Fine‐scale spatial patterns in populations of the clonal moss Hylocomium splendens partly reflect structuring processes in the boreal forest floor

Abstract

Spatial patterns of important population or individual characteristics are expected to reflect structuring ecosystem processes to the extent that populations with strikingly different spatial patterns most likely have been structured by different processes. Few studies of spatial pattern in plant assemblages, bryophyte assemblages in particular, still exist and pattern has only vaguely been linked with process. In this study we describe fine‐scale spatial variation in size and the occurrence of terminated (non‐branching) segments in 21 Norwegian populations of the dominant boreal forest moss Hylocomium splendens, in search for general patterns of spatial structure.All Hylocomium splendens segments in the investigated plots were followed for a 10‐year period; their size (dry mass, estimated from in situ measurements) and fate (terminated or ramifying) were recorded annually. The spatial structure of size and terminating segments at scales from 0.5 to 16 cm was described for each population as fractal dimension profiles, derived from semi‐variograms. For about one half of the populations size and terminating segments could be assigned to a specific spatial pattern, the predicted outcome of one of three main structuring processes: (i) negative spatial dependence (fractal dimension F>3.0) on the scale of individuals (below 2 cm), indicative of negative interactions; (ii) positive spatial dependence (F<3.0) on the scale of individuals, indicative of positive interactions; and (iii) positive spatial dependence on broader (4–8 cm) scales, indicative of structuring by environmental factors or patchy disturbance. Patterns (i) and (iii) were observed both for size and terminated segments more often than expected. Fractal dimension profiles for size obtained separately for each year revealed temporal patterns of spatial structure that tended to be invariant over years. Negative spatial dependence of size, possibly due to large size difference between buried segments and other segments during self thinning, was typical of populations rapidly growing in number. Positive spatial dependence with a range of influence by the spatial process of 4–8 cm was observed in plots with sparse bryophyte cover and high cover of deciduous litter, probably due to accumulation of litter in depressions between shoots or groups of shoots. The main finding of this study is that different populations of the same species in the same type of ecosystem can be spatially structured in different ways. This accords with a model for the studied system as consisting of a mosaic of cells, of spatial extent of a few cm or larger, intergrading in space and temporally dynamic, the population characteristics of each cell being shaped by several structuring processes that vary, temporally and spatially, in their relative importance.