Conditional responses of organisms to habitat structure: an example from intertidal mudflats.

Habitat structure is often assumed to be a predictor of habitat function. However, habitat structure may be insufficient to predict the functional significance of a habitat if the level of resources in the habitat is a consequence of the interaction between the habitat structure and physical or biological factors. In this study, we investigated whether depressions in tidal flat sediments generated by stout razor clams, Tagelus plebeius, affect the spatial patterns of pit digging by deposit-feeding burrowing crabs, Chasmagnathus granulata. The pits dug by crabs while feeding overlapped with clam siphon holes at a frequency higher than expected at random, and measurements of pit-digging by crabs integrated over several days indicated a higher frequency of feeding in the sediment of depressions. The daily frequency of pit-digging by crabs in depressions was positively related to the organic matter content of their sediments, but was significantly higher than the frequency of pit-digging away from clam siphon holes only after events of high bedload sediment transport, when the organic matter in the sediments of these depressions peaked. This example demonstrates the conditional nature of the relationship between habitat structure and function by illustrating how a physical process--bedload sediment transport--may introduce variation in the function that depressions play as feeding sites for burrowing crabs. Published information suggests that such conditional responses of organisms to habitat structure: (1) occur in a variety of habitats; (2) involve a variety of structures either of biotic or abiotic origin; and (3) are the consequence of either physical or biological controls that vary in importance according to the general mechanism through which habitat structure affect resources. This broad experimental evidence suggests that the accuracy of predictive models linking habitat structure and function can be improved by incorporating a mechanistic perspective that allows recognition of the potential for conditional responses of organisms to habitat structure.