Abbreviations GLMM = generalized linear mixed modelling or generalized linear mixed models; DEM = Dynamic equilibrium model; IDH = Intermediate disturbance hypothesis.
AbstractQuestions: What are the interactive effects of succession time, disturbance and productivity on the fine-scale species richness of beach vegetation? How do these effects differ between functional groups that are known to represent different adaptive strategies?Location: Boreal beach and dune systems (ca. 60°-65°N) of the Baltic Sea coast in Finland, characterized by steep environmental gradients, post-glacial land uplift and resulting primary succession.Methods: The analysis was based on an extensive and systematic survey of vascular plant, bryophyte and lichen species and environmental factors. Total species richness and the species richness of seven functional groups were modelled as functions of succession time, disturbance, productivity and their statistical interactions. The effects of the local environmental variability were taken into account by applying generalized linear mixed models (GLMM).Results: All three environmental factors and their statistical interactions, including the three-way interaction, were highly significant in explaining total species richness. The effect of disturbance on total species richness changed from negative to positive along succession time and productivity gradients. The response of beach specialist richness to succession time and disturbance was unimodal, while the responses of the generalist functional groups were monotonic. Beach specialist richness showed a strong positive response to productivity, whereas the richness of the generalist groups was not related to changes in productivity.
Conclusions:The patterns of species richness on uplifting beaches are determined by the interplay of primary succession, disturbance and productivity. When both succession time and productivity increase, competitive exclusion starts to limit total species richness. In these circumstances, disturbance favours diversity by creating gaps in the vegetation. A combination of long succession time, low disturbance and low productivity result in maximum species richness. The richness of functional groups, particularly the opportunistic and disturbance-tolerant beach specialist group and the competitive but disturbance-sensitive groups, reach their peak in divergent environmental conditions. Consequently, the complex richness patterns observed on uplifting beaches are created not only by strongly interacting environmental drivers but also by differences in species' adaptive strategies.