Landscape heterogeneity and the biodiversity of Arctic stream communities: a habitat template analysis

Abstract
We predicted that substratum freezing and instability are major determinants of the variability of stream community structure in Arctic Alaska. Their effects were conceptualized as a two-dimensional habitat template that was assessed using a natural experiment based on five stream types (mountain-spring, tundra-spring, tundra, mountain, glacier). Detrended correspondence analysis (DCA) indicated distinct macroinvertebrate assemblages for each stream type. The contribution of functional feeding groups to assemblage biomass varied systematically among stream types, indicating that structure and function are linked. Assemblage position within a DCA biplot was used to assess factors controlling its structure. Springs separated from other stream types along a gradient of nutrient concentration and freezing probability. Glacier and mountain streams separated from springs and tundra streams along a gradient of substratum instability and freezing probability. Owing to differences in sources of discharge to streams, the effects of nutrients and substratum stability could not be separated from freezing. Although many factors likely contribute to the variability of Arctic stream communities, the major determinants may be conceptualized as a template structured by gradients in (i) nutrient supply and substratum freezing and (ii) substratum instability and substratum freezing. This template provides a basis for predicting the response of Arctic stream communities to climate change.