Biological modifiers of marine benthic seascapes: their role as ecosystem engineers

Benthic organisms in marine ecosystems modify the environment on different spatial and temporal scales. These modifications, many of which are initially at a microscale, are likely to have large scale effects on benthic seascapes. This is especially so if the species are ecosystem engineers. Most species of infaunal and epifaunal invertebrates and macrophytes contribute at a geophysical or geochemical level. Microorganisms also play a key but currently neglected role. In the intertidal and immediately sublittoral zone, algae and seagrasses, and mussels in mussel beds have received considerable attention. A substantial fossil record also exists. Mathematical modelling of these systems is still in its infancy, although several sophisticated mathematical tools have been applied. The effects of bioturbation and of microorganisms have been less studied, and little is known about the activities of benthic organisms in the deep sea. This paper addresses all these effects, and places them in the context of large scale benthic seascapes and of the extensive literature on species defined as ecosystem engineers in the sea