Seasonality in coastal macrobenthic biomass and its implications for estimating secondary production using empirical models

Macrobenthic secondary production is widely used to assess the trophic capacity, health, and functioning of marine and freshwater ecosystems. Annual production estimates are often calculated using empirical models and based on data collected during a single period of the year. Yet, many ecosystems show seasonal variations. Although ignoring seasonality may lead to biased and inaccurate estimates of annual secondary production, it has never been tested at the community level. Using time series of macrobenthic data collected seasonally at three temperate marine coastal soft-bottom sites, we assessed seasonal variations in biomass of macrobenthic invertebrates at both population and community levels. We then investigated how these seasonal variations affect the accuracy of annual benthic production when assessed using an empirical model and data from a single sampling event. Significant and consistent seasonal variations in biomass at the three study sites were highlighted. Macrobenthic biomass was significantly lower in late winter and higher in summer/early fall for 18 of the 30 populations analyzed and for all three communities studied. Seasonality led to inaccurate and often biased estimates of annual secondary production at the community level when based on data from a single sampling event. Bias varied by site and sampling period, but reached similar to 50% if biomass was sampled at its annual minimum or maximum. Since monthly sampling is rarely possible, we suggest that ecologists account for uncertainty in annual production estimates caused by seasonality.Agência financiadora EDF French Ministry of Higher Education, Research and Innovation French Ministry for the Ecological and Inclusive Transition through the Marine Strategy Framework Directive Agreement French Biodiversity Agency (Agence francaise pour la biodiversite) as part of the CAPANOUR projectinfo:eu-repo/semantics/publishedVersio

Trophic interactions within bentho-demersal communities : influence of the food availability on the carrying capacity of juvenile fish coastal nurseries.

Les habitats côtiers à fonds meubles sont essentiels dans le cycle de vie de nombreuses espèces de poissons. Ces habitats ont pour caractéristique d’être peu étendus mais sont aussi très productifs et jouent à ce titre un rôle de nourricerie pour les juvéniles de poissons bentho-démersaux. Les fortes abondances de proies semblent être l’une des raisons principales de la présence des juvéniles au sein de ces habitats. Bien que déjà étudié, le caractère limitant de la nourriture disponible fait toujours l’objet de débats dans la communauté scientifique. Une des raisons principales est que l’étude des milieux côtiers est rendue complexe par la diversité des facteurs entrant en jeu dans le développement des jeunes stades de poissons. A partir du cas d’étude de la baie de Vilaine, une des nourriceries les plus productives du golfe de Gascogneles recherches menées dans cette thèse visent à définir le rôle du facteur alimentaire dans l’organisation de la nourricerie et dans sa capacité à soutenir le développement des juvéniles de poissons. Le fil conducteur de cette thèse est donc de déterminer si le facteur alimentaire limite la production de juvéniles. Les résultats montrent une forte exploitation de la production alimentaire ainsi qu'une superposition spatiale entre les densités de juvéniles de poissons et de leurs proies. La cohérence de ces résultats tend à soutenir l'hypothèse que la capacité d'accueil de la baie de Vilaine est atteinte et donc que le facteur alimentaire est limitant. Les implications de ce mécanisme de régulation sur la dynamique des populations nourricerSoft sediment coastal habitats are essential in the life cycle of numerous fishes. These habitats are spatially-limited but very productive, and play a key role of nursery for the juveniles of benthic and demersal fishes. High abundance of prey seems to be one of the main reasons of the presence of juvenile fish within these habitats. Although widely studied, the limiting aspect of the feeding factor is still debated in the scientific community. One of the main reasons is that studying coastalhabitats is complex because numerous factors influence the development of the first life stages of fish. From the study case of the Bay of Vilaine, one of the most productive nurseries of the Bay of Biscay, research conducted in this thesis aims to define the role of the feeding factor in the organisation of the nursery and in its capacity to support the development of juvenile fish. The common thread of this thesis is thus to determine if the feeding factor limits the juvenile fish production. The results show a strong exploitation of the food production by the juvenile fish community and a spatial match between the densities of juvenile fish and their prey. The consistency of these results tends to support that the hypothesis that the carrying capacity of the Bay of Vilaine is reached, and that the feeding factor is limiting. The consequences of this regulation process on the dynamics of nursery-dependent fish populations are discussed

Interactions trophiques au sein des communautés bentho-demersales : Influence de la disponibilité alimentaire sur la capacité d’accueil des nourriceries côtières de juvéniles de poissons

Soft sediment coastal habitats are essential in the life cycle of numerous fishes. These habitats are spatially-limited but very productive, and play a key role of nursery for the juveniles of benthic and demersal fishes. High abundance of prey seems to be one of the main reasons of the presence of juvenile fish within these habitats. Although widely studied, the limiting aspect of the feeding factor is still debated in the scientific community. One of the main reasons is that studying coastalhabitats is complex because numerous factors influence the development of the first life stages of fish. From the study case of the Bay of Vilaine, one of the most productive nurseries of the Bay of Biscay, research conducted in this thesis aims to define the role of the feeding factor in the organisation of the nursery and in its capacity to support the development of juvenile fish. The common thread of this thesis is thus to determine if the feeding factor limits the juvenile fish production. The results show a strong exploitation of the food production by the juvenile fish community and a spatial match between the densities of juvenile fish and their prey. The consistency of these results tends to support that the hypothesis that the carrying capacity of the Bay of Vilaine is reached, and that the feeding factor is limiting. The consequences of this regulation process on the dynamics of nursery-dependent fish populations are discussed.Les habitats côtiers à fonds meubles sont essentiels dans le cycle de vie de nombreuses espèces de poissons. Ces habitats ont pour caractéristique d’être peu étendus mais sont aussi très productifs et jouent à ce titre un rôle de nourricerie pour les juvéniles de poissons bentho-démersaux. Les fortes abondances de proies semblent être l’une des raisons principales de la présence des juvéniles au sein de ces habitats. Bien que déjà étudié, le caractère limitant de la nourriture disponible fait toujours l’objet de débats dans la communauté scientifique. Une des raisons principales est que l’étude des milieux côtiers est rendue complexe par la diversité des facteurs entrant en jeu dans le développement des jeunes stades de poissons. A partir du cas d’étude de la baie de Vilaine, une des nourriceries les plus productives du golfe de Gascogneles recherches menées dans cette thèse visent à définir le rôle du facteur alimentaire dans l’organisation de la nourricerie et dans sa capacité à soutenir le développement des juvéniles de poissons. Le fil conducteur de cette thèse est donc de déterminer si le facteur alimentaire limite la production de juvéniles. Les résultats montrent une forte exploitation de la production alimentaire ainsi qu'une superposition spatiale entre les densités de juvéniles de poissons et de leurs proies. La cohérence de ces résultats tend à soutenir l'hypothèse que la capacité d'accueil de la baie de Vilaine est atteinte et donc que le facteur alimentaire est limitant. Les implications de ce mécanisme de régulation sur la dynamique des populations nourrice

Changes in the productivity of US West Coast fish stocks

The California Current ecosystem is highly dynamic at interannual to interdecadal time scales. Variability has been documented in pelagic and other fish species, but climate change may be altering the historical models of variation. This study investigates changes in productivity of 31 fish stocks in the California Current ecosystem. Productivity was measured from recent stock assessments, as the per-capita recruitment rate, estimated with a dynamic stock-recruitment model. Contrary to expectations, the dynamic stock-recruitment model fit better than the corresponding stationary model for only seven of the 31 stocks. There was little evidence of linear drift in productivity that might be expected to result from climate change. Climate variables improved forecast accuracy for a few stocks, but there was no common climate signal in productivity. One explanation of these results is that most of the west coast stocks are above their biomass levels for maximum sustainable yield, making them less susceptible to environmental variation. On the other hand, the dynamic recruitment models improved short-term forecasts for all stocks, which may be useful for quota setting. Finally, results for the subset of stocks with dynamic recruitment models could be used to establish dynamic biological reference points

Available Benthic Energy Coefficient (ABEC): a generic tool to estimate the food profitability in coastal fish nurseries

The benthic production of prey seems to be one of the main drivers among many environmental factors that influence the quality of fish nurseries and potentially limit their carrying capacity. However, the contribution of food availability in the growth and survival of juveniles is still controversial. The Available Benthic Energy Coefficient (ABEC) aims to assess the trophic profitability of benthic invertebrate prey; this concept reflects the combination of energy richness and availability of prey. A value of the coefficient was associated with each prey species. This value was calculated from the product of 4 components: (1) mass energy, (2) productivity, (3) regeneration, and (4) accessibility. Thus, this coefficient is expressed as a quantity of energy per unit of weight and per year. From this coefficient, it is simple to calculate the annual production of profitable energy for the fish community in a delimited nursery; it only needs information about the biomass of benthic invertebrates via a standard sampling method. This tool appears to be decisive in properly estimating the carrying capacity of such a fish nursery. Prey classifications based on taxonomy or trophic guilds are widely used in predator-prey studies; comparison with a classification based on ABEC highlighted the energetic heterogeneity of these groups. ABEC can also be used as an index of profitable energy, thereby substituting the usual classifications of prey in trophic studies

Influence of food availability on the spatial distribution of juvenile fish within soft sediment nursery habitats

Soft sediments in coastal shallow waters constitute nursery habitats for juveniles of several flatfishes. The quality of a nursery is defined by its capacity to optimize the growth and the survival of juvenile fish. The influence of biotic factors, such as food availability, is poorly studied at the scale of a nursery ground. Whether food availability limits juvenile survival is still uncertain. A spatial approach is used to understand the influence of food availability on the distribution of juvenile fish of various benthic and demersal species in the Bay of Vilaine (France), a productive nursery ground. We quantified the spatial overlap between benthic macro-invertebrates and their predators (juvenile fish) to assess if the latter were spatially covering the most productive areas of the Bay. Three scenarios describing the shapes of the predator–prey spatial relationship were tested to quantify the strength of the relationship and consequently the importance of food availability in determining fish distribution. Our results underline that both food availability and fish densities vary greatly over the nursery ground. When considering small organisational levels (e.g., a single fish species), the predator–prey spatial relationship was not clear, likely because of additional environmental effects not identified here; but at larger organisational level (the whole juvenile fish community), a strong overlap between the fish predators and their prey was identified. The evidence that fish concentrate in sectors with high food availability suggests that either food is the limiting factor in that nursery or/and fish display behavioural responses by optimising their energetic expenditures associated with foraging. Further investigations are needed to test the two hypotheses and to assess the impact of benthic and demersal juvenile fish in the food web of coastal nurseries

Decadal changes in the productivity of New England fish populations

The northwest Atlantic continental shelf is a large ecosystem undergoing rapid environmental changes, which are expected to modify the productivity of natural marine resources. Current management of most fished species assumes stationary production relationships or time-invariant recruitment rates. With linear state-space models, we examined the evidence of dynamic productivity for 25 stocks of the Northeast US shelf. We expanded the suite of options available within the state-space approach to produce robust estimates. Fifteen of the stocks exhibited time-varying productivity or changes in their maximum reproductive rate. Few productivity time series are related across the whole region, though adjacent stocks of the same species exhibited similar trends. Some links to region-wide environmental variables were observed. We demonstrate that fish recruitment can often be better predicted over a short-term horizon by accounting for dynamic productivity, which could be valuable for fisheries management. Improving predictions by incorporating environmental covariates or covariance among the stocks must be considered case by case and with caution, as their relationships may change over time.The accepted manuscript in pdf format is listed with the files at the bottom of this page. The presentation of the authors' names and (or) special characters in the title of the manuscript may differ slightly between what is listed on this page and what is listed in the pdf file of the accepted manuscript; that in the pdf file of the accepted manuscript is what was submitted by the author

Food limitation of juvenile marine fish in a coastal and estuarine nursery

International audienceDespite their importance for species conservation and sound management of exploited living resources, the density-dependent mechanisms that regulate wild populations are among the least understood process in ecology. In many marine fish species, there is strong evidence that regulation occurs at the juvenile stage, when individuals concentrate in spatially restricted nurseries. However, little is known about the underlying mechanisms. Whether competition for food resources determines fish growth and survival is particularly controversial. We investigated whether food supply may have limited juvenile fish production (integrating both growth and survival) in a coastal and estuarine nursery in western Europe. Using a recent bioenergetics-based approach, we calculated annual macrobenthic food production (FP) and annual food consumption (FC) by juvenile fish and predatory invertebrates for three consecutive years (2008–2010). We also calculated exploitation efficiency (FC:FP) and used it as an index of food limitation. Results revealed substantial interannual variations in FP (FP ~2–3 times higher in 2008 and 2010 than in 2009). FC by young-of-the-year fish followed a fairly similar pattern. In addition, predatory invertebrates consumed as much food as juvenile fish, highlighting the need to consider all dominant epibenthic predators when estimating the overall predation pressure on macrobenthic prey. Lastly, exploitation efficiency of the entire epibenthic predator community reached ~30% in 2009, which is relatively high despite the conservative modeling approach. Overall, these results suggest that food supply may have limited juvenile fish production during the study period, at least in 2009. Nonetheless, further studies based on longer time-series and/or other study sites are required to strengthen these findings

Novel approach for testing the food limitation hypothesis in estuarine and coastal fish nurseries

Survival success of early life stages of fish is regulated by density-dependent effects, but the limiting factors explaining these effects have not been well identified. In coastal habitats, juveniles of many fish species occur in high concentrations and possibly compete for food resources. This study compared the ratio of food consumption to benthic prey production to test whether food availability is a major factor defining the carrying capacity of fish nurseries. We developed a method to quantify the exploitation efficiency (also called ecotrophic efficiency) of the juvenile fish feeding on benthic prey, expressed as a ratio of food consumption to food production. This method includes many sources of uncertainty and a key parameter of prey accessibility. Applied to the case study of the Bay of Vilaine (north Bay of Biscay), results suggest food limitation for juvenile fish. The generic nature of this method supports its wide application in various nursery habitats. As food limitation is a density-dependent process having a dampening effect on recruitment variability, examining its occurrence over time and space will improve our comprehension of nursery-dependent fish dynamics