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Evol Ecol Res 2: 791-802 (2000)     Full PDF if your library subscribes.

Species richness, species–area curves and Simpson’s paradox

Samuel M. Scheiner,1 Stephen B. Cox,2 Michael Willig,2 Gary G. Mittelbach,3 Craig Osenberg4 and Michael Kaspari5

1Department of Life Sciences (2352), Arizona State University West, P.O. Box 37100, Phoenix, AZ 85069, 2Program in Ecology and Conservation Biology, Department of Biological Sciences and The Museum, Texas Tech University, Lubbock, TX 79409, 3W.K. Kellogg Biological Station, 3700 E. Gull Lake Drive, Michigan State University, Hickory Corners, MI 49060, 4Department of Zoology, University of Florida, Gainesville, FL 32611 and 5Department of Zoology, University of Oklahoma, Norman, OK 73019, USA

Address all correspondence to Samuel M. Scheiner, National Science Foundation, 4201 Wilson Boulevard, Arlington, VA 22230, USA.


A key issue in ecology is how patterns of species diversity differ as a function of scale. The scaling function is the species–area curve. The form of the species–area curve results from patterns of environmental heterogeneity and species dispersal, and may be system-specific. A central concern is how, for a given set of species, the species–area curve varies with respect to a third variable, such as latitude or productivity. Critical is whether the relationship is scale-invariant (i.e. the species–area curves for different levels of the third variable are parallel), rank-invariant (i.e. the curves are non-parallel, but non-crossing within the scales of interest) or neither, in which case the qualitative relationship is scale-dependent. This recognition is critical for the development and testing of theories explaining patterns of species richness because different theories have mechanistic bases at different scales of action. Scale includes four attributes: sample-unit, grain, focus and extent. Focus is newly defined here. Distinguishing among these attributes is a key step in identifying the probable scale(s) at which ecological processes determine patterns.

Keywords: combining data, productivity, scale, species–area curve, species diversity, species richness.

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