Warming shelf seas drive the subtropicalization of European pelagic fish communities

Pelagic fishes are among the most ecologically and economically important fish species in European seas. In principle, these pelagic fishes have potential to demonstrate rapid abundance and distribution shifts in response to climatic variability due to their high adult motility, planktonic larval stages, and low dependence on benthic habitat for food or shelter during their life histories. Here, we provide evidence of substantial climate‐driven changes to the structure of pelagic fish communities in European shelf seas. We investigated the patterns of species‐level change using catch records from 57 870 fisheries‐independent survey trawls from across European continental shelf region between 1965 and 2012. We analysed changes in the distribution and rate of occurrence of the six most common species, and observed a strong subtropicalization of the North Sea and Baltic Sea assemblages. These areas have shifted away from cold‐water assemblages typically characterized by Atlantic herring and European sprat from the 1960s to 1980s, to warmer‐water assemblages including Atlantic mackerel, Atlantic horse mackerel, European pilchard and European anchovy from the 1990s onwards. We next investigated if warming sea temperatures have forced these changes using temporally comprehensive data from the North Sea region. Our models indicated the primary driver of change in these species has been sea surface temperatures in all cases. Together, these analyses highlight how individual species responses have combined to result in a dramatic subtropicalization of the pelagic fish assemblage of the European continental shelf.     

Physiological and ecological effects of increasing temperature on fish production in lakes of Arctic Alaska

Lake ecosystems in the Arctic are changing rapidly due to climate warming. Lakes are sensitive integrators of climate‐induced changes and prominent features across the Arctic landscape, especially in lowland permafrost regions such as the Arctic Coastal Plain of Alaska. Despite many studies on the implications of climate warming, how fish populations will respond to lake changes is uncertain for Arctic ecosystems. Least Cisco (Coregonus sardinella) is a bellwether for Arctic lakes as an important consumer and prey resource. To explore the consequences of climate warming, we used a bioenergetics model to simulate changes in Least Cisco production under future climate scenarios for lakes on the Arctic Coastal Plain. First, we used current temperatures to fit Least Cisco consumption to observed annual growth. We then estimated growth, holding food availability, and then feeding rate constant, for future projections of temperature. Projected warmer water temperatures resulted in reduced Least Cisco production, especially for larger size classes, when food availability was held constant. While holding feeding rate constant, production of Least Cisco increased under all future scenarios with progressively more growth in warmer temperatures. Higher variability occurred with longer projections of time mirroring the expanding uncertainty in climate predictions further into the future. In addition to direct temperature effects on Least Cisco growth, we also considered changes in lake ice phenology and prey resources for Least Cisco. A shorter period of ice cover resulted in increased production, similar to warming temperatures. Altering prey quality had a larger effect on fish production in summer than winter and increased relative growth of younger rather than older age classes of Least Cisco. Overall, we predicted increased production of Least Cisco due to climate warming in lakes of Arctic Alaska. Understanding the implications of increased production of Least Cisco to the entire food web will be necessary to predict ecosystem responses in lakes of the Arctic.     

Sea temperature is the primary driver of recent and predicted fish community structure across Northeast Atlantic shelf seas

Climate change has strongly influenced the distribution and abundance of marine fish species, leading to concern about effects of future climate on commercially harvested stocks. Understanding the key drivers of large-scale spatial variation across present-day marine assemblages enables predictions of future change. Here we present a unique analysis of standardised abundance data for 198 marine fish species from across the Northeast Atlantic collected by 23 surveys and 31,502 sampling events between 2005 and 2018. Our analyses of the spatially comprehensive standardised data identified temperature as the key driver of fish community structure across the region, followed by salinity and depth. We employed these key environmental variables to model how climate change will affect both the distributions of individual species and local community structure for the years 2050 and 2100 under multiple emissions scenarios. Our results consistently indicate that projected climate change will lead to shifts in species communities across the entire region. Overall, the greatest community-level changes are predicted at locations with greater warming, with the most pronounced effects at higher latitudes. Based on these results, we suggest that future climate-driven warming will lead to widespread changes in opportunities for commercial fisheries across the region.     

Lifeform indicators reveal large‐scale shifts in plankton across the North‐West European shelf

Increasing direct human pressures on the marine environment, coupled with climate‐driven changes, is a concern to marine ecosystems globally. This requires the development and monitoring of ecosystem indicators for effective management and adaptation planning. Plankton lifeforms (broad functional groups) are sensitive indicators of marine environmental change and can provide a simplified view of plankton biodiversity, building an understanding of change in lower trophic levels. Here, we visualize regional‐scale multi‐decadal trends in six key plankton lifeforms as well as their correlative relationships with sea surface temperature (SST). For the first time, we collate trends across multiple disparate surveys, comparing the spatially and temporally extensive Continuous Plankton Recorder (CPR) survey (offshore) with multiple long‐term fixed station‐based time‐series (inshore) from around the UK coastline. These analyses of plankton lifeforms showed profound long‐term changes, which were coherent across large spatial scales. For example, ‘diatom’ and ‘meroplankton’ lifeforms showed strong alignment between surveys and coherent regional‐scale trends, with the 1998–2017 decadal average abundance of meroplankton being 2.3 times that of 1958–1967 for CPR samples in the North Sea. This major, shelf‐wide increase in meroplankton correlated with increasing SSTs, and contrasted with a general decrease in holoplankton (dominated by small copepods), indicating a changing balance of benthic and pelagic fauna. Likewise, inshore‐offshore gradients in dinoflagellate trends, with contemporary increases inshore contrasting with multi‐decadal decreases offshore (approx. 75% lower decadal mean abundance), urgently require the identification of causal mechanisms. Our lifeform approach allows the collation of many different data types and time‐series across the NW European shelf, providing a crucial evidence base for informing ecosystem‐based management, and the development of regional adaptation plans.     

Interactions between invading benthivorous fish and native whitefish in subarctic lakes

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Consistent trophic amplification of marine biomass declines under climate change

The impact of climate change on the marine food web is highly uncertain. Nonetheless, there is growing consensus that global marine primary production will decline in response to future climate change, largely due to increased stratification reducing the supply of nutrients to the upper ocean. Evidence to date suggests a potential amplification of this response throughout the trophic food web, with more dramatic responses at higher trophic levels. Here we show that trophic amplification of marine biomass declines is a consistent feature of the Coupled Model Intercomparison Project Phase 5 (CMIP5) Earth System Models, across different scenarios of future climate change. Under the business‐as‐usual Representative Concentration Pathway 8.5 (RCP8.5) global mean phytoplankton biomass is projected to decline by 6.1% ± 2.5% over the twenty‐first century, while zooplankton biomass declines by 13.6% ± 3.0%. All models project greater relative declines in zooplankton than phytoplankton, with annual zooplankton biomass anomalies 2.24 ± 1.03 times those of phytoplankton. The low latitude oceans drive the projected trophic amplification of biomass declines, with models exhibiting variable trophic interactions in the mid‐to‐high latitudes and similar relative changes in phytoplankton and zooplankton biomass. Under the assumption that zooplankton biomass is prey limited, an analytical explanation of the trophic amplification that occurs in the low latitudes can be derived from generic plankton differential equations. Using an ocean biogeochemical model, we show that the inclusion of variable C:N:P phytoplankton stoichiometry can substantially increase the trophic amplification of biomass declines in low latitude regions. This additional trophic amplification is driven by enhanced nutrient limitation decreasing phytoplankton N and P content relative to C, hence reducing zooplankton growth efficiency. Given that most current Earth System Models assume that phytoplankton C:N:P stoichiometry is constant, such models are likely to underestimate the extent of negative trophic amplification under projected climate change.     

Exploitation and habitat degradation as agents of change within coral reef fish communities

Over‐exploitation and habitat degradation are the two major drivers of global environmental change and are responsible for local extinctions and declining ecosystem services. Here we compare the top‐down effect of exploitation by fishing with the bottom‐up influence of habitat loss on fish communities in the most diverse of ecological systems, coral reefs. Using a combination of multivariate techniques and path analyses, we illustrate that the relative importance of coral cover and fishing in controlling fish abundance on remote Fijian reefs varies between species and functional groups. A decline in branching Acropora coral is strongly associated with a decline in abundance of coral‐feeding species, and a decrease in coral‐associated habitat complexity, which has indirectly contributed to reduced abundance of small‐bodied damselfish. In contrast, reduced fishing pressure, brought about by declining human populations and a shift to alternate livelihoods, is associated with increased abundance of some piscivores and fisheries target species. However, availability of prey is controlled by coral‐associated habitat complexity and appears to be a more important driver of total piscivore abundance compared with fishing pressure. Effects of both fishing and coral loss are stronger on individual species than functional groups, as variation in the relative importance of fishing or coral loss among species within the same functional group attenuated the impact of either of these potential drivers at the functional level. Overall, fishing continues to have an influence on Fijian fish communities; however, habitat loss is currently the overriding agent of change. The importance of coral loss mediated by climate change is expected to have an increasing contribution to fish community dynamics, particularly in remote locations or where the influence of fishing is waning.     

The impact of climate change on the structure of Pleistocene food webs across the mammoth steppe

Species interactions form food webs, impacting community structure and, potentially, ecological dynamics. It is likely that global climatic perturbations that occur over long periods of time have a significant influence on species interaction patterns. Here, we integrate stable isotope analysis and network theory to reconstruct patterns of trophic interactions for six independent mammalian communities that inhabited mammoth steppe environments spanning western Europe to eastern Alaska (Beringia) during the Late Pleistocene. We use a Bayesian mixing model to quantify the contribution of prey to the diets of local predators, and assess how the structure of trophic interactions changed across space and the Last Glacial Maximum (LGM), a global climatic event that severely impacted mammoth steppe communities. We find that large felids had diets that were more constrained than those of co-occurring predators, and largely influenced by an increase in Rangifer abundance after the LGM. Moreover, the structural organization of Beringian and European communities strongly differed: compared with Europe, species interactions in Beringian communities before—and possibly after—the LGM were highly modular. We suggest that this difference in modularity may have been driven by the geographical insularity of Beringian communities.     

Climate change in size-structured ecosystems

One important aspect of climate change is the increase in average temperature, which will not only have direct physiological effects on all species but also indirectly modifies abundances, interaction strengths, food-web topologies, community stability and functioning. In this theme issue, we highlight a novel pathway through which warming indirectly affects ecological communities: by changing their size structure (i.e. the body-size distributions). Warming can shift these distributions towards dominance of small- over large-bodied species. The conceptual, theoretical and empirical research described in this issue, in sum, suggests that effects of temperature may be dominated by changes in size structure, with relatively weak direct effects. For example, temperature effects via size structure have implications for top-down and bottom-up control in ecosystems and may ultimately yield novel communities. Moreover, scaling up effects of temperature and body size from physiology to the levels of populations, communities and ecosystems may provide a crucially important mechanistic approach for forecasting future consequences of global warming.     

Elevated carbon dioxide affects behavioural lateralization in a coral reef fish

Elevated carbon dioxide (CO(2)) has recently been shown to affect chemosensory and auditory behaviour, and activity levels of larval reef fishes, increasing their risk of predation. However, the mechanisms underlying these changes are unknown. Behavioural lateralization is an expression of brain functional asymmetries, and thus provides a unique test of the hypothesis that elevated CO(2) affects brain function in larval fishes. We tested the effect of near-future CO(2) concentrations (880 μatm) on behavioural lateralization in the reef fish, Neopomacentrus azysron. Individuals exposed to current-day or elevated CO(2) were observed in a detour test where they made repeated decisions about turning left or right. No preference for right or left turns was observed at the population level. However, individual control fish turned either left or right with greater frequency than expected by chance. Exposure to elevated-CO(2) disrupted individual lateralization, with values that were not different from a random expectation. These results provide compelling evidence that elevated CO(2) directly affects brain function in larval fishes. Given that lateralization enhances performance in a number of cognitive tasks and anti-predator behaviours, it is possible that a loss of lateralization could increase the vulnerability of larval fishes to predation in a future high-CO(2) ocean.     

Interactions between temperature and nutrients across levels of ecological organization

Temperature and nutrient availability play key roles in controlling the pathways and rates at which energy and materials move through ecosystems. These factors have also changed dramatically on Earth over the past century as human activities have intensified. Although significant effort has been devoted to understanding the role of temperature and nutrients in isolation, less is known about how these two factors interact to influence ecological processes. Recent advances in ecological stoichiometry and metabolic ecology provide a useful framework for making progress in this area, but conceptual synthesis and review are needed to help catalyze additional research. Here, we examine known and potential interactions between temperature and nutrients from a variety of physiological, community, and ecosystem perspectives. We first review patterns at the level of the individual, focusing on four traits – growth, respiration, body size, and elemental content – that should theoretically govern how temperature and nutrients interact to influence higher levels of biological organization. We next explore the interactive effects of temperature and nutrients on populations, communities, and food webs by synthesizing information related to community size spectra, biomass distributions, and elemental composition. We use metabolic theory to make predictions about how population‐level secondary production should respond to interactions between temperature and resource supply, setting up qualitative predictions about the flows of energy and materials through metazoan food webs. Last, we examine how temperature–nutrient interactions influence processes at the whole‐ecosystem level, focusing on apparent vs. intrinsic activation energies of ecosystem processes, how to represent temperature–nutrient interactions in ecosystem models, and patterns with respect to nutrient uptake and organic matter decomposition. We conclude that a better understanding of interactions between temperature and nutrients will be critical for developing realistic predictions about ecological responses to multiple, simultaneous drivers of global change, including climate warming and elevated nutrient supply.     

Scaling metabolism from individuals to reef‐fish communities at broad spatial scales

Fishes contribute substantially to energy and nutrient fluxes in reef ecosystems, but quantifying these roles is challenging. Here, we do so by synthesising a large compilation of fish metabolic‐rate data with a comprehensive database on reef‐fish community abundance and biomass. Individual‐level analyses support predictions of Metabolic Theory after accounting for significant family‐level variation, and indicate that some tropical reef fishes may already be experiencing thermal regimes at or near their temperature optima. Community‐level analyses indicate that total estimated respiratory fluxes of reef‐fish communities increase on average ~2‐fold from 22 to 28 °C. Comparisons of estimated fluxes among trophic groups highlight striking differences in resource use by communities in different regions, perhaps partly reflecting distinct evolutionary histories, and support the hypothesis that piscivores receive substantial energy subsidies from outside reefs. Our study demonstrates one approach to synthesising individual‐ and community‐level data to establish broad‐scale trends in contributions of biota to ecosystem dynamics.     

Warmer and browner waters decrease fish biomass production

Climate change studies have long focused on effects of increasing temperatures, often without considering other simultaneously occurring environmental changes, such as browning of waters. Resolving how the combination of warming and browning of aquatic ecosystems affects fish biomass production is essential for future ecosystem functioning, fisheries, and food security. In this study, we analyzed individual‐ and population‐level fish data from 52 temperate and boreal lakes in Northern Europe, covering large gradients in water temperature and color (absorbance, 420 nm). We show that fish (Eurasian perch, Perca fluviatilis) biomass production decreased with both high water temperatures and brown water color, being lowest in warm and brown lakes. However, while both high temperature and brown water decreased fish biomass production, the mechanisms behind the decrease differed: temperature affected the fish biomass production mainly through a decrease in population standing stock biomass, and through shifts in size‐ and age‐distributions toward a higher proportion of young and small individuals in warm lakes; brown water color, on the other hand, mainly influenced fish biomass production through negative effects on individual body growth and length‐at‐age. In addition to these findings, we observed that the effects of temperature and brown water color on individual‐level processes varied over ontogeny. Body growth only responded positively to higher temperatures among young perch, and brown water color had a stronger negative effect on body growth of old than on young individuals. Thus, to better understand and predict future fish biomass production, it is necessary to integrate both individual‐ and population‐level responses and to acknowledge within‐species variation. Our results suggest that global climate change, leading to browner and warmer waters, may negatively affect fish biomass production, and this effect may be stronger than caused by increased temperature or water color alone.     

Effects of altered temperature and precipitation on desert protozoa associated with biological soil crusts

Biological soil crusts are diverse assemblages of bacteria, cyanobacteria, algae, fungi, lichens, and mosses that cover much of arid land soils. The objective of this study was to quantify protozoa associated with biological soil crusts and test the response of protozoa to increased temperature and precipitation as is predicted by some global climate models. Protozoa were more abundant when associated with cyanobacteria/lichen crusts than with cyanobacteria crusts alone. Amoebae, flagellates, and ciliates originating from the Colorado Plateau desert (cool desert, primarily winter precipitation) declined 50-, 10-, and 100-fold, respectively, when moved in field mesocosms to the Chihuahuan Desert (hot desert, primarily summer rain). However, this was not observed in protozoa collected from the Chihuahuan Desert and moved to the Sonoran desert (hot desert, also summer rain, but warmer than Chihuahuan Desert). Protozoa in culture began to encyst at 37 degrees C. Cysts survived the upper end of daily temperatures (37-55 degrees C), and could be stimulated to excyst if temperatures were reduced to 15 degrees C or lower. Results from this study suggest that cool desert protozoa are influenced negatively by increased summer precipitation during excessive summer temperatures, and that desert protozoa may be adapted to a specific desert's temperature and precipitation regime.     

Climate change and freshwater ecosystems: impacts across multiple levels of organization

Fresh waters are particularly vulnerable to climate change because (i) many species within these fragmented habitats have limited abilities to disperse as the environment changes; (ii) water temperature and availability are climate-dependent; and (iii) many systems are already exposed to numerous anthropogenic stressors. Most climate change studies to date have focused on individuals or species populations, rather than the higher levels of organization (i.e. communities, food webs, ecosystems). We propose that an understanding of the connections between these different levels, which are all ultimately based on individuals, can help to develop a more coherent theoretical framework based on metabolic scaling, foraging theory and ecological stoichiometry, to predict the ecological consequences of climate change. For instance, individual basal metabolic rate scales with body size (which also constrains food web structure and dynamics) and temperature (which determines many ecosystem processes and key aspects of foraging behaviour). In addition, increasing atmospheric CO₂ is predicted to alter molar CNP ratios of detrital inputs, which could lead to profound shifts in the stoichiometry of elemental fluxes between consumers and resources at the base of the food web. The different components of climate change (e.g. temperature, hydrology and atmospheric composition) not only affect multiple levels of biological organization, but they may also interact with the many other stressors to which fresh waters are exposed, and future research needs to address these potentially important synergies.     

Effects of ocean acidification on the early life history of a tropical marine fish

Little is known about how fishes and other non-calcifying marine organisms will respond to the increased levels of dissolved CO₂ and reduced sea water pH that are predicted to occur over the coming century. We reared eggs and larvae of the orange clownfish, Amphiprion percula, in sea water simulating a range of ocean acidification scenarios for the next 50–100 years (current day, 550, 750 and 1030 ppm atmospheric CO₂). CO₂ acidification had no detectable effect on embryonic duration, egg survival and size at hatching. In contrast, CO₂ acidification tended to increase the growth rate of larvae. By the time of settlement (11 days post-hatching), larvae from some parental pairs were 15 to 18 per cent longer and 47 to 52 per cent heavier in acidified water compared with controls. Larvae from other parents were unaffected by CO₂ acidification. Elevated CO₂ and reduced pH had no effect on the maximum swimming speed of settlement-stage larvae. There was, however, a weak positive relationship between length and swimming speed. Large size is usually considered to be advantageous for larvae and newly settled juveniles. Consequently, these results suggest that levels of ocean acidification likely to be experienced in the near future might not, in isolation, significantly disadvantage the growth and performance of larvae from benthic-spawning marine fishes.     

Predicting ecosystem shifts requires new approaches that integrate the effects of climate change across entire systems

Most studies that forecast the ecological consequences of climate change target a single species and a single life stage. Depending on climatic impacts on other life stages and on interacting species, however, the results from simple experiments may not translate into accurate predictions of future ecological change. Research needs to move beyond simple experimental studies and environmental envelope projections for single species towards identifying where ecosystem change is likely to occur and the drivers for this change. For this to happen, we advocate research directions that (i) identify the critical species within the target ecosystem, and the life stage(s) most susceptible to changing conditions and (ii) the key interactions between these species and components of their broader ecosystem. A combined approach using macroecology, experimentally derived data and modelling that incorporates energy budgets in life cycle models may identify critical abiotic conditions that disproportionately alter important ecological processes under forecasted climates.     

Large but uneven reduction in fish size across species in relation to changing sea temperatures

Ectotherms often attain smaller body sizes when they develop at higher temperatures. This phenomenon, known as the temperature–size rule, has important consequences for global fisheries, whereby ocean warming is predicted to result in smaller fish and reduced biomass. However, the generality of this phenomenon and the mechanisms that drive it in natural populations remain unresolved. In this study, we document the maximal size of 74 fish species along a steep temperature gradient in the Mediterranean Sea and find strong support for the temperature–size rule. Importantly, we additionally find that size reduction in active fish species is dramatically larger than for more sedentary species. As the temperature dependence of oxygen consumption depends on activity levels, these findings are consistent with the hypothesis that oxygen is a limiting factor shaping the temperature–size rule in fishes. These results suggest that ocean warming will result in a sharp, but uneven, reduction in fish size that will cause major shifts in size‐dependent interactions. Moreover, warming will have major implications for fisheries as the main species targeted for harvesting will show the most substantial declines in biomass.     

Competition and facilitation in multispecies plant-herbivore systems of productive environments

We develop a multispecies plant-herbivore model to explore how plant competition for light and the selectivity of herbivores affect abundance patterns of plants and herbivores along productivity gradients. The model considers a small and a tall plant species, a generalist herbivore, and a selective herbivore. The selective herbivore feeds only on the small plant species. In the absence of the generalist herbivore, the tall plant species becomes increasingly dominant with increasing productivity, and the small plant and its selective herbivore disappear. The model shows that generalist herbivores can facilitate selective herbivores by suppressing competition for light. This favours the small plant species, and thereby the selective herbivores. The model predictions are qualitatively consistent with field studies of multispecies plant-herbivore systems.