Spatial patterns of invertebrate species richness in a river: the relationship between riffles and microhabitats

This study describes the pattern of invertebrate species richness in a river reach with large differences in habitat complexity at two, hierarchically nested, spatial scales. The aim was to determine whether the mass effect was likely to be increasing invertebrate species richness in epilithic microhabitats in this river. The mass effect is the process by which the species richness of a patch is increased when it acts as a ‘sink’ for species generated by ‘source’ patches. Microhabitat patch types in Mountain River, Tasmania, were distinguished on the basis of physical structure and orientation on the river bed. They were nested within two types of riffle with contrasting structural complexity: bedrock and boulder-cobble riffles. It was hypothesized that microhabitats with high species richness would act as source patches, contributing species to other microhabitats (sinks) and thereby increasing their species richness. Microhabitat sampling was carried out in four consecutive seasons and rarefaction was used to estimate riffle-scale species richness. Analysis of variance (ANOVA) was used to compare the identical microhabitats present in the contrasting riffle types, to detect evidence of the mass effect in either riffle type. The more structurally complex boulder-cobble riffles had higher species richness than did bedrock riffles. Amongst the microhabitats, the spaces beneath the cobbles had the most species. Microhabitats accounted for a higher percentage of the variation in species richness than did differences between riffles of the same type. No evidence was found for the operation of the mass effect in either riffle type. The majority of species found only in boulder-cobble riffles were unique to the beneathcobble microhabitat and appeared to be unable to colonize other microhabitats, even as transients. In Mountain River, smallscale habitat characteristics appeared to be more important than larger-scale effects in determining microhabitat species richness.