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CHAPTER 2<br />

THE SMALL- To MESO-SCALE SPATIAL DISTRIBUTION OF<br />

MARINE BENTHIC INVERTEBRATES ON DRUM SANDS, WITH<br />

PARTICULAR REFERENCE TO PYGOSPIO ELEGANS<br />

INTRODUCTION<br />

Spatial heterogeneity is a fundamental characteristic of animal and plant populations<br />

(Connell, 1963; Taylor, 1984), as well as the physical environment which the<br />

organisms inhabit. The assessment of the spatial variability of some variable, e.g.,<br />

abundance of an organism, is often the starting point from which questions and<br />

hypotheses about important processes on the population or community level are<br />

generated (Andrew and Mapstone, 1987; Myers and Giller, 1988; Levin, 1992).<br />

Furthermore, Hall et al. (1993) recommended that an analysis of spatial pattern in<br />

both field experiments and survey programmes should be a priority for benthic<br />

ecologists.<br />

There are numerous techniques available for the identification of spatial pattern, such<br />

as spectral analysis (Renshaw and Ford, 1984), quadrat-variance (Greig-Smith, 1952),<br />

spatial autocorrelation analysis (Cliff and Ord, 1973; Legendre and Fortin, 1989),<br />

dispersion indices (Morisita, 1962; Elliot, 1977; Krebs, 1989) and nested hierarchical<br />

analysis of variance (Morrisey et al., 1992; Lindegarth et al., 1995). These different<br />

techniques all provide different information on spatial pattern, require different<br />

sampling strategies and are more or less suitable at different spatial scales. For<br />

example, while a hierarchical nested analysis of variance requires a nested sampling<br />

design and is most suitable for determining large-scale (100m-kms) variation<br />

(Lindegarth et al., 1995), spatial autocorrelation analysis, which requires a more<br />

regular sampling design (Thrush et al., 1989) determines the underlying spatial<br />

structure and is more suitable for smaller-scale investigations (Morrisey et al., 1992).<br />

Methods such as Greig-Smith's (1952) quadrat-variance technique, or related<br />

techniques such as the paired-quadrat variance (PQV) and two-term local quadrat<br />

15

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