Size Assessment of the Gray Reef Shark Carcharhinus amblyrhynchos Inferred from Teeth Marks on Human Wounds

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Noémie JUBLIER and Eric CLUA
Jublier, N. and Clua, E. (2018). Size Assessment of the Gray Reef Shark Carcharhinus amblyrhynchos Inferred from Teeth Marks on Human Wounds. Journal of Forensic Science, doi: 10.1111/1556-4029.13738.

An accurate assessment of the biting shark size is paramount for better understanding the agonistic behavior of a species toward humans. The gray reef shark is involved in many accidental bites. Based on the capture of 35 gray sharks, we calculated the algorithm that allows the assessment of the shark size, through the interdental distance (IDD) inferred from teeth marks on human wounds. Our results show a negative allometric relationship and that IDD calculated from imprints perpetrated by the upper jaw are globally similar with those from the lower jaw, in spite of heterodonty. We applied our findings to two cases of accidental bites by this species. Both of the victims had declared that the shark length was “at least 2 m”. Based on our algorithm, the assessment of the shark TL was approx. 180 and 160 cm, respectively, which correspond to an overestimation of ≥20% by the victims.

Managing bite risk for divers in the context of shark feeding ecotourism: A case study from French Polynesia (Eastern Pacific)

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Eric EG CLUA
Clua, E. (2018) Managing bite risk for divers during shark feeding ecotourism: a case study from French Polynesia. Tourism management. 68, 275-283.
Shark-based ecotourism has been recently expanding around the world. Provisioning sharks, however, is not
without risk and accidental bites on humans are regularly reported. Such events may jeopardise the shark
ecotourism industry and local economies. Through a case study from French Polynesia, I investigated whether
changes in feeding practices of lemon sharks between the early 1990s and 2013 are related to recent accidental
bites of divers. Hand-feeding, ‘smelling’ and surface feeding facilitated the development of agonistic behaviour in sharks, which resulted in accidental bites. The level of interaction between sharks and humans appears to be the most important driver for analysing bite-risk. This paper presents the very first framework for risk-analysis and will allow managers to better manage the risk associated with current practices in shark-feeding around the world. The paper also provides technical guidelines for the design of adequate legal frameworks that will support any sustainable shark feeding operation.

Individual shark profiling: An innovative and environmentally responsible approach for selectively managing human fatalities

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Eric EG Clua and John DC Linnell
Clua and Linnell (2018)Individual shark profiling: An innovative and environmentally responsible approach for selectively managing human fatalities. Conservation Letters. 2018;e12612. https://doi.org/10.1111/conl.12612

Most shark-induced human fatalities are followed by widespread and unselective culling campaigns that have limited effectiveness and may have high ecological costs for threatened species. The blanket culling strategy implicitly assumes that incident risk is directly correlated with shark density, an assumption that has yet to be demonstrated. We present the alternative hypothesis that incidents are more likely to be caused by behavioral variability among individual sharks than due to shark density. Throughout their ontogenetic development, large species of sharks opportunistically establish a diet that is rarely, if ever, inclusive of humans as a food source. We propose that, some animals with specific behaviors (including boldness) may potentially pose a higher risk than conspecifics. Under this scenario, the risk of a shark attack in a given area would relate to the presence of a limited number of high-risk individuals rather than shark density.

In terms of management of human fatalities, such a hypothesis would favor abandoning general culling campaigns and replacing them with approaches that profile and selectively remove the potential problem individuals, as is done in the terrestrial realm when managing predators that attack humans or livestock.

Evidence of economic benefits for public investment in MPAs

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Pascal, N., Brathwaite, A., Philip, M., Brander, L., Seidl, A. & Clua, E.
Pascal, N., Brathwaite, A., Philip, M., Brander, L., Seidl, A. & Clua, E. (2018). Evidence of economic benefits for public investment in MPAs. Ecosystem Services, 30: 3–13. 10.1016/j.ecoser.2017.10.017

MPAs enhance some of the Ecosystem Services (ES) provided by coral reefs and clear, robust valuations of these impacts may help to improve stakeholder support and better inform decision-makers. Pursuant to this goal, Cost-Benefit Analyses (CBA) of MPAs in 2 different contexts were analysed: a community based MPA with low tourism pressure in Vanuatu, and a government managed MPA with relatively high tourism pressure, in Saint Martin. Assessments were made on six ES: fish biomass, scenic beauty, protection against coastal erosion, bequest and existence values, social capital and CO2 sequestration, which were quantified via different approaches that included experimental fishery, surveys and benefit transfer. Total operating costs for each MPA were collected and the benefit-cost ratio and return on investment based on 25-year discounted projections computed. Sensitivity analyses were conducted on MPA impacts, and discount rates (5%, 7% and 10%). The investment indicators all showed positive results with the impact on the tourism ES being the largest estimated for all MPAs, highlighting the importance of this relationship. The study also demonstrated a relatively high sensitivity of the results to different levels of impacts on ES, which highlights the need for reducing scientific knowledge gaps.

Convergence of marine megafauna movement patterns in coastal and open oceans

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Sequeira, A.M.M., Rodríguez, J.P., Eguíluz, V.M., Clua, E. et al.
Sequeira, A.M.M., Rodríguez, J.P., Eguíluz, V.M., Clua, E. et al. (2018) Convergence of movement patterns of marine megafauna in coastal and open oceans. Proceedings of the National Academy of Science, https://doi.org/10.1073/pnas.1716137115.

The extent of increasing anthropogenic impacts on large marine vertebrates partly depends on the animals’ movement patterns. Effective conservation requires identification of the key drivers of movement including intrinsic properties and extrinsic constraints associated with the dynamic nature of the environments the animals inhabit. However, the relative importance of intrinsic versus extrinsic factors remains elusive. We analyze a global dataset of ∼2.8 million locations from >2,600 tracked individuals across 50 marine vertebrates evolutionarily separated by millions of years and using different locomotion modes (fly, swim,walk/paddle). Strikingly, movement patterns show a remarkable convergence, being strongly conserved across species and independent of body length and mass, despite these traits ranging over 10 orders of magnitude among the species studied. This represents a fundamental difference between marine and terrestrial vertebrates not previously identified, likely linked to the reduced costs of locomotion in water. Movement patterns were primarily explained by the interaction between species-specific traits and the habitat(s) they
move through, resulting in complex movement patterns when moving close to coasts compared with more predictable patterns when moving in open oceans. This distinct difference may be associated with greater complexity within coastal microhabitats, highlighting a critical role of preferred habitat in shaping marine vertebrate global movements. Efforts to develop understanding of the characteristics of vertebrate movement should consider the habitat(s) through which they move to identify how movement patterns will alter with forecasted severe ocean changes, such as reduced Arctic sea ice cover, sea level rise, and declining oxygen content.

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

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Pierpaolo F. Brena, Johann Mourier, Serge Planes and Eric E. Clua
Brena PF, Mourier J, Planes S, Clua EE. 2018 Concede or clash? Solitary sharks competing for food assess rivals to decide. Proc. R. Soc. B 285: 20180006. http://dx.doi.org/10.1098/rspb.2018.0006

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.

Extreme Inverted Trophic Pyramid of Reef Sharks Supported by Spawning Groupers

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Johann Mourier, Jeffrey Maynard, Valeriano Parravicini, Laurent Ballesta, Eric Clua, Michael L. Domeier, Serge Planes
Mourier, J., Maynard, J., Parravicini, V., Ballesta, L., Clua, E., Domeier, M. L. and Planes, S. (2016) Extreme Inverted Trophic Pyramid of Reef Sharks Supported by Spawning Groupers. Current Biology 26: 1-6.

The extent of the global human footprint limits our understanding of what is natural in the marine environment. Remote, near-pristine areas provide some baseline expectations for biomass and suggest that predators dominate, producing an inverted biomass pyramid. The southern pass of Fakarava atoll—a biosphere reserve in French Polynesia—hosts an average of 600 reef sharks, two to three times the biomass per hectare documented for any other reef shark aggregations. This huge biomass of predators makes the trophic pyramid inverted. Bioenergetics models indicate that the sharks require ∼90 tons of fish per year, whereas the total fish production in the pass is ∼17 tons per year. Energetic theory shows that such trophic structure is maintained through subsidies, and empirical evidence suggests that sharks must engage in wide-ranging foraging excursions to meet energy needs. We used underwater surveys and acoustic telemetry to assess shark residency in the pass and feeding behavior and used bioenergetics models to understand energy flow. Contrary to previous findings, our results highlight that sharks may overcome low local energy availability by feeding on fish spawning aggregations, which concentrate energy from other local trophic pyramids. Fish spawning aggregations are known to be targeted by sharks, but they were previously believed to play a minor role representing occasional opportunistic supplements. This research demonstrates that fish spawning aggregations can play a significant role in the maintenance of local inverted pyramids in pristine marine areas. Conservation of fish spawning aggregations can help conserve shark populations, especially if combined with shark fishing bans.

Economic valuation of coral reef ecosystem service of coastal protection: A pragmatic approach

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Nicolas Pascal, Michel Allenbach, Angelique Brathwaite, Lauretta Burke, Guillaume Le Port, Eric Clua
Pascal, N., Allenbach, M., Brathwaite, A., Burke, L., Le Port, G. and Clua, E. (2016) Economic valuation of coral reef ecosystem service of coastal protection: A pragmatic approach. Ecosystem Services 21: 72-80.

The identification and economic valuation of ecosystem services (ES) are becoming important components of coral reef management. In many contexts, protection of human assets against coastal floods is one of the most important ES provided by coral reefs. The methods utilized to characterize this ES should be able to accommodate situations with low data availability, without sacrificing robustness. In this paper, we suggest such an approach that utilizes expert opinion and does not require copious amounts of data. Our primary objective is to find a balance between simple and complex models that can be used in a data scarce environment, to produce an economic valuation of the coral reef ES of protection against coastal floods. The approach has three steps: (i) identify geographic zones and assets at risk, (ii) identify the contributing role of coral reefs in the protection of coasts and, (iii) value the annual repair costs of assets through the avoided damage cost approach. The proposed method seems appropriate for advocacy with policy makers, but appears to be less effective for small scale approaches, such as those required for Payment for ES negotiations or marine spatial planning.

First record of giant trevally (Caranx ignobilis) around the Clipperton Atoll-La Passion Island (North-Eastern Tropical Pacific)

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Eric Clua, Sandra Bessudo, Michel Labrecque, Mauricio Hoyos-Padilla
Clua, E., Bessudo, S., Labrecque, M. and Hoyos-Padilla, M. (2016) First record of giant trevally (Caranx ignobilis) around the Clipperton Atoll-La Passion Island (North-Eastern Tropical Pacific). Cybium 40(3): 259-260.

As the giant trevally is absent from the list by Fourrière et al. (2014), we present here the first record of C. ignobilis in the waters surrounding the Clipperton atoll.

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

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Guillaume Costalat, Jeremy Coquart, Ingrid Castres, Fabrice Joulia, Olivier Sirost, Eric Clua, Frédéric Lemaître
Costalat, G., Coquart, J., Castres, I., Joulia, F., Sirost, O., Clua, E. and Lemaître, F. (2016) The oxygen-conserving potential of the diving response: A kinetic-based analysis. Journal of Sports Sciences. DOI: 10.1080/02640414.2016.1183809.

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.

Species identification of the shark involved in the 2007 Lifou fatal attack on a swimmer: A reply to Tirard et al. (2015)

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Eric Clua, Bernard Séret
Clua, E. and Séret, B. (2016) Species identification of the shark involved in the 2007 Lifou fatal attack on a swimmer: A reply to Tirard et al. (2015). Journal of Forensic and Legal Medicine 40: 58-60.

Based on new photographs of the wound, Tirard et al. (2015) tried to demonstrate that the shark involved in a fatal attack on a human in Lifou in 2007 had homodont teeth and that it sawed the femur instead of directly cutting it, promoting the hypothesis that it was a tiger shark instead of a white shark. They also contested the data provided by the direct witness of the attack about the behaviour of the shark, specific to this former species. The evidences they provide are not convincing and, based on the absence of tissue loss and description of a jumping behaviour, we still believe that it was a single bite-and-spit attack by a white shark.

Misidentification of video clips and missing sequences among supplementary online material from Martin (2007)

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Eric Clua
Clua, E. (2016) Misidentification of video clips and missing sequences among supplementary online material from Martin (2007). Marine and Freshwater Behaviour and Physiology.

Sharks are marine predators that constitute a potential threat to humans and their specific behaviours often play a critical role in triggering fatal attacks (Clua & Séret 2010; Clua & Reid 2013; Clua et al. 2014). Several authors have recently outlined the economic importance of shark-based ecotourism which far outweighs the single-use income obtained from fishing (Clua et al. 2011; Gallagher & Hammerschlag 2011; Vianna et al. 2012). However, the development of such activities increases the potential interactions between sharks and humans and the correlated risk of accidental bites (Brena et al. 2015), in particular when unsuitable provisioning practices such as hand-feeding are implemented (Clua & Torrente 2015). In such a context, a better understanding of shark agonistic behaviours is indispensable.

In a valuable response to this challenge, Aidan Martin published a paper entitled ‘A review of shark agonistic displays: comparison of display features and implications for shark-human interactions’ (Martin 2007). This paper focuses on the behaviour of 23 shark species and includes supplementary online material (SOM) composed of several videos displaying specific shark behaviours (see Figure 1). These are available at: http://www.tandfonline.com/ doi/suppl/10.1080/10236240601154872. The nine files that can be downloaded include the following common text ‘gmfw_a_215414_sup_’ with an additional number ranging from 001 to 009; a short description of the video clip is also provided. One would expect these files that are ranked by increasing numbers to correspond to the video clips in their citation order in the text (as video clip 1, 2, etc.). There are, however, some discrepancies between the text and the SOM as well as some missing material.

Possible collapse of reef shark populations in remote coral reef ecosystems in the Coral Sea (Western Pacific)

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Eric Clua, Thomas Vignaud
Clua, E. and Vignaud, T. (2016) Possible collapse of reef shark populations in remote coral reef ecosystems in the Coral Sea (Western Pacific). Cybium, International Journal of Ichthyology.

The decline of meso-predators such as reef sharks is a concern as such species can have important ecological roles in maintaining reef ecosystem resilience. Two field trips conducted in August 2010 and November 2011 to the Chesterfield archipelago (Coral Sea) allowed us to assess the abundances and average sizes of medium-bodied Carcharhinidae with a specific focus on grey reef shark (Carcharhinus amblyrhynchos), through fishing (46 hours of accumulated effort) and underwater visual censuses (25 hours of accumulated effort). We found low abundance and small average total length (TL) for all reef shark species, and in the case of the grey reef shark, an average abundance of 2.1 individuals/dive with the majority of animals less than 110 cm TL. We compared our findings with historical data and, given our low sampling effort, we so far hypothesise that a general strong decline in the reef shark populations may have occurred in this area, probably due to recent overfishing. The enforcement of conservation measures is strongly recommended among these remote reefs as well as complementary studies for confirming this hypothesis.

Determining the Role of Hand Feeding Practices in Accidental Shark Bites on Scuba Divers

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Eric Clua, Frédéric Torrente
Clua, E. and Torrente, F. (2015) Determining the Role of Hand Feeding Practices in Accidental Shark Bites on Scuba Divers. Journal of Forensic Science and Criminology 5: 502.

Shark-based ecotourism is significantly developping around the world, often without appropriate management of risk. This activity involves a risk of accidental bites on divers that can be quite severe or even fatal. Objectives: To determine if ecotourism companies’ liability can be engaged in the context of bites on scuba divers in presence of hand-feeding practices, supporting the legitimacy of financial compensation for the victims. Methods: We analyzed the development from the mid-eighties to 2010 of shark-based ecotourism through artificial provisioning practices in Moorea island (French Polynesia) and more specifically the features and motivation of two bites on divers by Sicklefin Lemon sharks. Results: The specific practice of hand-feeding can be considered as a facilitating factor for accidental bites on divers, potentially involving the diving operator’s responsability. Conclusions: Our findings should support the technical work of experts that might be called in such cases.

Pattern of movements from a home reef in the Chesterfield Islands (Coral Sea) by the endangered Giant Grouper, Epinephelus lanceolatus

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Eric Clua, Claude Chauvet, Johann Mourier, Jonathan Mark Werry, John E. Randall
Clua, E., Chauvet, C., Mourier, J., Werry, J. M. and Randall, J. E. (2015) Pattern of Movements from a home reef in the Chesterfield Islands (Coral sea) by the Giant Grouper, Epinephelus lanceolatus. Aquatic Living Resources, 28: 53-58.

This study determined the movements of a Giant Grouper, Epinephelus lanceolatus, in which an acoustic tag was surgically implanted and monitored by an array of six VR2W acoustic receiver units from August 2010 to January 2013 in the remote, uninhabited Chesterfield Islands, Coral Sea (800 km West of New Caledonia). Our data revealed a home reef area (residency rate of 44.9%) with an increased activity revealed by movements at dawn and dusk toward and between two adjacent reef passages, probably for foraging. The fish was absent from its resident reef between October and December 2010 and 2012, corresponding to the time known for spawning aggregations of this species in New Caledonia. A skipped spawning seems to have occurred in 2011. We hope these data will be complemented in the future by locating the spawning site or sites and thus provide adequate conservation measures. The Coral Sea links two World Heritage Sites, the Australian Great Barrier Reefs and the New Caledonian coral reefs. It would be fitting to create a Marine Protected Area for the Chesterfield Islands between these two major conservation areas of the sea.

Shark and ray provisioning: functional insights into behavioral, ecological and physiological responses across multiple scales

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Pierpaolo F. Brena, Johann Mourier, Serge Planes, Eric Clua
Brena, P. F., Mourier, J., Planes, S. and Clua, E. (2015) Shark provisioning: functional insights into behavioral, ecological and physiological responses across multiple scales. Mar. Ecol. Prog. Ser. 538: 273-283.

The use of olfactory stimuli and the provision of food are a common practice to prompt artificial aggregations of emblematic wild species and ensure the economic viability of the wildlife-watching industry. Several elasmobranch species have been targeted by such operations in a variety of locations for over four decades. A recent review succinctly addressed the potential effects of shark diving tourism, including shark provisioning, on shark individual behavior and ecology, but the general paucity of data on the ecology of elasmobranchs precluded general statements. By using a functional framework, we reviewed the findings of the 22 available studies that investigated the behavioral, physiological, and ecological response of 14 shark and three ray species targeted by artificial provisioning. Focusing on the underlying processes that rule the response of targeted elasmobranch species, we report further effects acting beyond the individual-scale and their cross-scale relationships. We suggest that the most commonly described alterations of individual movement patterns have cascading effects through the group and community-scale, ultimately resulting in altered health condition and individual behavior toward humans. We conclude by stressing the potential for provisioning activities to support the investigation of complex ecological and behavioral processes in elasmobranchs.

Insight into cumulative intra-guild and intra-specific depredation among sharks

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Eric Clua, Jean-Baptiste Galves, Jonathan Mark Werry
Clua, E., Galves, J-B. and Werry, J. M. (2014) Insight into cumulative intra-guild and intra-specific depredation among sharks. Cybium 38(4): 311-313.

Intra-guild predation (IGP), depredation of hooked sharks and cannibalism by large individuals on smaller conspecifics have been documented for both tiger shark Galeocerdo cuvier and bull shark Carcharhinus leucas. In this paper we report a case study of cumulative inter and intra-depredation from the Indian and the Pacific Ocean involving large G. cuvier and C. leucas adults at the final stage. These findings further the hypothesis that IGP and cannibalism among adult sharks occurs in the wild and potentially influences predator-prey relationships in tropical marine ecosystems.

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

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Thomas M. Vignaud, Johann Mourier, Jeffrey A. Maynard, Raphael Leblois, Julia Spaet, Eric Clua, Valentina Neglia, Serge Planes
Vignaud, T. M., Mourier, J., Maynard, J. A., Leblois, R., Spaet, J., Clua, E., Neglia, V. and Planes, S. (2014) Blacktip reef sharks, Carcharhinus melanopterus, have high genetic structure and varying demographic histories in their Indo-Pacific range. Molecular ecology 23, 5193–5207.

For free-swimming marine species like sharks, only population genetics and demo-graphic 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. Fourteen 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.

Fatal attack by a juvenile tiger shark, Galeocerdo cuvier, on a kitesurfer in New Caledonia (South Pacific)

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Eric Clua, Pierre-Marie Bescond, Dennis Reid
Clua, E., Bescond, P-M. and Reid, D. (2014) Fatal attack by a juvenile tiger shark, Galeocerdo cuvier, on a kitesurfer in New Caledonia (South Pacific). Journal of Forensic and Legal Medicine 25: 67-70.

We present a case of a non-provoked fatal shark attack on a 15-year old male kitesurfer in New Caledonia. The victim lost his board and was pulled by the sail along the water surface in a reef passage when a shark attacked. The shark inflicted at least two bites on the left leg, including a severe one around the knee, resulting in a quick hypovolemic shock that was fatal. The analysis of one of these bites indicated that a 2.8 m TL (est. length) tiger shark was responsible for this attack. The features of the attack are consistent with those of a predator response to a surface feeding stimulus.

Biological Data of Pygmy Killer Whale (Feresa attenuata) from a Mass Stranding in New Caledonia (South Pacific) Associated with Hurricane Jim in 2006

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Eric Clua, Charles A. Manire, Claire Garrigue
Clua, E., Manire, C. A. and Garrigue, C. (2014) Biological Data of Pygmy Killer Whale (Feresa attenuata) from a Mass Stranding in New Caledonia (South Pacific) Associated with Hurricane Jim in 2006. Marine Mammals, 40(2): 162-172. DOI: 10.1578/AM.40.2.2014.162.

Despite its distribution throughout the tropics and subtropics, the pygmy killer whale (Feresa attenuata) is one of the most poorly known species of odontocetes (Cetacea: Delphinidae). We used the opportunity of a mass stranding of six animals in New Caledonia (early February 2006) to gather information about their biology. Four animals, including three males and one female, were found dead, and morphometrics, including dental counts, were collected. Two live mature males of 236 and 246 cm total length (TL), respectively, were closely monitored and sampled via blood analysis. As it was not likely to survive, the second animal was euthanized and necropsied. Following the euthanasia of the larger animal, the smaller one, which was probably staying out of social solidarity, returned on its own to the open sea. The necropsy revealed the presence of cardiopulmonary collapse and enlarged and congested testes. Blood parameters confirmed a deteriorating health status for both animals, enhanced by starvation. Some of the relative morphometric measurements of all six stranded pygmy killer whales seemed to be larger for these animals living in the southwest Pacific as compared to the literature for this species. We hypothesize that this group of pygmy killer whales was probably pushed through the Coral Sea toward the New Caledonian lagoon by Hurricane Jim, which occurred in the area from 26 January until 2 February. These observations reveal January as a potential part of the mating season in this area for this rare, elusive, and unknown species. It also supports the notion that early sacrifice of distressed, terminal animals could be a way to improve the survival rate of other less traumatized individuals during cetacean mass strandings.

Shark-watching ecotourism in the Pacific islands: A move towards “payments for ecosystem services”?

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Eric Clua, Nicolas Pascal
Clua, E. and Pascal, N. (2014) Shark-watching ecotourism in the Pacific islands: A move towards “payments for ecosystem services”?. SPC Fisheries Newsletter #144: 30-34.

Ecotourism based on observing iconic animal species in their natural habitats has become increasingly popular around the world and the Pacific Islands are no exception to this trend. Among these iconic species, sharks hold a special place as an attraction for tourists, including tourists who are not divers. One of the characteristics of this activity is the need for artificial feeding to ensure that there are enough animals present in a specific spot to be observed. So, while shark watching undeniably provides significant levels of income to local economies, it does, however, raise a certain number of problems in terms of its impact on the ecosystem, human safety and even a legitimate distribution of the dividends it generates (Clua et al. 2011). Another advantage of this activity is that it strengthens sharks’ economic value in the eyes of decision-makers at a time when these animals are generally being overfished throughout the world (Clarke et al. 2006), in spite of scientific warnings about the need to preserve these super-predators within their marine ecosystems. Against a backdrop in which environmental arguments have shown their limits over the past few decades in terms of providing any real protection, an economic approach appears to be both complementary and necessary to ensure the sustainable development of shark populations in the South Pacific (Vianna et al. 2012). The purpose of this article is to present the general outlines of such an economic approach, highlighting ecotourism as a virtuous use of sharks that makes it possible to generate income while maintaining them in their ecosystem. Nevertheless, this approach is not totally virtuous unless it respects the three fundamental aspects of sustainable development: 1) environmental, 2) social, and 3) economic. This goal will only be reached through the implementation of “payments for ecosystem services” as we will attempt to demonstrate.

Écotourisme basé sur l’observation des requins dans le Pacifique insulaire : vers des « paiements pour services écosystémiques » ?

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Eric Clua, Nicolas Pascal
Clua, E. et Pascal, N. (2014) Écotourisme basé sur l’observation des requins dans le Pacifique insulaire : vers des « paiements pour services écosystémiques » ?. Lettre d’information sur les pêches de la CPS n°144 : 30-34.

L’écotourisme basé sur l’observation dans leur milieu d’animaux emblématiques est en plein essor à travers le Monde, et le Pacifique insulaire n’échappe pas à la tendance. Parmi ces espèces, les requins occupent une place privilégiée en tant qu’attraction pour les touristes, sans qu’ils soient forcément plongeurs. Une des caractéristiques de cette activité repose sur la nécessité d’avoir recours au nourrissage artificiel (feeding en anglais) afin d’assurer la présence d’animaux en quantité importante et en un lieu précis pour favoriser l’observation. Aussi, s’il permet indéniablement de dégager des revenus conséquents pour les économies locales, l’écotourisme basé sur l’observation des requins n’est pas sans poser certains problèmes en terme d’impact sur les écosystèmes, de sécurité pour l’Homme, voire de répartition légitime des dividendes qu’il génère (Clua et al. 2011). Un autre atout de cette activité repose sur le fait qu’elle renforce la valeur économique des requins aux yeux des décideurs politiques et ce, dans un contexte où ces animaux sont globalement surpêchés à l’échelle planétaire (Clarke et al. 2006), malgré les mises en gardes des scientifiques sur la nécessité de conserver ces super-prédateurs au sein des écosystèmes marins. Dans un contexte où les arguments écologiques ont montré leur limite sur les dernières décennies pour obtenir une vraie protection, une approche économique semble complémentaire et nécessaire pour assurer un développement durable des populations de requins dans le Pacifique Sud (Vianna et al. 2012). L’objet de cet article est de présenter les grands axes d’une telle approche économique, en mettant l’accent sur l’écotourisme en tant qu’utilisation vertueuse des requins, permettant de dégager des revenus tout en les maintenant dans leur écosystème. Cette approche ne peut néanmoins être totalement vertueuse que si elle respecte les trois volets fondateurs du développement durable, à savoir le volet économique, mais aussi les volets écologique et social. Cet objectif ne sera atteint que via la mise en place de « paiements pour services écosystémiques », comme nous allons tenter de le démontrer.

Reef-Fidelity and Migration of Tiger Sharks, Galeocerdo cuvier, across the Coral Sea

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Jonathan M. Werry, Serge Planes, Michael L. Berumen, Kate A. Lee, Camrin D. Braun, Eric Clua
Werry, J. M., Planes, S., Berumen, M. L., Lee, K. A., Braun, C. D. and Clua, E. (2014) Reef-Fidelity and Migration of Tiger Sharks, Galeocerdo cuvier, across the Coral Sea. PLoS ONE 9(1): e83249.

Knowledge of the habitat use and migration patterns of large sharks is important for assessing the effectiveness of large predator Marine Protected Areas (MPAs), vulnerability to fisheries and environmental influences, and management of shark-human interactions. Here we compare movement, reef-fidelity, and ocean migration for tiger sharks, Galeocerdo cuvier, across the Coral Sea, with an emphasis on New Caledonia. Thirty-three tiger sharks (1.54 to 3.9 m total length) were tagged with passive acoustic transmitters and their localised movements monitored on receiver arrays in New Caledonia, the Chesterfield and Lord Howe Islands in the Coral Sea, and the east coast of Queensland, Australia. Satellite tags were also used to determine habitat use and movements among habitats across the Coral Sea. Sub-adults and one male adult tiger shark displayed year-round residency in the Chesterfields with two females tagged in the Chesterfields and detected on the Great Barrier Reef, Australia, after 591 and 842 days respectively. In coastal barrier reefs, tiger sharks were transient at acoustic arrays and each individual demonstrated a unique pattern of occurrence. From 2009 to 2013, fourteen sharks with satellite and acoustic tags undertook wide-ranging movements up to 1114 km across the Coral Sea with eight detected back on acoustic arrays up to 405 days after being tagged. Tiger sharks dove 1136 m and utilised three-dimensional activity spaces averaged at 2360 km(3). The Chesterfield Islands appear to be important habitat for sub-adults and adult male tiger sharks. Management strategies need to consider the wide-ranging movements of large (sub-adult and adult) male and female tiger sharks at the individual level, whereas fidelity to specific coastal reefs may be consistent across groups of individuals. Coastal barrier reef MPAs, however, only afford brief protection for large tiger sharks, therefore determining the importance of other oceanic Coral Sea reefs should be a priority for future research.

Sex-based spatial segregation of adult bull sharks, Carcharhinus leucas, in the New Caledonian great lagoon

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Jonathan Mark Werry, Eric Clua
Werry, J. M. and Clua, E. (2013) Sex-based spatial segregation of adult bull sharks, Carcharhinus leucas, in the New Caledonian great lagoon. Aquatic Living Resources, 26(04), 281-288.

Conservation of threatened large sharks and management of shark-human interactions requires an understanding of shark occurrence and movement patterns. Here, we present the first catch, movement and behaviour data of adult bull sharks, Carcharhinus leucas, in New Caledonia. Amongst six adult C. leucas tagged with passive acoustic tags, four females were caught in coastal waters while males were only found at an isolated oceanic barrier coral reef over 100 km from the nearest river mouth. Two females were monitored in the southern New Caledonia lagoon for 707 and 208 days respectively and displayed classical transient behaviour and sporadic short-term residency around a coastal reef bay, with movements in and out a river detected prior to spring. Adult C. leucas in New Caledonia may develop a sex-based spatial segregation with an atypical presence of adult males in oceanic environments, probably influenced by the unique estuarine-marine continuum of the New Caledonian great lagoon.

Genetic Network and Breeding Patterns of a Sicklefin Lemon Shark (Negaprion acutidens) Population in the Society Islands, French Polynesia

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Johann Mourier, Nicolas Buray, Jennifer K. Schultz, Eric Clua, Serge Planes
Mourier, J., Buray, N., Schultz, J. K., Clua, E. and Planes, S. (2013) Genetic Network and Breeding Patterns of a Sicklefin Lemon Shark (Negaprion acutidens) Population in the Society Islands, French Polynesia. PLoS ONE 8(8): e73899.

Human pressures have put many top predator populations at risk of extinction. Recent years have seen alarming declines in sharks worldwide, while their resilience remains poorly understood. Studying the ecology of small populations of marine predators is a priority to better understand their ability to withstand anthropogenic and environmental stressors. In the present study, we monitored a naturally small island population of 40 adult sicklefin lemon sharks in Moorea, French Polynesia over 5 years. We reconstructed the genetic relationships among individuals and determined the population’s mating system. The genetic network illustrates that all individuals, except one, are interconnected at least through one first order genetic relationship. While this species developed a clear inbreeding avoidance strategy involving dispersal and migration, the small population size, low number of breeders, and the fragmented environment characterizing these tropical islands, limits its complete effectiveness.

Features and motivation of a fatal attack by a juvenile white shark, Carcharodon carcharias, on a young male surfer in New Caledonia (South Pacific)

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Eric Clua, Dennis Reid
Clua, E. and Reid, D. (2013) Features and motivation of a fatal attack by a juvenile white shark, Carcharodon carcharias, on a young male surfer in New Caledonia (South Pacific). Journal of Forensic and Legal Medicine. DOI: 10.1016/j.jflm.2013.03.009.

We present the case of a non provoked fatal shark attack on a 19-year old male surfer in New Caledonia. Several severe bites removed the right arm and all flesh from the right thigh, provoking a quick hypovolemic shock that was fatal. The information provided by a witness and the analysis of a partial bite on the right calf allowed us to identify a 2.7 m TL (est. length) white shark as responsible for this attack. The features of the attack are consistent with a young predator motivated by hunger and the development of its predatory skills.

Microsatellite Analyses of Blacktip Reef Sharks (Carcharhinus melanopterus) in a Fragmented Environment Show Structured Clusters

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Thomas Vignaud, Eric Clua, Johann Mourier, Jeffrey Maynard, Serge Planes
Vignaud, T., Clua, E., Mourier, J., Maynard, J. and Planes, S. (2013) Microsatellite Analyses of Blacktip Reef Sharks (Carcharhinus melanopterus) in a Fragmented Environment Show Structured Clusters. PLoS ONE 8(4): e61067.

The population dynamics of shark species are generally poorly described because highly mobile marine life is challenging to investigate. Here we investigate the genetic population structure of the blacktip reef shark (Carcharhinus melanopterus) in French Polynesia. Five demes were sampled from five islands with different inter-island distances (50-1500 km). Whether dispersal occurs between islands frequently enough to prevent moderate genetic structure is unknown. We used 11 microsatellites loci from 165 individuals and a strong genetic structure was found among demes with both F-statistics and Bayesian approaches. This differentiation is correlated with the geographic distance between islands. It is likely that the genetic structure seen is the result of all or some combination of the following: low gene flow, time since divergence, small effective population sizes, and the standard issues with the extent to which mutation models actually fit reality. We suggest low levels of gene flow as at least a partial explanation of the level of genetic structure seen among the sampled blacktip demes. This explanation is consistent with the ecological traits of blacktip reef sharks, and that the suitable habitat for blacktips in French Polynesia is highly fragmented. Evidence for spatial genetic structure of the blacktip demes we studied highlights that similar species may have populations with as yet undetected or underestimated structure. Shark biology and the market for their fins make them highly vulnerable and many species are in rapid decline. Our results add weight to the case that total bans on shark fishing are a better conservation approach for sharks than marine protected area networks.

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

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Yann Laurans, Nicolas Pascal, Thomas Binet, Luke Brander, Eric Clua, Gilbert David, Dominique Rojat, Andrew Seidl
Laurans, Y., Pascal, N., Binet, T., Brander, L., Clua, E., David, G., Rojat, D. and Seidl, A. (2013) Economic valuation of ecosystem services from coral reefs in the South Pacific: Taking stock of recent experience. Journal of Environmental Management 116: 135-144.

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.

Behavioural patterns of a Tiger Shark (Galeocerdo cuvier) feeding aggregation at a blue whale carcass in Prony Bay, New Caledonia

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Eric Clua, Claude Chauvet, Tyffen Read, Jonathan M. Werry, Shing Y. Lee
Clua, E., Chauvet, C., Read, T., Werry, J. M. and Lee, S. Y. (2013) Behavioural patterns of a Tiger Shark (Galeocerdo cuvier) feeding aggregation at a blue whale carcass in Prony Bay, New Caledonia. Marine and Freshwater Behaviour and Physiology, 46:1, 1-20.

Tiger Sharks, Galeocerdo cuvier, are large top-level predators usually solitary as adults. Observation of their scavenging activity on the carcass of a dead whale offered a rare opportunity for better understanding the pattern of intra-specific behaviour within the aggregations of these large predators. In January 2002, the stranding, subsequent death and consumption of a 17.4m total length (TL) blue whale, Balaenoptera musculus, was observed and filmed in Prony Bay, southern New Caledonia. After three weeks of confinement in the bay, the cetacean was killed by adult bullsharks Carcharhinus leucas. The first adult Tiger Shark was subsequently observed around the carcass after 36h. The fat slicks from the carcass attracted further Tiger Sharks which arrived after an additional 24h. The use of photo-identification on video footage collected during four observation sessions over an eight-day period identified 46 individual Tiger Sharks (primarily adult females between 3.3 and 4m TL) participating in the feeding aggregation. Only four animals were identified in two seperate observation sessions (over two consecutive days), suggesting a short-term residency pattern of several hours (<36h) around the carcass. As the arrival time of Tiger Sharks to the carcass differed, most arrivals of a new participant were followed by a frenzied period of intense intra-specific interaction. Different biting and agonistic behaviours were demonstrated by the Tiger Sharks on the carcass, including three new behaviours previously undescribed for this species. Size and level of aggressiveness appeared to be the determining factors of dominance amongst Tiger Sharks. These observations and analysis demonstrate that systematic study of feeding aggregations supported by photo-identification could contribute to knowledge of large shark ecology when coupled with capture-recapture, genetic fingerprinting and tagging techniques.

Prevalence and proposal for cost-effective management of the ciguatera risk in the Noumea fish market, New Caledonia (South Pacific)

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Eric Clua, Pierpaolo F. Brena, Côme Lecasble, Reine Ghnassia, Claude Chauvet
Clua, E., Brena, P. F., Lecable, C., Ghnassia, R. and Chauvet, C. (2011) Prevalence and proposal for cost-effective management of the ciguatera risk in the Noumea fish market, New Caledonia (South Pacific). Toxicon 58: 591-601.

Ciguatera fish poisoning (CFP) is a common intoxication associated with the consumption of reef fish, which constitutes a critical issue for public health in many countries. The complexity of its epidemiology is responsible for the poor management of the risk in tropical fish markets. We used the example of the Noumea fish market in New Caledonia to develop a cost-effective methodology of assessing the CFP risk. We first used published reports and the knowledge of local experts to define a list of potentially poisonous local species, ranked by their ciguatoxic potential. Based on two 1-month surveys in the market, conducted in winters 2008 and 2009, we then calculated the consolidated ratio of biomass of potentially poisonous species vs. total biomass of fish sold on the market. The prevalence of high CFP-risk species in the market was 16.1% and 18.9% in 2008 and 2009, respectively. The most common high CFP risk species were groupers (serranids), king mackerels (scombrids), snappers (lutjanids), barracudas (sphyaraenids), emperors (lethrinids) and wrasses (labrids). The size (age) of the fish also plays a critical role in the potential ciguatoxic risk. According to proposals of average size thresholds provided by experts for high-risk species, we were also able to assess the additional risk induced by the sale of some large fish on the market. The data collected both from experts and from the market allowed us to develop a cost-effective proposal for improving the management of the CFP risk in this market. However, the successful implementation of any regulation aiming to ban some specific species and sizes from the market, with an acceptable economical impact, will require the improvement of the expertise in fish identification by public health officers and, ideally, the commitment of retailers.

Business partner or simple catch? The economic value of the sicklefin lemon shark in French Polynesia

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Eric Clua, Nicolas Buray, Pierre Legendre, Johann Mourier, Serge Planes
Clua, E., Buray, N., Legendre, P., Mourier, J. and Planes, S. (2011) Business partner or simple catch? The economic value of the sicklefin lemon shark in French Polynesia. Marine and Freshwater Research 62: 764-770.

Most arguments invoked so far by the scientific community in favour of shark conservation rely on the ecological importance of sharks, and have little impact on management policies. During a 57-month study, we were able to individually recognise 39 sicklefin lemon sharks that support a shark-feeding ecotourism activity in Moorea Island, French Polynesia. We calculated the direct global revenue generated by the provisioning site, based on the expenses of local and international divers. The total yearly revenue was around USD5.4 million and the 13 sharks most often observed at the site had an average contribution each of around USD316 699. Any one of these sharks represents a potential contribution of USD2.64 million during its life span. We argue that publicising economic values per individual will be more effective than general declarations about their ecological importance for convincing policy makers and fishers that a live shark is more valuable than a dead shark for the local economy. Studies monitoring the potential negative ecological effects of long-term feeding of sharks should, however, be conducted to ensure these are also considered. Besides declarations about the nonconsumptive direct-use value of sharks, as promoted by ecotourism, the calculation of their other economic values should also benefit shark conservation.

Effects of provisioning on shark behaviour: Reply to Brunnschweiler & McKenzie (2010)

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Eric Clua, Nicolas Buray, Pierre Legendre, Johann Mourier, Serge Planes
Clua, E., Buray, N., Legendre, P., Mourier, J. and Planes, S. (2010) Effects of provisioning on shark behaviour: Reply to Brunnschweiler & McKenzie (2010). Marine Ecology Progress Series 420: 285-288.

Brunnschweiler & McKenzie (2010; Mar Ecol Prog Ser 420:283–284) expressed reservations over the findings of Clua et al. (2010; Mar Ecol Prog Ser 414:257–266), mostly related to the lack of a reference site or a control group in the methodology. In our study, we distinguished between 39 individuals of sicklefin lemon sharks Negaprion acutidens, mainly based on photo-identification. Our study was based on the field-survey approach, with time (a continuous variable) as the source of variation, and thus a control group was not necessary. We provide here additional data that support the notion that abundance of lemon sharks on the provisioning site was increasing, both in their number and fidelity. We maintain our conclusion that sicklefin lemon shark provisioning off Moorea Island can continue, but should be more intensely controlled.

Unprovoked Fatal Shark Attack in Lifou Island (Loyalty Islands, New Caledonia, South Pacific) by a Great White Shark, Carcharodon carcharias

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Eric Clua, Bernard Séret
Clua, E. and Séret, B. (2010) Unprovoked Fatal Shark Attack in Lifou Island (Loyalty Islands, New Caledonia, South Pacific) by a Great White Shark, Carcharodon carcharias. American Journal of Forensic Medecine and Pathology 31: 281-286.

The case of a fatal, unprovoked shark attack is reported and analyzed. The incident took place on the 30th of September 2007, in the lagoon of Luengoni Bay, Lifou Island (Loyalty Islands, New Caledonia). A young French woman who was snorkeling was severely bitten on the right thigh and died of hemorrhage. An analysis based in particular on the size and color of the shark, the characteristics of the wounds, and the behavior of the shark before and after the bite suggests that the aggressor was a great white shark, Carcharodon carcharias.

Behavioural response of sicklefin lemon sharks Negaprion acutidens to underwater feeding for ecotourism purposes

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Eric Clua, Nicolas Buray, Pierre Legendre, Johann Mourier, Serge Planes
Clua, E., Buray, N., Legendre, P., Mourier, J. and Planes, S. (2010) Behavioural response of sicklefin lemon sharks (Negaprion acutidens) to underwater feeding for ecotourism purposes. Marine Ecology Progress Series 414: 257-266.

The feeding of marine predators is a popular means by which tourists and tour operators can facilitate close observation and interaction with wildlife. Shark-feeding has become the most developed provisioning activity around the world, despite its controversial nature. Amongst other detrimental effects, the long-term aggregation of sharks can modify the natural behaviour of the animals, potentially increase their aggression toward humans, and favour inbreeding. During 949 diving surveys conducted over 44 mo, we investigated the ecology and residence patterns of 36 photoidentified adult sicklefin lemon sharks Negaprion acutidens. The group contained 20 females and 16 males. From this long-term survey, we identified 5 different behavioural groups that we described as ‘new sharks’ (7), ‘missing sharks’ (4), ‘resident sharks’ (13), ‘unpredictable sharks’ (5) and ‘ghost sharks’ (7). In spite of movements in and out of the area by some males and females, which were probably related to mating, the general trend was that residency significantly increased during the study, particularly in males, showing a risk of inbreeding due to the reduction of shark mobility. Intraand interspecific aggression was also witnessed, leading to an increased risk of potentially severe bites to humans. Our findings suggest the need for a revision of the legal framework of the provisioning activity in French Polynesia, which could include a yearly closure period to decrease shark behavioural modifications due to long-term shark-feeding activities.

Ventilatory function in breath-hold divers: effect of glossopharyngeal insufflation

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Frederic Lemaître, Eric Clua, Bernard Andréani, Ingrid Castres, Didier Chollet
Lemaître, F., Clua, E., Andreani, B., Castres, I. and Chollet, D. (2010) Ventilatory function in breath-hold divers: effect of glossopharyngeal insufflation. European Journal of Applied Physiology 108: 741-747.

This study was conducted to determine whether ventilatory parameters would change in breath-hold divers (BHDs) after they performed the glossopharyngeal technique for lung insufflation. Fifteen elite BHDs, 16 non-expert BHDs and 15 control subjects participated in this cross-sectional study. Volumes and expiratory flow rates were measured twice, before and after the glossopharyngeal technique performed at rest. Before the technique, greater forced vital capacity (FVC) and forced expiratory volume in 1 s (FEV(1)) and lower FEV(1)/FVC were noted in the elite and non-expert BHDs compared with controls. No difference was noted regarding the other pulmonary parameters. After the technique, increases were noted in FVC, FEV(1) and maximal voluntary ventilation in the elite BHDs (P < 0.001, respectively). The FEF(25-75%)/FVC ratios were lower in the BHDs both before and after the technique, indicating possible dysanapsis. The ventilatory parameters observed after the glossopharyngeal technique indicated (1) higher lung volumes in expert BHDs and (2) a correlation with BHD performance (maximal dynamic BH performance). This correlation became more significant after the technique, indicating a positive effect of glossopharyngeal insufflation on performance.

Underwater photo-identification of sicklefin lemon sharks, Negaprion acutidens, at Moorea (French Polynesia)

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Nicolas Buray, Johann Mourier, Serge Planes, Eric Clua
Buray, N., Mourier, J., Planes, S. and Clua, E. (2009) Underwater photo-identification of sicklefin lemon sharks, Negaprion acutidens, at Moorea (French Polynesia). Cybium, International Journal of Ichthyology; 33, 21-27.

Shark feeding is a controversial recreational activity that may alter shark behaviour. In order to investigate possible behavioural changes at the level of the individual, it is necessary to recognise each shark underwater and in a nonintrusive way. In this study, we tested a protocol based on natural marks on fins, and coloured spots and scars on the body to differentiate individual sicklefin lemon sharks. We found that a feeding group, aggregated for 26 months at a northern location off Moorea Island, comprised 32 animals (19 females and 13 males), identified from 2589 observations made over 541 dives. Post-dive photo-identification of individual sharks was a reliable technique, whereas a high level of skill was required to ensure an instantaneous identification underwater. However, direct underwater identification of individual sharks can be of potential use in shark behavioural studies.

The capture and culture of postlarval fish and invertebrates for the marine ornamental trade

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Johann D. Bell, Eric Clua, Cathy A. Hair, René Galzin, Peter J. Doherty
Bell, J. D., Clua, E., Hair, C. A., Galzin, R. and Doherty, P. J. (2009) The capture and culture of postlarval fish and invertebrates for the marine ornamental trade. Reviews in Fisheries Science. 17(2): 223-240.

Governments, non-government organizations, and other stakeholders are striving to develop practices, policies, and vehicles to make the tropical marine ornamental trade sustainable. Small-scale fisheries based on post-larval capture and culture (PCC) promise to contribute to this goal by (1) removing the risk of damaging corals (inherent in harvesting adults of target species established on reefs) by collecting post-larvae with light traps, nets, and purpose-built temporary shelters as they settle from the plankton to the substrate; and (2) translating the high mortality of post-larvae at settlement into high rates of survival in culture. Possible concerns about overfishing of post-larvae, harvesting the juveniles after they have run the gauntlet of predation at settlement, and the large proportion of bycatch can be eliminated or greatly alleviated by restricting the size and quantity of fishing gear, designing it to retain bycatch alive, and releasing bycatch at times and places that minimize predation. However, special caution is needed when PCC is used at small, isolated islands with self-replenishing populations. Although PCC is environmentally friendly, its contribution to the ornamental trade is expected to be limited. Large variation in the abundance and species composition of settling post-larvae among years, the logistics and costs of operating labor-intensive operations in remote locations, and competition with responsible enterprises harvesting wild adults or producing ornamentals in hatcheries are expected to constrain the viability and market share of dedicated PCC enterprises. PCC is expected to have the greatest uptake by part-time artisanal fishers in developing countries with infrastructure for exporting marine ornamentals. Such fishers are more immune to temporal variation in the supply of post-larvae—they can engage in PCC when valuable post-larvae are abundant and switch to other sources of income when they are scarce. Livelihood opportunities for smallholders could be enhanced through promotion of the environmental benefits of PCC among hobbyists maintaining marine ornamentals.

Shifting dominance among Scarid species on reefs representing a gradient of fishing pressure

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Eric Clua, Pierre Legendre
Clua, E. and Legendre, P. (2008) Shifting dominance among Scarid species on reefs representing a gradient of fishing pressure. Aquatic Living Resources, Vol. 21 (3): 339-348.

The present paper analyzes data collected between 2001 and 2002 on 81 reef fish species targeted by fishers at 5 sites in the Kingdom of Tonga (South Pacific). We first ranked the sites with respect to fishing pressure using two independent methods: (i) Tongan demography and reef surfaces available for fishing, and (ii) the differential effects of fishing on the whole set of 81 species grouped by their life history traits (LHT). We then focused on Parrotfish (Scaridae), which are heavily targeted in coral reef fisheries. We used the identified gradient of fishing pressure to study the effect of fishing on the community structure and test the hypothesis of “shifting dominance” amongst the 20 Scarid species present in the surveys. In addition to the classical effect of decreasing fish size in a family strongly targeted by fishers, the shifting dominance phenomenon includes a decrease in the abundance of the large-bodied and highly targeted species, favouring their replacement by smaller-bodied species from the same family, which are less impacted by fishing. In a context of interspecific competition amongst Scarids, the stress of fishing appears as a factor favouring the replacement of species with large maximum size, and LHT promoting low resilience, by smaller species with the opposite attributes. The discussion focuses on the various processes that can explain the shifting dominance phenomenon. The total density of resilient species, which increased along the gradient of increasing fishing pressure, can be used as an indicator of the over-exploitation of fish communities for reef fisheries management.

Medium scale approach (MSA) for improved assessment of coral reef fish habitat

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Eric Clua, Pierre Legendre, Laurent Vigliola, Franck Magron, Michel Kulbicki, Sébastien Sarramegna, Pierre Labrosse, René Galzin
Clua, E., Legendre, P., Vigliola, L., Magron, F., Kulbicki, M., Sarramegna, S., Labrosse, P. and Galzin, R. (2006) Medium scale approach (MSA) for improved assessment of coral reef fish habitat. Journal of Experimental Marine Biology and Ecology 333: 219-230.

Habitat characteristics play a critical role in structuring reef fish communities subjected to fishing pressure. The line intercept transect (LIT) method provides an accurate quantitative description of the habitat, but in a very narrow corridor less than 1 m wide. Such a scale is poorly adapted to the wide-ranging species that account for a significant part of these assemblages. We developed an easy-to-use medium scale approach (MSA), based on a semi-quantitative description of 20 quadrats of 25 m2 (500 m2 in total). We then simulated virtual reef landscapes of different complexities in a computer, on which we computed MSA using different methods of calculation. These simulations allowed us to select the best method of calculation, obtaining quantitative estimates with acceptable accuracy (comparison with the original simulated landscapes: R2 ranging from 0.986 to 0.997); they also showed that MSA is a more efficient estimator than LIT, generating percentage coverage estimates that are less variable. A mensurative experiment based on thirty 50-m transects, conducted by three teams of two divers, was used to empirically compare the two estimators and assess their ability to predict fish–habitat relationships. Three-factor multivariate ANOVAs (Teams, Reef, Methods) revealed again that LIT produced habitat composition estimates that were more variable than MSA. Canonical analyses conducted on fish biomass data successively aggregated by mobility patterns, trophic groups, and size classes, showed the higher predictive power of MSA habitat data over LIT. The MSA enriches the toolbox of methods available for reef habitat description at intermediate scale (< 1000 m2), between the scale where LIT is appropriate (< 100 m2) and the landscape approach (> 1000 m2).

Towards multidisciplinary indicator dashboards for coral reef fisheries management

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Eric Clua, Benoit Beliaeff, Claude Chauvet, Gilbert David, Jocelyne Ferraris, Mekhi Kronen, Michel Kulbicki, Pierre Labrosse, Yves Letourneur, Dominique Pelletier, Olivier Thébaud, Marc Léopold
Clua, E., Beliaeff, B., Chauvet, C., David, G., Ferraris, J., Kronen, M., Kulbicki, M., Labrosse, P., Letourneur, Y., Pelletier, D., Thébaud, O. and Léopold, M. (2005) Towards multidisciplinary indicator dashboards for coral reef fisheries management. Aquatic Living Resources 18: 199-213.

The diversity of reef ecosystems, the multiplicity of reef resource uses and the breadth of the range of the island socio-cultural contexts concerned make coral reef fisheries (CRF) management in the South Pacific a complex task. The health and state of the targeted resources depend both on ecosystem characteristics (as determined by ecological and biological factors) and on fishing pressure, whose effects are only partly known. Increasing harvests from commercial and recreational fishing increasingly overlap with traditional Subsistence activity, creating an important CRF management challenge. This paper presents a new approach to CRF assessment and monitoring by providing a set of multidisciplinary indicators. The fisheries system is assessed from three different viewpoints: ecology of targeted populations, exploitation and the broader socio-econornic fishery context. The use of complementary indicators chosen from each of these fields could balance the chronic lack of human and financial resources for the management of these fisheries. We suggest the use of these indicators through an assessment grid or an indicator dashboard specifically adapted to given situations and management objectives determined through a participatory approach. The operational efficiency of this dashboard depends on i) dialogue between users, ii) the objectivity of the proposed monitoring, iii) the Visual transcription of divergent/convergent interests amongst stakeholders, and iv) stakeholder involvement in the decision-making process. The use and constraints of such a tool are described with reference to Ouvea atoll (New-Caledonia, South Pacific) for which an analysis of available indicators for assessing fisheries status is presented.

Checklist of the shore and epipelagic fishes of Tonga

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Eric Clua, John E. Randall, David G. Smith, Jeffrey T. Williams, Michel Kulbicki, Gerald Mou Tham, Pierre Labrosse, Mecki Kronen, Bradley S. Mann
Clua, E., Randall, J. E., Smith, D. G., Williams, J. T., Kulbicki, M., Mou Tham, G., Labrosse, P., Kronen, M. and Mann, B. S. (2002) Checklist of the shore and epipelagic fishes of Tonga, Atoll Research Bulletin.

A checklist is given below of 1162 species of shore and epipelagic fishes belonging to 111 families that occur in the islands of Tonga, South Pacifie Ocean; 40 of these are epipelagic species. As might be expected, the fish fauna of Tonga is most similar to those of Samoa and Fiji; at least 658 species of the fishes found in Tonga are also known from Fiji and the islands of Samoa. Twelve species of shore fishes are presently known only from Tonga. Specimens of Tongan fishes are housed mainly in the fish collections of the National Museum of Natural History, Washington D.C.; Bernice P. Bishop Museum, Honolulu; Museum National d’Histoire Naturelle, Paris; and the Australian Museum, Sydney. Native Tongan names of fishes, when known, are presented after species names.

Mixed-species feeding aggregation of dolphins, large tunas and seabirds in the Azores

PUBLICATIONS SCIENTIFIQUESArticle de rang A
Eric Clua, François Grosvalet
Clua, E. and Grosvalet, F. (2001) Mixed-species feeding aggregation of dolphins, large tunas and seabirds in the Azores, Aquatic Living Resources, Vol. 14, N°1, pp: 11-18.

Each summer the presence of large concentrations of bait fish in the area of the central Azores Islands gives rise to mixed-species feeding aggregations usually at dawn and dusk. The encircling of prey initiated by common dolphins (Delphinus delphis), often mixed with spotted dolphins (Stenella frontalis), results in the formation of a compact ‘ball’ of several thousands prey fish close to the surface. Other dolphins, in particular the bottlenose (Tursiops truncatus), also eat the prey fish, whose high concentration makes them easy to capture. Large tunas (Thunnus thynnus, Thunnus albacares) sometimes participate in the phenomenon. Seabirds (mainly cory’s sheawaters, Calonectris diomedea borealis) are always present throughout the few minutes during which the entire collective food hunt takes place. A model of the phenomenon, based on 15 observations, is proposed. It comprises 4 stages: a preparation phase, an intensification phase, a mature phase, and a dispersion phase. These observations allow a better understanding of the tuna-dolphin aggregation process. They show that it is the tunas that generate and beneficiate from the aggregation with dolphins, rather than the contrary.