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There are many sample designs which can be employed to assess the spatial patterns<br />

of marine benthic populations and communities, each of them being more or less<br />

suitable for different habitats and different scales (Andrew and Mapstone, 1987).<br />

Contiguous sample designs give less equivocal assessments of spatial patterns since<br />

the abundances of all individuals within the extent of the survey are determined.<br />

When the extent of the survey is too large for contiguous sample designs, assessment<br />

of spatial patterns becomes more inexact.<br />

McArdle and Blackwell (1989) proposed that a systematic (grid) sample design was<br />

the most suitable for the detection of spatial patterns. This design will usually provide<br />

the smallest estimate of the mean (Milne, 1959) and patterns can be detected and<br />

displayed easily. The present results suggest that when the lag is large relative to the<br />

average patch size, however, this design may lead to equivocal results. Grid sample<br />

designs were used in this study to determine the spatial patterns of P. elegans on<br />

Drum Sands. Using mapping and spatial autocorrelation analysis, the size and spatial<br />

distribution of P. elegans patches could only be determined using a survey in which<br />

the lag, or distance between samples, was less than the mean size of the patches and<br />

less than the average distance between patches. Consequently, the 1 m survey<br />

provided important information about the size and spatial arrangement of small-scale<br />

P. elegans patches. The 8m and 40m surveys, however, revealed little information<br />

about the spatial distribution of P. elegans because the results obtained from these two<br />

surveys depended upon the actual spatial arrangement of the small-scale P. elegans<br />

patches. For example, if two neighbouring plots were located on separate P. elegans<br />

patches in either the 8m or the 40m surveys, mapping and spatial autocorrelation<br />

analysis may have erroneously indicated the presence of larger scale patches. This<br />

was revealed by the present study because P. elegans is a tube-building polychaete<br />

and, therefore, high-density areas were very visible on the sandflat. For subsurface<br />

invertebrate species, or species in which high-density areas are less readily observed, a<br />

grid-sample design can potentially lead to misleading results when patches smaller in<br />

size than the lag are present. For such species, a hierarchical sampling design may be<br />

more suitable since patchiness at a larger range of scales can be detected<br />

simultaneously (Morrisey et al., 1992). This sample design lacks the resolving power<br />

of grid-sample designs and, therefore, does not give information about the exact size<br />

236

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