Résumé: The Mediterranean Sea (MS) is a semi-enclosed sea characterized by a zonal west-east gradient of oligotrophy, where microbial growth is controlled by phosphate availability in most situations. External inputs of nutrients including Gibraltar inputs, river inputs and atmospheric deposition are therefore of major importance for the biogeochemistry of the MS. The latter has long been considered to be driven mainly by nutrient exchanges at Gibraltar. However, recent studies indicate that river inputs significantly affect nutrients concentrations in the Mediterranean Sea, although their resulting impact on its biogeochemistry remains poorly understood. In this study, our aim was to help fill this knowledge gap by addressing the large-scale and long-term impact of variations in river inputs on the biogeochemistry of the Mediterranean Sea over the last decades, using a coupled physical-biogeochemical 3D model (NEMO-MED12/Eco3M-Med). As a first result, it has been shown by the model that the strong diminution (60%) of phosphate (PO4) in river inputs into the Mediterranean Sea since the end of the 1980s induced a significant lowering of PO4 availability in the sub-surface layer of the Eastern Mediterranean Basin (EMB). One of the main consequences of PO4 diminution is the rise, never previously documented, of dissolved organic carbon (DOC) concentrations in the surface layer (by 20% on average over the EMB). Another main result concerns the gradual deepening of the top of the phosphacline during the period studied, thus generating a shift between the top of the nitracline and the top of the phosphacline in the EMB. This shift has already been observed in situ and documented in literature, but we propose here a new explanation for its occurrence in the EMB. The last main result is the evidence of the decline in abundance and the reduction of size of copepods calculated by the model over the years 1985-2010, that could partially explain the reduction in size of anchovy and sardine recently recorded in the MS. In this study, it is shown for the first time that the variations in river inputs that occurred in the last decades may have significantly altered the biogeochemical cycles of two key elements (P and C), in particular in the EMB. To conclude, the magnitude of the biogeochemical changes induced by river inputs and runoff alone over the last thirty years clearly calls for the use of realistic scenarios of river inputs along with climate scenarios for coupled physical-biogeochemical forecasts in the MS.

Résumé: Micronekton distributions and assemblages were investigated at two shallow seamounts of the south-western Indian Ocean using a combination of trawl data and a multi-frequency acoustic visualisation technique. La Pa rouse seamount (summit depth similar to 60 m) is located on the outskirts of the oligotrophic Indian South Subtropical Gyre (ISSG) province with weak mesoscale activities and low primary productivity all year round. The “MAD-Ridge” seamount (thus termed in this study; similar to 240 m) is located in the productive East African Coastal (EAFR) province with high mesoscale activities to the south of Madagascar. Higher micronekton species richness was recorded at MAD-Ridge compared to La Perouse. Resulting productivity at MAD-Ridge seamount was likely due to the action of mesoscale eddies advecting productivity and larvae from the Madagascar shelf rather than local dynamic processes such as Taylor column formation. Mean micronekton abundance/biomass, as estimated from mesopelagic trawl catches, were lower over the summit compared to the vicinity of the seamounts, due to net selectivity and catchability and depth gradient on micronekton assemblages. Mean acoustic densities in the night shallow scattering layer (SSL: 10-200 m) over the summit were not significantly different compared to the vicinity (within 14 nautical miles) of MAD-Ridge. At La Perouse and MAD-Ridge, the night and day SSL were dominated by common diel vertically migrant and non-migrant micronekton species respectively. While seamount-associated mesopelagic fishes such as Diaphus suborbitalis (La Perouse and MAD-Ridge) and Benthosema fibula= performed diel vertical migrations (DVM) along the seamounts' flanks, seamount-resident benthopelagic fishes, including Cookeolus japonicus (MAD-Ridge), were aggregated over MAD-Ridge summit. Before sunrise, mid-water migrants initiated their vertical migration from the intermediate to the deep scattering layer (DSL, La Perouse: 500-650 m; MAD-Ridge: 400-700 m) or deeper. During sunrise, the other taxa contributing to the night SSL exhibited a series of vertical migration events from the surface to the DSL or deeper until all migrants have reached the DSL before daytime. Possible mechanisms leading to the observed patterns in micronekton vertical and horizontal distributions are discussed. This study contributes to a better understanding of how seamounts influence the DVM, horizontal distribution and community composition of micronekton and seamount-associated/resident species at two poorly studied shallow topographic features in the south-western Indian Ocean.

Résumé: The present study describes for the first time the spatial distribution of five macrourid species throughout the Mediterranean Sea and analyses depth, geographical and time-related trends regarding their abundance, biomass and mean fish weight. The data were collected as part of the MEDITS annual bottom trawl survey carried out by several European Mediterranean countries from 1994 to 2015, using the same standardized gear and sampling protocol. The most represented species in terms of abundance and biomass was Coelorinchus caelorhincus. The bathymetric trend was different for each species. The shallowest occurring species was C. caelorhincus, followed by Hymenocephalus italicus and Nezumia sclerorhynchus, while Nezumia aequalis and Trachyrincus scabrus were the deepest. Overall, the mean weight of all the species increased with depth. C. caelorhincus and H italicus occurred in the entire study area: the first species showed relatively high catches in most areas, while the second was more abundant in the central and easternmost areas. N. aequalis and T. scabrus were mainly reported in the western basin, and N. sclerorhynchus in the central-eastern areas of the Mediterranean. An increasing inter-annual trend in abundance was only detected for C. caelorhincus and N. sclerorhynchus, while variable fluctuations were observed in the other species.

Résumé: In the Gulf of Guinea, bigeye tuna (Thunnus obesus; BET) and yellowfin tuna (Thunnus albacares; YFT) are an important part of commercial fisheries and play a prominent ecological role as top predators. Using fatty acid profiles and carbon and nitrogen stable isotopes, we examined their trophic niche partitioning in this understudied region. Trophic niche overlap was high (> 70%), similar to percentages in other ocean basins. BET occupied a higher trophic position than YFT and fed on deeper prey (high delta N-15 values and high proportions of monounsaturated fatty acids). The trophic position of YFT decreased slightly in the last 15 years (delta N-15 values decreased by similar to 0.5 parts per thousand), suggesting a change in epipelagic communities, as observed in the eastern Pacific Ocean. Ontogenic changes were limited to BET. For both species, the dietary proportion of the diatom marker 20:5(n-3) increased in the seasonal upwelling area, highlighting the influence of seasonal habitat on the diet of tuna. The relatively lipid-rich muscle (similar to 6% dry mass) of Atlantic tropical tuna suggests a richer diet in this region than that of Indian Ocean tropical tuna and (or) differences in energy allocation strategies.

Résumé: Spatio-temporally stable prey distributions coupled with individual foraging site fidelity are predicted to favour individual resource specialisation. Conversely, predators coping with dynamic prey distributions should diversify their individual diet and/or shift foraging areas to increase net intake. We studied individual specialisation in Scopoli's shearwaters (Calonectris diomedea) from the highly dynamic Western Mediterranean, using daily prey distributions together with resource selection, site fidelity and trophic-level analyses. As hypothesised, we found dietary diversification, low foraging site fidelity and almost no individual specialisation in resource selection. Crucially, shearwaters switched daily foraging tactics, selecting areas with contrasting prey of varying trophic levels. Overall, information use and plastic resource selection of individuals with reduced short-term foraging site fidelity allow predators to overcome prey field lability. Our study is an essential step towards a better understanding of individual responses to enhanced environmental stochasticity driven by global changes, and of pathways favouring population persistence.

Résumé: Understanding the environmental drivers of interactions between predators and humans is critical for public safety and management purposes. In the marine environment, this issue is exemplified by shark-human interactions. The annual shark bite incidence rate (SBIR) in La Reunion (Indian Ocean) is among the highest in the world (up to 1 event per 24,000 hours of surfing) and has experienced a 23-fold increase over the 2005-2016 period. Since 1988, 86% of shark bite events on ocean-users involved surfers off the leeward coast, where 96% of surfing activities took place. We modeled the SBIR as a function of environmental variables, including benthic substrate, sea temperature and period of day. The SBIR peaked in winter, during the afternoon and dramatically increased on coral substrate since the mid-2000s. Seasonal patterns of increasing SBIR followed similar fluctuations of large coastal shark occurrences (particularly the bull shark Carcharhinus leucas), consistent with the hypothesis that higher shark presence may result in an increasing likelihood of shark bite events. Potential contributing factors and adaptation of ocean-users to the increasing shark bite hazard are discussed. This interdisciplinary research contributes to a better understanding of shark-human interactions. The modeling method is relevant for wildlife hazard management in general.

Résumé: Understanding the relationship between environmental variables and pelagic species concentrations and dynamics is helpful to improve fishery management, especially in a changing environment. Drifting fish aggregating device (DFAD)-associated tuna and non-tuna biomass data from the fishers' echo-sounder buoys operating in the Atlantic Ocean have been modelled as functions of oceanographic (Sea Surface Temperature, Chlorophyll-a, Salinity, Sea Level Anomaly, Thermocline depth and gradient, Geostrophic current, Total Current, Depth) and DFAD variables (DFAD speed, bearing and soak time) using Generalized Additive Mixed Models (GAMMs). Biological interaction (presence of non-tuna species at DFADs) was also included in the tuna model, and found to be significant at this time scale. All variables were included in the analyses but only some of them were highly significant, and variable significance differed among fish groups. In general, most of the fish biomass distribution was explained by the ocean productivity and DFAD-variables. Indeed, this study revealed different environmental preferences for tunas and non-tuna species and suggested the existence of active habitat selection. This improved assessment of environmental and DFAD effects on tuna and non-tuna catchability in the purse seine tuna fishery will contribute to transfer of better scientific advice to regional tuna commissions for the management and conservation of exploited resources.

Résumé: This study examined the fatty acid composition of three sympatric tropical tuna species (bigeye Thunnus obesus, yellowfin T. albacares and skipjack tuna Kastuwonus pelamis) sampled in the Western Indian Ocean in 2013. The fatty acid compositions of neutral and polar lipids, respectively involved in energy storage and cell membrane structure, were explored and compared in four tissues (red and white muscles, liver and gonads), according to biological (size, sex and maturity) and environmental (season and area) factors. The liver and the red muscle were the fattest tissues (i.e., higher levels of storage lipids) in all species and polar lipids were the lowest in the white muscle. Species and tissue types explained most differences in fatty acid compositions, while environmental factors had limited effects, except in the hepatic cell membrane where fatty acid composition varied with monsoons. Docosahexaenoic acid (22:6n-3) was the major fatty acid in both polar and neutral lipid fractions, especially in muscles. Eicosapentaenoic acid (20:5n-3) and oleic acid (18:1n-9) were in higher proportion in neutral than in polar lipids. Arachidonic acid (20:4n-6) and 22:6n-3, together with docosapentaenoic acid (22:5n-6) and stearic acid (18:0), showed preferential accumulation in polar lipids. 20:4n-6 was particularly involved in cell membranes of ovary and white muscle. Overall, an important inter-individual variability in fatty acid compositions of structural lipids was found within tissue types despite considering biological factors that are most likely to influence this type of lipids. It suggests that fatty acid profiles are influenced by individual-specific behaviors.

Résumé: Stomach contents and stable carbon and nitrogen isotope analyses (delta C-13 and delta N-15) were used to investigate the trophic ecology of two apex predators, tiger sharks (Galeocerdo cuvier) and bull sharks (Carcharhinus leucas), from Reunion Island to describe their dietary habits at both the population and individual levels. In this oceanic island, the tiger and bull sharks were more piscivorous and teutophagous than noted in previous research from other localities. The delta C-13 values suggested that bull sharks depended on more neritic organic matter sources than tiger sharks, confirming a coastal habitat preference for bull sharks. Moreover, the total length of the bull shark influenced delta C-13 values, with smaller individuals being more coastal than larger individuals. All indicators suggest that there is a higher degree of similarity between individual tiger sharks compared with the more heterogeneous bull shark population, which is composed of individuals who specialize on different prey. These results suggest that the two species have different functions in these coastal habitats, and thus, they must be considered independently in terms of conservation and management.

Résumé: Two stocks of bluefin tuna (Thunnus thynnus) inhabit the north Atlantic; the western and eastern stocks spawn in the Gulf of Mexico and the Mediterranean Sea respectively. Trans-Atlantic movements occur outside spawning time whereas natal homing maintains stock structure. Commercial fisheries may exploit a mixed assemblage of both stocks. The incorporation of mixing rates into stock assessment is precluded by uncertainties surrounding stock discrimination. Otolith shape descriptors were used to characterise western and eastern stocks of Atlantic bluefin tuna in the present study and to estimate stock composition in catches of unknown origin. Otolith shape varied with length and between locations and years. Within a restricted size range (200-297-cm fork length (FL)) the two stocks were distinguished with an accuracy of 83%. Bayesian stock mixture analysis indicated that samples from the east Atlantic and Mediterranean were predominantly of eastern origin. The proportion assigned to the eastern stock showed slight spatial variation; however, overlapping 95% credible intervals indicated no significant difference (200-297 cm FL: central Atlantic, 73-100%; Straits of Gibraltar, 73-100%; Morocco, 50-99%; Portugal 64-100%). Otolith shape could be used in combination with other population markers to improve the accuracy of mixing rate estimates for Atlantic bluefin tuna.

Résumé: The lipid composition of somatic and reproductive tissues was determined for female skipjack tuna Katsuwonus pelamis caught in the western Indian Ocean between latitude 10 degrees N and 20 degrees S and longitude 40 degrees and 70 degrees E. The highest total lipid (TL) contents were in the liver and gonads, with white muscle levels approximately three-fold lower. Three lipid classes dominated: triacylglycerols (TAG), sterol esters and wax esters (SE-WE) and phospholipids (PL). Collectively, these accounted for between 70 and 80% of TLs. Changes in lipid concentrations were evaluated over the maturation cycle. Immature fish had the lowest gonad and liver TL levels; concentrations of TL, TAG, SE-WE and PL accumulated from immature to mature (spawning-capable) phase, reflecting sustained vitellogenic activity of the liver and a transfer of lipids to developing oocytes from the onset of vitellogenesis. Gonado-somatic and hepato-somatic indices were positively correlated with each other and positively related to TL in the gonads and liver. Fulton's condition index and lipid concentrations in muscle did not vary significantly over the maturation cycle; fat content in the main storage tissues was undepleted as the ovary developed. Hence, K. pelamis apparently supports reproduction directly from food intake over the breeding season. In the gonads, reserve lipids (SE-WE and TAG) and sterols were related to batch fecundity but this was not the case for somatic and hepatic tissues. These results suggest that K. pelamis utilizes an income breeding strategy.

Résumé: Tunas are highly specialized predators that have evolved numerous adaptations for a lifestyle that requires large amounts of energy consumption. Here we review our understanding of the bioenergetics and feeding dynamics of tunas on a global scale, with an emphasis on yellowfin, bigeye, skipjack, albacore, and Atlantic bluefin tunas. Food consumption balances bioenergetics expenditures for respiration, growth (including gonad production), specific dynamic action, egestion, and excretion. Tunas feed across the micronekton and some large zooplankton. Some tunas appear to time their life history to take advantage of ephemeral aggregations of crustacean, fish, and molluscan prey. Ontogenetic and spatial diet differences are substantial, and significant interdecadal changes in prey composition have been observed. Diet shifts from larger to smaller prey taxa highlight ecosystem-wide changes in prey availability and diversity and provide implications for changing bioenergetics requirements into the future. Where tunas overlap, we show evidence of niche separation between them; resources are divided largely by differences in diet percentages and size ranges of prey taxa. The lack of long-term data limits the ability to predict impacts of climate change on tuna feeding behaviour. We note the need for systematic collection of feeding data as part of routine monitoring of these species, and we highlight the advantages of using biochemical techniques for broad-scale analyses of trophic relations. We support the continued development of ecosystem models, which all too often lack the regional-specific trophic data needed to adequately investigate climate and fishing impacts.

Résumé: The “condition” is used as an indicator of fish health and is generally equated with the quantity of energy reserves. Biometric condition factors have been widely used and preferred over costly and time-consuming biochemical condition. Here, we investigated the relevance of four common condition factors based on biometric measurements (Le Cren's index, girth -length index, gonado-somatic index and hepato-somatic index) and of size- and weight -based empirical models to describe the physiological condition of tropical tunas. Biometric condition factors of bigeye (Thunnus obesus), skipjack (Katsuwonus pelamis) and yellowfin (Thunnus albacares) tunas sampled throughout 2013 in the western Indian Ocean region were assessed against benchmark biochemical indices (lipid content, protein content, triacylglycerol:sterol ratio and energy density) estimated in tissues with different physiological functions, i.e. red muscle, white muscle, liver, and gonads. Our findings suggest that tropical tunas do not store lipids in white muscle and that protein content is less variable than lipid content, which largely varies with ontogeny and the seasons according to tissue and species. This variability induced inconsistency between biometric factors, including the empirically adjusted ones, and biochemical indices, with the exception of the gonado-somatic index that fitted well to the composition of the gonads in the three species, and especially in females. (C) 2016 Elsevier Ltd. All rights reserved.

Résumé: Increasing abundance of non-commercial sprats and decreasing biomass and landings of commercial anchovies and sardines justify the need to study the feeding ecology and trophic niche overlap of these planktivorous species in the Gulf of Lions. Their diet has been investigated on the basis of stomach content and stable isotope analyses in 2011 and 2012 according to different depths and regions in the study area. The main prey were Corycaeidae copepods, Clauso/Paracalanus, Euterpina acutifrons and Microsetella, for sprats and small copepods, such as Microsetella, Oncaea and Corycaeidae, for anchovies and sardines. This is the first time that the diet of sprats is described in the Gulf of Lions. Sprats fed on a larger size spectrum of prey and seem to be more generalist feeders compared to anchovies and sardines. Ontogenetic changes as well as spatial and temporal variations of the diet occurred in the three species. Stable isotope analysis revealed mobility of sardines and sprats among feeding areas while anchovies exhibited preferred feeding areas. Sprats showed a higher relative condition assessed by C/N ratios than sardines and anchovies. Our results showed an overlap of the trophic niches for the three species, indicating a potential trophic competition in the Gulf of Lions.

Résumé: The Mozambique Channel (western Indian Ocean) is a dynamic environment characterised by strong mesoscale features, which influence all biological components of the pelagic ecosystem. We investigated the distribution, abundance and feeding behaviour of seabirds in the Mozambique Channel in relation to physical and biological environmental variables, with a specific interest in mesoscale features. Seabird censuses were conducted in summer and winter during 7 cruises in the southern and northern Mozambique Channel. Tropical species accounted for 49% of the 37 species identified and 97% of the individuals, and species from the sub-Antarctic region constituted 30% of the identifications. The typically tropical sooty tern (Onychoprion fuscata) was the dominant species during all cruises, and overall accounted for 74% of the species observations and 85% of counted birds. Outputs of Generalised Linear Models at the scale of the Mozambique Channel suggested that higher densities of flying and feeding birds occurred in areas with lower sea surface temperatures and lower surface chlorophyll a concentrations. Most of the flocks of feeding birds did not associate with surface schools of fish or marine mammals, but when they did, these flocks were larger, especially when associated with tuna. While tropical species seemed to favour cyclonic eddies, frontal and divergence zones, non-tropical species were more frequently recorded over shelf waters. Sooty terns foraged preferentially in cyclonic eddies where zooplankton, micronelcton and tuna schools were abundant. Among other major tropical species, frigatebirds (Fregata spp.) predominated in frontal zones between eddies, where tuna schools also frequently occurred and where geostrophic currents were the strongest. Red-footed boobies (Sula sub) concentrated in divergence zones characterised by low sea level anomalies, low geostrophic currents, and high zooplanlcton biomass close to the surface. Our results highlight the importance of mescoscale features in structuring the tropical seabird community in the Mozambique Channel, in addition to segregating tropical and non-tropical species. The mechanisms underlying the segregation of tropical seabirds seem to partially differ from that of other tropical regions, and this may be a consequence of the strong local mesoscale activity, affecting prey size and availability schemes. Beyond characterising the foraging habitats of the seabird community of the Mozambique Channel, this study highlights the importance of this region as a hot spot for seabirds; especially the southern part, where several endangered sub-Antarctic species over-winter.

Résumé: We analyzed stomach contents and muscle isotopic composition (delta C-13, delta N-15) of yellowstripe goatfish Mulloidichthys flavolineatus from a coral reef to investigate size-related dietary changes and resource or habitat partitioning. Juveniles (< 12 cm total length [TL]), young adults (12 <= TL < 17 cm) and adults (>= 17 cm TL) showed a high diet overlap, especially between juveniles and young adults. According to stomach contents analysis, M. flavolineatus widens its prey spectrum with increasing size from a common prey pool that includes polychaetes, tanaids and harpacticoid copepods. We observed a significant increase in delta C-13 values (from -17.1 +/- 0.5 parts per thousand for juveniles to -10.7 +/- 0.8 parts per thousand for adults), which were correlated to fish size. Adults (delta N-15 mean of 11.1 +/- 1.8 parts per thousand) were one trophic level above juveniles and young adults (7.7 +/- 0.5 and 7.4 +/- 0.5 parts per thousand, respectively). These patterns of isotopic changes confirmed ontogenetic dietary shifts. However, trophodynamics can be influenced by physiological factors such as growth and sexual maturity. M flavolineatus shift from a pelagic to a macrobenthic diet, which is equilibrated at the adult stage. Results from combined stomach contents (prey volume) and stable isotope analyses suggested a dominant contribution of polychaetes (macrofauna), possibly through selective feeding. Conversely, on the basis of prey volume and stable isotope data, meiofauna did not feature significantly in the diet despite their high abundance in stomachs. Two adult groups were distinguished based on their delta N-15 values (11.9 +/- 0.8 and 7.8 +/- 0.6 parts per thousand), indicating possible stage-specific partitioning in habitat use inside the reef.