Kelp carbon sink potential decreases with warming due to accelerating decomposition.

Cycling of organic carbon in the ocean has the potential to mitigate or exacerbate global climate change, but major questions remain about the environmental controls on organic carbon flux in the coastal zone. Here, we used a field experiment distributed across 28° of latitude, and the entire range of 2 dominant kelp species in the northern hemisphere, to measure decomposition rates of kelp detritus on the seafloor in relation to local environmental factors. Detritus decomposition in both species were strongly related to ocean temperature and initial carbon content, with higher rates of biomass loss at lower latitudes with warmer temperatures. Our experiment showed slow overall decomposition and turnover of kelp detritus and modeling of coastal residence times at our study sites revealed that a significant portion of this production can remain intact long enough to reach deep marine sinks. The results suggest that decomposition of these kelp species could accelerate with ocean warming and that low-latitude kelp forests could experience the greatest increase in remineralization with a 9% to 42% reduced potential for transport to long-term ocean sinks under short-term (RCP4.5) and long-term (RCP8.5) warming scenarios. However, slow decomposition at high latitudes, where kelp abundance is predicted to expand, indicates potential for increasing kelp-carbon sinks in cooler (northern) regions. Our findings reveal an important latitudinal gradient in coastal ecosystem function that provides an improved capacity to predict the implications of ocean warming on carbon cycling. Broad-scale patterns in organic carbon decomposition revealed here can be used to identify hotspots of carbon sequestration potential and resolve relationships between carbon cycling processes and ocean climate at a global scale.

Multidecadal changes in coastal benthic species composition and ecosystem functioning occur independently of temperature-driven community shifts.

Rising global temperatures are often identified as the key driver impacting ecosystems and the services they provide by affecting biodiversity structure and function. A disproportionate amount of our understanding of biodiversity and function is from short-term experimental studies and static values of biodiversity indices, lacking the ability to monitor long-term trends and capture community dynamics. Here, we analyse a biennial dataset spanning 32 years of macroinvertebrate benthic communities and their functional response to increasing temperatures. We monitored changes in species' thermal affinities to examine warming-related shifts by selecting their mid-point global temperature distribution range and linking them to species' traits. We employed a novel weighted metric using Biological Trait Analysis (BTA) to gain better insights into the ecological potential of each species by incorporating species abundance and body size and selecting a subset of traits that represent five ecosystem functions: bioturbation activity, sediment stability, nutrient recycling and higher and lower trophic production. Using biodiversity indices (richness, Simpson's diversity and vulnerability) and functional indices (richness, Rao's Q and redundancy), the community structure showed no significant change over time with a narrow range of variation. However, we show shifts in species composition with warming and increases in the abundance of individuals, which altered ecosystem functioning positively and/or non-linearly. Yet, when higher taxonomic groupings than species were excluded from the analysis, there was only a weak increase in the measured change in community-weighted average thermal affinities, suggesting changes in ecosystem functions over time occur independently of temperature increase-related shifts in community composition. Other environmental factors driving species composition and abundance may be more important in these subtidal macrobenthic communities. This challenges the prevailing emphasis on temperature as the primary driver of ecological response to climate change and emphasises the necessity for a comprehensive understanding of the temporal dynamics of complex systems.

Shoreline sentinels of global change show the consequences of extreme events.

Anthropogenic climate change along with the more frequent extreme weather it prompts, are having direct and indirect effects on distributions and abundance of species with consequence for community structure-especially if habitat providers are lost. Rocky shores have long been recognized as tractable experimental arenas for ecology contributing to theory. They have also emerged as important sentinel systems for tracking climate change responses of marine biodiversity and ecosystems, capitalizing on both historic broadscale surveys and time series. Combining these twin traditions is a powerful approach for better understanding and forecasting climate change impacts. Sustained observing allows extreme events to be detected and explored by in-parallel experimentation.

Applying landscape metrics to species distribution model predictions to characterize internal range structure and associated changes.

Distributional shifts in species ranges provide critical evidence of ecological responses to climate change. Assessments of climate-driven changes typically focus on broad-scale range shifts (e.g. poleward or upward), with ecological consequences at regional and local scales commonly overlooked. While these changes are informative for species presenting continuous geographic ranges, many species have discontinuous distributions-both natural (e.g. mountain or coastal species) or human-induced (e.g. species inhabiting fragmented landscapes)-where within-range changes can be significant. Here, we use an ecosystem engineer species (Sabellaria alveolata) with a naturally fragmented distribution as a case study to assess climate-driven changes in within-range occupancy across its entire global distribution. To this end, we applied landscape ecology metrics to outputs from species distribution modelling (SDM) in a novel unified framework. SDM predicted a 27.5% overall increase in the area of potentially suitable habitat under RCP 4.5 by 2050, which taken in isolation would have led to the classification of the species as a climate change winner. SDM further revealed that the latitudinal range is predicted to shrink because of decreased habitat suitability in the equatorward part of the range, not compensated by a poleward expansion. The use of landscape ecology metrics provided additional insights by identifying regions that are predicted to become increasingly fragmented in the future, potentially increasing extirpation risk by jeopardising metapopulation dynamics. This increased range fragmentation could have dramatic consequences for ecosystem structure and functioning. Importantly, the proposed framework-which brings together SDM and landscape metrics-can be widely used to study currently overlooked climate-driven changes in species internal range structure, without requiring detailed empirical knowledge of the modelled species. This approach represents an important advancement beyond predictive envelope approaches and could reveal itself as paramount for managers whose spatial scale of action usually ranges from local to regional.

Shape of species climate response curves affects community response to climate change.

Understanding how community composition is reshaped by changing climate is important for interpreting and predicting patterns of community assembly through time or across space. Community composition often does not perfectly correspond to expectations from current environmental conditions, leading to community-climate mismatches. Here, we combine data analysis and theory development to explore how species climate response curves affect the community response to climate change. We show that strong mismatches between community and climate can appear in the absence of demographic delays or limited species pools. Communities simulated using species response curves showed temporal changes of similar magnitude to those observed in natural communities of fishes and plankton, suggesting no overall delays in community change despite substantial unexplained variation from community assembly and other processes. Our approach can be considered as a null model that will be important to use when interpreting observed community responses to climate change and variability.

Global-scale species distributions predict temperature-related changes in species composition of rocky shore communities in Britain.

Changes in rocky shore community composition as responses to climatic fluctuations and anthropogenic warming can be shown by changes in average species thermal affinities. In this study, we derived thermal affinities for European Atlantic rocky intertidal species by matching their known distributions to patterns in average annual sea surface temperature. Average thermal affinities (the Community Temperature Index, CTI) tracked patterns in sea surface temperature from Portugal to Norway, but CTI for communities of macroalgae and plant species changed less than those composed of animal species. This reduced response was in line with the expectation that communities with a smaller range of thermal affinities among species would change less in composition along thermal gradients and over time. Local-scale patterns in CTI over wave exposure gradients suggested that canopy macroalgae allow species with ranges centred in cooler than local temperatures ('cold-affinity') to persist in otherwise too-warm conditions. In annual surveys of rocky shores, communities of animal species in Shetland showed a shift in dominance towards warm-affinity species ('thermophilization') with local warming from 1980 to 2018 but the community of plant and macroalgal species did not. From 2002 to 2018, communities in southwest Britain showed the reverse trend in CTI: declining average thermal affinities over a period of modest temperature decline. Despite the cooling, trends in species abundance were in line with the general mechanism of direction and magnitude of long-term trends depending on the difference between species thermal affinities and local temperatures. Cold-affinity species increased during cooling and warm-affinity ones decreased. The consistency of responses across different communities and with general expectations based on species thermal characteristics suggests strong predictive accuracy of responses of community composition to anthropogenic warming.

Geographical limits to species-range shifts are suggested by climate velocity.

The reorganization of patterns of species diversity driven by anthropogenic climate change, and the consequences for humans, are not yet fully understood or appreciated. Nevertheless, changes in climate conditions are useful for predicting shifts in species distributions at global and local scales. Here we use the velocity of climate change to derive spatial trajectories for climatic niches from 1960 to 2009 (ref. 7) and from 2006 to 2100, and use the properties of these trajectories to infer changes in species distributions. Coastlines act as barriers and locally cooler areas act as attractors for trajectories, creating source and sink areas for local climatic conditions. Climate source areas indicate where locally novel conditions are not connected to areas where similar climates previously occurred, and are thereby inaccessible to climate migrants tracking isotherms: 16% of global surface area for 1960 to 2009, and 34% of ocean for the 'business as usual' climate scenario (representative concentration pathway (RCP) 8.5) representing continued use of fossil fuels without mitigation. Climate sink areas are where climate conditions locally disappear, potentially blocking the movement of climate migrants. Sink areas comprise 1.0% of ocean area and 3.6% of land and are prevalent on coasts and high ground. Using this approach to infer shifts in species distributions gives global and regional maps of the expected direction and rate of shifts of climate migrants, and suggests areas of potential loss of species richness.

Are fish outside their usual ranges early indicators of climate-driven range shifts?

Shifts in species ranges are a global phenomenon, well known to occur in response to a changing climate. New species arriving in an area may become pest species, modify ecosystem structure, or represent challenges or opportunities for fisheries and recreation. Early detection of range shifts and prompt implementation of any appropriate management strategies is therefore crucial. This study investigates whether 'first sightings' of marine species outside their normal ranges could provide an early warning of impending climate-driven range shifts. We examine the relationships between first sightings and marine regions defined by patterns of local climate velocities (calculated on a 50-year timescale), while also considering the distribution of observational effort (i.e. number of sampling days recorded with biological observations in global databases). The marine trajectory regions include climate 'source' regions (areas lacking connections to warmer areas), 'corridor' regions (areas where moving isotherms converge), and 'sink' regions (areas where isotherms locally disappear). Additionally, we investigate the latitudinal band in which first sightings were recorded, and species' thermal affiliations. We found that first sightings are more likely to occur in climate sink and 'divergent' regions (areas where many rapid and diverging climate trajectories pass through) indicating a role of temperature in driving changes in marine species distributions. The majority of our fish first sightings appear to be tropical and subtropical species moving towards high latitudes, as would be expected in climate warming. Our results indicate that first sightings are likely related to longer-term climatic processes, and therefore have potential use to indicate likely climate-driven range shifts. The development of an approach to detect impending range shifts at an early stage will allow resource managers and researchers to better manage opportunities resulting from range-shifting species before they potentially colonize.

Improving the interpretability of climate landscape metrics: An ecological risk analysis of Japan's Marine Protected Areas.

Conservation efforts strive to protect significant swaths of terrestrial, freshwater and marine ecosystems from a range of threats. As climate change becomes an increasing concern, these efforts must take into account how resilient-protected spaces will be in the face of future drivers of change such as warming temperatures. Climate landscape metrics, which signal the spatial magnitude and direction of climate change, support a convenient initial assessment of potential threats to and opportunities within ecosystems to inform conservation and policy efforts where biological data are not available. However, inference of risk from purely physical climatic changes is difficult unless set in a meaningful ecological context. Here, we aim to establish this context using historical climatic variability, as a proxy for local adaptation by resident biota, to identify areas where current local climate conditions will remain extant and future regional climate analogues will emerge. This information is then related to the processes governing species' climate-driven range edge dynamics, differentiating changes in local climate conditions as promoters of species range contractions from those in neighbouring locations facilitating range expansions. We applied this approach to assess the future climatic stability and connectivity of Japanese waters and its network of marine protected areas (MPAs). We find 88% of Japanese waters transitioning to climates outside their historical variability bounds by 2035, resulting in large reductions in the amount of available climatic space potentially promoting widespread range contractions and expansions. Areas of high connectivity, where shifting climates converge, are present along sections of the coast facilitated by the strong latitudinal gradient of the Japanese archipelago and its ocean current system. While these areas overlap significantly with areas currently under significant anthropogenic pressures, they also include much of the MPA network that may provide stepping-stone protection for species that must shift their distribution because of climate change.

Ecological and methodological drivers of species' distribution and phenology responses to climate change.

Climate change is shifting species' distribution and phenology. Ecological traits, such as mobility or reproductive mode, explain variation in observed rates of shift for some taxa. However, estimates of relationships between traits and climate responses could be influenced by how responses are measured. We compiled a global data set of 651 published marine species' responses to climate change, from 47 papers on distribution shifts and 32 papers on phenology change. We assessed the relative importance of two classes of predictors of the rate of change, ecological traits of the responding taxa and methodological approaches for quantifying biological responses. Methodological differences explained 22% of the variation in range shifts, more than the 7.8% of the variation explained by ecological traits. For phenology change, methodological approaches accounted for 4% of the variation in measurements, whereas 8% of the variation was explained by ecological traits. Our ability to predict responses from traits was hindered by poor representation of species from the tropics, where temperature isotherms are moving most rapidly. Thus, the mean rate of distribution change may be underestimated by this and other global syntheses. Our analyses indicate that methodological approaches should be explicitly considered when designing, analysing and comparing results among studies. To improve climate impact studies, we recommend that (1) reanalyses of existing time series state how the existing data sets may limit the inferences about possible climate responses; (2) qualitative comparisons of species' responses across different studies be limited to studies with similar methodological approaches; (3) meta-analyses of climate responses include methodological attributes as covariates; and (4) that new time series be designed to include the detection of early warnings of change or ecologically relevant change. Greater consideration of methodological attributes will improve the accuracy of analyses that seek to quantify the role of climate change in species' distribution and phenology changes.

Temperature tracking by North Sea benthic invertebrates in response to climate change.

Climate change is a major threat to biodiversity and distributions shifts are one of the most significant threats to global warming, but the extent to which these shifts keep pace with a changing climate is yet uncertain. Understanding the factors governing range shifts is crucial for conservation management to anticipate patterns of biodiversity distribution under future anthropogenic climate change. Soft-sediment invertebrates are a key faunal group because of their role in marine biogeochemistry and as a food source for commercial fish species. However, little information exists on their response to climate change. Here, we evaluate changes in the distribution of 65 North Sea benthic invertebrate species between 1986 and 2000 by examining their geographic, bathymetric and thermal niche shifts and test whether species are tracking their thermal niche as defined by minimum, mean or maximum sea bottom (SBT) and surface (SST) temperatures. Temperatures increased in the whole North Sea with many benthic invertebrates showing north-westerly range shifts (leading/trailing edges as well as distribution centroids) and deepening. Nevertheless, distribution shifts for most species (3.8-7.3 km yr(-1) interquantile range) lagged behind shifts in both SBT and SST (mean 8.1 km yr(-1)), resulting in many species experiencing increasing temperatures. The velocity of climate change (VoCC) of mean SST accurately predicted both the direction and magnitude of distribution centroid shifts, while maximum SST did the same for contraction of the trailing edge. The VoCC of SBT was not a good predictor of range shifts. No good predictor of expansions of the leading edge was found. Our results show that invertebrates need to shift at different rates and directions to track the climate velocities of different temperature measures, and are therefore lagging behind most temperature measures. If these species cannot withstand a change in thermal habitat, this could ultimately lead to a drop in benthic biodiversity.

Environmental factors have stronger effects than biotic processes in patterns of intertidal populations along the southeast coast of Brazil.

Rocky shore communities are shaped by complex interactions among environmental drivers and a range of biological processes. Here, we investigated the importance of abiotic and biotic drivers on the population structure of key rocky intertidal species at 62 sites, spanning ∼50% of the Brazilian rocky shoreline (i.e., ∼500 km). Large-scale population patterns were generally explained by differences in ocean temperature and wave exposure. For the gastropod species Lottia subrugosa, differences at smaller scales (i.e., 0.1-1 km) were better explained by other abiotic influences such as freshwater discharge and substrate roughness. Based on the general population patterns of intertidal species identified, three main oceanographic groups were observed: a cold-oligotrophic grouping at northern sites (Lakes sub-region), a eutrophic group associated with large estuaries and urban zones (Santos and Guanabara bays); and a transitional warm-water group found between the two more productive areas. Larger individuals of Stramonita brasiliensis, L. subrugosa and Echinolittorina lineolata were generally found in the cold-oligotrophic system (i.e., upwelling region), while small suspension feeders dominate the warm-eutrophic systems. Evidence of bottom-up regulation was not observed, and top-down regulation effects were only observed between the whelk S. brasiliensis and its mussel prey Pernaperna. Environmental drivers as compared to biotic interactions, therefore, play a key role determining the population structure of multiple intertidal species, across a range of spatial scales along the SW Atlantic shores.

Global impacts of marine heatwaves on coastal foundation species.

With increasingly intense marine heatwaves affecting nearshore regions, foundation species are coming under increasing stress. To better understand their impacts, we examine responses of critical, habitat-forming foundation species (macroalgae, seagrass, corals) to marine heatwaves in 1322 shallow coastal areas located across 85 marine ecoregions. We find compelling evidence that intense, summer marine heatwaves play a significant role in the decline of foundation species globally. Critically, detrimental effects increase towards species warm-range edges and over time. We also identify several ecoregions where foundation species don't respond to marine heatwaves, suggestive of some resilience to warming events. Cumulative marine heatwave intensity, absolute temperature, and location within a species' range are key factors mediating impacts. Our results suggest many coastal ecosystems are losing foundation species, potentially impacting associated biodiversity, ecological function, and ecosystem services provision. Understanding relationships between marine heatwaves and foundation species offers the potential to predict impacts that are critical for developing management and adaptation approaches.

Beyond climate change attribution in conservation and ecological research.

There is increasing pressure from policymakers for ecologists to generate more detailed 'attribution' analyses aimed at quantitatively estimating relative contributions of different driving forces, including anthropogenic climate change (ACC), to observed biological changes. Here, we argue that this approach is not productive for ecological studies. Global meta-analyses of diverse species, regions and ecosystems have already given us 'very high confidence' [sensu Intergovernmental Panel on Climate Change (IPCC)] that ACC has impacted wild species in a general sense. Further, for well-studied species or systems, synthesis of experiments and models with long-term observations has given us similarly high confidence that they have been impacted by regional climate change (regardless of its cause). However, the role of greenhouse gases in driving these impacts has not been estimated quantitatively. Should this be an ecological research priority? We argue that development of quantitative ecological models for this purpose faces several impediments, particularly the existence of strong, non-additive interactions among different external factors. However, even with current understanding of impacts of global warming, there are myriad climate change adaptation options already developed in the literature that could be, and in fact are being, implemented now.

Dynamic species distribution models from categorical survey data.

1. Species distribution models are static models for the distribution of a species, based on Hutchinson's niche concept. They make probabilistic predictions about the distribution of a species, but do not have a temporal interpretation. In contrast, density-structured models based on categorical abundance data make it possible to incorporate population dynamics into species distribution modelling. 2. Using dynamic species distribution models, temporal aspects of a species' distribution can be investigated, including the predictability of future abundance categories and the expected persistence times of local populations, and how these may respond to environmental or anthropogenic drivers. 3. We built density-structured models for two intertidal marine invertebrates, the Lusitanian trochid gastropods Phorcus lineatus and Gibbula umbilicalis, based on 9 years of field data from around the United Kingdom. Abundances were recorded on a categorical scale, and stochastic models for year-to-year changes in abundance category were constructed with winter mean sea surface temperature (SST) and wave fetch (a measure of the exposure of a shore) as explanatory variables. 4. Both species were more likely to be present at sites with high SST, but differed in their responses to wave fetch. Phorcus lineatus had more predictable future abundance and longer expected persistence times than G. umbilicalis. This is consistent with the longer lifespan of P. lineatus. 5. Where data from multiple time points are available, dynamic species distribution models of the kind described here have many applications in population and conservation biology. These include allowing for changes over time when combining historical and contemporary data, and predicting how climate change might alter future abundance conditional on current distributions.

The relationship between salmon (Salmo salar) farming and cell abundance of harmful algal taxa.

The effects of nutrient effluents from fish cage aquaculture are an important eutrophication concern. It has been proposed that marine fish farm derived nutrients have the potential to increase phytoplankton abundance and lead to intensification of Harmful Algal Blooms (HABs), and that these blooms may negatively impact both the finfish and the shellfish industry. This study addressed this hypothesis using farmed salmon biomass in Scottish marine waters (as a proxy for nutrient load added to the water column as a consequence of fish farming) cell abundance of HAB taxa that most frequently impact shellfish farms and human health in the region (Dinophysis spp., Alexandrium spp. and Pseudo-nitzschia spp.), and cell abundance of one phytoplankton species of particular concern to the salmon farming industry (Karenia mikimotoi). Data from a 15-year weekly HAB monitoring programme and parallel national monitoring data relating to salmon farm stocking biomass were summarised in 5 km per 5 km aggregation boxes. Linear regression models were used to assess (i) inter-annual variation in cell abundance and total annual farmed salmon biomass; (ii) intra-annual (monthly) variation in harmful phytoplankton cell abundance and salmon biomass; (iii) a further analysis included seasonal effects within the intra-annual analysis. Farmed salmon biomass alone had a non-significant effect on cell abundance of any of the studied phytoplankton taxa. In contrast, a significant effect on cell abundance was found when using location, month or season as the predictive variable. Despite the non-significant impact of fish biomass on phytoplankton counts, the relationship varied seasonally, with a different response of Dinophysis spp. indicating a taxa specific interaction. A possible explanation for the lack of a significant relationship between farmed salmon and harmful phytoplankton cell abundance is that aquaculture farms are generally located in hydrodynamically energetic locations where recurrent flushing likely allows efficient dilution of nutrients. Overall, the analyses suggest that current levels of salmon farming activities do not markedly impact the abundance of routinely monitored biotoxin producing or fish killing phytoplankton taxa in Scottish waters.

Seasonal and spatial variability in rates of primary production and detritus release by intertidal stands of Laminaria digitata and Saccharina latissima on wave-exposed shores in the northeast Atlantic.

Coastal habitats are increasingly recognized as fundamentally important components of global carbon cycles, but the rates of carbon flow associated with marine macrophytes are not well resolved for many species in many regions. We quantified density, rates of primary productivity, and detritus production of intertidal stands of two common intertidal kelp species-Laminaria digitata (oarweed) and Saccharina latissima (sugar kelp)-on four NE Atlantic rocky shores over 22 months. The density of L. digitata was greater at exposed compared to moderately exposed shores but remained consistently low for S. latissima throughout the survey period. Individual productivity and erosion rates of L. digitata did not differ between exposed and moderately exposed shores but differed across exposure levels throughout the year at moderately exposed sites only. Productivity and erosion of S. latissima remained low on moderately exposed shores and showed no clear seasonal pattern. Patterns of productivity and total detrital production (erosion and dislodgement) per m2 of both L. digitata and S. latissima followed closely that of densities per m2, peaking in May during both survey years. Temperature and light were key factors affecting the productivity rates of L. digitata and S. latissima. Erosion rates of L. digitata were affected by wave exposure, temperature, light, grazing, and epiphyte cover, but only temperature-affected erosion of S. latissima. Production of biomass and detritus was greater in L. digitata than in S. latissima and exceeded previous estimates for subtidal and warmer-water affinity kelp populations (e.g., Laminaria ochroleuca). These biogenic habitats are clearly important contributors to the coastal carbon cycle that have been overlooked previously and should be included in future ecosystem models. Further work is required to determine the areal extent of kelp stands in intertidal and shallow subtidal habitats, which is needed to scale up local production estimates to entire coastlines.

Biological Impacts of Marine Heatwaves.

Climatic extremes are becoming increasingly common against a background trend of global warming. In the oceans, marine heatwaves (MHWs)-discrete periods of anomalously warm water-have intensified and become more frequent over the past century, impacting the integrity of marine ecosystems globally. We review and synthesize current understanding of MHW impacts at the individual, population, and community levels. We then examine how these impacts affect broader ecosystem services and discuss the current state of research on biological impacts of MHWs. Finally, we explore current and emergent approaches to predicting the occurrence andimpacts of future events, along with adaptation and management approaches. With further increases in intensity and frequency projected for coming decades, MHWs are emerging as pervasive stressors to marine ecosystems globally. A deeper mechanistic understanding of their biological impacts is needed to better predict and adapt to increased MHW activity in the Anthropocene.