Tree dispersion in oak-dominated forests along an environmental gradient.

Spatial pattern was analyzed in seventeen stands of oak-dominated forest to address the hypothesis that species tended to be aggregated under favorable conditions and widely spaced in xeric, nutrient poor conditions. Trees were sampled at 80-100 points in each stand with the distance-to-nearest neighbor method. Soil samples were collected in each stand for analysis of total nitrogen, total phosphorus, total potassium, soil pH, soil texture, and soil organic matter. Growing season precipitation was also recorded from climate stations near each stand. Quercus stellata (Wang.) dominated 10 stands, Q. marilandica (Muenchh.) dominated three stands and these species were codominant in four stands. Principal components analysis identified a soil texture/fertility gradient across the study area. Quercus stellata and all species combined were aggregated in most stands, whereas Q. marilandica was mostly randomly distributed within a stand. Small trees of all species combined tended to be aggregated and large trees were randomly dispersed in all but two stands, suggesting competition. Mean distance between large-large pairs was always greater than mean distance between small-small pairs in all stands, but this difference was only significant in one stand. Correlations between nearest neighbor distance and combined size of nearest neighbors were significant and positive in 12 of 17 stands. In all cases, however, slopes were shallow suggesting that competition is weak in these communities and has a limited effect on spacing of neighboring trees. Contrary to our hypothesis, trees were more aggregated on coarse-textured soils with low organic matter content. For all species combined, degree of aggregation was unrelated to growing season precipitation. Aggregation appears to be common in these forests because environmental stress in many stands reduces growth rates. Trees have not yet reached a size at which competition or other interactions can greatly increase interplant distances and reduce the degree of aggregation. A simple graphical model is developed to describe the relationship between patterns, stress and competition in plant communities.