Rijnsdorp, A. D., Peck, M. A., Engelhard, G. H., Möllmann, C., and Pinnegar, J. K. 2009. Resolving the effect of climate change on fish populations. – ICES Journal of Marine Science, 66: 1570–1583. This paper develops a framework for the study of climate on fish populations based on first principles of physiology, ecology, and available observations. Environmental variables and oceanographic features that are relevant to fish and that are likely to be affected by climate change are reviewed. Working hypotheses are derived from the differences in the expected response of different species groups. A review of published data on Northeast Atlantic fish species representing different biogeographic affinities, habitats, and body size lends support to the hypothesis that global warming results in a shift in abundance and distribution (in patterns of occurrence with latitude and depth) of fish species. Pelagic species exhibit clear changes in seasonal migration patterns related to climate-induced changes in zooplankton productivity. Lusitanian species have increased in recent decades (sprat, anchovy, and horse mackerel), especially at the northern limit of their distribution areas, while Boreal species decreased at the southern limit of their distribution range (cod and plaice), but increased at the northern limit (cod). Although the underlying mechanisms remain uncertain, available evidence suggests climate-related changes in recruitment success to be the key process, stemming from either higher production or survival in the pelagic egg or larval stage, or owing to changes in the quality/quantity of nursery habitats.

Marine ecosystems such as the Baltic Sea are currently under strong atmospheric and anthropogenic pressure. Besides natural and human-induced changes in climate, major anthropogenic drivers such as overfishing and anthropogenic eutrophication are significantly affecting ecosystem structure and function. Recently, studies demonstrated the existence of alternative stable states in various terrestrial and aquatic ecosystems. These so-called ecosystem regime shifts have been explained mainly as a result of multiple causes, e.g. climatic regime shifts, overexploitation or a combination of both. The occurrence of ecosystem regime shifts has important management implications, as they can cause significant losses of ecological and economic resources. Because of hysteresis in ecosystem responses, restoring regimes considered as favourable may require drastic and expensive management actions. Also the Baltic Sea, the largest brackish water body in the world ocean, and its ecosystems are strongly affected by atmospheric and anthropogenic drivers. Here, we present results of an analysis of the state and development of the Central Baltic Sea ecosystem integrating hydroclimatic, nutrient, phyto-and zooplankton as well as fisheries data. Our analyses of 52 biotic and abiotic variables using multivariate statistics demonstrated a major reorganization of the ecosystem and identified two stable states between 1974 and 2005, separated by a transition period in 1988-1993. We show the change in Baltic ecosystem structure to have the characteristics of a discontinuous regime shift, initiated by climate-induced changes in the abiotic environment and stabilized by fisheries-induced feedback loops in the food web. Our results indicate the importance of maintaining the resilience of an ecosystem to atmospherically induced environmental change by reducing the anthropogenic impact.

The index of the North Atlantic Oscillation, the dominant mode of climatic variability in the North Atlantic region, changed in the late 1980s (1987–1989) from a negative to a positive phase. This led to regime shifts in the ecology of the North Sea (NS) and the central Baltic Sea (CBS), which involved all trophic levels in the pelagial of these two neighbouring continental shelf seas. Increasing air and sea surface temperatures, which affected critical physical and biological processes, were the main direct and indirect driving forces. After 1987, phytoplankton biomass in both systems increased and the growing season was extended. The composition of phyto- and zooplankton communities in both seas changed conspicuously, e.g. dinoflagellate abundance increased and diatom abundance decreased in the CBS. Key copepod species that are essential in fish diets experienced pronounced changes in biomass. Abundance of Calanus finmarchicus (NS) and Pseudocalanus sp. (CBS) fell to low levels, whereas C. helgolandicus (NS) and Temora longicornis and Acartia spp. (CBS) were persistently abundant. These changes in biomass of different copepod species had dramatic consequences on biomass, fisheries, and landings of key fish species: North Sea cod declined, cod in the CBS remained at low levels, and CBS sprat reached unprecedented high biomass levels resulting in high yields. The synchronous regime shifts in NS and CBS resulted in profound changes in both marine ecosystems. However, the reaction of fish populations to the bottom-up mechanisms caused by the same climatic shift was very different for the three fish stocks.

Möllmann, C., Müller-Karulis, B., Kornilovs, G., and St John, M. A. 2008. Effects of climate and overfishing on zooplankton dynamics and ecosystem structure: regime shifts, trophic cascade, and feeback loops in a simple ecosystem. – ICES Journal of Marine Science, 65: 302–310. The Central Baltic Sea is the largest brackish waterbody in the world ocean, containing a highly productive but low-diversity ecosystem. Climate-induced changes in hydrography recently caused an ecosystem regime shift with changes at all trophic levels. The most pronounced changes in the ecosystem occurred at the zooplankton and fish trophic levels. In the zooplankton, dominance changed between the copepods Pseudocalanus acuspes and Acartia spp., a result of reduced salinities and increased temperatures. The change in hydrography also affected the reproductive success of the major fish species, resulting in a change in dominance from the piscivorous cod (Gadus morhua) to the planktivorous sprat (Sprattus sprattus). First, we investigate statistically the occurrence of regime shifts in time-series of key hydrographic variables and the biomass time-series of key species. Second, we demonstrate a three-level trophic cascade involving zooplankton. Finally, we model the ecosystem effects of the abiotic and biotic changes on copepod biomass and recruitment of fish stocks. Our results demonstrate the linkage between climate-induced zooplankton and fish regime changes, and how overfishing amplified the climate-induced changes at both trophic levels. Hence, our study demonstrates (i) the multiple pathways along which climatic and anthropogenic pressures can propagate through the foodweb; (ii) how both effects act synergistically to cause and stabilize regime changes; and (iii) the crucial role of zooplankton in mediating these ecosystem changes.

Modern fisheries research and management must understand and take account of the interactions between climate and fishing, rather than try to disentangle their effects and address each separately. These interactions are significant drivers of change in exploited marine systems and have ramifications for ecosystems and those who depend on the services they provide. We discuss how fishing and climate forcing interact on individual fish, marine populations, marine communities, and ecosystems to bring these levels into states that are more sensitive to (i.e. more strongly related with) climate forcing. Fishing is unlikely to alter the sensitivities of individual finfish and invertebrates to climate forcing. It will remove individuals with specific characteristics from the gene pool, thereby affecting structure and function at higher levels of organisation. Fishing leads to a loss of older age classes, spatial contraction, loss of sub-units, and alteration of life history traits in populations, making them more sensitive to climate variability at interannual to interdecadal scales. Fishing reduces the mean size of individuals and mean trophic level of communities, decreasing their turnover time leading them to track environmental variability more closely. Marine ecosystems under intense exploitation evolve towards stronger bottom–up control and greater sensitivity to climate forcing. Because climate change occurs slowly, its effects are not likely to have immediate impacts on marine systems but will be manifest as the accumulation of the interactions between fishing and climate variability — unless threshold limits are exceeded. Marine resource managers need to develop approaches which maintain the resilience of individuals, populations, communities and ecosystems to the combined and interacting effects of climate and fishing. Overall, a less-heavily fished marine system, and one which shifts the focus from individual species to functional groups and fish communities, is likely to provide more stable catches with climate variability and change than would a heavily fished system

Large-scale climatic conditions prevailing over the central Baltic Sea resulted in declining salinity and oxygen concentrations in spawning areas of the eastern Baltic cod stock. These changes in hydrography reduced the reproductive success and, combined with high fishing pressure, caused a decline of the stock to the lowest level on record in the early 1990s. The present study aims at disentangling the interactions between reproductive effort and hydrographic forcing leading to variable recruitment. Based on identified key processes, stock dynamics is explained using updated environmental and life stage-specific abundance and production time-series. Declining salinities and oxygen concentrations caused high egg mortalities and indirectly increased egg predation by clupeid fish. Low recruitment, despite enhanced hydrographic conditions for egg survival in the mid-1990s, was due to food limitation for larvae, caused by the decline in the abundance of the copepod Pseudocalanus sp. The case of the eastern Baltic cod stock exemplifies the multitude effects climatic variability may have on a fish stock and underscores the importance of knowledge of these processes for understanding stock dynamics.

Climate change and resource exploitation have been shown to modify the importance of bottom-up and top-down forces in ecosystems. However, the resulting pattern of trophic control in complex food webs is an emergent property of the system and thus unintuitive. We develop a statistical nondeterministic model, capable of modeling complex patterns of trophic control for the heavily impacted North Sea ecosystem. The model is driven solely by fishing mortality and climatic variables and based on timeseries data covering >40 y for six plankton and eight fish groups along with one bird group (>20 y). Simulations show the outstanding importance of top-down exploitation pressure for the dynamics of fish populations. Whereas fishing effects on predators indirectly altered plankton abundance, bottom-up climatic processes dominate plankton dynamics. Importantly, we show planktivorous fish to have a central role in the North Sea food web initiating complex cascading effects across and between trophic levels. Our linked model integrates bottom-up and top-down effects and is able to simulate complex long-term changes in ecosystem components under a combination of stressor scenarios. Our results suggest that in marine ecosystems, pathways for bottomup and top-down forces are not necessarily mutually exclusive and together can lead to the emergence of complex patterns of control.trophic control | ecosystem modeling | marine food web functioning | wasp-waist | regime shifts T he question of whether food webs are resource-(bottom-up) or predation-(top-down) controlled is one of the most fundamental research questions in ecology (1-3). Marine ecosystems, originally thought to be mainly steered by bottom-up control, have recently been shown to exhibit periods of top-down control due to the extraction of large predators through fishing (4-7) or climate oscillations (8). Furthermore, experimental evidence shows climate warming may exert a host of indirect effects on aquatic food webs mediated through shifts in the magnitudes of top-down and bottom-up forcing (9, 10). However, for large marine ecosystems that are not amenable to experimentation studies, investigations of how interactions in their complex food webs mediate the influence of both top-down (e.g., fishing) and bottom-up (e.g., climate change) control are lacking or are based on aggregated species complexes. We model an extensive historical dataset for the North Sea (over 45 y) at the lowest possible resolution (often species) to determine key interactions between species and estimate their responses to pressures. The model reveals both simple (direct) and complex (indirect) pathways linking plankton to seabirds and can highlight the wider effects of climate change and potential actions by fishery managers.The North Sea is one of the most anthropogenically impacted marine ecosystem and is thought to be fundamentally driven from the bottom-up through climatic (temperature-related) influences on plankton, planktivorous fish, and the pelagic stages of demersal fish (11-13). Som...

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