ORGANISMS AS ECOSYSTEM ENGINEERS

Ecosystem engineers are organisms that directly or indirectly modulate the availability of resources to other species, by causing physical state changes in biotic or abiotic materials. In so doing they modify, maintain and create habitats. Autogenic engineers (e.g. corals, or trees) change the environment via their own physical structures (i.e. their living and dead tissues). Allogenic engineers (e.g. woodpeckers, beavers) change the environment by transforming living or non-living materials from one physical state to another, via mechanical or other means. The direct provision of resources to other species, in the form of living or dead tissues is not engineering. Organisms act as engineers when they modulate the supply of a resource or resources other than themselves. We recognise and define five types of engineering and provide examples. Humans are allogenic engineers par excellence, and also mimic the behaviour of autogenic engineers, for example by constructing glasshouses. We explore related concepts including the notions of extended phenotypes and keystone species. Some (but not all) products of ecosystem engineering are extended phenotypes. Many (perhaps most) impacts of keystone species include not only trophic effects, but also engineers and engineering. Engineers differ in their impacts. The biggest effects are attributable to species with large per capita impacts, living at high densities, over large areas for a long time, giving rise to structures that persist for millennia and that modulate many resource flows (e.g. mima mounds created by fossorial rodents). The ephemeral nests constructed by small, passerine birds lie at the opposite end of this continuum. We provide a tentative research agenda for an exploration of the phenomenon of organisms as ecosystem engineers, and suggest that all habitats on earth support, and are influenced by, ecosystem engineers.