Detecting two-dimensional spatial structure in biological data.

Cliff and Ord (1973) made versatile methods available for the direct utilization of location data in the analysis of dispersion patterns, but their monograph has as yet seen little use in the ecological literature. Application of their weighted forms of Geary's c and Moran's I indices of spatial autocorrelation to some marine benthos data demonstrates a diversity of population structure not anticipated on the basis of more common measures of pattern. These indices provide objective means to evaluate numerous recent spatial models and hypotheses in geographical ecology and genetics. The procedures are particularly attractive because (1) they efficiently utilize data which are often wasted (i.e., sample coordinates), (2) their application puts few constraints on sampling designs which would otherwise be employed, and (3) they reveal and quantify pattern differences which are not obvious to the untrained eye.