Paine, Robert Treat, 3d, 1933–2016, American ecologist, b. Cambridge, Mass., Ph.D. Univ. of Michigan, 1961. He was on the faculty of the Univ. of Washington from 1962 to 1998. Paine's major contribution to the field of ecology was his concept of “keystone species,” species have much more of an impact on their ecosystems than others, an impact that is often independent of the species predominance or productivity in the ecosystem. Paine conducted much of his research on intertidal systems along the shores of the Olympic Peninsula in Washington state, where he studied ochre sea stars and sea otters and observed how their removal from the environment encouraged the proliferaton of other species, leading to a disruption of the abundance, diversity, and distribution of species within that ecosystem.
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