An Assessment of the Mechanistic Basis for the High Endemism and Landscape-Scale Biodiversity in Headwater Streams

Observations suggest that headwater streams have lower species diversity within a site than larger streams, but higher beta diversity, and thus gamma diversity, across a catchment. This pattern of diversity includes taxonomic richness and genetic diversity, as well as a high degree of endemism. I review several mechanisms that potentially contribute to the overall high diversity of freshwater organisms in headwaters, although these mechanisms are interdependent. These include the high numbers of headwater streams, heterogeneity of habitats and resources, founder effects, colonization dynamics, isolation, and strong selection, all leading to diversification of forms. However, riverscape diversity patterns vary across taxonomic and functional groups, highlighting that patterns of diversity are driven by different processes for different organisms. More explicitly structured sampling designs will better address patterns of taxonomic richness and for a broader range of taxa. It will be interesting to find ways to partition the relative importance of different mechanisms in contributing to the variation in diversity among headwaters. The great importance of headwater streams to global biodiversity conservation is clear, but will be more evident when better assessments of diversity patterns across these small systems are available.