Structural complexity in mussel beds: the fractal geometry of surface topography.

Seafloor topographic complexity is ecologically important because it provides habitat structure and alters boundary-layer flow over the bottom. Despite its importance, there is little agreement on how to define and measure surface complexity. The purpose of this investigation was to utilize fractal geometry of vertical cross-section profiles to characterize the surface topography of the soft-bottom mussel bed (Mytilus edulis L.) at Bob's Cove, ME, USA. Mussels there have been shown previously to have spatially ordered fractal characteristics in the horizontal plane. Two hypotheses were tested. The first was that the bed surface is fractal over the spatial scale of 1.44-200 mm, with fractal dimension less than or equal to 1.26, the value for the Koch curve, our model for bed profiles. The second was that bed surface topography (i.e., in vertical profile) is less complex than the mussel bed spatial pattern (i.e., aerial view in the horizontal plane). Both hypotheses were supported. Cross-sections of plaster casts of the bed produced 88 surface profiles, all of which were fractal over the entire spatial scale of more two orders of magnitude employed in the analysis. Fractal dimension values (D) for individual profiles ranged from 1.031 to 1.310. Fractal dimensions of entire casts ranged up to mean (1.242+/-0.046) and median (1.251) values similar to 1.26, the theoretical value of the Koch curve. The bed surface was less complex than the bed spatial pattern because every profile had D<1.36, the smallest value previously obtained from aerial views of the bed. The investigation demonstrated for the first time that surface topography of a soft-bottom mussel bed was fractal at a spatial scale relevant to hydrodynamic processes and habitat structure important for benthic organisms. The technique of using cross-section profiles from casts of the bed surface avoided possible underestimates of fractal dimension that can result from other profiling methods reported in the literature. The results demonstrate that fractal dimension can be useful in the analysis of habitat space and water flow over any irregular seafloor surface because it incorporates the size, shape, and scale of roughness elements into a simple, numerical metric.