivals. Sharks produced more agonistic displays and spent more time around the bait as competitors were more abundant. Moreover, the morphological attributes of competitors had very limited influence on the structure of shark social interactions. Instead, sharks appeared to establish tolerance relationships with competitors according to their individual behaviour. Furthermore, the more two sharks were observed together at a given study site, the fewer agonistic interactions they exchanged. We discuss these findings as evidence of the use of social cues in a non-gregarious predatory species and suggest directions for future research.

The oxygen-conserving potential of the diving response: A kinetic-based analysis.

We investigated the oxygen-conserving potential of the human diving response by comparing trained breath-hold divers (BHDs) to non-divers (NDs) during simulated dynamic breath-holding (BH). Changes in haemodynamics [heart rate (HR), stroke volume (SV), cardiac output (CO)] and peripheral muscle oxygenation [oxyhaemoglobin ([HbO2]), deoxyhaemoglobin ([HHb]), total haemoglobin ([tHb]), tissue saturation index (TSI)] and peripheral oxygen saturation (SpO2) were continuously recorded during simulated dynamic BH. BHDs showed a breaking point in HR kinetics at mid-BH immediately preceding a more pronounced drop in HR (-0.86 bpm.%-1) while HR kinetics in NDs steadily decreased throughout BH (-0.47 bpm.%-1). By contrast, SV remained unchanged during BH in both groups (all P > 0.05). Near-infrared spectroscopy (NIRS) results (mean ± SD) expressed as percentage changes from the initial values showed a lower [HHb] increase for BHDs than for NDs at the cessation of BH (+24.0 ± 10.1 vs. +39.2 ± 9.6%, respectively; P < 0.05). As a result, BHDs showed a [tHb] drop that NDs did not at the end of BH (-7.3 ± 3.2 vs. -3.0 ± 4.7%, respectively; P < 0.05). The most striking finding of the present study was that BHDs presented an increase in oxygen-conserving efficiency due to substantial shifts in both cardiac and peripheral haemodynamics during simulated BH. In addition, the kinetic-based approach we used provides further credence to the concept of an "oxygen-conserving breaking point" in the human diving response.

Blacktip reef sharks, Carcharhinus melanopterus, have high genetic structure and varying demographic histories in their Indo-Pacific range.

For free-swimming marine species like sharks, only population genetics and demographic history analyses can be used to assess population health/status as baseline population numbers are usually unknown. We investigated the population genetics of blacktip reef sharks, Carcharhinus melanopterus; one of the most abundant reef-associated sharks and the apex predator of many shallow water reefs of the Indian and Pacific Oceans. Our sampling includes 4 widely separated locations in the Indo-Pacific and 11 islands in French Polynesia with different levels of coastal development. Four-teen microsatellite loci were analysed for samples from all locations and two mitochondrial DNA fragments, the control region and cytochrome b, were examined for 10 locations. For microsatellites, genetic diversity is higher for the locations in the large open systems of the Red Sea and Australia than for the fragmented habitat of the smaller islands of French Polynesia. Strong significant structure was found for distant locations with FST values as high as ~0.3, and a smaller but still significant structure is found within French Polynesia. Both mitochondrial genes show only a few mutations across the sequences with a dominant shared haplotype in French Polynesia and New Caledonia suggesting a common lineage different to that of East Australia. Demographic history analyses indicate population expansions in the Red Sea and Australia that may coincide with sea level changes after climatic events. Expansions and flat signals are indicated for French Polynesia as well as a significant recent bottleneck for Moorea, the most human-impacted lagoon of the locations in French Polynesia.

Behavioural heterogeneity across killer whale social units in their response to feeding opportunities from fisheries.

Intra-population heterogeneity in the behavioural response of predators to changes in prey availability caused by human activities can have major evolutionary implications. Among these activities, fisheries, while extracting resources, also provide new feeding opportunities for marine top predators. However, heterogeneity in the extent to which individuals have responded to these opportunities within populations is poorly understood. Here, we used 18 years of photo-identification data paired with statistical models to assess variation in the way killer whale social units within a subantarctic population (Crozet Islands) interact with fisheries to feed on fish caught on fishing gear (i.e., depredation behaviour). Our results indicate large heterogeneity in both the spatial and temporal extents of depredation across social units. While some frequently depredated on fishery catches over large areas, others sporadically did so and in small areas consistently over the years. These findings suggest that killer whale social units are exposed to varying levels of impacts of depredation, both negative (potential retaliation from fishers) and positive (food provisioning), on their life history traits, and may explain the contrasted demographic patterns observed within the declining population at Crozet but also potentially within the many other killer whale populations documented depredating on fisheries catches worldwide.

Directed conservation of the world's reef sharks and rays.

Many shark populations are in decline around the world, with severe ecological and economic consequences. Fisheries management and marine protected areas (MPAs) have both been heralded as solutions. However, the effectiveness of MPAs alone is questionable, particularly for globally threatened sharks and rays ('elasmobranchs'), with little known about how fisheries management and MPAs interact to conserve these species. Here we use a dedicated global survey of coral reef elasmobranchs to assess 66 fully protected areas embedded within a range of fisheries management regimes across 36 countries. We show that conservation benefits were primarily for reef-associated sharks, which were twice as abundant in fully protected areas compared with areas open to fishing. Conservation benefits were greatest in large protected areas that incorporate distinct reefs. However, the same benefits were not evident for rays or wide-ranging sharks that are both economically and ecologically important while also threatened with extinction. We show that conservation benefits from fully protected areas are close to doubled when embedded within areas of effective fisheries management, highlighting the importance of a mixed management approach of both effective fisheries management and well-designed fully protected areas to conserve tropical elasmobranch assemblages globally.

Economic valuation of ecosystem services from coral reefs in the South Pacific: taking stock of recent experience.

The economic valuation of coral reefs ecosystem services is currently seen as a promising approach to demonstrate the benefits of sustainable management of coral ecosystems to policymakers and to provide useful information for improved decisions. Most coral reefs economic studies have been conducted in the United States, Southeast Asia and the Caribbean, and only a few have covered the South Pacific region. In this region, coral reefs are essential assets for small island developing states as well as for developed countries. Accordingly, a series of ecosystem services valuations has been carried out recently in the South Pacific, to try and supply decision-makers with new information. Applying ecosystem services valuation to the specific ecological, social, economic and cultural contexts of the South Pacific is however not straightforward. This paper analyses how extant valuations address the various management challenges of coral reef regions in general and more specifically for the South Pacific. Bearing in mind that economic valuation has to match policy-making contexts, we emphasize a series of specific considerations when conducting and applying ecosystem services valuation in South Pacific ecological and social contexts. Finally, the paper examines the decision-making situations in which extant valuations took place. We conclude that, although ecosystem valuations have been effectively used as a means to raise awareness with respect to coral reef conservation, methodologies will have to be further developed, with multidisciplinary inputs, if they are to provide valuable inputs in local and technical decision-making.

Provisioning ecotourism does not increase tiger shark site fidelity.

A perennial criticism of provisioning ecotourism is that it alters the natural behavior and ecology of the target species by providing an artificial food source. Here we evaluate its impact on the long-term site fidelity patterns of tiger sharks in French Polynesia. We hypothesized that a significant impact of provisioning would lead to (1) increases in individual site fidelity over time, and (2) an increase in the number of resident individuals over time. Of 53 individuals photo-identified and monitored during > 500 dives over five years, 10 individuals accounted for > 75% of all sightings, whereas 35 sharks were sighted very infrequently. Even the most frequently observed tiger sharks exhibited overall low fidelity at the site and showed no increase in site fidelity over time. Furthermore, the number of tiger sharks sighted during each dive did not increase. The observed patterns of tiger shark sightings were best explained by natural movements, including general roaming within home ranges along the coastline and seasonal migrations. Despite the apparent lack of impact of provisioning ecotourism on tiger shark ecology in Tahitian waters, it would be prudent to implement a strict code of conduct during any future provisioning activities to maximize the safety of participants and animals involved.

Citizen science provides valuable data to evaluate elasmobranch diversity and trends throughout the French Polynesia's shark sanctuary.

Observers of the Polynesian Shark Observatory (ORP), a citizen science network organized mainly through the Polynesian dive centers, collected an unprecedented amount of data from more than 13,916 dives spanning 43% of the islands of French Polynesia between July 8, 2011, and April 11, 2018. The objective for this type of data collection, which is not accessible within the standard research context, was to provide a unique dataset, and the opportunity to explore the specific diversity, distribution, seasonality and abundance of many elasmobranch species spread out throughout the territory of French Polynesia. Since the data are based on random citizen observations, the spatial distribution was biased toward the most frequented sites and islands where scuba diving is most developed. Overall, the increase in observed abundance of rays and sharks observed in French Polynesia, and the three most sampled islands as well as the high specific diversity recorded for the region, provide first evidence on the effectiveness of the French Polynesia's Shark Sanctuary, established in 2006. These data, collected randomly by the volunteers, also provide insights into potential movement patterns and site fidelity of some of the more commonly observed species. While no final conclusions can be drawn, it is clear that the network of volunteers that regularly contributes information to the Polynesian Shark Observatory plays a very important role in the delivery of much needed data for conservation and management action, as well as providing perspectives for new directions in research on sharks and rays in French Polynesia.

First tiger shark Galeocerdo cuvier bite in 75 years in French Polynesia (Eastern Central Pacific).

We analyzed a tiger shark (estimated 2.8 m total length) bite on a snorkeler. The removal of the terminal part of the leg suggests a predatory motivation for the bite. This is the first documented bite by a tiger shark in French Polynesia waters for the past 75 years.

Like a rolling stone: Colonization and migration dynamics of the gray reef shark (Carcharhinus amblyrhynchos).

Designing appropriate management plans requires knowledge of both the dispersal ability and what has shaped the current distribution of the species under consideration. Here, we investigated the evolutionary history of the endangered gray reef shark (Carcharhinus amblyrhynchos) across its range by sequencing thousands of RADseq loci in 173 individuals in the Indo-Pacific (IP). We first bring evidence of the occurrence of a range expansion (RE) originating close to the Indo-Australian Archipelago (IAA) where two stepping-stone waves (east and westward) colonized almost the entire IP. Coalescent modeling additionally highlighted a homogenous connectivity (Nm ~ 10 per generation) throughout the range, and isolation by distance model suggested the absence of barriers to dispersal despite the affinity of C. amblyrhynchos to coral reefs. This coincides with long-distance swims previously recorded, suggesting that the strong genetic structure at the IP scale (F ST ~ 0.56 between its ends) is the consequence of its broad current distribution and organization in a large number of demes. Our results strongly suggest that management plans for the gray reef shark should be designed on a range-wide rather than a local scale due to its continuous genetic structure. We further contrasted these results with those obtained previously for the sympatric but strictly lagoon-associated Carcharhinus melanopterus, known for its restricted dispersal ability. Carcharhinus melanopterus exhibits a similar RE dynamic but is characterized by a stronger genetic structure and a nonhomogeneous connectivity largely dependent on local coral reefs availability. This sheds new light on shark evolution, emphasizing the roles of IAA as source of biodiversity and of life-history traits in shaping the extent of genetic structure and diversity.

Widespread diversity deficits of coral reef sharks and rays.

A global survey of coral reefs reveals that overfishing is driving resident shark species toward extinction, causing diversity deficits in reef elasmobranch (shark and ray) assemblages. Our species-level analysis revealed global declines of 60 to 73% for five common resident reef shark species and that individual shark species were not detected at 34 to 47% of surveyed reefs. As reefs become more shark-depleted, rays begin to dominate assemblages. Shark-dominated assemblages persist in wealthy nations with strong governance and in highly protected areas, whereas poverty, weak governance, and a lack of shark management are associated with depauperate assemblages mainly composed of rays. Without action to address these diversity deficits, loss of ecological function and ecosystem services will increasingly affect human communities.

Effects of a COVID-19 lockdown-induced pause and resumption of artificial provisioning on blacktip reef sharks (Carcharhinus melanopterus) and pink whiprays (Pateobatis fai) in French Polynesia (East-Pacific).

The tourism activities linked to artificial provisioning of blacktip reef sharks (Carcharhinus melanopterus) and pink whiprays (Pateobatis fai) on a specific site in French Polynesia were suddenly and completely stopped due to a COVID-19 lockdown that lasted 6 weeks from March 20 until April 30, 2020. Using both drone footage and underwater counting, we were able to track the abundance of those two species before, during, and after reopening and thus investigate the impact of provisioning on wild shark populations. The absence of any stimulus during this long period resulted in almost total desertion of the site by the elasmobranchs. However, 1 day prior to reopening, some individuals of both species positively reacted to the single acoustic stimulus of an engine boat, showing the resilience of conditioning, and some elasmobranchs reacted to acoustic and olfactive stimuli linked to the provisioning practice from the first day after reopening. During the first 2 weeks after reopening, the abundance of both species remained at reduced levels comparable to those observed between 2008 and 2010 for sharks; i.e., around 9 animals in the presence of local tourists. Pre-lockdown abundance levels, reaching approximatively 15 individuals for sharks and 10 for rays, were considered restored 1 and 2 months after reopening for blacktip reef sharks and pink whiprays, respectively. These findings improve our capacity to better understand the potential effects of artificial provisioning tourism on the abundance of elasmobranchs by showing that conditioning is resilient for several weeks, suggesting that intermittent interruption of elasmobranchs feeding would not really help to decrease its impact on animal welfare.