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Individual metabolism generally scales with body mass with an exponent around 3/4. From dimensional arguments it follows that maximum population growth rate (rmax) scales with a −1/4 exponent. However, the dimensional argument implicitly assumes that offspring size is proportional to adult size. Here we calculate rmax from metabolic scaling at the level of individuals within size-structured populations while explicitly accounting for offspring size. We identify four general patterns of how rmax scales with adult mass based on four empirical life-history patterns employed by groups of species. These life-history patterns are determined by how traits of somatic growth rate and/or offspring mass relate to adult mass. One life-history pattern – constant adult:offspring mass ratio and somatic growth rate independent of adult mass – leads to the classic −1/4 scaling of rmax. The other three life-history patterns lead either to non-metabolic population growth scaling with adult mass or do not follow a power-law relationship at all. Using life-history data of five marine taxa and terrestrial mammals, we identify species groups that belong to one of each case. We predict that elasmobranchs, copepods, and mammals follow standard −1/4 power-law scaling, whereas teleost fish and bivalves do not have a pure power-law scaling. Our work highlights how taxa may deviate from the classic −1/4 metabolic scaling pattern of maximum population growth. The approach is generic and can be applied to any taxa.

Fisheries have reduced the abundances of large piscivores—such as gadids (cod, pollock, etc.) and tunas—in ecosystems around the world. Fisheries also target smaller species—such as herring, capelin, and sprat—that are important parts of the piscivores’ diets. It has been suggested that harvesting of these so-called forage fish will harm piscivores. Multispecies models used for fisheries assessments typically ignore important facets of fish community dynamics, such as individual-level bioenergetics and/or size structure. We test the effects of fishing for both forage fish and piscivores using a dynamic, multitrophic, size-structured, bioenergetics model of the Baltic Sea. In addition, we analyze historical patterns in piscivore-biomass declines and fishing mortalities of piscivores and forage fish using global fish-stock assessment data. Our community-dynamics model shows that piscivores benefit from harvesting of their forage fish when piscivore fishing mortality is high. With substantial harvesting of forage fish, the piscivores can withstand higher fishing mortality. On the other hand, when piscivore fishing mortality is low, piscivore biomass decreases with more fishing of the forage fish. In accordance with these predictions, our statistical analysis of global fisheries data shows a positive interaction between the fishing mortalities of forage-fish stocks and piscivore stocks on the strength of piscivore-biomass declines. While overfishing of forage fish must be prevented, our study shows that reducing fishing pressures on forage fish may have unwanted negative side effects on piscivores. In some cases, decreasing forage-fish exploitation could cause declines, or even collapses, of piscivore stocks. acceptedVersion

Bottom trawl fishing is a controversial activity. It yields about a quarter of the world's wild seafood, but also has impacts on the marine environment. Recent advances have quantified and improved understanding of large‐scale impacts of trawling on the seabed. However, such information needs to be coupled with distributions of benthic invertebrates (benthos) to assess whether these populations are being sustained under current trawling regimes. This study collated data from 13 diverse regions of the globe spanning four continents. Within each region, we combined trawl intensity distributions and predicted abundance distributions of benthos groups with impact and recovery parameters for taxonomic classes in a risk assessment model to estimate benthos status. The exposure of 220 predicted benthos‐group distributions to trawling intensity (as swept area ratio) ranged between 0% and 210% (mean = 37%) of abundance. However, benthos status, an indicator of the depleted abundance under chronic trawling pressure as a proportion of untrawled state, ranged between 0.86 and 1 (mean = 0.99), with 78% of benthos groups > 0.95. Mean benthos status was lowest in regions of Europe and Africa, and for taxonomic classes Bivalvia and Gastropoda. Our results demonstrate that while spatial overlap studies can help infer general patterns of potential risk, actual risks cannot be evaluated without using an assessment model that incorporates trawl impact and recovery metrics. These quantitative outputs are essential for sustainability assessments, and together with reference points and thresholds, can help managers ensure use of the marine environment is sustainable under the ecosystem approach to management., T. Mazor was supported during her research by a CSIRO Postdoctoral Fellowship and CSIRO Ruby Payne‐Scott Award.

Author(s): Petrik, Colleen M; Stock, Charles A; Andersen, Ken H; van Denderen, P Daniel; Watson, James R | Abstract: Global climate change is expected to impact ocean ecosystems through increases in temperature, decreases in pH and oxygen, increased stratification, with subsequent declines in primary productivity. These impacts propagate through the food chain leading to amplified effects on secondary producers and higher trophic levels. Similarly, climate change may disproportionately affect different species, with impacts depending on their ecological niche. To investigate how global environmental change will alter fish assemblages and productivity, we used a spatially explicit mechanistic model of the three main fish functional types reflected in fisheries catches (FEISTY) coupled to an Earth system model (GFDL-ESM2M) to make projections out to 2100. We additionally explored the sensitivity of projections to uncertainties in widely used metabolic allometries and their temperature dependence. When integrated globally, the biomass and production of all types of fish decreased under a high emissions scenario (RCP 8.5) compared to mean contemporary conditions. Projections also revealed strong increases in the ratio of pelagic zooplankton production to benthic production, a dominant driver of the abundance of large pelagic fish vs. demersal fish under historical conditions. Increases in this ratio led to a “pelagification” of ecosystems exemplified by shifts from benthic-based food webs toward pelagic-based ones. The resulting pelagic systems, however, were dominated by forage fish, as large pelagic fish suffered from increasing metabolic demands in a warming ocean and from declines in zooplankton productivity that were amplified at higher trophic levels. Patterns of relative change between functional types were robust to uncertainty in metabolic allometries and temperature dependence, though projections of the large pelagic fish had the greatest uncertainty. The same accumulation of trophic impacts that underlies the amplification of productivity trends at higher trophic levels propagates to the projection spread, creating an acutely uncertain future for the ocean’s largest predatory fish.

A recent meta-analysis by Barneche et al. (2018; Science, 360(6389): 642–645) shows that fish reproductive output scales hypergeometrically with female weight. This result challenges the common assumption that reproductive output is proportional to weight. The implication made is that current theory and practice severely underestimate the importance of larger females for population replenishment. Their example for cod (Gadus morhua) shows that current practice makes an error of 149%. By properly accounting for fish demography, we show that the error is maximally on the order of 10% and in most other fish stocks likely much less.

Large teleost (bony) fish are a dominant group of predators in the oceans and constitute a major source of food and livelihood for humans. These species differ markedly in morphology and feeding habits across oceanic regions; large pelagic species such as tunas and billfish typically occur in the tropics, whereas demersal species of gadoids and flatfish dominate boreal and temperate regions. Despite their importance for fisheries and the structuring of marine ecosystems, the underlying factors determining the global distribution and productivity of these two groups of teleost predators are poorly known. Here, we show how latitudinal differences in predatory fish can essentially be explained by the inflow of energy at the base of the pelagic and benthic food chain. A low productive benthic energy pathway favours large pelagic species, whereas equal productivities support large demersal generalists that outcompete the pelagic specialists. Our findings demonstrate the vulnerability of large teleost predators to ecosystem-wide changes in energy flows and hence provide key insight to predict the responses of these important marine resources under global change.

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.

Large-scale spatial heterogeneity in fisheries production is predominantly controlled by the availability of zooplankton and benthic organisms, which have a complex relationship with primary production. To investigate how cross-ecosystem differences in these drivers determine fish assemblages and productivity, we constructed a spatially explicit mechanistic model of three fish functional types: forage, large pelagic, and demersal fishes. The model is based on allometric scaling principles, includes basic life cycle transitions, and has trophic interactions between the fishes and with their pelagic and benthic food resources. The model was applied to the global ocean, with plankton food web estimates and ocean conditions from a high-resolution earth system model. Further, a simple representation of fishing was included, and led to moderate matches with total, large pelagic, and demersal catches, including re-creation of observed variations in fish catch spanning two orders of magnitude. Our results highlight several ecologically meaningful model sensitivities. First, coexistence between forage and large pelagic fish in productive regions occurred when forage fish survival is promoted via both favorable metabolic allometry and enhanced predator avoidance in adult forage fish. Second, the prominence of demersal fish is highly sensitive to the efficiency of energy transfer to benthic invertebrates. Third, the latitudinal distribution of the total catch is modulated by the temperature dependence of metabolic rates, with increased sensitivity pushing fish biomass toward the poles. Fourth, forage fish biomass is suppressed by strong top-down controls on temperate and subpolar shelves, where mixed assemblages of large pelagic and demersal fishes exerted high predation rates. Last, spatial differences in the dominance of large pelagics vs. demersals is strongly related to the ratio of pelagic zooplankton production to benthic production. We discuss the potential linkages between model misfits and unresolved processes including movement, spawning phenology, seabird and marine mammal predators, and socioeconomically driven fishing pressure, which are identified as priorities for future model development. Ultimately, the model and analyses herein are intended as a baseline for a robust, mechanistic tool to understand, quantify, and predict global fish biomass and yield, now and in a future dominated by climate change and improved fishing technology.

The relationship between biodiversity and ecosystem functioning (BEF) is a topic of considerable interest to scientists and managers because a better understanding of its underlying mechanisms may help us mitigate the consequences of biodiversity loss on ecosystems. Our current knowledge of BEF relies heavily on theoretical and experimental studies, typically conducted on a narrow range of spatio-temporal scales, environmental conditions, and trophic levels. Hence, whether a relationship holds in the natural environment is poorly understood, especially in exploited marine ecosystems. Using large-scale observations of marine fish communities, we applied a structural equation modelling framework to investigate the existence and significance of BEF relationships across northwestern European seas. We find that ecosystem functioning, here represented by spatial patterns in total fish biomass, is unrelated to species richness—the most commonly used diversity metric in BEF studies. Instead, community evenness, differences in species composition, and abiotic variables are significant drivers. In particular, we find that high fish biomass is associated with fish assemblages dominated by a few generalist species of a high trophic level, who are able to exploit both the benthic and pelagic energy pathway. Our study provides a better understanding of the mechanisms behind marine ecosystem functioning and allows for the integration of biodiversity into management considerations.

Bottom fishing such as trawling and dredging may pose serious risks to the seabed and benthic habitats, calling for a quantitative assessment method to evaluate the impact and guide management to develop mitigation measures. We provide a method to estimate the sensitivity of benthic habitats based on the longevity composition of the invertebrate community. We hypothesize that long‐lived species are more sensitive to trawling mortality due to their lower pace of life (i.e., slower growth, late maturation). We analyze data from box‐core and grab samples taken from 401 stations in the English Channel and southern North Sea to estimate the habitat‐specific longevity composition of the benthic invertebrate community and of specific functional groups (i.e., suspension feeders and bioturbators), and examine how bottom trawling affects the longevity biomass composition. The longevity biomass composition differed between habitats governed by differences in sediment composition (gravel and mud content) and tidal bed‐shear stress. The biomass proportion of long‐lived species increased with gravel content and decreased with mud content and shear stress. Bioturbators had a higher median longevity than suspension feeders. Trawling, in particular by gears that penetrate the seabed >2 cm, shifted the community toward shorter‐lived species. Changes from bottom trawling were highest in habitats with many long‐lived species (hence increasing with gravel content, decreasing with mud content). Benthic communities in high shear stress habitats were less affected by bottom trawling. Using these relationships, we predicted the sensitivity of the benthic community from bottom trawling impact at large spatial scale (the North Sea). We derived different benthic sensitivity metrics that provide a basis to estimate indicators of trawling impact on a continuous scale for the total community and specific functional groups. In combination with high resolution data of trawling pressure, our approach can be used to monitor and assess trawling impact and seabed status at the scale of the region or broadscale habitat and to compare the environmental impact of bottom‐contacting fishing gears across fisheries.

Trait-based approaches enable comparison of community composition across multiple organism groups. Yet, little is known about the degree to which empirical trait responses found for one taxonomic group can be generalized across organisms. In this study, we investigated the spatial variability ofmarine community-weighted mean traits and compared their environmental responses across multiple taxa and habitats, including pelagic zooplankton (copepods), demersal fish, and benthic infaunal invertebrates. We used extensive, spatially explicit datasets collected from scientific surveys in the North Sea and examined community composition of these groups using a trait-based approach. In order to cover the key biological characteristics of an organism, we considered three life-history traits (adult size, offspring size,and fecundity) and taxon-specific feeding traits. While many of the traits co-varied in space and notably demonstrated a south–north gradient, none of the traits showed a consistent spatial distribution across all groups. However, traits are often correlated as a result of trade-offs. When studying spatial patterns ofmultiple traits variability in fish and copepods, we showed a high spatial correlation. This also applied to a lesser extent to fish and benthic infauna, whereas no correlation was found between benthic infauna and copepods. The result suggested a decoupling in the community traits between strictly benthic and strictly pelagic species. The strongest drivers of spatial variability for many community traits are the gradients in temperature seasonality, primary productivity, fishing effort, and depth. Spatial variability in benthic traitsalso co-varied with descriptors of the seabed habitat. Overall, results showed that trait responses to environmental gradients cannot be generalized across organism groups, pointing toward potential complex responses of multi-taxa communities to environmental changes and highlighting the need for cross-habitat multi-trait analyses to foresee how environmental change will affect community structure and biodiversity at large

Marine protected areas (MPAs) are widely used to protect exploited fish species as well as to conserve marine habitats and their biodiversity. They have become a popular management tool also for bottom trawl fisheries, a common fishing technique on continental shelves worldwide. The effects of bottom trawling go far beyond the impact on target species, as trawls also affect other components of the benthic ecosystem and the seabed itself. This means that for bottom trawl fisheries, MPAs can potentially be used not only to conserve target species but also to reduce impact of these side-effects of the fishery. However, predicting the protective effects of MPAs is complicated because the side-effects of trawling potentially alter the food-web interactions between target and non-target species. These changes in predatory and competitive interactions among fish and benthic invertebrates may have important ramifications for MPAs as tools to manage or mitigate the effects of bottom trawling. Yet, in current theory regarding the functioning of MPAs in relation to bottom trawl fisheries, such predatory and competitive interactions between species are generally not taken into account. In this paper, we discuss how food-web interactions that are potentially affected by bottom trawling may alter the effectiveness of MPAs to protect (i) biodiversity and marine habitats, (ii) fish populations, (iii) fisheries yield and (iv) trophic structure of the community. We make the case that in order to be applicable for bottom trawl fisheries, guidelines for the implementation of MPAs must consider their potential food-web effects, at the risk of failing management

Understanding trawling impacts on the benthic ecosystem depends to a large extent on the ability to estimate trawling activity at the appropriate scale. Several studies have assessed trawling at fine spatial scales yearly, largely ignoring temporal patterns. In this study, we analysed these temporal patterns in beam trawl effort intensity at 90 stations of the Dutch continental shelf of the North Sea for a period of 10 years, at a fine temporal (weekly) and spatial (110 × 70 m) scale using Vessel Monitoring by Satellite (VMS) data. Our results show that trawling is aggregated in time and shows clear seasonality, related to the behavior of the fleet and migration patterns of the target fish species. The temporal patterns affect the overall impact on and the recovery of the benthic community, as is illustrated with a benthic population model. Our results imply that trawling impact studies using high-resolution data like VMS should take account of the possibility of temporal aggregation and seasonality in trawling to improve the assessment of the impact of trawling on the population dynamics of benthos.

Bottom trawls are a globally used fishing gear that physically disturb the seabed and kill non-target organisms, including those that are food for the targeted fish species. There are indications that ensuing changes to the benthic invertebrate community may increase the availability of food and promote growth and even fisheries yield of target fish species. If and how this occurs is the subject of ongoing debate, with evidence both in favour and against. We model the effects of trawling on a simple ecosystem of benthivorous fish and two food populations (benthos), susceptible and resistant to trawling. We show that the ecosystem response to trawling depends on whether the abundance of benthos is top-down or bottom-up controlled. Fishing may result in higher fish abundance, higher (maximum sustainable) yield and increased persistence of fish when the benthos which is the best-quality fish food is also more resistant to trawling. These positive effects occur in bottom-up controlled systems and systems with limited impact of fish feeding on benthos, resembling bottom-up control. Fishing leads to lower yields and fish persistence in all configurations where susceptible benthos are more profitable prey. Our results highlight the importance of mechanistic ecosystem knowledge as a requirement for successful management.

Size-specific competition and predation interactions often link the population dynamics of fish species in their response to exploitation. The effects of harvesting on interacting fish species is of increasing relevance as more and more fish populations worldwide are reduced by fishing. When stocks are harvested, effects of harvesting may percolate to populations of other species with which it interacts through competition, predation, etcetera. When multiple species are exploited, this can lead to interactions between fisheries, mediated by ecological interactions. Nevertheless, most fish stocks are managed using a single-species framework. We studied how single-species explanations of historical population dynamics work out when size-based interactions between harvested species are taken into account. We have taken as a case study the dynamics of cod (Gadus morhua) and herring (Clupea harengus) in the North Sea. These dynamics are generally considered to be shaped by fishing pressure on and food availability to single species. Our results indicate that the explanatory power of these factors is maintained with the inclusion of species interactions, but the processes leading to the observed patterns are altered as the fates of the species are interdependent. The sign and magnitude of the interaction between the species depends on the state of the populations, their exploitation history and environmental factors such as resource productivity. This context-dependent response to changing fishery intensity has important ramifications for management. We show that management plans for the exploitation of either one of these species, or for the recovery of North Sea cod, which do not account for these subtle interactions, may fail or backfire. Hence, such interactions link the fate of these species in complex ways, which must be taken into consideration for successful management of their exploitation, including harvesting at maximum sustainable yield, as we move towards an ecosystem-based management of marine fisheries.

Aim: Understanding how fish food webs emerge from planktonic and benthic production that sustain them is an important challenge for predicting fisheries production under climate change and quantifying the role of fish in carbon and nutrient cycling. We examine if a trait-based fish community model using the fish traits of maximum body weight and vertical habitat strategy can meet this challenge by globally representing fish food web diversity.Location: Global oceansTime period: Predictions are representative of the early 1990sMajor taxa studied: Marine teleost fishMethod: We present a size- and trait-based fish community model that explicitly resolves the dependence of fish on pelagic and benthic energy pathways to globally predict fish food web biogeography. The emergent food web structures are compared with regionally-calibrated models in three different ecosystem types and used to estimate two fish ecosystem functions: potential fisheries production and benthic-pelagic coupling.Results: Variations in pelagic-benthic energy pathways and seafloor depth drive the emergent biogeography of fish food webs from shelf systems to the open ocean, and across the global ocean. Most shelf regions have high benthic production, which favors demersal fish that feed on pelagic and benthic pathways. Continental slopes also show a coupling of benthic and pelagic pathways, sustained through vertically migrating and interacting mesopelagic and deep-sea demersal fish. Open ocean fish communities are primarily structured around the pelagic pathway. Global model results compare favorably with data-driven regional food web models, suggesting that maximum weight and vertical behavior can capture large-scale variations in food web structure.Main conclusion: Mechanistically linking ocean productivity with upper trophic levels using a size- and trait-based fish community model results in spatial variations in food web structure. Energy pathways vary with ocean productivity and seabed depth, thereby shaping the dominant traits and fish communities across ocean biomes.

All additional text, figures, and tables cited in the main analysis are present in this document

All additional text, figures, and tables cited in the main analysis are present in this document