Human activities are fundamentally altering biodiversity. Projections of declines at the global scale are contrasted by highly variable trends at local scales, suggesting that biodiversity change may be spatially structured. Here, we examined spatial variation in species richness and composition change using more than 50,000 biodiversity time series from 239 studies and found clear geographic variation in biodiversity change. Rapid compositional change is prevalent, with marine biomes exceeding and terrestrial biomes trailing the overall trend. Assemblage richness is not changing on average, although locations exhibiting increasing and decreasing trends of up to about 20% per year were found in some marine studies. At local scales, widespread compositional reorganization is most often decoupled from richness change, and biodiversity change is strongest and most variable in the oceans.
Biodiversity enhances many of nature's benefits to people, including the regulation of climate and the production of wood in forests, livestock forage in grasslands and fish in aquatic ecosystems. Yet people are now driving the sixth mass extinction event in Earth's history. Human dependence and influence on biodiversity have mainly been studied separately and at contrasting scales of space and time, but new multiscale knowledge is beginning to link these relationships.
A rich body of knowledge links biodiversity to ecosystem functioning (BEF), but it is primarily focused on small scales. We review the current theory and identify six expectations for scale dependence in the BEF relationship: (1) a nonlinear change in the slope of the BEF relationship with spatial scale; (2) a scale‐dependent relationship between ecosystem stability and spatial extent; (3) coexistence within and among sites will result in a positive BEF relationship at larger scales; (4) temporal autocorrelation in environmental variability affects species turnover and thus the change in BEF slope with scale; (5) connectivity in metacommunities generates nonlinear BEF and stability relationships by affecting population synchrony at local and regional scales; (6) spatial scaling in food web structure and diversity will generate scale dependence in ecosystem functioning. We suggest directions for synthesis that combine approaches in metaecosystem and metacommunity ecology and integrate cross‐scale feedbacks. Tests of this theory may combine remote sensing with a generation of networked experiments that assess effects at multiple scales. We also show how anthropogenic land cover change may alter the scaling of the BEF relationship. New research on the role of scale in BEF will guide policy linking the goals of managing biodiversity and ecosystems.
The metacommunity concept has the potential to integrate local and regional dynamics within a general community ecology framework. To this end, the concept must move beyond the discrete archetypes that have largely defined it (e.g. neutral vs. species sorting) and better incorporate local scale species interactions and coexistence mechanisms. Here, we present a fundamental reconception of the framework that explicitly links local coexistence theory to the spatial processes inherent to metacommunity theory, allowing for a continuous range of competitive community dynamics. These dynamics emerge from the three underlying processes that shape ecological communities: (1) density-independent responses to abiotic conditions, (2) density-dependent biotic interactions and (3) dispersal. Stochasticity is incorporated in the demographic realisation of each of these processes. We formalise this framework using a simulation model that explores a wide range of competitive metacommunity dynamics by varying the strength of the underlying processes. Using this model and framework, we show how existing theories, including the traditional metacommunity archetypes, are linked by this common set of processes. We then use the model to generate new hypotheses about how the three processes combine to interactively shape diversity, functioning and stability within metacommunities.
The effects of global and local environmental changes are transmitted through networks of interacting organisms to shape the structure of communities and the dynamics of ecosystems. We tested the impact of elevated temperature on the top-down and bottom-up forces structuring experimental freshwater pond food webs in western Canada over 16 months. Experimental warming was crossed with treatments manipulating the presence of planktivorous fish and eutrophication through enhanced nutrient supply. We found that higher temperatures produced top-heavy food webs with lower biomass of benthic and pelagic producers, equivalent biomass of zooplankton, zoobenthos and pelagic bacteria, and more pelagic viruses. Eutrophication increased the biomass of all organisms studied, while fish had cascading positive effects on periphyton, phytoplankton and bacteria, and reduced biomass of invertebrates. Surprisingly, virus biomass was reduced in the presence of fish, suggesting the possibility for complex mechanisms of top-down control of the lytic cycle. Warming reduced the effects of eutrophication on periphyton, and magnified the already strong effects of fish on phytoplankton and bacteria. Warming, fish and nutrients all increased whole-system rates of net production despite their distinct impacts on the distribution of biomass between producers and consumers, plankton and benthos, and microbes and macrobes. Our results indicate that warming exerts a host of indirect effects on aquatic food webs mediated through shifts in the magnitudes of top-down and bottom-up forcing.
Climate warming is occurring in concert with other anthropogenic changes to ecosystems. However, it is unknown whether and how warming alters the importance of top-down vs. bottom-up control over community productivity and variability. We performed a 16-month factorial experimental manipulation of warming, nutrient enrichment, and predator presence in replicated freshwater pond mesocosms to test their independent and interactive impacts. Warming strengthened trophic cascades from fish to primary producers, and it decreased the impact of eutrophication on the mean and temporal variation of phytoplankton biomass. These impacts varied seasonally, with higher temperatures leading to stronger trophic cascades in winter and weaker algae blooms under eutrophication in summer. Our results suggest that higher temperatures may shift the control of primary production in freshwater ponds toward stronger top-down and weaker bottom-up effects. The dampened temporal variability of algal biomass under eutrophication at higher temperatures suggests that warming may stabilize some ecosystem processes.
Increases in atmospheric temperature and nutrients from land are thought to be promoting the expansion of harmful cyanobacteria in lakes worldwide, yet to date there has been no quantitative synthesis of long-term trends. To test whether cyanobacteria have increased in abundance over the past ~ 200 years and evaluate the relative influence of potential causal mechanisms, we synthesised 108 highly resolved sedimentary time series and 18 decadal-scale monitoring records from north temperate-subarctic lakes. We demonstrate that: (1) cyanobacteria have increased significantly since c. 1800 ce, (2) they have increased disproportionately relative to other phytoplankton, and (3) cyanobacteria increased more rapidly post c. 1945 ce. Variation among lakes in the rates of increase was explained best by nutrient concentration (phosphorus and nitrogen), and temperature was of secondary importance. Although cyanobacterial biomass has declined in some managed lakes with reduced nutrient influx, the larger spatio-temporal scale of sedimentary records show continued increases in cyanobacteria throughout the north temperate-subarctic regions.
Habitat loss fragments metacommunities, altering the movement of species between previously connected habitat patches. The consequences of habitat loss for ecosystem functioning depend, in part, on how these changes in connectivity alter the spatial insurance effects of biodiversity. Spatial insurance is the maintenance of biodiversity and stable ecosystem functioning in changing environments that occurs when species are able to move between local habitat patches in order to track conditions to which they are adapted. Spatial insurance requires a combination of species sorting dynamics, which allow species to disperse to habitats where they are productive, and mass effect dynamics, where dispersal allows species to persist in marginal habitats where environmental conditions do not support growth. Here we use a spatially explicit metacommunity model to show that the relative contribution of species sorting and mass effects to spatial insurance changes with the rate of dispersal. We then simulate different sequences of habitat loss by removing habitat patches based on their betweenness centrality (the degree to which a patch serves as a connection between other patches in the metacommunity). We demonstrate that the sequence of habitat loss has a large, non‐linear impact on diversity, ecosystem functioning and stability. Spatial insurance is lost because habitat fragmentation impedes species sorting, while promoting mass effects and dispersal limitation. We find that species sorting dynamics, and thus spatial insurance, are most robust to the removal of habitat patches with low betweenness centrality. These findings advance our understanding of how habitat connectivity facilitates the maintenance of biodiversity and ecosystem functioning, and may prove useful for the design of habitat networks.
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