Plot shape effects on plant species diversity measurements

Abstract. Question: Do rectangular sample plots record more plant species than square plots as suggested by both empirical and theoretical studies?

Location: Grasslands, shrublands and forests in the Mediterranean-climate region of California, USA.

Methods: We compared three 0.1-ha sampling designs that differed in the shape and dispersion of 1-m² and 100-m² nested subplots. We duplicated an earlier study that compared the Whittaker sample design, which had square clustered subplots, with the modified Whittaker design, which had dispersed rectangular subplots. To sort out effects of dispersion from shape we used a third design that overlaid square subplots on the modified Whittaker design. Also, using data from published studies we extracted species richness values for 400-m² subplots that were either square or 1:4 rectangles partially overlaid on each other from desert scrub in high and low rainfall years, chaparral, sage scrub, oak savanna and coniferous forests with and without fire.

Results: We found that earlier empirical reports of more than 30% greater richness with rectangles were due to the confusion of shape effects with spatial effects, coupled with the use of cumulative number of species as the metric for comparison. Average species richness was not significantly different between square and 1:4 rectangular sample plots at either 1- or 100-m². Pairwise comparisons showed no significant difference between square and rectangular samples in all but one vegetation type, and that one exhibited significantly greater richness with squares. Our three intensive study sites appear to exhibit some level of self-similarity at the scale of 400 m², but, contrary to theoretical expectations, we could not detect plot shape effects on species richness at this scale.

Conclusions: At the 0.1-ha scale or lower there is no evidence that plot shape has predictable effects on number of species recorded from sample plots. We hypothesize that for the mediterranean-climate vegetation types studied here, the primary reason that 1:4 rectangles do not sample greater species richness than squares is because species turnover varies along complex environmental gradients that are both parallel and perpendicular to the long axis of rectangular plots. Reports in the literature of much greater species richness recorded for highly elongated rectangular strips than for squares of the same area are not likely to be fair comparisons because of the dramatically different periphery/area ratio, which includes a much greater proportion of species that are using both above and below-ground niche space outside the sample area.