Abstract
Metacommunity theory emphasizes that local communities are jointly affected by environmental filtering and spatial processes. However, the roles of spatial processes are often given insufficient attention in bioassessment practices, which may bias the assessments of ecological status based on biotic metrics. Here, we quantified the relative importance and the seasonal stability of spatial processes, natural conditions and human-induced factors in structuring variation in different bioassessment metrics based on macroinvertebrate communities. Our study systems were two extensively sampled large and shallow lakes with strong nutrient gradients related to human disturbance. The roles of different drivers were examined for three kinds of indicators: general diversity, trait-based and taxonomic distinctness metrics, and their performance in characterizing human disturbance was evaluated. Overall, human-induced and spatial factors were all important in explaining variation in the three types of bioassessment metrics. Contrary to our expectations, however, we found that the importance of spatial processes on bioassessment metrics can be comparable to the effects of local environmental conditions at the within-lake scale. Furthermore, the results showed substantial seasonal variability in the relative roles of different drivers, which might be linked to life-cycle seasonality of macroinvertebrates. As expected, trait-based metrics generally were best associated with human-induced variables in both lakes, whereas general diversity and taxonomic distinctness metrics performed poorly. The low effectiveness of taxonomic distinctness metrics might due to low species richness associated with high nutrient levels. To conclude, our results suggest that bioassessment cannot exclusively rely on the idea of environmental filtering even if we focus on fine spatial scales. We hence strongly urge that spatial processes, natural drivers and temporal variability should be better considered in combination in the development and application of bioassessment approaches. In addition, taxonomic distinctness measures should be used with caution, especially for the ecosystems and organism groups typically characterized by low species richness.
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