Résumé: The stable carbon (delta C-13) and nitrogen (delta N-15) isotope values of soft tissues of micronekton (crustaceans, squid, mesopelagic fish) and zooplankton were measured from organisms collected on the RV Antea at two seamounts located in the south-western Indian Ocean: La Perouse (summit depth similar to 60 m) and “MAD-Ridge” (thus named in this study; summit depth similar to 240 m). Surface particulate organic matter (POM-Surf) showed higher delta C-13 at the more productive MAD-Ridge than at the oligotrophic La Perouse seamount. Particulate organic matter and zooplankton were depleted in N-15 at La Pemuse pinnacle compared with MAD-Ridge. Gelatinous organisms and crustaceans occupied the lowest and intermediate tmphic levels (TL similar to 2 and 3 respectively) at both seamounts. Mesopelagic fish and smaller-sized squid sampled at both seamounts occupied TL similar to 3 to 4, whereas the large nektonic squid, Ommastrephes bartramii, collected at MAD-Ridge only, exhibited a TL of similar to 5. The delta N-15 values of common open-water mesopelagic taxa were strongly influenced by specimen size and feeding habits at both seamounts, with an increase in delta N-15 values with increasing size. Carnivorous fish species sampled exclusively over the seamounts' flanks and summits exhibited TL values of similar to 4, irrespective of their wide size ranges. The work could not demonstrate any differences in delta C-13 values of mesopelagic fish between the seamounts and the surrounding oceanic areas. The study segregated clusters of mesopelagic organisms according to their delta C-13 and delta N-15 values, with variations in stable isotope values reflecting a complex range of processes possibly linked to productivity as well as biological and ecological traits of the species (size and feeding mode).

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: Spatial distribution, community composition and trophic roles of micronekton (crustaceans, fishes and squids) were investigated in the Indian South Subtropical Gyre (ISSG) province and the East African Coastal province (EAFR), by combining acoustic surveys, mid-water trawls and stable isotope analyses from scientific cruises conducted in 2009 and 2010. Mesopelagic micronekton performed diel vertical migrations in both provinces, from deep (400-740 m) to surface (0-200 m) layers at dusk and in the opposite direction at dawn, with some species migrating below 740 m. The EAFR province was more dynamic than the oligotrophic ISSG province, with enhanced eddy activity and enhanced yearly productivity. The active enrichment mechanisms in the EAFR, in terms of available primary production, led to high micronekton acoustic density (as a proxy of micronekton abundance) and large micronekton weight and abundance estimates from trawl data. Particulate organic matter in the EAFR exhibited greater enrichment in C-13 and N-15 compared to the ISSG and, consequently, tissues of selected micronekton organisms in the EAFR were more enriched in N-15 (higher delta N-15 values). In both provinces, micronekton encompassed a wide range of isotopic niches, with large overlaps between species. Micronekton and swordfish in the EAFR had an overlapping range of delta N-15 values, contrasting with the ISSG province where swordfish were two trophic levels higher than the sampled micronekton. Our results provide some evidence that the combined action of riverine input and the dynamics of eddies might influence productivity in the EAFR, and hence the abundance of micronekton and the enrichment of tissues in N-15, compared to the oligotrophic ISSG province.

Résumé: Coral reef environments are generally recognized as being the most threatened of marine ecosystems. However, it is extremely difficult to distinguish the effects of climate change from other forcing factors, mainly because it is difficult to study ecosystems that are isolated from human pressure. The five Iles Eparses (Scattered Islands) are located in the Western Indian Ocean (WIO) and can be considered to be “pristine” ecosystems not subject to anthropogenic pressure. This study characterized their plankton assemblages for the first time, by determining the abundances of microbial (virus, bacteria, heterotrophic protists and phytoplankton) and metazooplankton communities in various lagoon and ocean sites around each island. The Europa lagoon has extensive, productive mangrove forests, which have the highest nutrient concentrations (nitrogen forms, dissolved organic carbon) and whose microbial communities present a peculiar structure and functioning. By means of bioassay experiments, we observed that bacterial production and growth rates are higher in Europa than those reported for the other islands. Tromelin, which lies outside the Mozambique Channel, had the lowest biological productivity, nutrient concentrations, and bacterial growth rates. Multivariate analysis indicated that distinct microbial assemblages occur in association to varying nutrient concentrations. Molecular fingerprinting showed clear discrimination of the structure of the archaea, bacteria and eukaryotes community between the sites. Our results suggest that the geographical distance can influence the diversity of dominant microbial taxa in the WIO.

Résumé: Delphinid populations show highly variable patterns of genetic diversity and population structure. Previous studies indicate that habitat discontinuities and geographic isolation are major drivers of population division in cetaceans. Spinner dolphins (Stenella longirostris) are distributed in all tropical oceans, but they are particularly common around islands and atolls. This species occurs in shallow waters at daytime to rest and socialise, and feeds on offshore mesopelagic prey overnight. Here, we investigated the genetic population structure of spinner dolphins in the Southwest Indian Ocean along a west-east geographic gradient, from eastern Africa to the Mascarene archipelago. We combined analyses of 12 microsatellite loci, mtDNA control region sequences, and sighting data to assess genetic differentiation and characterise habitat preferences of these populations. Significant genetic structure among the three sampled sites (Zanzibar, Mayotte and La Reunion) was observed using both types of molecular markers. Overall, our results indicate that geographic isolation and potentially other factors, such as shallow-water habitats to rest and socialise, may be important drivers of the genetic population structure of insular spinner dolphins in this region.

Résumé: The pattern of surface circulation in the Mozambique Channel was elucidated from the trajectories of 82 satellite-tracked drifters over the period 2000-2010 and complementary satellite-derived altimetry. Overall, the trajectories indicated that anticyclonic activity was mostly observed on the western side of the Channel, with cyclonic activity being more prevalent in the east. A lack of eddy activity was noted in the southeast corner of the Channel (i.e. SW of Madagascar). Drifter behaviour illustrated that surface water from the Comoros Basin, entrained into anticyclonic eddies during formation, can be retained and isolated for months whilst being transported southwards through the Channel. During a tropical cyclone weather event, a drifter was observed to switch between counter-rotating eddies indicating that horizontal mixing of the Ekman layer does occur. The drifters also illustrated and emphasised the flow field and transport between eddies (i.e. the interstitial flow) in the Mozambique Channel. Despite the dominance of southward propagating anticyclones, drifters were able to move north and south through the Channel in the frontal flow field between eddies within periods of 51-207 days. Cross-channel transport in both directions between the Madagascan and Mozambique shelf regions was similarly observed, with time spans of 19-30 days. Surprisingly, drifters from the southern limb of the East Madagascar Current were transported westward across the channel to the Mozambique shelf. This transport was similarly facilitated by the frontal flow field between eddies. It is hypothesised that the frontal zones between eddies and interstitial waters play an important role in distributing biota in the Mozambique Channel.

Résumé: The MESOBIO programme investigated mesoscale dynamics using an integrated ecosystem approach, linking physical and biogeochemical processes with different trophic levels. Observation and modeling were used in combination to explain the main processes occurring in the mesoscale eddy field. The particular shape of the Mozambique Channel, composed of two basins interconnected through a narrow zone, favours the generation of mesoscale eddies and increases the opportunity for eddy-shelf interactions. Phytoplankton abundance peaked in areas of nutrient enrichment that are often found in the core of cyclonic eddies, as well as on the continental shelf. Grazers in zooplankton communities exhibited high biovolume in cyclonic eddies, but their abundance was lower in fronts and divergence zones, with lowest biovolume in anticyclones. Biovolume was highest at shelf stations, but very variable and similar to phytoplankton. Age of eddies, their subsequent maturation stage and the dynamics of the eddy field played a major role effecting zooplankton abundance. Micronekton presented abundance patterns coherent with zooplankton distribution, however this was only demonstrated by acoustic methods, whereas mid-water trawl collection and predators stomach contents (predators being used as biological samplers) did not reveal significant relationships with mesoscale features. For upper trophic levels, the average density of foraging seabirds was lowest in anticyclones, highest in cyclones and at intermediate levels in divergence, shelf and frontal zones. However, multifaceted behavioral responses were observed in such a highly variable environment. Swordfish was clearly associated with divergence zones, and to a lesser extent with fronts, suggesting that the higher density in divergences was related to the presence of its main prey, essentially large squids. Although tunas tended to be more abundant in areas with weak geostrophic currents, their relationship to mesoscale features was not straightforward as adult tunas caught by longline have the ability to explore different foraging habitats over a broad range of depths. Several suggestions for advancing eddy-related research from the current state of knowledge are proposed in the second part of the paper.

Résumé: We investigated the diversity and distribution of two communities, micronekton organisms and large predatory fishes, sampled in mesoscale features of the Mozambique Channel from 2003 to 2009, by combining mid-water trawls, stomach contents of fish predators and instrumented longline fishing surveys. The highest species richness for assemblages was found in divergences and fronts rather than in the core of eddies. Despite an unbalanced scheme, diversity indices did not differ significantly between cyclonic and anticyclonic eddies, divergences and fronts. We found that eddies and associated physical cues did not substantially affect the distribution of micronektonic species which are mainly driven by the diel vertical migration pattern. Top predators exhibited a more complex response. Swordfish (Xiphias gladius) associated better with mesoscale features than tunas, with a clear preference for divergences which is consistent with the diel vertical migrations and occurrence of its main prey, the flying squids Sthenoteuthis oualaniensis (Ommastrephidae). On the other hand, the probability of presence of yellowfin tuna was not tied to any specific eddy structure. However, the highest values of positive yellowfin CPUEs were associated with low horizontal gradients of sea-level anomalies. We also showed a non-linear response of positive yellowfin CPUEs with respect to the depth of the minimal oxygen content. The larger the distance between the hooks and the minimal oxygen layer, towards the surface or at greater depths, the higher the CPUE, highlighting that yellowfin congregated in well-oxygenated waters. Micronekton sampled by mid-water trawls and stomach contents exhibited different species composition. The highly mobile organisms were not caught by trawling whereas they remain accessible to predators. The combination of stomach contents and mid-water trawls undoubtedly improved our understanding of the micronekton assemblage distribution. Our results provide some evidence that mesoscale features in the Mozambique Channel do not strongly affect the distribution of the mid-trophic level organisms such as micronekton and most of the large predatory fishes, and hypotheses are proposed to support this result.

Résumé: A multidisciplinary programme, MESOBIO (Influence of mesoscale dynamics on biological productivity at multiple trophic levels in the Mozambique Channel) was undertaken in the Mozambique Channel within the framework of a scientific partnership between France and South Africa. MESOBIO focused on the signature of the highly energetic eddy dynamics in the Mozambique Channel. The Channel, which is known to be one of the most turbulent areas in the world ocean, has a great diversity of marine organisms and is the site of active pelagic fisheries. MESOBIO was mostly based on observations at sea during 12 multidisciplinary cruises between 2002 and 2010. Hydrographic measurements, sampling of biological organisms ranging from phytoplankton to top predators, and experiments on primary production and energy transfer through the food web, were conducted onboard various research vessels. The data were analysed in relation to eddy field characteristics for the periods of the cruises, including seasonal or inter-annual variability in mesoscale activity. A modelling approach was also developed within MESOBIO for both the circulation in the Channel and the biogeochemical response to eddy forcing. This paper introduces the suite of articles on the MESOBIO investigations by summarizing background knowledge for the different disciplines and the key issues that were addressed within the programme.

Résumé: We analyse the coupling between sea surface chlorophyll concentration (CC) and the physical environment in the Mozambique Channel (MZC) using statistical models. Seasonal and interannual patterns are studied along with the role of mesoscale dynamics on enhancement and concentration processes for phytoplankton. We use SeaWifs data for CC and two other remotely sensed data sets, TMMI for sea surface temperature (SST) and merged altimetry products for sea level anomaly and geostrophic current. Empirical Orthogonal Functions (EOF) on SSC and SST show strong seasonality and partition the MZC into three distinct sub-areas. The chlorophyll variability is mostly driven by seasonality, but more in the North (10 degrees S-16 degrees S) and South (24 degrees S-30 degrees S), and explains respectively 64% and 82% of the CC variance. In the Central part (16 degrees S-24 degrees S), the seasonal signal has less influence (60% variance). There, complex EOFs on Sea Level Anomaly (SLA) highlight the role of mesoscale activity (i.e. eddies and filament structures) in the spatial distribution of chlorophyll. Five mesoscale descriptors (shear, stretch, vorticity, deformation and eddy kinetic energy) are derived from the altimetry data to quantify the eddies-related physical patterns in the central region of the MZC. We use generalized Additive Models to explain the effect of those features on phytoplankton enhancement. The best model fit (r(2) = 0.73) includes shear, stretch, vorticity and the latitude-longitude interaction as eddies are well structured in space. Cyclonic eddies associated with negative vorticity are conductive to phytoplankton enhancement by the effect of upwelling in the core notably during the spin-up phase. The interaction between eddies generate strong frontal mixing favourable to the production and aggregation of organic matter. The mesoscale activity is also affected by interannual variability with consequences on CC. We highlight a substantial reduction of the SLA pattern in 2000-2001 when the SOI positive phase is peaking (Nina-type pattern). The strong relationship between mesoscale eddies and SOI suggests that primary productivity in the MZC is also under the influence of distant forcing at a basin scale.