Resilience in river ecosystems requires that organisms must persist in the face of highly dynamic hydrological and geomorphological variations. Disturbance events such as floods and droughts are postulated to shape life history traits that support resilience, but river management and conservation would benefit from greater understanding of the emergent effects in communities of river organisms. We unify current knowledge of taxonomic‐, phylogenetic‐, and trait‐based aspects of river communities that might aid the identification and quantification of resilience mechanisms. Temporal variations in river productivity, physical connectivity, and environmental heterogeneity resulting from floods and droughts are highlighted as key characteristics that promote resilience in these dynamic ecosystems. Three community‐wide mechanisms that underlie resilience are (a) partitioning (competition/facilitation) of dynamically varying resources, (b) dispersal, recolonization, and recruitment promoted by connectivity, and (c) functional redundancy in communities promoted by resource heterogeneity and refugia. Along with taxonomic and phylogenetic identity, biological traits related to feeding specialization, dispersal ability, and habitat specialization mediate organism responses to disturbance. Measures of these factors might also enable assessment of the relative contributions of different mechanisms to community resilience. Interactions between abiotic drivers and biotic aspects of resource use, dispersal, and persistence have clear implications for river conservation and management. To support these management needs, we propose a set of taxonomic, phylogenetic, and life‐history trait metrics that might be used to measure resilience mechanisms. By identifying such indicators, our proposed framework can enable targeted management strategies to adapt river ecosystems to global change.

Aquatic species living in running waters are widely acknowledged to be vulnerable to climate-induced, thermal and hydrological fluctuations. Climate changes can interact with other environmental changes to determine structural and functional attributes of communities. Although such complex interactions are most likely to occur in a multiple-stressor context as frequently encountered in large rivers, they have received little attention in such ecosystems. In this study, we aimed at specifically addressing the issue of relative long-term effects of global and local changes on benthic macroinvertebrate communities in multistressed large rivers. We assessed effects of hydroclimatic vs. water quality factors on invertebrate community structure and composition over 30 years (1979-2008) in the Middle Loire River, France. As observed in other large European rivers, water warming over the three decades (+0.9 °C between 1979-1988 and 1999-2008) and to a lesser extent discharge reduction (-80 m(3) s(-1) ) were significantly involved in the disappearance or decrease in taxa typical from fast running, cold waters (e.g. Chloroperlidae and Potamanthidae). They explained also a major part of the appearance and increase of taxa typical from slow flowing or standing waters and warmer temperatures, including invasive species (e.g. Corbicula sp. and Atyaephyra desmarestii). However, this shift towards a generalist and pollution tolerant assemblage was partially confounded by local improvement in water quality (i.e. phosphate input reduction by about two thirds and eutrophication limitation by almost one half), explaining a significant part of the settlement of new pollution-sensitive taxa (e.g. the caddisfly Brachycentridae and Philopotamidae families) during the last years of the study period. The regain in such taxa allowed maintaining a certain level of specialization in the invertebrate community despite climate change effects.

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Global change is increasing biotic homogenization globally, which modifies the functioning of ecosystems. While tendencies towards taxonomic homogenization in biological communities have been extensively studied, functional homogenization remains an understudied facet of biodiversity. Here, we tested four hypotheses related to long-term changes (1991-2016) in the taxonomic and functional arrangement of freshwater macroinvertebrate assemblages across space and possible drivers of these changes. Using data collected annually at 64 river sites in mainland New Zealand, we related temporal changes in taxonomic and functional spatial β-diversity, and the contribution of individual sites to β-diversity, to a set of global, regional, catchment and reach-scale environmental descriptors. We observed long-term, mostly climate-induced, temporal trends towards taxonomic homogenization but functional differentiation among macroinvertebrate assemblages. These changes were mainly driven by replacements of species and functional traits among assemblages, rather than nested species loss. In addition, there was no difference between the mean rate of change in the taxonomic and functional facets of β-diversity. Climatic processes governed overall population and community changes in these freshwater ecosystems, but were amplified by multiple anthropogenic, topographic and biotic drivers of environmental change, acting widely across the landscape. The functional diversification of communities could potentially provide communities with greater stability, resistance and resilience capacity to environmental change, despite ongoing taxonomic homogenization. Therefore, our study highlights a need to further understand temporal trajectories in both taxonomic and functional components of species communities, which could enable a clearer picture of how biodiversity and ecosystems will respond to future global changes. K E Y W O R D S β-diversity, biotic homogenization, climate change, freshwater macroinvertebrates, functional diversity, human disturbance | 6905 MOUTON eT al.

When investigating the fields of biogeography and macroecology, climate-and productivity-related variables are frequently identified as the strongest correlates of species-diversity patterns. These variables have been usually merged under the climate/ productivity hypothesis and describe the direct and indirect actions of climate on species. Being among the most vulnerable ecosystems to climate change, streams and rivers are expected to be influenced both by climatic and trophic (i.e. productivity-related) factors. We propose here to distinguish the relative influence of the two processes on large-scale, long-term changes in the functional diversity of freshwater invertebrate communities over two decades in France. To this end, we designed two functional indices using invertebrate traits to surrogate the respective mechanisms: climate vulnerability and feeding specialisation. Using geographically weighted regression (GWR) models, we showed that trends in both indices, along with the initial regional speciespools, have significantly contributed to the overall long-term increase in functional diversity of invertebrate communities. In addition, we highlighted a strong geographical differentiation in the contribution patterns with the climate vulnerability effect decreasing with latitude and the feeding specialisation effect being higher in headwaters than in large rivers. Finally, taking into account this non-stationarity in the ecological processes and responses using GWR models allowed explaining about 75% of the long-term changes in the community diversity. Consequently, this study offers sound perspectives in predicting the future patterns of trends in functional diversity of communities under different scenarios of environmental changes, like climate and/or land-use.

Although species traits have the potential to disentangle long-term effects of multiple, potentially confounded drivers in ecosystems, this issue has received very little attention in the literature. We aimed at filling this gap by assessing the relative effects of hydroclimatic and water quality factors on the trait composition of invertebrate assemblages over 30 years in the Middle Loire River (France). Using a priori predictions on the long-term variation of trait-based adaptations over the three decades, we evaluated the ability of invertebrate traits to indicate the effects of warming, discharge reduction and water quality improvement. Hydroclimatic and water quality factors contributed to up to 65% of the variation in trait composition. More than 70% of the initial trait response predictions made according to observed long-term hydroclimatic changes were confirmed. They supported a general climate-induced trend involving adapted resistance and resilience strategies. A partial confounding effect of water quality improvement acting on trophic processes was also highlighted, indicating that improved water quality management can significantly help to reduce some adverse effects of climate change. This trait-based approach can have wider implications for investigating long-term changes driven by multiple, potentially confounded factors, as frequently encountered in the context of global change.

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