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American Zoologist 1981 21(4):845-852; doi:10.1093/icb/21.4.845
© 1981 by The Society for Integrative and Comparative Biology
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Environs: The Superniches of Ecosystems1

BERNARD C. PATTEN
Department of Zoology and Institute of Ecology, University of Georgia Athens, Georgia 30602

Evolution proceeds by natural selection of heritable variations of individual organisms based on direct influences of environment. However, indirect effects probably vastly outweigh direct ones in ecosystems. Therefore, why is evolution based on direct effects only? The ecological niche represents the point of direct contact between organisms and their environments. To encompass indirect influences, niches are extended to new structures, environs, which are units of organism-environment coevolution. The motive force for coevolution is closure of outputs back upon inputs of the organism members of ecosystems. Closure is achieved by biogeochemical cycling and feedback interactions, direct and indirect, between organisms. To the extent that closure does not occur, there is no imperative for organism-environment coevolution. Coevolution at the system level based on indirect effects is compatible with normal evolution at the individual organism level based on direct effects. The organism is the unit of the latter, but environs are the unit of coevolution.


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