GROWTH AND COMPETITION IN CLONAL PLANTS - PERSISTENCE OF SHOOT POPULATIONS AND SPECIES-DIVERSITY

This paper reviews studies on growth and size-structure dynamics of shoots and clones in clonal plants in comparison with those in non-clonal plants, and discusses the characteristics of clonal plants. The mode of competition between individuals (symmetric versus asymmetric, degree of competitive asymmetry), growth dynamics of individuals, allocation pattern between organs and spatial pattern of individuals are closely correlated with each other in non-clonal plant populations. Theoretical and field studies based on the diffusion model revealed that plants of ''height-growth'' type (mostly early-successional tree species) and plants of ''diameter-growth'' type (mostly late-successional tree species) tend to exhibit asymmetric competition and symmetric competition respectively. Moreover, asymmetrically competing plants show smaller effects of variation in individual growth rate and spatial pattern on the size-structure dynamics of the population than symmetrically competing plants. Therefore, the spatial pattern of individuals should be considered especially for plants undergoing symmetric competition. These results for non-clonal plants should have a significant implication also for the growth dynamics and competition in clonal plants. The mean growth rate of shoots [G(t,x) function] and hence the mode of competition between shoots differs among clonal plant species as in non-clonal plants. However, a large magnitude and size-independence (or slightly negative size-dependence) of the variation in growth rate of shoots [D(t,x) function], especially at the early stage in a growing season is a common characteristic of many clonal plant species, in contrast to the positively size-dependent variation in individual growth rate in non-clonal plants. This type of variation in shoot growth rate leads to the persistence of stable shoot populations even when the mean growth rate function is changed, and also in cases where the shoot population structure would be unstable in the absence of variation in growth rate. It is suggested that competition between clones is symmetric in most clonal plant species, which brings about small-scale spatio-temporal changes in species abundance and hence species diversity.