Hierarchical large-scale to local-scale influence of abiotic factors in summer-fragmented Mediterranean rivers: structuring effects on fish distributions, assemblage composition and species richness

Abstract

Summer rainfall shortage and low or no flows in Mediterranean rivers make it difficult for fish to attain their final preferenda and hinder the assembling of fish communities. We investigated in this study the degree to which large-scale features of the watersheds and reach-scale habitats correlated with a non-random structure of fish species' richness, composition and distribution under these conditions by intensively sampling a river system during summer and using Geographical Information Systems data. The results of multivariate ordination and association analyses showed that multi-scale factors were hierarchically related and that they correlated with non-random fish distributions and assemblage structure. They also highlighted large-scale factors shaping reach-scale water availability for fish during summer. Fish species' richness, composition and distribution varied along a gradient of water shortage and connectivity loss which were more severe with the smaller drainage area into the reaches upstream. Fish species' distributions along these gradients were strongly correlated with both life-history traits (adult size) and biogeographical origin (native vs. exotic). The results for the summer environmental segregation of exotic versus small endemic richness characteristics indicate strong species' interactions forcing the latter into the upstream, harsher seasonal habitats. Special fitness costs are to be expected in drought years, in regulated rivers, and in general, in a climate change scenario in Mediterranean river networks. The study emphasizes that the conservation of the threatened Mediterranean freshwater fish faunas requires operating at the proper spatial scale. In particular, reach-scale habitat improvements are adequate but not enough during the summer in fragmenting river networks, unless there is proper watershed-scale management of the flow regimes.