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Although zooplankton were extensively studied in the North Sea, knowledge about winter zooplankton assemblages is still scarce, despite potential influence of zooplankton overwintering stocks on seasonal plankton succession and productivity. Furthermore, several economically and ecologically important fish species reproduce during winter contributing to the zooplankton community as passive members (eggs) or predators (larvae). To elucidate on winter zooplankton distribution, abundance and composition in the Southern North Sea and Eastern English Channel, we defined assemblages based on mesozoo- and ichthyoplankton data sampled between January and February 2008 using fuzzy-clustering and indicator species. Mesozoo- and ichthyoplankton (eggs+larvae) were integrated in a common analysis by using a spatial grid adapted to the datasets and defined by means of a geostatistical method developed in agronomics. Potential environmental drivers of assemblage distribution were evaluated by means of GLMM and comparison with data from 2022 facilitated insight about the inter-annual representativeness of the assemblages. Five zooplankton assemblages were found varying with regard to total zooplankton abundance, dominant and indicator taxa. Spatial variability of abiotic (dissolved nutrients, salinity, depth, temperature, organic matter in suspension, chlorophyll a), biotic variables (phyto- and microplankton composition), water masses and fish spawning grounds were revealed as potential drivers of assemblage distribution. Assemblages off the Rhine-Scheldt estuary and in the German Bight harbored the biggest zooplankton overwintering stocks that might influence the grazing pressure on phytoplankton spring production. Assemblages off the Rhine-Scheldt estuary and covering the English Channel and the Southern Bight were found to be of high importance for herring and plaice larvae. Although further analyses suggested inter-annual representativeness of the assemblages found (2008 vs 2022), the assessment of further years would be necessary to account for potential inter-annual variability. Future studies could profit from the assessment of microzooplankton facilitating insight in fish larvae feeding potential and zooplankton overwintering strategies. [ABSTRACT FROM AUTHOR]

Knowledge of the trophic structure and variability of planktonic communities is a key factor in understanding food-web dynamics and energy transfer from zooplankton to higher trophic levels. In this study, we investigated how stable isotopes of mesozooplankton species varied seasonally (winter, spring, autumn) in relation to environmental factors and plankton size classes in a temperate coastal ecosystem. Our results showed that spring is characterized by the strongest vertical and size-structured plankton food-web, mainly fueled by the phytoplankton bloom. As a result, spring displayed the largest isotopic niche space and trophic divergence among species. On the contrary, both pelagic and benthic-derived carbon influenced low productive seasons (winter and autumn), resulting in more generalist strategies (trophic redundancy). Stable isotope mixing models were used to explore how different seasonal structures influenced the overall food web up to predatory plankton (i.e., mysids, chaetognaths, and fish larvae). Different feeding strategies were found in spring, with predators having either a clear preference for larger prey items (> 1 mm, for herring and dab larvae) or a more generalist diet (sprat and dragonets larvae). During low productive seasons, predators seemed to be more opportunistic, feeding on a wide range of size classes but focusing on smaller prey. Overall, the food-web architecture of plankton displayed different seasonal patterns linked to components at the base of the food web that shaped the main energy fluxes, either from phytoplankton or recycled material. Additionally, these patterns extended to carnivorous plankton, such as fish larvae, emphasizing the importance of bottom-up processes. [ABSTRACT FROM AUTHOR]

Among all human-induced pressures, ocean warming is expected to be one of the major drivers of change in marine ecosystems. Fish species are particularly vulnerable during embryogenesis. Here, the impact of temperature was assessed on embryonic stages of Atlantic herring (Clupea harengus), a species of high socio-economic interest, with a particular focus on the under-studied eastern English Channel winter-spawning component (Downs herring). Key traits linked to growth and development were experimentally evaluated at three temperatures (8°C, 10°C and 14°C), from fertilization to hatching, in standardized controlled conditions. Overall negative impacts of increased temperature were observed on fertilization rate, mean egg diameter at eyed stage, hatching rate and yolk sac volume. A faster developmental rate and a change in development stage frequency of newly hatched larvae were also observed at higher temperature. Potential parental effects were detected for four key traits (i.e. fertilization rate, eyed survival rate, mean egg diameter and hatching rate), despite a limited number of families. For instance, a large variability among families was shown in survival rate at eyed stage (between 0 and 63%). Potential relationships between maternal characteristics and embryo traits were therefore explored. We show that a substantial proportion of variance (between 31 and 70%) could be explained by the female attributes considered. More particularly, age, traits linked to life history (i.e. asymptotic average length and Brody growth rate coefficient), condition and length were important predictors of embryonic key traits. Overall, this study constitutes a stepping-stone to investigate potential consequences of warming on Downs herring recruitment and provides first insights on potential parental effects. [ABSTRACT FROM AUTHOR]

Digestive system functionality is a key process linked to larval recruitment and survival. However, little is known about organ development and enzyme maturation of the digestive system of North Sea Atlantic herring (Clupea harengus). In this study, herring larvae were reared at 13 °C from hatching to 69 day post hatch, covering four developmental stages: (1) yolk sac (8–9 mm), (2) pre-flexion (9–14 mm), (3) flexion (12–18 mm) and (4) post-flexion stages (15–30 mm). Combined histological (semi-quantitative scoring) and enzyme analyses (pancreatic and intestinal) showed that developmental stages are strongly linked to physiological changes. The larvae lack a functional stomach and use the intestine as the primary site of digestion which is mainly supported by pancreatic enzyme activity. The intestine acquired adult enzymatic digestive features with a functional brush border at the end of the flexion stage and pyloric ceca started to develop during the post-flexion stage. The transition from pre-flexion to flexion stage and the end of the post-flexion stage are energetically taxing periods as indicated by a reduced number and size of liver vacuoles. Based on these findings, we consider these moments as critical periods, where herring larvae could be dramatically affected by suboptimal feeding conditions in the field. This implies that pre-flexion stage larvae with low or no liver reserves may not be able to proceed to the next developmental stage. Hence, the level of energy storage in first-feeding larvae needs to be examined for its use as a field indicator of survival and development. [ABSTRACT FROM AUTHOR]

Condition indices aim to evaluate the physiological status of fish larvae by estimating both the level of starvation and potential of survival. Histological indices reveal direct effects of starvation whereas biochemical indices such as lipid classes or RNA:DNA ratios are used as proxies of condition, giving information on the amount of energy reserves and growth rate, respectively. We combined these three indices to evaluate ontogenetic variations of growth performance, lipid dynamics and nutritional condition of plaice larvae caught in the field during winter 2017 in the eastern English Channel and the Southern Bight of the North Sea. RNA:DNA ratios showed that larvae at the beginning of metamorphosis (stage 4) had a lower growth rate than younger individuals (stages 2 and 3). A significant increase in the proportion of triglycerides also occurred at stage 4, indicating energy storage. Histological indices indicated that most of the larvae were in good condition, even younger ones with low lipid reserves. There was, however, an increase in the proportion of healthy individuals over ontogeny, especially with respect to liver vacuoles which were larger and more numerous for stage 4 larvae. Combined together, these condition indices revealed the ontogenetic shift in the energy allocation strategy of plaice larvae. Young larvae (stages 2 and 3) primarily allocate energy towards somatic growth. The decrease in growth performance for stage 4 was not related to poor condition, but linked to a higher proportion of energy stored as lipids. Since the quantity of lipid reserves is particularly important for plaice larvae to withstand starvation during metamorphosis, this could be considered as a second critical period after the one of exogenous feeding for larval survival and recruitment success. [ABSTRACT FROM AUTHOR]

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Evaluating fish larval condition in terms of nutrition and growth is essential as it will influence their development and survival capacity. The present study aims to investigate larval condition of Downs herring ( Clupea harengus L.) during winter in the Eastern English Channel and Southern Bight of the North Sea. Four condition indices including ingestion rate based on gut fluorescence, instantaneous growth based on RNA/DNA, DNA/C ratios, and otolith microstructure were combined at an individual scale on herring larvae collected during the 2015 International Bottom Trawl Survey-MIK sampling. The four indices demonstrated a clear shift in the larval condition occurring at a larval size of 13 mm. While smaller larvae were shown to feed and grow, larger larvae exhibited a slower growth rate though actively feeding. This suggests that 13 mm could be a critical size for Downs herring larvae. This ontogenetic shift in the larval condition is discussed regarding environmental conditions, diet shift, and growth strategies. It is concluded that the shift from an omnivorous to a carnivorous diet constitutes an additional critical step besides such as the shift from endogenous to exogenous nutrition. [ABSTRACT FROM AUTHOR]

The invasive ctenophore Mnemiopsis leidyi has been reported in various coastal locations in the southern North Sea in the past years. Since 2009, International Bottom Trawl Surveys have recorded this species each winter in open waters. As this species, well-known for its dramatic disturbance of ecosystems, was expected not to be able to overwinter offshore it is crucial to understand its distribution dynamics. Two modelling methods, a quantile regression and a particle tracking model, were used (1) to identify habitats where the invasive ctenophore M. leidyi could survive the North Sea cold winters and (2) to investigate the dispersal of individuals between these different habitats, emphasizing favorable areas where sustainable populations could have been established. Temperature was found to be the crucial factor controlling the winter distribution of M. leidyi in the North Sea. High abundance predictions in winter were associated with low values of temperature, which characterise south-eastern coastal areas and estuaries influenced by riverine runoff. A retention-based M. leidyi population was indicated along the northern Dutch coast and German Bight and a transport-based population offshore from the western Danish coast. Individuals found in the open waters were transported from southern coasts of the North Sea, thus the open water population densities depend on the flux of offspring from these areas. This study provides the first estimates of the overwinter areas of this invasive species over the cold winters in the North Sea. Based on the agreement of habitat and dispersal model results, we conclude that M. leidyi has become established along south-eastern coasts of the North Sea where the environment conditions allows overwintering and it can be retained for later blooms. [ABSTRACT FROM AUTHOR]

Spatial interactions between saithe (Pollachius virens) and hake (Merluccius merluccius) were investigated in the North Sea. Saithe is a well-established species in the North Sea, while occurrence of the less common hake has recently increased in the area. Spatial dynamics of these two species and their potential spatial interactions were explored using binomial generalized linear models (GLM) applied to the International Bottom Trawl Survey (IBTS) data from 1991 to 2012. Models included different types of variables: (i) abiotic variables including sediment types, temperature, and bathymetry; (ii) biotic variables including potential competitors and potential preys presence; and (iii) spatial variables. The models were reduced and used to predict and map probable habitats of saithe, hake but also, for the first time in the North Sea, the distribution of the spatial overlap between these two species. Changes in distribution patterns of these two species and of their overlap were also investigated by comparing species' presence and overlap probabilities predicted over an early (1991–1996) and a late period (2007–2012). The results show an increase in the probability over time of the overlap between saithe and hake along with an expansion towards the southwest and Scottish waters. These shifts follow trends observed in temperature data and might be indirectly induced by climate changes. Saithe, hake, and their overlap are positively influenced by potential preys and/or competitors, which confirms spatial co-occurrence of the species concerned and leads to the questions of predator–prey relationships and competition. Finally, the present study provides robust predictions concerning the spatial distribution of saithe, hake, and of their overlap in the North Sea, which may be of interest for fishery managers. [ABSTRACT FROM PUBLISHER]

Planque, B., Bellier, E., and Loots, C. 2011. Uncertainties in projecting spatial distributions of marine populations. – ICES Journal of Marine Science, 68: 1045–1050.Projection of future spatial distributions of marine populations is a central issue for ecologists and managers. The measure of projection uncertainty is particularly important, because projections can only be useful if they are given with a known and sufficiently high level of confidence. Uncertainties can arise for the observation process, conceptual and numerical model formulations, parameter estimates, model evaluation, appropriate consideration of spatial and temporal scales, and finally the potential of adaptation of living systems. Comprehensive analyses of these multiple sources of uncertainty have not been carried out so far, and how these uncertainties are considered in current studies has not yet been described. To analyse how these different sources of uncertainty are currently considered in marine research, we did a survey of published literature during the period 2005–2009. From the 75 publications selected, we calculated how frequently each type of uncertainty was considered. We found that little attention is given to most sources of uncertainty, except for uncertainty in parameter estimates. As a result, most current projections are expected to be far less reliable than usually assumed. The conclusion is that, unless uncertainty can be better accounted for, such projections may be of limited use, or even risky to use for management purposes. [ABSTRACT FROM PUBLISHER]

The processes that control the spatial distribution of North Sea whiting ( Merlangius merlangus) spawning adults are investigated using a statistical multi-model approach. Models of external and internal controls on the population, such as environmental conditions, spatial constraints, present or past spatial distribution, and demographic state of the population, are evaluated, compared and ranked to select those that are the best able to predict the observed distribution of spawning adults. Model selection is greatly influenced by the selection method, either based on data fitting or prediction, as well as by the threshold value used to stop the selection. Model selection based on prediction tends to select simpler models than selection based on data fitting. The hypotheses underlying the selected models are inferred to play a significant role in controlling North Sea whiting spatial distribution. The multi-model inference approach developed in this study enables comparison of several theoretical concepts and hypotheses and the results provide important clues on the processes involved in the control of the spatial distribution of whiting. We conclude that whiting has a high spatial fidelity to spawning site which can be linked to either geographical attachment or year-to-year persistence of the spatial distribution of the population. Environmental factors - temperature and salinity - appear to influence the geographical extent of spawning whiting distribution, whereas local abundance levels are primarily controlled by internal factors, i.e., population size and spatial segregation between ages. [ABSTRACT FROM AUTHOR]

Understanding and predicting the distribution of organisms in heterogeneous environments lies at the heart of ecology. The spatial distribution of fish populations observed in the wild results from the complex interactions of multiple controls both external or internal to the fish populations. Whilst species distribution models (SDMs) have been mostly concerned with static description of species distribution as a function of environmental constraints, models of animal movements (MAMs) have focussed on the dynamic nature of spatial distribution of groups of individuals under a number of constraints external and internal to the population. Besides SDMs and MAMs, modelling the spatial distribution of fish populations can be achieved by models that are fundamentally static (like SDMs) but can also incorporate many hypotheses on the control of fish spatial distribution (like MAMs). The hypotheses underlying these models need to make sense at the population level - rather than at the individual or species level -we term these 'population distribution models' (PDMs). PDMs are statistical models that rely on several hypotheses, which include: (i) control through geographical attachment, (ii) environmental conditions, (iii) density-dependent habitat selection, (iv) spatial dependency, (v) population demographic structure, (vi) species interactions and (vii) population memory. We review the basis behind each of these conceptual models and we examine corresponding numerical applications. We argue that the conceptual models are complementary rather than competing, that existing numerical applications are still rarely compared and combined, and that PDMs can offer a statistical framework to achieve this. We recommend that the numerical models associated with different hypotheses be constructed within such a common general framework. This will permit evaluation, comparison and combination of the multiple hypotheses on fish spatial distribution. It will ultimately lead to a more comprehensive understanding of the factors controlling the spatial distribution of fish populations and to more accurate predictions in which model uncertainty is accounted for. [ABSTRACT FROM AUTHOR]

Loots, C., Vaz, S., Planque, B., and Koubbi, P. 2010. What controls the spatial distribution of the North Sea plaice spawning population? Confronting ecological hypotheses through a model selection framework. – ICES Journal of Marine Science, 67: 244–257.The spatial dynamics of spawning fish are crucial because they influence the survival rates of eggs and larvae and ultimately impact the reproductive success of populations. The factors that control these dynamics are complex and potentially many, and they interact. A model-selection-based approach was developed to confront various hypotheses of control of the spatial distribution of spawning population of North Sea plaice (Pleuronectes platessa). For each hypothesis or combination thereof, statistical models were constructed. These were then ranked and selected based on their ability to adjust and predict observed spatial distributions. The North Sea plaice population seems to have developed strong attachment to specific spawning sites, where geographic location and population memory are important controlling factors. Temporal changes in spatial distribution patterns appear to be influenced primarily by population size and demography. Variations in hydrographic conditions such as temperature and salinity do not appear to control interannual fluctuations in spatial distribution. This means that, for reproduction, applying conventional habitat models may falsely attribute major controlling effects to environmental conditions. It is concluded that a multiple-hypothesis approach is essential to understanding and predicting the present and future distribution of the North Sea plaice population during its spawning season. [ABSTRACT FROM PUBLISHER]

Electrona antarctica is one of the most abundant mesopelagic fishes in the oceanic zone surrounding the Kerguelen Archipelago in the Indian sector of the Southern Ocean. Generalized additive models (GAM) combined with geographical information systems (GIS) were used to predict and map the abundance of this species according to three environmental variables: sea surface temperature, bathymetry and surface chlorophyll a. The model was applied on the Antarctic Polar Front in the eastern part of Kerguelen Archipelago. E. antarctica seems to be linked to areas presenting low chlorophyll a concentrations, depths greater than 500 m and temperatures lower than 5°C. The model was then applied to the Kerguelen’s plateau for three different years: 1998, 1999 and 2000. The position of Antarctic Polar Front and the intensity of an upwelling play an important role in the abundance variability of E. antarctica. Furthermore, the model allows the understanding of the habitat of E. antarctica and its trophic place in the pelagic ecosystem. [ABSTRACT FROM AUTHOR]