Johann Mourier

Temperature and oxygen limit the distribution of marine ectotherms. Haematological traits underlying blood-oxygen carrying capacity are thought to be correlated with thermal tolerance in certain fishes, and this relationship is hypothesised to be explained by oxygen supply capacity. We tested this hypothesis using reef shark neonates as experimental models because they live near their upper thermal limits and are physiologically sensitive to low oxygen conditions. We first described in situ associations between temperature and oxygen at the study site (Moorea, French Polynesia) and found that the habitats for reef shark neonates (Carcharhinus melanopterus and Negaprion acutidens) were hyperoxic at the maximum recorded temperatures. Next, we tested for in situ associations between thermal habitat characteristics and haematological traits of neonates. Contrary to predictions, we only demonstrated a negative association between haemoglobin concentration and maximum habitat temperatures...

In recent decades, network analyses have become ubiquitous in ecology, facilitating our understanding of linkages between paired entities, whether it be genes, proteins, individuals, species, or habitats (Bluthgen et al., 2008; Croft et al., 2008; Krause et al., 2007; Proulx et al., 2005; Wey et al., 2008). Network theory (also known as graph theory) originates from the mathematical and social sciences but has developed concurrently across many disciplines, including computational science, physics, management, genetics, and epidemiology (Newman, 2010), to name but a few.

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 non- intrusive 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 loca - tion 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. resUMe. - Photo-identification des requins limon faucille, Negaprion acutidens, a Moorea...

Globally, the frequency of shark bites is rising, resulting in an increasing demand for shark deterrents and measures to lessen the impact of shark bites on humans. Most existing shark protection measures are designed to reduce the probability of a bite, but fabrics that minimise injuries when a shark bite occurs can also be used as mitigation devices. Here, we assessed the ability of the Ocean Guardian Scuba7 and Kevlar material to reduce the likelihood of blacktip reef sharks, Carcharhinus melanopterus, from feeding, and to minimise injuries from shark bites. Sharks were enticed to consume a small piece of local reef fish (bait) placed between the two Scuba7 electrodes with the deterrents randomly being turned on or kept off. In the second experiment, the bait was attached to a small pouch made of either standard neoprene or neoprene with a protective layer of Kevlar around it. The Scuba7 reduced the proportion of baits being taken by 67%, (from 100% during control trials to 33%)....

An animal's energy landscape considers the power requirements associated with residing in or moving through habitats. Within marine environments, these landscapes can be dynamic as water currents will influence animal power requirements and can change rapidly over diel and tidal cycles. In channels and along slopes with strong currents, updraft zones may reduce energy expenditure of negatively buoyant fishes that are also obligate swimmers. Despite marine predators often residing within high-current area, no study has investigated the potential role of the energetic landscape in driving such habitat selectivity. Over 500 grey reef sharks Carcharhinus amblyrhynchos reside in the southern channel of Fakarava Atoll, French Polynesia. We used diver observations, acoustic telemetry and biologging to show that sharks use regions of predicted updrafts and switch their core area of space use based on tidal state (incoming versus outgoing). During incoming tides, sharks form tight groups and display shuttling behaviour (moving to the front of the group and letting the current move them to the back) to maintain themselves in these potential updraft zones. During outgoing tides, group dispersion increases, swimming depths decrease and shuttling behaviours cease. These changes are likely due to shifts in the nature and location of the updraft zones, as well as turbulence during outgoing tides. Using a biomechanical model, we estimate that routine metabolic rates for sharks may be reduced by 10%-15% when in updraft zones. Grey reef sharks save energy using predicted updraft zones in channels and 'surfing the slope'. Analogous to birds using wind-driven updraft zones, negatively buoyant marine animals may use current-induced updraft zones to reduce energy expenditure. Updrafts should be incorporated into dynamic energy landscapes and may partially explain the distribution, behaviour and potentially abundance of marine predators.

Over the past decade, drones have become a popular tool for wildlife management and research. Drones have shown significant value for animals that were often difficult or dangerous to study using traditional survey methods. In the past five years drone technology has become commonplace for shark research with their use above, and more recently, below the water helping to minimise knowledge gaps about these cryptic species. Drones have enhanced our understanding of shark behaviour and are critically important tools, not only due to the importance and conservation of the animals in the ecosystem, but to also help minimise dangerous encounters with humans. To provide some guidance for their future use in relation to sharks, this review provides an overview of how drones are currently used with critical context for shark monitoring. We show how drones have been used to fill knowledge gaps around fundamental shark behaviours or movements, social interactions, and predation across multipl...

ABSTRACTGenetic relatedness in animal societies is often a factor that drives the structure of social groups. In the marine world, most studies which have investigated this question have focused on marine mammals such as whales and dolphins. For sharks, recent studies have demonstrated preferential associations among individuals from which social communities emerge. Assortment patterns have been found according to phenotypic or behavioural traits but the role of genetic relatedness in shaping the social structure of adult shark populations has, to the best our knowledge, never been investigated. Here, we used a social network analysis crossed with DNA microsatellite genotyping to investigate the role of the genetic relatedness in the social structure of a blacktip reef shark (Carcharhinus melanopterus) population. Based on data from 156 groups of sharks, we used generalized affiliation indices to isolate social preferences from non-social associations, controlling for the contributi...

Animal movement patterns are increasingly analysed as spatial networks. Currently, structures of complex movements are typically represented as a single-layer (or monoplex) network. However, aggregating individual movements, to generate population-level inferences, considerably reduces information on how individual or species variability influences spatial connectivity and thus identifying the mechanisms driving network structure remains difficult. Here, we propose incorporating the recent conceptual advances in multilayer network analyses with the existing movement network approach to improve our understanding of the complex interaction between spatial and/or social drivers of animal movement patterns. Specifically, we explore the application and interpretation of this framework using an empirical example of shark movement data gathered using passive remote sensors in a coral reef ecosystem. We first show how aggregating individual movement networks can lead to the loss of informat...

Distinguishing the factors that influence activity within a species advances understanding of their behavior and ecology. Continuous observation in the marine environment is not feasible but biotelemetry devices provide an opportunity for detailed analysis of movements and activity patterns. This study investigated the detail that calibration of accelerometers measuring root mean square (RMS) acceleration with video footage can add to understanding the activity patterns of male and female Port Jackson sharks (Heterodontus portusjacksoni) in a captive environment. Linear regression was used to relate RMS acceleration output to time‐matched behavior captured on video to quantify diel activity patterns. To validate captive data, diel patterns from captive sharks were compared with diel movement data from free‐ranging sharks using passive acoustic tracking. The RMS acceleration data showed captive sharks exhibited nocturnal diel patterns peaking during the late evening before midnight and decreasing before sunrise. Correlation analysis revealed that captive animals displayed similar activity patterns to free‐ranging sharks. The timing of wild shark departures for migration in the late breeding season corresponded with elevated diel activity at night within the captive individuals, suggesting a form of migratory restlessness in captivity. By directly relating RMS acceleration output to activity level, we show that sex, time of day, and sex‐specific seasonal behavior all influenced activity levels. This study contributes to a growing body of evidence that RMS acceleration data are a promising method to determine activity patterns of cryptic marine animals and can provide more detailed information when validated in captivity.

Ontogenetic niche shifts are widespread. However, individual differences in size at birth, morphology, sex, and personalities can cause variability in behavior. As such, inherent inter-individual differences within populations may lead to context-dependent changes in behavior with animal body size, which is of concern for understanding population dynamics and optimizing ecological monitoring. Using stable carbon and nitrogen isotope values from concurrently sampled tissues, we quantified the direction and magnitude of intraspecific variation in trophic shifts among three shark species, and how these changed with body size: spurdogs (Squalus spp.) in deep-sea habitats off La Réunion, bull sharks (Carcharhinus leucas) in estuarine habitats of the Florida Everglades, and blacktip reef sharks (Carcharhinus melanopterus) in coral reef ecosystems of Moorea, French Polynesia. Intraspecific variation in trophic shifts was limited among spurdogs, and decreased with body size, while bull sharks exhibited greater individual differences in trophic shifts, but also decreased in variability through ontogeny. In contrast, blacktip reef sharks exhibited increased intraspecific variation in trophic interactions with body size. Variability in trophic interactions and ontogenetic shifts are known to be associated with changes in energetic requirements, but can vary with ecological context. Our results suggest that environmental stability may affect variability within populations , and ecosystems with greater spatial and/or temporal variability in environmental conditions, and those with more diverse food webs may facilitate greater individual differences in trophic interactions, and thus ontogenetic trophic shifts. In light of concerns over environmental disturbance, elucidating the contexts that promote or dampen phenotypic variability is invaluable for predicting population-and community-level responses to environmental changes.

A novel image analysis-based technique applied to unmanned aerial vehicle (UAV) survey data is described to detect and locate individual free-ranging sharks within aggregations. The method allows rapid collection of data and quantification of fine-scale swimming and collective patterns of sharks. We demonstrate the usefulness of this technique in a small-scale case study exploring the shoaling tendencies of blacktip reef sharks Carcharhinus melanopterus in a large lagoon within Moorea, French Polynesia. Using our approach, we found that C. melanopterus displayed increased alignment with shoal companions when distributed over a sandflat where they are regularly fed for ecotourism purposes as compared with when they shoaled in a deeper adjacent channel. Our case study highlights the potential of a relatively low-cost method that combines UAV survey data and image analysis to detect differences in shoaling patterns of free-ranging sharks in shallow habitats. This approach offers an alternative to current techniques commonly used in controlled settings that require time-consuming post-processing effort. K E Y W O R D S blacktip reef shark, Carcharhinus melanopterus, image analysis, shoaling behaviour, UAV Significance Statement. The difficulty in observing and quantifying natural behaviours of wild marine organisms without disturbance limits our understanding of their behavioural responses to a changing environment. We developed an approach combining image analysis and unmanned aerial vehicle surveys to observe free-ranging large group-living epipelagic animals and rapidly quantify their movements and behaviour. We demonstrate the potential of this approach in a study exploring shoaling patterns and swimming dynamics of blacktip reef sharks in shallow lagoon microhabitats.

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.

Food web structure is shaped by interactions within and across trophic levels. As such, understanding how the presence and absence of predators, prey, and competitors affect species foraging patterns is important for predicting the consequences of changes in species abundances, distributions, and behaviors. Here, we used plasma d 13 C and d 15 N values from juvenile blacktip reef sharks (Carcharhinus melanopterus) and juvenile sicklefin lemon sharks (Negaprion acutidens) to investigate how species co-occurrence affects their trophic interactions in littoral waters of Moorea, French Polynesia. Co-occurrence led to isotopic niche partitioning among sharks within nurseries, with significant increases in d 15 N values among sicklefin lemon sharks, and significant decreases in d 15 N among blacktip reef sharks. Niche segregation likely promotes coexistence of these two predators during early years of growth and development, but data do not suggest coexistence affects life history traits, such as body size, body condition, and ontogenetic niche shifts. Plasticity in trophic niches among juvenile blacktip reef sharks and sicklefin lemon sharks also suggests these predators are able to account for changes in community structure, resource availability, and intra-guild competition, and may fill similar functional roles in the absence of the other species, which is important as environmental change and human impacts persist in coral reef ecosystems.

Individuals can play different roles in maintaining connectivity and social cohesion in animal populations and thereby influence population robustness to perturbations. We performed a social network analysis in a reef shark population to assess the vulnerability of the global network to node removal under different scenarios. We found that the network was generally robust to the removal of nodes with high centrality. The network appeared also highly robust to experimental fishing. Individual shark catchability decreased as a function of experience, as revealed by comparing capture frequency and site presence. Altogether, these features suggest that individuals learnt to avoid capture, which ultimately increased network robustness to experimental catch-and-release. Our results also suggest that some caution must be taken when using capture–recapture models often used to assess population size as assumptions (such as equal probabilities of capture and recapture) may be violated by individual learning to escape recapture.

Knowledge of the broad-scale movement patterns of sharks is essential to developing effective management strategies. Currently there is a large bias in studies focusing on species that are either large apex predators or found in tropical to subtropical regions. There is limited knowledge of the movements and migrations of benthic and temperate shark species. The present study used passive acoustic telemetry to investigate the movement patterns of a benthic shark species, the Port Jackson shark (Heterodontus portusjacksoni). Individuals were tagged with acoustic transmitters between 2012 and 2014 and their movements were monitored within Jervis Bay and along the east Australian coastline for up to 4 years. Male and female Port Jackson sharks demonstrated high levels of philopatry to both Jervis Bay and their tagging location across multiple years. Although males and females did not differ in their arrival times, females departed from Jervis Bay later than males. Approximately half the tagged individuals migrated in a southward direction, with individuals being detected at Narooma, Bass Strait and Cape Barron Island. This study provides conclusive evidence of bisexual philopatry in a benthic temperate shark species, confirming previous hypotheses, and presents the most detailed migration route for Port Jackson sharks to date.

The extent of the global human footprint [ 1 ] limits our understanding of what is natural in the marine environment. Remote, near-pristine areas provide some baseline expectations for biomass [ 2, 3 ] 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 [ 4 ]. 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 [ 5–9 ], and empirical evidence suggests that sharks must engage in wide-ranging foraging excursions to meet energy needs [ 9, 10 ]. 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.