Trophic niche overlap of sprat and commercial small pelagic teleosts in the Gulf of Lions (NW Mediterranean Sea)

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.

Influence of environmental variability and age on the body condition of small pelagic fish in the Gulf of Lions

Endogenous and environmental variables are fundamental in explaining variations in fish condition. Based on more than 20 yr of fish weight and length data, relative condition indices were computed for anchovy and sardine caught in the Gulf of Lions. Classification and regression trees (CART) were used to identify endogenous factors affecting fish condition, and to group years of similar condition. Both species showed a similar annual cycle with condition being minimal in February and maximal in July. CART identified 3 groups of years where the fish populations generally showed poor, average and good condition and within which condition differed between age classes but not according to sex. In particular, during the period of poor condition (mostly recent years), sardines older than 1 yr appeared to be more strongly affected than younger individuals. Time-series were analyzed using generalized linear models (GLMs) to examine the effects of oceanographic abiotic (temperature, Western Mediterranean Oscillation [WeMO] and Rhone outflow) and biotic (chlorophyll a and 6 plankton classes) factors on fish condition. The selected models explained 48 and 35% of the variance of anchovy and sardine condition, respectively. Sardine condition was negatively related to temperature but positively related to the WeMO and mesozooplankton and diatom concentrations. A positive effect of mesozooplankton and Rhone runoff on anchovy condition was detected. The importance of increasing temperatures and reduced water mixing in the NW Mediterranean Sea, affecting planktonic productivity and thus fish condition by bottom-up control processes, was highlighted by these results. Changes in plankton quality, quantity and phenology could lead to insufficient or inadequate food supply for both species.

Live−dead comparison of benthic foraminiferal faunas from the Rhône prodelta (Gulf of Lions, NW Mediterranean): Development of a proxy for palaeoenvironmental reconstructions

Dead benthic foraminiferal faunas (> 150 μm) from the Rhône prodelta (Gulf of Lions, NW Mediterranean) were analysed at 41 stations (15–100 m water depth) sampled in June 2005 and September 2006, and compared to the living faunas investigated during previous studies at the same stations. The comparison between dead and living assemblages enhances the understanding of taphonomic processes that may modify the composition of the dead faunas in this area. We observed a loss of individuals from living to dead assemblages of species characterised by a fairly fragile test and therefore more prone to fragmentation or dissolution (e.g., Bolivina alata, Quinqueloculina tenuicollis). Allochthonous dead and/or live specimens may be transported to some parts of the prodelta, particularly the shallowest sites where hydrodynamic processes (i.e., river flood, storm swells, longshore currents) are more intense. These specimens may originate from relict deltaic structures (e.g., Elphidium spp. from the lobe of Bras de Fer) or from surrounding areas (e.g., Ammonia beccarii forma beccarii from the river). Opportunistic species (e.g., Bulimina marginata, Cassidulina carinata) characterised by high reproductive rates have much higher relative abundances in the dead than in the living fauna. Cluster analyses based on dead foraminiferal assemblages divide our study area into four main thanatofacies directly related to distinct local environmental conditions prevailing in the prodelta. Close to the river mouth, Ammonia beccarii forma beccarii and Ammonia tepida are found in sediments subject to a high riverine influence (i.e., bottom currents, high organic and inorganic material input of continental origin). Elphidium species are abundant in the silty-sandy relict deltaic lobe west of the river mouth which is characterised by strong longshore currents that disturb the benthic environment. Nonion fabum, Rectuvigerina phlegeri and Valvulineria bradyana are found along the coast west of the Rhône River mouth, in the area defined as the “river plume” thanatofacies. In the more stable and deeper prodeltaic area, species known to feed on fresh phytodetritus (e.g., Bulimina aculeata/marginata, C. carinata, Hyalinea balthica) dominate the faunas. Since only minor variations in species relative abundances and spatial distributional patterns are observed between the living and the dead faunas, we consider that our thanatofacies have not been influenced by substantial transport of dead tests. This suggests that fossil benthic foraminifera can provide a reliable tool for investigating the development of the palaeo-Rhône prodelta

A multitracer approach to assess the spatial contamination pattern of hake (Merluccius merluccius) in the French Mediterranean

Chemical contamination levels and stable isotope ratios provide integrated information about contaminant exposure, trophic position and also biological and environmental influences on marine organisms. By combining these approaches with otolith shape analyses, the aim of the present study was to document the spatial variability of Hg and PCB contamination of the European hake (Merluccius merluccius) in the French Mediterranean, hypothesizing that local contaminant sources, environmental conditions and biological specificities lead to site-specific contamination patterns. High Hg concentrations discriminated Corsica (average: 1.36 ± 0.80 μg g− 1 dm) from the Gulf of Lions (average values < 0.5 μg g− 1 dm), where Rhône River input caused high PCB burdens. CB 153 average concentrations ranged between 4.00 ± 0.64 and 18.39 ± 12.38 ng g− 1 dm in the Gulf of Lions, whatever the sex of the individuals, whereas the highest values in Corsica were 6.75 ± 4.22 ng g− 1 dm. Otolith shape discriminated juveniles and adults, due to their different habitats. The use of combined ecotracers was revealed as a powerful tool to discriminate between fish populations at large and small spatial scale, and to enable understanding of the environmental and biological influences on contamination patterns.

Aerial surveys to monitor bluefin tuna abundance and track efficiency of management measures

Conservation and management measures for exploited fish species rely on our ability to monitor variations in population abundance. In the case of the eastern stock of Atlantic bluefin tuna (ABFT), recent changes in management policies have strongly affected the reliability of fishery-dependent indicators due to drastic changes in fishing season/area, fisheries selectivity and strategy. However, fishery-independent indices of abundance are rare for large pelagic fish, and obtaining them is often costly and labor intensive. Here, we show that scientific aerial surveys are an appropriate tool for monitoring juvenile bluefin tuna abundance in the Mediterranean. We present an abundance index based on 62 aerial surveys conducted since 2000, using 2 statistical approaches to deal with the sampling strategy: line and strip transects. Both approaches showed a significant increase in juvenile ABFT abundance in recent years, resulting from the recovery plan established in 2007. Nonetheless, the estimates from the line transect method appear to be more robust and stable. This study provides essential information for fisheries management. Expanding the spatial coverage to other nursery grounds would further increase the reliability and representativeness of this index.

Post-rift evolution of the Gulf of Lion margin tested by stratigraphic modelling

The sedimentary architecture of basins and passive margins is determined by a complex interaction of parameters, including subsidence, eustasy, and sediment supply. A quantification of the post-rift (20 Ma-0 Ma) vertical movements of the Gulf of Lion (West Mediterranean) is proposed here based on the stratigraphic study of sedimentary paleomarkers using a large 3D grid of reflection seismic data, correlations with existing drillings, and refraction data. Post-rift subsidence was measured by the direct use of sedimentary geometries analysed in 3D and validated by numerical stratigraphic modelling. Three domains of subsidence were found: on the continental shelf and slope, subsidence corresponds to a seaward tilting with different amplitudes, whereas the deep basin subsides purely vertically. We show that these domains fit with the deeper crustal domains highlighted by previous geophysical data, and that post-break-up subsidence follows the initial hinge lines of the rifting phase. Subsidence rates are quantified on each domain for each stratigraphic interval. At a constant distance from the rotational hinge line, the Plio-Quaternary subsidence rate is constant on the shelf overall. Conversely, Miocene subsidence rates are very different on the eastern and western shelves. Stratigraphic simulations focused on the Messinian salinity crisis (MSC) were also performed. Their results are discussed together with our post-rift subsidence estimates in order to provide ideas and hypotheses for future detailed quantifications of Miocene subsidence, including isostatic readjustments linked to the MSC.

A four-dimensional analysis of the thermal structure in the Gulf of Lion

The Theoretical and Experimental Tomography in the Sea Experiment (THETIS 1) took place in the Gulf of Lion to observe the evolution of the temperature field and the process of deep convection during the 1991-1992 winter. The temperature measurements consist, of moored sensors, conductivity-temperature-depth and expendable bathythermograph surveys, ana acoustic tomography. Because of this diverse data set and since the field evolves rather fast, the analysis uses a unified framework, based on estimation theory and implementing a Kalman filter. The resolution and the errors associated with the model are systematically estimated. Temperature is a good tracer of water masses. The time-evolving three-dimensional view of the field resulting from the analysis shows the details of the three classical convection phases: preconditioning, vigourous convection, and relaxation. In all phases, there is strong spatial nonuniformity, with mesoscale activity, short timescales, and sporadic evidence of advective events (surface capping, intrusions of Levantine Intermediate Water (LIW)). Deep convection, reaching 1500 m, was observed in late February; by late April the field had not yet returned to its initial conditions (strong deficit of LIW). Comparison with available atmospheric flux data shows that advection acts to delay the occurence of convection and confirms the essential role of buoyancy fluxes. For this winter, the deep. mixing results in an injection of anomalously warm water (Delta T similar or equal to 0.03 degrees) to a depth of 1500 m, compatible with the deep warming previously reported.

Statistical Analysis of Sea Surface Temperature and Chlorophyll-a Concentration Patterns in the Gulf of Tadjourah (Djibouti)

The sea surface temperature (SST) and chlorophyll-a concentration (CHL-a) were analysed in the Gulf of Tadjourah from two set of 8-day composite satellite data, respectively from 2008 to 2012 and from 2005 to 2011. A singular spectrum analysis (SSA) shows that the annual cycle of SST is strong (74.3% of variance) and consists of warming (April-October) and cooling (November-March) of about 2.5C than the long-term average. The semi-annual cycle captures only 14.6% of temperature variance and emphasises the drop of SST during July-August. Similarly, the annual cycle of CHL-a (29.7% of variance) depicts high CHL-a from June to October and low concentration from November to May. In addition, the first spatial empirical orthogonal function (EOF) of SST (93% of variance) shows that the seasonal warming/cooling is in phase across the whole study area but the southeastern part always remaining warmer or cooler. In contrast to the SST, the first EOF of CHL-a (54.1% of variance) indicates the continental shelf in phase opposition with the offshore area in winter during which the CHL-a remains sequestrated in the coastal area particularly in the south-east and in the Ghoubet Al-Kharab Bay. Inversely during summer, higher CHL-a quantities appear in the offshore waters. In order to investigate processes generating these patterns, a multichannel spectrum analysis was applied to a set of oceanic (SST, CHL-a) and atmospheric parameters (wind speed, air temperature and air specific humidity). This analysis shows that the SST is well correlated to the atmospheric parameters at an annual scale. The windowed cross correlation indicates that this correlation is significant only from October to May. During this period, the warming was related to the solar heating of the surface water when the wind is low (April-May and October) while the cooling (November-March) was linked to the strong and cold North-East winds and to convective mixing. The summer drop in SST followed by a peak of CHL-a, seems strongly correlated to the upwelling. The second EOF modes of SST and CHL-a explain respectively 1.3% and 5% of the variance and show an east-west gradient during winter that is reversed during summer. This work showed that the seasonal signals have a wide spatial influence and dominate the variability of the SST and CHL-a while the east-west gradient are specific for the Gulf of Tadjourah and seem induced by the local wind modulated by the topography.

Characterizing, modelling and understanding the climate variability of the deep water formation in the North-Western Mediterranean Sea

Observing, modelling and understanding the climate-scale variability of the deep water formation (DWF) in the North-Western Mediterranean Sea remains today very challenging. In this study, we first characterize the interannual variability of this phenomenon by a thorough reanalysis of observations in order to establish reference time series. These quantitative indicators include 31 observed years for the yearly maximum mixed layer depth over the period 1980–2013 and a detailed multi-indicator description of the period 2007–2013. Then a 1980–2013 hindcast simulation is performed with a fully-coupled regional climate system model including the high-resolution representation of the regional atmosphere, ocean, land-surface and rivers. The simulation reproduces quantitatively well the mean behaviour and the large interannual variability of the DWF phenomenon. The model shows convection deeper than 1000 m in 2/3 of the modelled winters, a mean DWF rate equal to 0.35 Sv with maximum values of 1.7 (resp. 1.6) Sv in 2013 (resp. 2005). Using the model results, the winter-integrated buoyancy loss over the Gulf of Lions is identified as the primary driving factor of the DWF interannual variability and explains, alone, around 50 % of its variance. It is itself explained by the occurrence of few stormy days during winter. At daily scale, the Atlantic ridge weather regime is identified as favourable to strong buoyancy losses and therefore DWF, whereas the positive phase of the North Atlantic oscillation is unfavourable. The driving role of the vertical stratification in autumn, a measure of the water column inhibition to mixing, has also been analyzed. Combining both driving factors allows to explain more than 70 % of the interannual variance of the phenomenon and in particular the occurrence of the five strongest convective years of the model (1981, 1999, 2005, 2009, 2013). The model simulates qualitatively well the trends in the deep waters (warming, saltening, increase in the dense water volume, increase in the bottom water density) despite an underestimation of the salinity and density trends. These deep trends come from a heat and salt accumulation during the 1980s and the 1990s in the surface and intermediate layers of the Gulf of Lions before being transferred stepwise towards the deep layers when very convective years occur in 1999 and later. The salinity increase in the near Atlantic Ocean surface layers seems to be the external forcing that finally leads to these deep trends. In the future, our results may allow to better understand the behaviour of the DWF phenomenon in Mediterranean Sea simulations in hindcast, forecast, reanalysis or future climate change scenario modes. The robustness of the obtained results must be however confirmed in multi-model studies.

Nutrients limiting the Algal Growth Potential (AGP) in the Gulf of Riga, eastern Baltic Sea, in spring and early summer 1996

In May, June and July 1996, samples wcre collected along one transect greatly influenced by river discharge (eastern side of the gulf), along one transect slightly influence by river discharge (western side), at one station Iocated in the mouth of the main river (River Daugava), at one station located in the center of the Gulf and at several nearshore locations of the western side. Ratios of rnolecular concentrations of in situ dissolved ioorganic nitrogen, phosphorus and silicon, as weIl as enrichment bioassays were llsed to dctcrrnine which nutrient (s) lirnited the potential biomass of phytoplankton. Both comparison of (NO.d-N02+NJ.L): P04 (DIN: DIP) values with Redfic1d's ratio and bioassay inspection led to the sarne conclusions. Phosphorus was clearly the nutrient most limiting for the potcntial biornass of test species in nitrogen- rich waters, which occurred in mid spring, in the upper layer of the southern-eastern part of the Gulf which is greatly influenced by river discharge. In late spring, with the decrease of the total DIN reserve, nitrogen and phosphorus showed an equallimiting role. In deeper layers of this area and out of the river plume (western side and central part of the gulf), nitrogen was the limiting nutrient. In summer, whcn river discharge was the lowest, a11 DIN concentrations but one ranged between 1.6 and 2.6 µM, and the whole area was nitrogen-limited for both the cyanobacterial and the algal test strains. In 74% of the samples for which nitrogen was the limiting nutrient, phosphorus was recorded to be the second potentially limiting nutrient. In contrast, silicon never appeared as limiting the growth potential of either Microcystis aeruginosa or Phaeodactylum tricornutum; phosphorus was the limiting nutrient when DIN: Si03 values were >1 (in May), but DIN: Si03 was <1 when nitrogen was limiting (June and July). The authors conclude that the recently reported decrease of silicon loading in coastal waters and its subsequent enhanced importance in pushing the outcome of species competition towards harmful species may not yet be the most important factor for the Gulf of Riga. Iron appeared for 12% of the tests in the list of nutrients limiting the potential biomass. Tentative results also indicated that a significant fraction of the nitrogen (~,4 µg-atom N 1(-1) taken up by Microcystis aeruginosa may have been in the form of dissolved organic nitrogen (DON). It is thus also suggested tentatively that more attention be paid to these nitrients during further research in the Gulf of Riga.

Body reserves mediate trade-offs between life-history traits: new insights from small pelagic fish reproduction

Limited resources in the environment prevent individuals from simultaneously maximizing all life-history traits, resulting in trade-offs. In particular, the cost of reproduction is well known to negatively affect energy investment in growth and maintenance. Here, we investigated these trade-offs during contrasting periods of high versus low fish size and body condition (before/after 2008) in the Gulf of Lions. Female reproductive allocation and performance in anchovy (Engraulis encrasicolus) and sardine (Sardina pilchardus) were examined based on morphometric historical data from the 1970s and from 2003 to 2015. Additionally, potential maternal effects on egg quantity and quality were examined in 2014/2015. After 2008, the gonadosomatic index increased for sardine and remained steady for anchovy, while a strong decline in mean length at first maturity indicated earlier maturation for both species. Regarding maternal effects, for both species egg quantity was positively linked to fish size but not to fish lipid reserves, while the egg quality was positively related to lipid reserves. Atresia prevalence and intensity were rather low regardless of fish condition and size. Finally, estimations of total annual numbers of eggs spawned indicated a sharp decrease for sardine since 2008 but a slight increase for anchovy during the last 5 years. This study revealed a biased allocation towards reproduction in small pelagic fish when confronted with a really low body condition. This highlights that fish can maintain high reproductive investment potentially at the cost of other traits which might explain the present disappearance of old and large individuals in the Gulf of Lions.