Are we ready to track climate-driven shifts in marine species across international boundaries? - A global survey of scientific bottom trawl data.

Marine biota are redistributing at a rapid pace in response to climate change and shifting seascapes. While changes in fish populations and community structure threaten the sustainability of fisheries, our capacity to adapt by tracking and projecting marine species remains a challenge due to data discontinuities in biological observations, lack of data availability, and mismatch between data and real species distributions. To assess the extent of this challenge, we review the global status and accessibility of ongoing scientific bottom trawl surveys. In total, we gathered metadata for 283,925 samples from 95 surveys conducted regularly from 2001 to 2019. We identified that 59% of the metadata collected are not publicly available, highlighting that the availability of data is the most important challenge to assess species redistributions under global climate change. Given that the primary purpose of surveys is to provide independent data to inform stock assessment of commercially important populations, we further highlight that single surveys do not cover the full range of the main commercial demersal fish species. An average of 18 surveys is needed to cover at least 50% of species ranges, demonstrating the importance of combining multiple surveys to evaluate species range shifts. We assess the potential for combining surveys to track transboundary species redistributions and show that differences in sampling schemes and inconsistency in sampling can be overcome with spatio-temporal modeling to follow species density redistributions. In light of our global assessment, we establish a framework for improving the management and conservation of transboundary and migrating marine demersal species. We provide directions to improve data availability and encourage countries to share survey data, to assess species vulnerabilities, and to support management adaptation in a time of climate-driven ocean changes.

Biodiversity-ecosystem functioning relationships in fish communities: biomass is related to evenness and the environment, not to species richness.

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.

Growth and food consumption of whiting Merlangius merlangus.

In the western Baltic Sea (WBS), whiting Merlangius merlangus is the main piscivorous fish together with cod Gadus morhua. In the present study, we investigate the growth and food consumption rates of WBS M. merlangus and compare the growth rates of males and females with those of M. merlangus in the North Sea (NS). Food consumption rates are estimated directly from sampled stomach contents in the WBS using a gastric evacuation rate model and indirectly by using a static energy-budget model together with the growth rates. The results indicate that male and female M. merlangus in the WBS have similar feeding and growth strategies, while in the NS M. merlangus show more pronounced differences in food consumption and growth dynamics between the sexes. Female WBS M. merlangus grow significantly slower than their conspecifics in the NS, but there is no significant difference for males. Sexual size dimorphism is seen in both areas, but for M. merlangus in the WBS the difference is less pronounced. Food consumption rates in the WBS differ between seasons, with the lowest food intake in the first 2 quarters of the year and the highest in the 3rd quarter. No differences in consumption rates were seen between males and females, which could be related to the more similar growth pattern seen for M. merlangus in the WBS.

Differences in density-dependence drive dual offspring size strategies in fish.

Offspring size reflects the optimal balance between female fecundity and allocation of energy to each offspring. Most fish, in particular teleost species, produce many small eggs, while others, notably elasmobranch species, have low fecundity and large offspring. No general explanation has yet been put forwards to explain these different strategies between species which occupy similar habitats. We approach the problem by (1) examining the differences between life history parameters of teleost fish and elasmobranchs and (2) an evolutionary model. We show that life history parameters characterising growth, mortality and reproductive output are almost similar between teleosts and elasmobranchs. We find that a model which accounts for density-dependence predicts dual offspring size strategies: either invariant with adult size or proportional to adult size. The model predicts that the invariant strategy is associated with an absence of density-dependence in early life whereas proportional offspring are subject to density-dependence throughout life. Parameterising the model using life history data regenerates the observed dual offspring size pattern. We conjecture that the life stage where density-dependent competition occurs is an important factor behind the observed difference in offspring size strategies.

The consequences of balanced harvesting of fish communities.

Balanced harvesting, where species or individuals are exploited in accordance with their productivity, has been proposed as a way to minimize the effects of fishing on marine fish communities and ecosystems. This calls for a thorough examination of the consequences balanced harvesting has on fish community structure and yield. We use a size- and trait-based model that resolves individual interactions through competition and predation to compare balanced harvesting with traditional selective harvesting, which protects juvenile fish from fishing. Four different exploitation patterns, generated by combining selective or unselective harvesting with balanced or unbalanced fishing, are compared. We find that unselective balanced fishing, where individuals are exploited in proportion to their productivity, produces a slightly larger total maximum sustainable yield than the other exploitation patterns and, for a given yield, the least change in the relative biomass composition of the fish community. Because fishing reduces competition, predation and cannibalism within the community, the total maximum sustainable yield is achieved at high exploitation rates. The yield from unselective balanced fishing is dominated by small individuals, whereas selective fishing produces a much higher proportion of large individuals in the yield. Although unselective balanced fishing is predicted to produce the highest total maximum sustainable yield and the lowest impact on trophic structure, it is effectively a fishery predominantly targeting small forage fish.

The marine diversity spectrum.

Distributions of species body sizes within a taxonomic group, for example, mammals, are widely studied and important because they help illuminate the evolutionary processes that produced these distributions. Distributions of the sizes of species within an assemblage delineated by geography instead of taxonomy (all the species in a region regardless of clade) are much less studied but are equally important and will illuminate a different set of ecological and evolutionary processes. We develop and test a mechanistic model of how diversity varies with body mass in marine ecosystems. The model predicts the form of the 'diversity spectrum', which quantifies the distribution of species' asymptotic body masses, is a species analogue of the classic size spectrum of individuals, and which we have found to be a new and widely applicable description of diversity patterns. The marine diversity spectrum is predicted to be approximately linear across an asymptotic mass range spanning seven orders of magnitude. Slope -0.5 is predicted for the global marine diversity spectrum for all combined pelagic zones of continental shelf seas, and slopes for large regions are predicted to lie between -0.5 and -0.1. Slopes of -0.5 and -0.1 represent markedly different communities: a slope of -0.5 depicts a 10-fold reduction in diversity for every 100-fold increase in asymptotic mass; a slope of -0.1 depicts a 1.6-fold reduction. Steeper slopes are predicted for larger or colder regions, meaning fewer large species per small species for such regions. Predictions were largely validated by a global empirical analysis. Results explain for the first time a new and widespread phenomenon of biodiversity. Results have implications for estimating numbers of species of small asymptotic mass, where taxonomic inventories are far from complete. Results show that the relationship between diversity and body mass can be explained from the dependence of predation behaviour, dispersal, and life history on body mass, and a neutral assumption about speciation and extinction.

Multi-decadal responses of a cod (Gadus morhua) population to human-induced trophic changes, fishing, and climate.

Understanding how human impacts have interacted with natural variability to affect populations and ecosystems is required for sustainable management and conservation. The Baltic Sea is one of the few large marine ecosystems worldwide where the relative contribution of several key forcings to changes in fish populations can be analyzed with empirical data. In this study we investigate how climate variability and multiple human impacts (fishing, marine mammal hunting, eutrophication) have affected multi-decadal scale dynamics of cod in the Baltic Sea during the 20th century. We document significant climate-driven variations in cod recruitment production at multi-annual timescales, which had major impacts on population dynamics and the yields to commercial fisheries. We also quantify the roles of marine mammal predation, eutrophication, and exploitation on the development of the cod population using simulation analyses, and show how the intensity of these forcings differed over time. In the early decades of the 20th century, marine mammal predation and nutrient availability were the main limiting factors; exploitation of cod was still relatively low. During the 1940s and subsequent decades, exploitation increased and became a dominant forcing on the population. Eutrophication had a relatively minor positive influence on cod biomass until the 1980s. The largest increase in cod biomass occurred during the late 1970s, following a long period of hydrographically related above-average cod productivity coupled to a temporary reduction in fishing pressure. The Baltic cod example demonstrates how combinations of different forcings can have synergistic effects and consequently dramatic impacts on population dynamics. Our results highlight the potential and limitations of human manipulations to influence predator species and show that sustainable management can only be achieved by considering both anthropogenic and naturally varying processes in a common framework.

The Baltic Sea Atlantis: An integrated end-to-end modelling framework evaluating ecosystem-wide effects of human-induced pressures.

Achieving good environmental status in the Baltic Sea region requires decision support tools which are based on scientific knowledge across multiple disciplines. Such tools should integrate the complexity of the ecosystem and enable exploration of different natural and anthropogenic pressures such as climate change, eutrophication and fishing pressures in order to compare alternative management strategies. We present a new framework, with a Baltic implementation of the spatially-explicit end-to-end Atlantis ecosystem model linked to two external models, to explore the different pressures on the marine ecosystem. The HBM-ERGOM initializes the Atlantis model with high-resolution physical-chemical-biological and hydrodynamic information while the FISHRENT model analyses the fisheries economics of the output of commercial fish biomass for the Atlantis terminal projection year. The Baltic Atlantis model composes 29 sub-areas, 9 vertical layers and 30 biological functional groups. The balanced calibration provides realistic levels of biomass for, among others, known stock sizes of top predators and of key fish species. Furthermore, it gives realistic levels of phytoplankton biomass and shows reasonable diet compositions and geographical distribution patterns for the functional groups. By simulating several scenarios of nutrient load reductions on the ecosystem and testing sensitivity to different fishing pressures, we show that the model is sensitive to those changes and capable of evaluating the impacts on different trophic levels, fish stocks, and fisheries associated with changed benthic oxygen conditions. We conclude that the Baltic Atlantis forms an initial basis for strategic management evaluation suited for conducting medium to long term ecosystem assessments which are of importance for a number of pan-Baltic stakeholders in relation to anthropogenic pressures such as eutrophication, climate change and fishing pressure, as well as changed biological interactions between functional groups.

Population structure of Atlantic mackerel (Scomber scombrus).

Atlantic mackerel (Scomber scombrus) occurs on both sides of the north Atlantic and has traditionally been grouped into 5 spawning components, some of which were thought to be isolated natal homing stocks. Previous studies have provided no evidence for cross Atlantic migration and no or weak support for isolated spawning components within either side of the North Atlantic. We question the de-facto accepted hypothesis of isolation between spawning components on the basis of spawning and age distribution data. The spawning intensities, proxied by larval abundances, are negatively correlated between the North Sea and Celtic Sea, which indicates that the two spawning components may be connected by straying individuals. This finding is based on unique larvae samples collected before the collapse of North Sea component, thus showing that the exchange is not a recent phenomenon due to the collapse. The analyses of old as well as more recent age distributions show that strong year classes spread into other areas where they spawn as adults ("twinning"). Our findings are in accordance with the lack of solid evidence for stock separation from previous analyses of tagging data, genetics, ectoparasite infections, otolith shapes, and blood phenotypes. Because no method has been able to identify the origin of spawning mackerel unequivocally from any of the traditional spawning components, and in the light of our results, we conclude that straying outweighs spatial segregation. We propose a new model where the population structure of mackerel is described as a dynamic cline, rather than as connected contingents. Temporal changes in hydrography and mackerel behavior may affect the steepness of the cline at various locations. The new interpretation of the population structure of Atlantic mackerel has important implications for research, assessment and management.