Evolutionary Consequences of Food Chain Length in Kelp Forest Communities (Biogeography/Coevolution/Herbivory/Phlorotannins/Predation) PETER D
Proc. Natl. Acad. Sci. USA Vol. 92, pp. 8145-8148, August 1995 Ecology Evolutionary consequences of food chain length in kelp forest communities (biogeography/coevolution/herbivory/phlorotannins/predation) PETER D. STEINBERG*, JAMES A. ESTEStt, AND FRANK C. WINTER§ *School of Biological Sciences, University of New South Wales, P.O. Box 1, Kensington, New South Wales, 2033, Australia; tNational Biological Service, A-316 Earth and Marine Sciences Building, University of California, Santa Cruz, CA 95064; and §University of Auckland, Leigh Marine Laboratory, P.O. Box 349, Warkworth, New Zealand Communicated by Robert T. Paine, University of Washington, Seattle, WA, May 12, 1995 ABSTRACT Kelp forests are strongly influenced by mac- consistently important structuring processes throughout the roinvertebrate grazing on fleshy macroalgae. In the North food web. Under these conditions, we would predict that Pacific Ocean, sea otter predation on macroinvertebrates top-level consumers are resource limited. Consequently, the substantially reduces the intensity of herbivory on macroal- next lower trophic level should be consumer limited, in turn gae. Temperate Australasia, in contrast, has no known pred- causing the level below that (if one exists) to again be resource ator of comparable influence. These ecological and biogeo- limited. Looking downward through the food web from this graphic patterns led us to predict that (i) the intensity of very generalized perspective, a pattern emerges of strongly herbivory should be greater in temperate Australasia than in interacting couplets of adjacent trophic levels. Given these the North Pacific Ocean; thus (ii) Australasian seaweeds have circumstances, the interactive coupling between plants and been under stronger selection to evolve chemical defenses and herbivores should be strong in even-numbered systems and (iii) Australasian herbivores have been more strongly selected weak in odd-numbered systems, a prediction recently substan- to tolerate these compounds.
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