Production efficiency may differ considerably among individuals and populations. Here we theoretically analyze to what extent differences in efficiency among populations relate to basic physiological processes. We derive characteristics for a stationary population of constant size using (1) an extended Bertalanffy growth equation and (2) Dynamic Energy Budget (DEB) theory to describe individual energetics and mortality combined with an additional thinning rule to provide a more realistic description of survival than the one usually made by only assuming constant background mortality. The production-assimilation ratio as predicted by DEB theory appeared to be strongly correlated with the Bertalanffy growth coefficient and the ultimate reproduction rate. Birds and to a lesser extent mammals have a much lower ratio than other taxa. Benthic marine invertebrate grazers, which are mainly molluscs, showed a higher ratio than pelagic grazers, mostly arthropods, which may have consequences for overall ecological efficiency when marine coastal systems get more dominated by the benthic compartment as a result of human impacts such as the construction of windfarms.

Shallow coastal zones are intensely used by humans but simultaneously are biodiversity hotspots, with a crucial role in the life cycle of many marine species. The competition for food or space between humans and protected wildlife intensifies under pressure of an increased need for marine resources for human consumption. For successful management it is important to establish the key components driving such conflicts of interest. Here we focus on the protected common scoter (Melanitta nigra), a sea duck wintering in coastal habitats that are rich in food, but also among the most disturbed marine systems worldwide. Due to the scoters' shyness disturbance impacts the birds' ability to forage and poses a conflict for balancing bird conservation and economics, including a fishery on its main bivalve prey Spisula subtruncata. In this study, we use an energy budget model to quantify the consequences of depth, currents and disturbance on scoter energetics and carrying capacity. Energetics were described using physical parameters and field data on food availability and disturbance. Results reveal non-linear relationships and a threshold value for when a scoter can no longer maintain its energy balance. This is caused by limited foraging time, rather than food availability. From a conservation perspective, this implies that a precautionary principle should be used, because there will be no warning when an area becomes unsuitable. In addition, the model was applied to study the effects of disturbance from different kinds of shipping in a coastal area of the North Sea, north of The Netherlands. Cargo shipping has the largest impact on the carrying capacity, where there is spatial overlap of prey and an intensively used shipping lane. In other prey distribution situations shrimp vessels may cause most disturbance. Spisula-Ensis fisheries did not limit the potential carrying capacity due to the limited catches and number of fishing trips. Scoter protection should be aimed at flexible spatial management and on only those vessel types above a Spisula bed with a large number of trips, and above all should work from a precautionary principle given the critical thresholds for scoter presence.

A minimum stoichiometric carbon and nitrogen model of an entire ecosystem based on Dynamic Energy Budget (DEB) theory is presented. The ecosystem contains nutrients, producers, consumers, decomposers and detritus. All three living groups consist of somatic structure and either one (consumers and decomposers) or two (producers) reserve compartments, hence the living matter is described by seven state variables. Four types of detritus are distinguished. As the system is closed for matter, the dynamics of the nutrients carbon dioxide and ammonium follow automatically from the dynamics of the other 11 state variables. All DEB organisms in the model are V1-morphs, which means that surface area of each organism is proportional to volume. The resulting ontogenetic symmetry implies that complicated modelling of size structure is not required. The DEB V1-morph model is explained in detail, and the same holds for the idea of synthesizing units, which plays a key role in DEB modelling. First results of system dynamics are presented.

This study shows the effect of climate change on the growth and survival of early life history stages of common sole (Solea solea) in different nursery areas of the North Sea, by combining a larval transport model with an individual-based growth model (Dynamic Energy Budget) to assess the fate from egg to young of the year at the end of the first growth season. Three scenarios of climate change, inspired by the 2040 Intergovernmental Panel on Climate Change projections, are tested and results are compared to a reference situation representative of current climate conditions. Under climate change scenarios where wind changes, water temperature increases and earlier spawning are considered, the early arrival of fish larvae in their nurseries results in larger young of the year at the end of summer. However, early arrival leads to higher mortality due to initially slow growth in spring. Future climate scenarios result in higher biomass and reduced first-year survival. How this result translates into changes at population level and stock management needs further investigation. Nonetheless, this study illustrates that processes linking life stages are paramount to understand and predict possible consequences of future climate conditions on population dynamics.

The aphorism, ‘All models are wrong, but some models are useful’, originally referred to statistical models, but is now used for scientific models in general. When presenting results from a marine simulation model, this statement effectively stops discussions about the quality of the model, as there is always another observation to mismatch, and thereby another confirmation why the model cannot be trusted. It is common that observations are less challenged and are often viewed as a ‘gold standard’ for judging models, whereas proper interpretations and the true value of models are often overlooked. Models are not perfect, and there are many examples where models are used improperly to provide misleading answers with great confidence, but to what extent does an observation represent the truth? The precision of the observational gear may be high, but what about representativeness? The interpretation of observations is simply another model, but this time not coded in a computer language but rather formed by the individual observer. We submit that it would be more productive to initiate a process where the norm is that models and observations are joined to strengthen both. In the end, neither method is the goal, but both are useful tools for disclosing the truth. Biased views on either observational or modeling approaches would limit us from achieving this goal.

Different fish species and life stages depend not only on food abundance, but also on the size of planktonic prey, and (mis-)matches in time and space with suitable prey may influence the growth and survival of fish during their lifetime. We explored the sensitivity of a fish community to spatial-temporal differences in plankton prey fields. Data from 5 different lower trophic level models in the North Sea (Delft3D-WAQ, ECOHAM, ECOSMO, HBM-ERGOM and NORWE-COM) were used to force the food web model OSMOSE which simulates spatially and temporally explicit higher trophic level fish dynamics. The estimated fish biomass levels were clearly and positively linked to zooplankton biomass, and sensitivity studies varying zooplankton biomass revealed that spatial and temporal variation in zooplankton drives the differences in absolute fish biomass. More zooplankton size bins resulted in less fish biomass due to size-based foraging constraints (i.e. a smaller proportion of bins falls within the prey size range of a fish, resulting in a decrease in available food). Nevertheless, we found a consistent response across models in the relative biomass contribution and spatial patterns of selected fish groups, indicating low sensitivity of the composition of the simulated fish community to the zooplankton input. The robustness of the outcome will aid model acceptance and implementation into management action. Relative, not absolute, changes in primary and secondary production may therefore be used to study the effects of management scenarios on fish communities.

This study investigates the effect of water temperature on the development rate of eggs and larvae, the duration of the endogenous feeding period and its consequences for recruitment of smelt (Osmerus eperlanus) in Dutch lakes IJsselmeer and Markermeer. This study measured temperature-dependent egg and larval development rates as well as mortality rates from fertilization till the moment of absorption of the yolk-sac and from yolk-sac depletion onwards in temperature-controlled indoor experiments. Using multinomial modelling the authors found significant differences in development time of egg development stages under different temperature regimes. Based on historic water temperatures, the model predicted that the larval endogenous feeding period has advanced at a rate of about 2.9 days per decade in a more than 50 year period since 1961, yet there was no change in the duration of the endogenous feeding period. As zooplankton is more responsive to daylight than water temperature cues, a mismatch between the peak of the onset of exogenous feeding of smelt and the peak of zooplankton blooms could lead to high mortality and therefore low recruitment of smelt. Such a mismatch might contribute to a decline in the smelt population in Lake IJsselmeer and Lake Markermeer.

We reviewed the use of survey-based pre-recruit abundance indices in short-term recruitment forecasts for fish species relying on coastal habitats at the juvenile stage and that are assessed by ICES. We collated information from stock assessment reports and from a questionnaire filled out by the stock assessors. Among the 78 stocks with juvenile coastal dependence, 49 use short-term forecasts in stock assessment. Survey-based pre-recruit abundance indices were available for 35 of these stocks, but only 14 were used to forecast recruitment. The questionnaire indicated that the limited use of survey-based pre-recruit abundance indices was primarily due to sampling inefficiency, which may preclude reliable recruitment estimates. The sampling is inefficient because the juvenile coastal distribution is outside the geographical area covered by large-scale surveys or targeted coastal surveys are conducted on limited spatial and temporal scales. However, our analysis of the relationship between survey-based pre-recruit indices and assessment-generated recruitment indices revealed that survey-based pre-recruit abundance indices were sufficiently accurate to provide useful information for predicting future recruitment. We recommend expansion of the use of survey-based indices of pre-recruit abundance in stock assessment and recruitment forecasting, and consideration of how to include juveniles in ongoing and future surveys.

The pulse trawl is presented as an alternative for the traditional tickler chain beam trawl. The beam trawl has a direct lethal effect on the benthic community, which is much less for the pulse trawl. However, the pulse trawl, due to its electrical field has non-lethal effects on the benthic community. The existence of such non-lethal effects have been demonstrated for different organisms and most often involve a short period of inactivity. Such a period of inactivity may have consequences at the population level as it may limit feeding. Yet the population dynamical consequences, nor food web level consequences, have been studied in experimental or field settings. To study non-lethal and lethal effects of trawling on the benthic community a model was developed which is based on a trait-based characterization of the food web. This model describes the equilibrium dynamics of predators, filter feeders and deposit feeders. The model includes direct mortality as a function of fishing mortality and a non-lethal effect, which was incorporated as a decrease of the maximum food intake rate. This approach was based on experiments with electric pulse and benthic invertebrates. We conducted a comparative analysis of lethal and non-lethal impact of trawling on the equilibrium dynamics of the benthic food web. Only lethal effects represent a tickler chain beam trawl and a combination of lethal and non-lethal effect represents a pulse trawl. We first studied the effects of lethal and non-lethal impacts on the food web separately. We then studied the trade-off between lethal and non-lethal effects for a given total impact on the community. This approach shows how much non-lethal impact is required to compensate for a decrease in lethal effects of trawling. The study shows that lethal effects need to be replaced by non-lethal effects with similar strength in order to maintain the effect of fishing intensity on the food web. Such strong non-lethal effects are unlikely to occur in the field based on experiments assessing the impact of electricity on benthic macro-invertebrates and trawling intensity. In other words, the combined non-lethal and lethal effects of the pulse trawl are unlikely to have a similar effect on the macro-invertebrate benthic community compared to the lethal effects of the traditional trawl.

Bottom trawl fisheries have significant effects on benthic habitats and communities, and these effects have been studied intensively in the last decades. Most of these studies have related the changes in benthic community composition to direct effect of trawl gears on benthos, through imposed mortality. This line of argumentation ignores the fact that benthic organisms themselves form a complex food web and that bottom trawling may trigger secondary effects through this food web. We studied the potential consequences of such food web effects using a model of benthic predators, filter feeders, deposit feeders and fish. Our analysis shows how inclusion of ecological interactions complicates the relationship between bottom trawling intensity and the state of the benthic community and causes a non-linear and non-monotonic response of the benthic community to trawling. This shows that indirect food web effects can fundamentally alter the response of a benthic ecosystem to bottom trawling, compared to the direct effects of mortality. In light of our results, we argue that indicators of fishing impact on benthos need to account for positive as well as negative effects of bottom trawling, in order to accurately quantify the impact. Our findings highlight that understanding the food web ecology of the benthic ecosystem is crucial for understanding and predicting the effects of trawling on the seafloor. Work that promotes such understanding of the food web ecology seems a more productive research strategy than conducting ever more empirical trawling effect measurements.

This desk study describes the biology of the European lobster H. gammarus.Using the obtained data a model was developed to describe the growth of the European lobster under assumed conditions on the anti-scouring of monopiles in Dutch OWFs.One of the main questions to answer was, if, theoretically, local productivity supports the continuous harvesting of lobsters with passive fishery methods.

Coastal habitats (e.g., seagrass beds, shallow mud, and sand flats) strongly influence survival, growth, and reproduction of marine fish and invertebrate species. Many of these species have declined over the past decades, coincident with widespread degradation of coastal habitats, such that an urgent need exists to model the quantitative value of coastal habitats to their population dynamics. For exploited species, demand for habitat considerations will increase as fisheries management contends with habitat issues in stock assessments and management in general moves toward a more ecosystem-based approach. The modeling of habitat function has, to date, been done on a case-by-case basis involving diverse approaches and types of population models, which has made it difficult to generalize about methods for incorporating habitat into population models. In this review, we offer guiding concepts for how habitat effects can be incorporated in population models commonly used to simulate the population dynamics of fish and invertebrate species. Many marine species share a similar life-history strategy as long-lived adults with indeterminate growth, high fecundity, a planktonic larval form, and benthic juveniles and adults using coastal habitats. This suite of life-history traits unites the marine species across the case studies, such that the population models can be adapted for other marine species. We categorize population models based on whether they are static or dynamic representations of population status, and for dynamic, further into unstructured, age/size class structured, and individual-based. We then use examples, with an emphasis on exploited species, to illustrate how habitat has been incorporated, implicitly (correlative) and explicitly (mechanistically), into each of these categories. We describe the methods used and provide details on their implementation and utility to facilitate adaptation of the approaches for other species and systems. We anticipate that our review can serve as a stimulus for more widespread use of population models to quantify the value of coastal habitats, so that their importance can be accurately realized and to facilitate cross-species and cross-system comparisons. Quantitative evaluation of habitat effects in population dynamics will increasingly be needed for traditional stock assessments, ecosystem-based management, conservation of at-risk habitats, and recovery of overexploited stocks that rely on critical coastal habitats during their life cycle.

The present study describes the responses of summer phytoplankton biomass to changes in top-down forcing (expressed as zooplankton mortality) in three ecosystems (the North Sea, the Baltic Sea and the Nordic Seas) across different 3D ecosystem models. In each of the model set-ups, we applied the same changes in the magnitude of mortality ( +/- 20%) of the highest trophic zooplankton level (Z1). Model results showed overall dampened responses of phytoplankton relative to Z1 biomass. Phytoplankton responses varied depending on the food web structure and trophic coupling represented in the models. Hence, a priori model assumptions were found to influence cascades and pathways in model estimates and, thus, become highly relevant when examining ecosystem pressures such as fishing and climate change. Especially, the different roles and parameterizations of additional zooplankton groups grazed by Z1, and their importance for the outcome, emphasized the need for better calibration data. Spatial variability was high within each model indicating that physics (hydrodynamics and temperature) and nutrient dynamics also play vital roles for ecosystem responses to top-down effects. In conclusion, the model comparison indicated that changes in top-down forcing in combination with the modelled food-web structure affect summer phytoplankton biomass and, thereby, indirectly influence water quality of the systems.

Het rapport “Toestand Vis en Visserij in de Zoete Rijkswateren” bestaat uit drie delen. Dit rapport (Deel II) is een achtergronddocument waarin de gebruikte monitoringsmethodieken in de verschillende vismonitoringen in de zoete Rijkswateren in 2016 in detail worden beschreven.

Het aalbeheerplan is geëvalueerd in het licht van de voornoemde “beheerdoelen” uit de Aalverordening. De methodiek die bij deze evaluatie is gehanteerd komt voort uit de ICES aal werkgroep (WGEEL). Hierdoor wordt in deze evaluatie alleen ingegaan op de effectiviteit van maatregelen in relatie tot beheerdoelen opgesteld door de Raad van de Europese Unie. De evaluatie is uitgevoerd door middel van modellen, vangstgegevens, veldwaarnemingen en statistische analyses, en worden uitvoerig beschreven in de rapportage. Het geheel van deze inspanning resulteerde in schattingen van voor (2005-2007) en na (2008-2010, 2011-2013 en 2014-2016) de implementatie van het Aalbeheerplan.

We review and compare four broad categories of spatially-explicit modelling approaches currently used to understand and project changes in the distribution and productivity of living marine resources including: 1) statistical species distribution models, 2) physiology-based, biophysical models of single life stages or the whole life cycle of species, 3) food web models, and 4) end-to-end models. Single pressures are rare and, in the future, models must be able to examine multiple factors affecting living marine resources such as interactions between: i) climate-driven changes in temperature regimes and acidification, ii) reductions in water quality due to eutrophication, iii) the introduction of alien invasive species, and/or iv) (over-)exploitation by fisheries. Statistical (correlative) approaches can be used to detect historical patterns which may not be relevant in the future. Advancing predictive capacity of changes in distribution and productivity of living marine resources requires explicit modelling of biological and physical mechanisms. New formulations are needed which (depending on the question) will need to strive for more realism in ecophysiology and behaviour of individuals, life history strategies of species, as well as trophodynamic interactions occurring at different spatial scales. Coupling existing models (e.g. physical, biological, economic) is one avenue that has proven successful. However, fundamental advancements are needed to address key issues such as the adaptive capacity of species/groups and ecosystems. The continued development of end-to-end models (e.g., physics to fish to human sectors) will be critical if we hope to assess how multiple pressures may interact to cause changes in living marine resources including the ecological and economic costs and trade-offs of different spatial management strategies. Given the strengths and weaknesses of the various types of models reviewed here, confidence in projections of changes in the distribution and productivity of living marine resources will be increased by assessing model structural uncertainty through biological ensemble modelling.

Modern approaches to Ecosystem-Based Management and sustainable use of marine resources must account for the myriad of pressures (interspecies, human and environmental) affecting marine ecosystems. The network of feeding interactions between co-existing species and populations (food webs) are an important aspect of all marine ecosystems and biodiversity. Here we describe and discuss a process to evaluate the selection of operational food-web indicators for use in evaluating marine ecosystem status. This process brought together experts in food-web ecology, marine ecology, and resource management, to identify available indicators that can be used to inform marine management. Standard evaluation criteria (availability and quality of data, conceptual basis, communicability, relevancy to management) were implemented to identify practical food-web indicators ready for operational use and indicators that hold promise for future use in policy and management. The major attributes of the final suite of operational food-web indicators were structure and functioning. Indicators that represent resilience of the marine ecosystem were less developed. Over 60 potential food-web indicators were evaluated and the final selection of operational food-web indicators includes: the primary production required to sustain a fishery, the productivity of seabirds (or charismatic megafauna), zooplankton indicators, primary productivity, integrated trophic indicators, and the biomass of trophic guilds. More efforts should be made to develop thresholds-based reference points for achieving Good Environmental Status. There is also a need for international collaborations to develop indicators that will facilitate management in marine ecosystems used by multiple countries.