Spatial and temporal interaction between sediment and microphytobenthos in a temperate estuarine macro-intertidal bay

Intertidal flats of the estuarine macro-intertidal Baie des Veys (France) were investigated to identify spatial features of sediment and microphytobenthos (MPB) in April 2003. Gradients occurred within the domain, and patches were identified close to vegetated areas or within the oyster-farming areas where calm physical conditions and biodeposition altered the sediment and MPB landscapes. Spatial patterns of chl a content were explained primarily by the influence of sediment features, while bed elevation and compaction brought only minor insights into MPB distribution regulation. The smaller size of MPB patches compared to silt patches revealed the interplay between physical structure defining the sediment landscape, the biotic patches that they contain, and that median grain-size is the most important parameter in explaining the spatial pattern of MPB. Small-scale temporal dynamics of sediment chl a content and grain-size distribution were surveyed in parallel during 2 periods of 14 d to detect tidal and seasonal variations. Our results showed a weak relationship between mud fraction and MPB biomass in March, and this relationship fully disappeared in July. Tidal exposure was the most important parameter in explaining the summer temporal dynamics of MPB. This study reveals the general importance of bed elevation and tidal exposure in muddy habitats and that silt content was a prime governing physical factor in winter. Biostabilisation processes seemed to behave only as secondary factors that could only amplify the initial silt accumulation in summer rather than primary factors explaining spatial or long-term trends of sediment changes.

Seep communities from two mud volcanoes in the deep eastern Mediterranean Sea: faunal composition, spatial patterns and environmental control

The Mediterranean Sea constitutes a unique environment to study cold-seep ecosystems due to the presence of different geodynamic settings, from an active margin along the Mediterranean Ridge (MR) to a passive margin in the Nile Deep-Sea Fan (NDSF). We attempted to identify the structure of benthic communities associated with the Napoli and Amsterdam mud volcanoes (MVs) located on the MR and to establish the links between faunal distribution and environmental conditions at different spatial scales. Comparison between the 2 MVs revealed that the faunal distribution seemed to be mainly controlled by the characteristics of the microhabitats. On both geological structures, the variability between the different microhabitats was higher than the variability observed between replicates of the same microhabitat, and the distribution of macro-fauna was apparently linked to gradients in physico-chemical conditions. The peripheral sites from Napoli were generally more oxygenated and harboured lower species richness than the active sites. The reduced sediment microhabitat from Amsterdam presented the highest methane concentrations and was mainly colonised by symbiont-bearing vesicomyid bivalves and heterotrophic dorvilleid polychaetes. Overall, a higher taxonomic diversity was observed on Napoli. Sub-stratum type was hypothesised to be the second factor influencing faunal distribution. The results of this study highlight the high heterogeneity of faunal communities associated with seep ecosystems within this region and the need to pursue investigations at various spatial and temporal scales.

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.

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.

Abrupt shifts in the Gironde fish community: an indicator of ecological changes in an estuarine ecosystem

For decades, global climate change has directly and indirectly affected the structure and function of ecosystems. Abrupt changes in biodiversity have been observed in response to linear or sudden modifications to the environment. These abrupt shifts can cause long-term reorganizations within ecosystems, with communities exhibiting new functional responses to environmental factors. Over the last 3 decades, the Gironde estuary in southwest France has experienced 2 abrupt shifts in both the physical and chemical environments and the pelagic community. Rather than describing these shifts and their origins, we focused on the 3 inter-shift periods, describing the structure of the fish community and its relationship with the environment during these periods. We described fish biodiversity using a limited set of descriptors, taking into account both species composition and relative species abundances. Inter-shift ecosystem states were defined based on the relationship between this description and the hydro-physico-chemical variables and climatic indices defining the main features of the environment. This relationship was described using generalized linear mixed models on the entire time series and for each inter-shift period. Our results indicate that (1) the fish community structure has been significantly modified, (2) environmental drivers influencing fish diversity have changed during these 3 periods, and (3) the fish-environment relationships have been modified over time. From this, we conclude a regime shift has occurred in the Gironde estuary. We also highlight that anthropogenic influences have increased, which re-emphasizes the importance of local management in maintaining fish diversity and associated goods and services within the context of climate change.

Temporal variation of delta C-13 in particulate organic matter and oyster Crassostrea gigas in Marennes-Oleron Bay (France): Effect of freshwater inflow

The temporal variability of delta(13)C in suspended particulate organic matter (POM) and oyster Crassostrea gigas along a salinity gradient was investigated from May 1992 to September 1993 within the estuarine bay of Marennes-Oleron (France). During this period the mean daily discharge of the Charente River exhibited large seasonal variation, with a high discharge from November 1992 to January 1993. Contrary to that at the river mouth and the marine littoral, delta(13)C in POM and in oysters at mid-estuary was affected by the high flood period. The delta(13)C values of POM decreased in mid-estuary and remained at low levels during the high discharge period, indicating an increasing contribution of terrestrial inputs to the estuarine POM pool. At the same site, a remarkable decrease of delta(13)C in oysters occurred between December 1992 and March 1993 (after a time lag compared to the ambient POM), indicating incorporation of terrestrial organic matter in oyster tissues during the high flood discharge. The lag between the delta(13)C decrease in POM and oysters is attributed to the time needed for oyster tissues to incorporate enough newly terrestrial light carbon to be recognized by the delta(13)C measure (about 1 to 2 mo). This time interval depends on tissue turnover time. The delta(13)C POM decrease (i.e. 1.3 parts per thousand) cannot explain entirely the decrease observed in oysters (i.e. 2.3 parts per thousand). In fact, the pattern exhibited by mid-estuarine oysters can be explained by the increasing contribution of terrestrial organic matter to their feeding, and the inability to preferentially utilize specific components of the estuarine POM that are C-13-enriched.

Impacts of high-nitrate freshwater inputs on macrotidal ecosystems. I. Seasonal evolution of nutrient limitation for the diatom-dominated phytoplankton of the Bay of Brest (France)

The chemical factors (inorganic nitrogen, phosphate, silicic acid) that potentially or actually control primary production were determined for the Bay of Brest, France, a macrotidal ecosystem submitted to high-nitrate-loaded freshwater inputs (winter nitrate freshwater concentrations >700 mu M, Si:N molar ratio as low as 0.2, i.e. among the lowest ever published). Intensive data collection and observations were carried out from February 1993 to March 1994 to determine the variations of physical [salinity, temperature, photosynthetically active radiation (PAR), freshwater discharges] and chemical (oxygen and nutrients) parameters and their impacts on the phytoplankton cycle (fluorescence, pigments, primary production). With insufficient PAR the winter stocks of nutrients were almost nonutilized and the nitrate excess was exported to the adjacent ocean, due to rapid tidal exchange. By early April, a diatom-dominated spring bloom developed (chlorophyll a maximum = 7.7 mu g l(-1); primary production maximum = 2.34 g C m(-2) d(-1)) under high initial nutrient concentrations. Silicic acid was rapidly exhausted over the whole water column; it is inferred to be the primary limiting factor responsible for the collapse of the spring bloom by mid-May. Successive phytoplankton developments characterized the period of secondary blooms during summer and fall (successive surface chlorophyll a maxima = 3.5, 1.6, 1.8 and 1.0 mu g l(-1); primary production = 1.24, 1.18 and 0.35 g C m(-2) d(-1)). Those secondary blooms developed under lower nutrient concentrations, mostly originating from nutrient recycling. Until August, Si and P most likely limited primary production, whereas the last stage of the productive period in September seemed to be N limited instead, this being a period of total nitrate depletion in almost the whole water column. Si limitation of spring blooms has become a common feature in coastal ecosystems that receive freshwater inputs with Si:N molar ratios <1. The peculiarity of Si Limitation in the Bay of Brest is its extension through the summer period.

Lipid and lipid carbon stable isotope composition of the hydrothermal vent shrimp Mirocaris fortunata: evidence for nutritional dependence on photosynthetically fixed carbon

Mirocaris fortunata were sampled from the Lucky Strike hydrothermal vent area (Eiffel Tower site) on the mid-Atlantic ridge during the French DIVA 2 cruise (June 1994). Small adults (17 to 22 mm total length), although morphologically identical, could be divided into 2 categories on the basis of pigmentation, lipid composition and C-13/C-12 stable isotope ratios of fatty acids. Highly pigmented small adults (8.6 to 9.2 mu g carotenoid shrimp(-1)) contained higher levels of total lipid than similar-sized individuals containing lower levels of pigment (0.9 to 2.9 mu g carotenoid shrimp(-1)). Lipid class analysis indicated that wax esters comprised 62.5% of total lipid in the former group. These pigmented shrimp also contained high proportions of polyunsaturated fatty acids (PUFA), particularly the phototrophic microplanktonic markers 20:5(n-3) and 22:6(n-3) (14.0 and 33.5% respectively). By contrast small adults (22 mm) and adult shrimp (25 to 26 mm) with low levels of carotenoid pigmentation contained lower amounts of total lipid, little or no wax ester and low levels of 20:5(n-3) and 22:6(n-3), but did contain 16:2(n-4) and 18:2(n-4) and the non-methylene interrupted dienes 20:2 Delta 5,13 and 22:2 Delta 7,15. GC-IRMS analysis of all fatty acids and fatty alcohols in the pigmented small adults indicated delta(13)C values of -18.2 to -27.7 parts per thousand, which is consistent with a photosynthetic carbon source for these compounds. The C-13/C-12 isotope composition of fatty acids from low-pigmented small adults and adults was more variable (-12.5 to -33.1 parts per thousand) and suggests a bimodal distribution which may be attributable to differing nutritional sources or the physiological/reproductive status of these shrimp. Samples of eggs, which are carried by the female on the pleopods, represented approximately 57% of total somatic lipid which indicates a substantial reproductive investment by this species. The egg lipids comprised high proportions of triacylglycerols (64.4 to 78.0% of total lipid) whilst the fatty acid composition was dominated by the monounsaturated fatty acids 16:1(n-7), 18:1(n-7) and 18:1(n-9), which accounted for 65.7 to 33.5% of total fatty acids. By contrast, PUFA were relatively minor components of egg lipids, particularly 20:5(n-3) and 22:6(n-3), which accounted for only 1.1 and 2.9% of total egg fatty acids respectively. This indicates that the reproductive investment by this species is supported mainly by material derived from bacterial chemosynthesis. The potential for M. fortunata hedge betting by producing larvae which either metamorphose at the vent site or adopt a bathypelagic lifestyle and delay metamorphosis to facilitate more widespread dispersal is discussed.

Explaining dinophysis cf. acuminata abundance in Antifer (Normandy, France) using dynamic linear regression

Classical regression analysis can be used to model time series. However, the assumption that model parameters are constant over time is not necessarily adapted to the data. In phytoplankton ecology, the relevance of time-varying parameter values has been shown using a dynamic linear regression model (DLRM). DLRMs, belonging to the class of Bayesian dynamic models, assume the existence of a non-observable time series of model parameters, which are estimated on-line, i.e. after each observation. The aim of this paper was to show how DLRM results could be used to explain variation of a time series of phytoplankton abundance. We applied DLRM to daily concentrations of Dinophysis cf. acuminata, determined in Antifer harbour (French coast of the English Channel), along with physical and chemical covariates (e.g. wind velocity, nutrient concentrations). A single model was built using 1989 and 1990 data, and then applied separately to each year. Equivalent static regression models were investigated for the purpose of comparison. Results showed that most of the Dinophysis cf. acuminata concentration variability was explained by the configuration of the sampling site, the wind regime and tide residual flow. Moreover, the relationships of these factors with the concentration of the microalga varied with time, a fact that could not be detected with static regression. Application of dynamic models to phytoplankton time series, especially in a monitoring context, is discussed.

Age estimation in common sole Solea solea larvae: validation of daily increments and evaluation of a pattern recognition technique

Close similarities have been found between the otoliths of sea-caught and laboratory-reared larvae of the common sole Solea solea (L.), given appropriate temperatures and nourishment of the latter. But from hatching to mouth formation. and during metamorphosis, sole otoliths have proven difficult to read because the increments may be less regular and low contrast. In this study, the growth increments in otoliths of larvae reared at 12 degrees C were counted by light microscopy to test the hypothesis of daily deposition, with some results verified using scanning electron microscopy (SEM), and by image analysis in order to compare the reliability of the 2 methods in age estimation. Age was first estimated (in days posthatch) from light micrographs of whole mounted otoliths. Counts were initiated from the increment formed at the time of month opening (Day 4). The average incremental deposition rate was consistent with the daily hypothesis. However, the light-micrograph readings tended to underestimate the mean ages of the larvae. Errors were probably associated with the low-contrast increments: those deposited after the mouth formation during the transition to first feeding, and those deposited from the onset of eye migration (about 20 d posthatch) during metamorphosis. SEM failed to resolve these low-contrast areas accurately because of poor etching. A method using image analysis was applied to a subsample of micrograph-counted otoliths. The image analysis was supported by an algorithm of pattern recognition (Growth Demodulation Algorithm, GDA). On each otolith, the GDA method integrated the growth pattern of these larval otoliths to averaged data from different radial profiles, in order to demodulate the exponential trend of the signal before spectral analysis (Fast Fourier Transformation, FFT). This second method both allowed more precise designation of increments, particularly for low-contrast areas, and more accurate readings but increased error in mean age estimation. The variability is probably due to a still rough perception of otolith increments by the GDA method, counting being achieved through a theoretical exponential pattern and mean estimates being given by FFT. Although this error variability was greater than expected, the method provides for improvement in both speed and accuracy in otolith readings.

Sampling design optimization for EROD measurements in fish

Activity of 7-ethoxyresorufin-O-deethylase (EROD) in fish is certainly the best-studied biomarker of exposure applied in the field to evaluate biological effects of contamination in the marine environment. Since 1991, a feasibility study for a monitoring network using this biomarker of exposure has been conducted along French coasts. Using data obtained during several cruises, this study aims to determine the number of fish required to detect a given difference between 2 mean EROD activities, i.e. to achieve an a priori fixed statistical power (l-beta) given significance level (alpha), variance estimations and projected ratio of unequal sample sizes (k). Mean EROD activity and standard error were estimated at each of 82 sampling stations. The inter-individual variance component was dominant in estimating the variance of mean EROD activity. Influences of alpha, beta, k and variability on sample sizes are illustrated and discussed in terms of costs. In particular, sample sizes do not have to be equal, especially if such a requirement would lead to a significant cost in sampling extra material. Finally, the feasibility of longterm monitoring is discussed.

Ingestion of a bacterivorous ciliate by the oyster Crassostrea gigas: Protozoa as a trophic link between picoplankton and benthic suspension-feeders

The linked concepts of 'microbial loop' and 'protozoan trophic link' have been very well documented in filter-feeding microzooplankton such as copepods, but have not been applied to energy transfer to benthic suspension-feeding macrofauna, with the exception of the recent demonstration of heterotrophic flagellate assimilation by mussels. The oyster Crassostrea gigas obtains energy resources by filtering microalgae (similar to 5 to 100 mu m). However, in turbid estuaries, light-limited phytoplanktonic production cannot entirely account for oyster energy requirements. Conversely, picoplankters (<2 mu m), which are main effecters of coastal energy flow and matter cycling, are not efficiently retained by oyster filtration. Ciliate protozoal as both micro-sized cells (similar to 5 to 100 run) and bacteria grazers, may represent a major intermediary in trophic transfer between picoplankton and metazoa. The ciliate Uronema was intensely cultured and labelled, using the cyanobacteria Synechococcus as an auto-fluorescent biomarker. The labelled ciliates were offered as potential prey to oysters. We report here the first experimental evidence of a significant retention and ingestion of ciliates by oysters, supporting the role of protozoa as a realistic trophic link between picoplankters and filter-feeding bivalves and thus enhancing their potential importance in estuarine microbial food webs.

Kinetics of tidal resuspension of microbiota: Testing the effects of sediment cohesiveness and bioturbation using flume experiments

Resuspension of the top few sediment layers of tidal mud flats is known to enhance planktonic biomass of microbiota (benthic diatoms and bacteria). This process is mainly controlled by tidal shear stress and cohesiveness of mud, and is also influenced by bioturbation activities. Laboratory experiments in a race track flume were performed to test the interactive effects of these factors on both the critical entrainment and resuspension kinetics of microbiota from silt-clay sediments from the Marennes-Oleron Bay, France. The marine snail Hydrobia ulvae was used to mimic surface bioturbation activities. As expected, the kinetics of microbial resuspension versus shear stress were largely controlled by the cohesiveness of silt-clay sediments. However, our results indicate that the effect of surface tracking by H. ulvae on microbial resuspension was clearly dependent on the interaction between sediment cohesiveness and shear velocity. Evidence was also found that microphytobenthos and bacteria are not simultaneously resuspended from silt-clay bioturbated sediments. This supports the theory that diatoms within the easily eroded mucus matrix behave actively and bacteria adhering to fine silt particles eroded at higher critical shear velocities behave passively.

Modelling the biogeochemical cycles of elements limiting primary production in the English channel .1. Role of thermohaline stratification

This paper presents the general framework of an ecological model of the English Channel. The model is a result of combining a physical sub-model with a biological one. in the physical submodel, the Channel is divided into 71 boxes and water fluxes between them are calculated automatically. A 2-layer, vertical thermohaline model was then linked with the horizontal circulation scheme. This physical sub-model exhibits thermal stratification in the western Channel during spring and summer and haline stratification in the Bay of Seine due to high flow rates from the river. The biological sub-model takes 2 elements, nitrogen and silicon, into account and divides phytoplankton into diatoms and dinoflagellates. Results from this ecological model emphasize the influence of stratification on chlorophyll a concentrations as well as on primary production. Stratified waters appear to be much less productive than well-mixed ones. Nevertheless, when simulated production values are compared with literature data, calculated production is shown to be underestimated. This could be attributed to a lack of refinement of the 2-layer box-model or processes omitted from the biological model, such as production by nanoplankton.

Modelling the biogeochemical cycles of elements limiting primary production in the English channel .2. Sensitivity analyses

A general framework for an ecological model of the English Channel was described in the first of this pair of papers. In this study, it was used to investigate the sensitivity of the model to various factors: model structure, parameter values, boundary conditions and forcing variables. These sensitivity analyses show how important quota formulation for phytoplankton growth is, particularly for growth of dinoflagellates. They also stress the major influence of variables and parameters related to nitrogen. The role played by rivers and particularly the river Seine was investigated. Their influence on global English Channel phytoplanktonic production seems to be relatively low, even though nutrient inputs determine the intensity of blooms in the Bay of Seine. The geographical position of the river Seine's estuary makes it important in fluxes through the Straits of Dover. Finally, the multi-annual study highlights the general stability of the English Channel ecosystem. These global considerations are discussed and further improvements to the model are proposed.