Spatiogenetic characteristics of beech stands with different degrees of autochthony

Autochthony in forest tree stands is characterized by a number of criteria, among which the range over which stands act as a population has been suggested to play a central role. Therefore, measures are needed for the delineation of populations or the detection of subpopulation structure. It is argued here that methods of population delineation must be based on the combined consideration of spatial distances and genetic differences between adult individuals. Conventional approaches and a set of newly developed methods are applied to seven isozyme loci in four beech stands which are distinguished by different types of forest management based on natural regeneration.

Permutation analyses show that correlations between spatial distances and genetic differences vary only little in the studied beech stands. In view of the popularity of this and related descriptors of spatiogenetic covariation, this result came as a surprise. The newly developed methods lead to a different conclusion. Significant spatiogenetic structure is indicated in all stands when considering the mean and variance of spatiogenetic separation, where separation is measured by the smallest spatiogenetic difference of an individual from all others. Spatiogenetic difference is measured here by a combination of the spatial distances and genetic differences between individuals. This descriptor indicates the existence of spatiogenetic clusters in the beech stands. In order to arrive at an explicit representation of cluster structure as a representation of subpopulation structure, two types of cluster structure (primary and alpha-isolated) are distinguished, both of which reflect desirable characteristics of subpopulation structure. Particularly in the alpha-isolated structure, the proportion of individuals organized in clusters, the effective size, and the effective number of clusters clearly distinguish and consistently rank the four stands with respect to their types of forest management and the associated criteria of autochthony.

The surprisingly high correspondence between our descriptors of spatiogenetic structure and forest management types confirms the appropriateness of the applied measure of cluster isolation and of the criterion for the choice of the level alpha of cluster isolation. The two types of cluster structure and their characteristic descriptors are thus suggested to be promising tools for the detection of subpopulation structure. To include the effects of long-distance gene flow, the presented methods can be extended as outlined to larger spatial scales in order to detect higher order population structure.