The integral structure of a benthic infaunal assemblage

It has been suggested that the scaling relationships of many features of the physical environment and biological traits are fractal-like, but for the marine benthic infauna certain aspects of the environment clearly are not. These include temporal features such as the cycles of annual climate, primary production and tides, and also some spatial features such as sediment granulometry and the size of the primary producers that constitute the food supply. We explicitly addressed these issues by determining the degree to which infaunal assemblage structure (diversity, species composition, spatial pattern) varies with mesh size, sample size and sample dispersion within an apparently homogeneous area of coarse intertidal sand in the Isles of Scilly, UK. Samples were extracted using a standard range of 5 mesh sizes (63, 125, 250, 500, 1000 μm), with the sample areas and distances between samples scaled to the mesh size. All metazoans were identified to species level. Diversity and species composition did not show a gradual and even degree of change over the size range. Instead, they showed a dramatic stepwise change between the 250 and 500 μm mesh size samples, being relatively constant in the < 500 and > 500 μm categories, with diversity higher in the former. Higher proportions of species in the < 500 μm categories had values of an index of dispersion significantly different to 1 than among species in the > 500 μm categories. This suggests a fractal structure within but not between the < 500 and > 500 μm body size categories. The implications of this for rapid diversity assessment by extrapolation between size classes are discussed. Although the interplay between 3 and 2 dimensional processes in what is a essentially a 2-D study may account for some of the observations, comparative studies suggest that these patterns do not simply correspond to the physical scaling of habitat complexity, and they must therefore relate to some more universal scaling relationships that are not fractal-like. We suggest that the important relationships are those between body size and various biological characteristics such as feeding behaviour, reproductive mode and life history as they are affected by the spatial and temporal structure of the environment.