The accelerating rate of global change has focused attention on the cumulative impacts of novel and extreme environmental changes (i.e., stressors), especially in marine ecosystems. As integrators of local catchment and regional processes, freshwater ecosystems are also ranked highly sensitive to the net effects of multiple stressors, yet there has not been a large-scale quantitative synthesis. We analysed data from 88 papers including 286 responses of freshwater ecosystems to paired stressors, and discovered that overall, their cumulative mean effect size was less than the sum of their single effects (i.e., an antagonistic interaction). Net effects of dual stressors on diversity and functional performance response metrics were additive and antagonistic, respectively. Across individual studies, a simple votecounting method revealed that the net effects of stressor pairs were frequently more antagonistic (41%) than synergistic (28%), additive (16%) or reversed (15%). Here, we define a reversal as occurring when the net impact of two stressors is in the opposite direction (negative or positive) from that of the sum of their single effects. While warming paired with nutrification resulted in additive net effects, the overall mean net effect of warming combined 2 with a second stressor was antagonistic. Most importantly, the mean net effects across all stressor pairs and response metrics were consistently antagonistic or additive, contrasting the greater prevalence of reported synergies in marine systems. Here, a possible explanation for more antagonistic responses by freshwater biota to stressors is that the inherent greater environmental variability of smaller aquatic ecosystems fosters greater potential for acclimation and co-adaptation to multiple stressors.

Species diversity is often an implicit source of biological insurance for communities against the impacts of novel perturbations, such as the introduction of an invasive species. High environmental heterogeneity (e.g., a mountainous gradient) is expected to beget greater regional species diversity and variation in functional traits related to environmental tolerances. Thus, heterogeneous metacommunities are expected to provide more tolerant colonists that buffer stressed local communities in the absence of dispersal limitation. We tested the hypothesis that importation of a regional zooplankton pool assembled from a diverse array of lakes and ponds lessens the impacts of a novel predator on local species-poor alpine communities by increasing response diversity (i.e., diversity of tolerances to environmental change) as mediated by variation in functional traits related to predator evasion. We also tested whether impacts varied with temperature, as warming may modify (e.g., dampen or amplify) invasion effects. An eight-week factorial experiment ([fishless vs. introduced Oncorhynchus mykiss (rainbow trout)] × [ambient temperature vs. heated] × [local vs. local + regional species pool]) was conducted using 32 1,000-L mesocosms. Associations between experimental treatments and species functional traits were tested by R-mode linked to Q-mode (RLQ) and fourth-corner analyses. Although the introduced predator suppressed local species richness and community biomass, colonization by several montane zooplankters reversed these negative effects, resulting in increased species diversity and production. Invasion resistance was unaffected by higher temperatures, which failed to elicit any significance impacts on the community. We discovered that the smaller body sizes of imported species drove functional overcompensation (i.e., increased production) in invaded communities. The observed ecological surprise showed how regionally sourced biodiversity from a highly heterogeneous landscape can offset, and even reverse, the local negative impacts of an invasive species. Further, prey body size was found to be a key species trait mediating the ecological impacts of the aquatic invasive predator. Our study highlights the novel application of a functional approach to understanding the impacts of biological invasions, using species traits that pertain directly to potential responses to exotic species.

Climate warming is a ubiquitous stressor in freshwater ecosystems, yet its interactive effects with other stressors are poorly understood. We address this knowledge gap by testing the ability of three contrasting null models to predict the joint impacts of warming and a range of other aquatic stressors using a new database of 296 experimental combinations. Despite concerns that stressors will interact to cause synergisms, we found that net impacts were usually best explained by the effect of the stronger stressor alone (the dominance null model), especially if this stressor was a local disturbance associated with human land use. Prediction accuracy depended on stressor identity and how asymmetric stressors were in the magnitude of their effects. These findings suggest we can effectively predict the impacts of multiple stressors by focusing on the stronger stressor, as habitat alteration, nutrients and contamination often override the biological consequences of higher temperatures in freshwater ecosystems.

Disentangling the environmental and spatial drivers of biological communities across large scales increasingly challenges modern ecology in a rapidly changing world. Here, we investigate the hierarchical and trait-based organization of regional and local factors of zooplankton communities at a macroscale of 1240 mountain lakes and ponds spanning western North America (California, USA, to Yukon Territory, Canada). Variation partitioning was used to test the hypothesized importance of climate, connectivity, catchment features, and exotic sportfish to zooplankton betadiversity in the context of key functional traits (body size and reproductive dispersal potential) given the pronounced environmental heterogeneity (e.g. thermal gradients), topographic barriers, and legacy of stocked fish in mountainous regions. Dispersal limitation was inferred from multispecies patch connectivity estimates based on nearest and average distances to occupied patches. Environmental heterogeneity best explained community composition as catchment/lake features (morphometry, land cover, and lithology) collectively captured greater variation than did climate (temperature, precipitation, and solar radiation), local stocking, or connectivity; however, single climatic variables captured the most variation individually. Macrospatial variation by larger obligate sexual species was better explained than that by smaller cyclically parthenogenetic asexual species. Our results provide several novel insights into the macroecology of zooplankton of the North American Cordillera, demonstrating their stronger associations to climatically driven aquatic-terrestrial habitat coupling than dynamics arising from introduced salmonids, human land-use, or species dispersal. These findings highlight the clear and important role of these communities as bioindicators of the limnological impacts of accelerating rates of climate change, as their responses appear relatively not confounded by local human perturbations or dispersal limitation.

Human-assisted introductions of exotic species are a leading cause of anthropogenic change in biodiversity; however, context dependencies and interactions with co-occurring stressors impede our ability to predict their ecological impacts. The legacy of historical sportfish stocking in mountainous regions of western North America creates a unique, natural quasiexperiment to investigate factors moderating invasion impacts on native communities across broad geographic and environmental gradients.