Hierarchical patterns of invertebrate assemblage structure in stony upland streams change with time and flow permanence

Summary 1. Studies in several parts of the world have examined variation in univariate descriptors of macroinvertebrate assemblage structure in perennially flowing stony streams across hierarchies of spatial scale using nested analyses of variance. However, few have investigated whether this spatial variation changes with time or whether these results are representative of habitats other than riffles or of other stream types, such as intermittently flowing streams. 2. We describe patterns in taxon richness and abundance from two sets of samples from stony streams in the Otway Range and the Grampians Range, Victoria, Australia, collected using hierarchical designs. Sampling of riffles was repeated in the Otways, to determine whether spatial patterns were consistent among times. In the Grampians, spatial patterns were compared between intermittent and perennially flowing streams (stream type) by sampling pools. 3. In the Otways streams, most variation in the dependent variables occurred between sample units. Patterns of variation among the other scales (streams, segments, riffles, groups of stones) were not consistent between sampling times, suggesting that they may have little ecological significance. 4. In the Grampians streams, variation in macroinvertebrate taxon richness and abundance differed significantly between replicate streams within each stream type but not between stream types or pools. The largest source of variation in taxon richness was stream type. Little variation occurred among sample units. 5. The pattern of most variation occurring among sample units is robust both to differences in the method of sampling and different dependent variables among studies and increasingly appears to be a property of riffles in stony, perennial upland streams. High variation among sample units (residual variation) limits the explanatory power of linear models and therefore, where samples are from a single sampling time, small but significant components of variation are unlikely to represent features of assemblage structure that will be stable over time.