Habitat structure and invertebrate assemblages on stream stones: A multivariate view from the riffle

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Abstract
Highly structured habitats, those with complex or heterogeneous physical structure, commonly contain more species than simply structured ones. However, tests of this relationship have been hampered by habitat‐specific definitions and measurements of habitat structure and, in stream studies, by insufficient information on spatial variation in faunal abundances. We sampled the fauna of 90 stones collected from an order 4 section of the Taggerty and Steavenson Rivers, southeastern Australia. The spatially nested sampling design encompassed three spatial scales (sites within the same stream order, riffles at the same site and groups of stones within the same riffle) but no temporal replication, as we were only able to sample once. Habitat structure of stones was quantified by measuring the amount and type of epilithon on stones, together with stone shapes, sizes, textures and surface complexities. For the latter two attributes, we used a new method, involving image analysis, which can be used to quantify habitat structure in comparable ways in other systems. Species richness (S) was related to some measures of habitat structure, being higher on stones with a rough texture and weakly related to the amount of epilithon present. Total numbers of individuals (N) were also higher on rough stones, and faunal composition (as quantified by hybrid multidimensional scaling) was related to abundance of epilithon. However, flow environments, as quantified by water velocities and depths, were also related to faunal composition. Riffles varied in both faunal densities and near‐bed flow environment, but not in any consistent way. Such variability means that individual riffles cannot be reasonably used to ‘represent’ stream sections or orders, an assumption commonly made in stream studies. Neglect of such issues has also resulted in poor conceptual integration between large‐ and small‐scale studies in stream ecology.