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Invariant scaling relations across tree-dominated communities

Nature volume 410, pages 655660 (05 April 2001) | Download Citation


  • A Corrigendum to this article was published on 16 October 2003


Organizing principles are needed to link organismal, community and ecosystem attributes across spatial and temporal scales. Here we extend allometric theory—how attributes of organisms change with variation in their size—and test its predictions against worldwide data sets for forest communities by quantifying the relationships among tree size–frequency distributions, standing biomass, species number and number of individuals per unit area. As predicted, except for the highest latitudes, the number of individuals scales as the -2 power of basal stem diameter or as the -3/4 power of above-ground biomass. Also as predicted, this scaling relationship varies little with species diversity, total standing biomass, latitude and geographic sampling area. A simulation model in which individuals allocate biomass to leaf, stem and reproduction, and compete for space and light obtains features identical to those of a community. In tandem with allometric theory, our results indicate that many macroecological features of communities may emerge from a few allometric principles operating at the level of the individual.

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We thank D. Ackerly, J. Ackerman, J. Brown, W. Chaloner, K. Harms, J. Haskell, D. Post, T. Phillips, P. Stoll, J. Valentine, R. Whittaker, J. Williams and M. Willig for discussions or comments on the manuscript; J. Miller, P. Raven and the Missouri Botanical Garden for help in disseminating the Gentry database; and J. Pringle and A. Landcaster for programming assistance with SWARM. This work stems in part from the Body Size in Ecology and Evolution Working Group sponsored by The National Center for Ecological Analysis and Synthesis (NCEAS, a centre funded by the NSF, the University of California Santa Barbara and the State of California). In particular we wish to acknowledge J. Haskell for critical help with the Gentry data; G. West and B. Tiffney for support; and F. Smith and the members of the body size working group for their input to this project. B.J.E. was supported by the NSF, the Santa Fe Institute and NCEAS. K.J.N. was supported by an Alexander von Humboldt Forschungspreis and NYS Hatch grant funds.

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  1. *Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, Arizona 85721, USA; National Center for Ecological Analysis and Synthesis, University of California, Santa Barbara, California, USA

    • Brian J. Enquist
  2. †Department of Plant Biology, Cornell University, Ithaca, New York 14853, USA

    • Karl J. Niklas


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Correspondence to Brian J. Enquist.

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