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The large increase in interest concerning the study of spatial patterns of species and<br />

communities has undoubtedly partly resulted from the expansion of techniques<br />

available for quantitatively and qualitatively assessing patterns. Most of these<br />

techniques, for example, quadrat-variance (Greig-Smith, 1952; Upton and Fingleton,<br />

1985), spatial autocorrelation analysis (Cliff and Ord, 1973), mapping (Burrough,<br />

1987), spatially-constrained clustering (Legendre and Fortin, 1989) and ordination<br />

(Wartenberg, 1985; Ter Braak, 1986) and semi-variogram analysis (see review by<br />

Legendre and Fortin, 1989) were developed by plant ecologists and have been adapted<br />

for use in other areas of ecology. Many of these techniques have been employed in<br />

the study of spatial patterns in the marine benthic environment.<br />

Pattern and observational scale are intrinsically linked (Addicott et al., 1987; Levin,<br />

1992; Platt and Sathyendranath, 1992; Hall et al., 1993; Lawrie, 1996). Patterns<br />

evident at one scale can either disappear or appear as noise when viewed at other<br />

scales (Eckman, 1979; Valiela, 1984; McArdle et al., 1990; Dutilleul and Legendre,<br />

1993; Thrush et al., 1997a). This is because the processes which give rise to patterns<br />

in nature are scale-dependent (Legendre et al., 1997; Thrush et al., 1997b). For<br />

example, non-overlapping distributions of two species at one scale may reflect<br />

interspecific competition while at larger scales, a positive association between the two<br />

species may result from habitat selection (Wiens, 1989). Consequently, studying the<br />

relationship between the pattern observed and the process creating that pattern is<br />

problematical (McArdle et al., 1997; Thrush et al., 1997a, 1997b). Levin (1992) and<br />

Thrush et al. (1997c) suggested that recognition that different processes are relevant at<br />

different scales precludes the reductionist approach of sampling at the 'right' scale<br />

(e.g., Wiens, 1989; Kotliar and Wiens, 1990). Consequently, they proposed that<br />

ecologists should ask the question of how we can draw conclusions from one scale to<br />

another. Often, however, the scale at which one samples depends upon the questions<br />

one is attempting to answer within a particular study or upon logistical constraints.<br />

McArdle and Blackwell (1989) and Legendre et al. (1997) suggested that a good<br />

starting point before planning an experiment is the identification of the patterns that<br />

can be detected at one or several spatial scales.<br />

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