Effects of water temperature on freshwater macroinvertebrates: a systematic review.

Water temperature is one of the main abiotic factors affecting the structure and functioning of aquatic ecosystems and its alteration can have important effects on biological communities. Macroinvertebrates are excellent bio-indicators and have been used for decades to assess the status of aquatic ecosystems as a result of environmental stresses; however, their responses to temperature are poorly documented and have not been systematically evaluated. The aims of this review are: (i) to collate and summarize responses of freshwater macroinvertebrates to different temperature conditions, comparing the results of experimental and theoretical studies; (ii) to understand how the focus of research on the effects of temperature on macroinvertebrates has changed during the last 51 years; and (iii) to identify research gaps regarding temperature responses, ecosystem types, organism groups, spatiotemporal scales, and geographical regions to suggest possible research directions. We performed a comparative assessment of 223 publications that specifically consider freshwater macroinvertebrates and address the effects of temperature. Short-term studies performed in the laboratory and focusing on insects exposed to a range of temperatures dominated. Field studies were carried out mainly in Europe, at catchment scale and almost exclusively in rivers; they mainly investigated responses to water thermal regime at the community scale. The most frequent biological responses tested were growth rate, fecundity and the time and length of emergence, whereas ecological responses mainly involved composition, richness, and distribution. Thermal research on freshwater macroinvertebrates has undergone a shift since the 2000s when studies involving extended spatiotemporal scales and investigating the effects of global warming first appeared. In addition, recent studies have considered the effects of temperature at genetic and evolutionary scales. Our review revealed that the effects of temperature on macroinvertebrates are manifold with implications at different levels, from genes to communities. However, community-level physiological, phenological and fitness responses tested on individuals or populations should be studied in more detail given their macroecological effects are likely to be enhanced by climate warming. In addition, most field studies at regional scales have used air temperature as a proxy for water temperature; obtaining accurate water temperature data in future studies will be important to allow proper consideration of the spatial thermal heterogeneity of water bodies and any effects on macroinvertebrate distribution patterns. Finally, we found an uneven number of studies across different ecosystems and geographic areas, with lentic bodies and regions outside the West underrepresented. It will also be crucial to include macroinvertebrates of high-altitude and tropical areas in future work because these groups are most vulnerable to climate warming for multiple reasons. Further studies on temperature-macroinvertebrate relationships are needed to fill the current gaps and facilitate appropriate conservation strategies for freshwater ecosystems in an anthropogenic-driven era.