Bioerosion increases the topographic complexity of soft rock habitats, thereby increasing species diversity. This increase in species diversity may either be associated with an increase in complexity or may simply be a consequence of the increase in available surface area for colonisation. The influence of habitat modification by piddocks on intertidal species richness was investigated through field survey using fractal geometry to assess topographical complexity. The relationship between topographical complexity and species diversity was examined using the species spacing technique, which uses fractal dimensions to normalise the species richness data in relation to topographical complexity. Six sites were chosen, comprising either clay or chalk substratum, which had a range of rock hardness. Through their rock-boring activities, piddocks significantly increased the topographical complexity of the shore. Associated with this increase was an increase in species richness at all sites. Using species spacing, at 5 of the 6 sites, the increased species richness was found to be area-independent, with more species being observed than would be expected for a simple increase in surface area alone. However, piddocks are also known to significantly increase the erosion of soft rock habitats, many of which are regarded as being of particular conservation importance because of their rarity within Europe. Piddocks thus increase intertidal biodiversity while at the same time significantly contributing to erosion of the substratum.
KEY WORDS: Biodiversity · Burrow morphology · Ecosystem engineers · Fractal dimensions · Habitat complexityResale or republication not permitted without written consent of the publisher Mar Ecol Prog Ser 355: 173-182, 2008 tures, i.e. living and dead tissues, whereas allogenic engineers change the environment through their behaviour and activity. However, some species can have both effects. For example, upright bivalve shells projecting from the substratum alter the water flow regime and can provide protection from predation (autogenic engineering), while their feeding activities remove particles from the water, and the production of faeces and pseudofaeces influences the organic matter content of the sediment and, consequently, its associated infauna (allogenic engineering;Commito & Rusignuolo 2000, Norkko et al. 2006, Spooner & Vaughn 2006.Although often overlooked because of their cryptic lifestyle, piddock bivalves belonging to the family Pholadidae are among the dominant organisms of many intertidal and subtidal soft rock habitats (chalk, limestone, clays, peat and sandstone). Three species of piddock commonly occur on the English south coast: the common piddock Pholas dactylus L., the white piddock Barnea candida (L.) and the little piddock B. parva (Pennant). Piddocks create conical burrows, with a narrow entrance and a larger rounded chamber, by using their shell to mechanically erode the substratum. These rock-boring activities modify soft rock environments by creating crevices ...