Abstract

Abstract The effects of mixed stands on biodiversity are increasingly being studied since they are supposed to offer higher habitat heterogeneity. Nevertheless, for tree-associated diversity, including epiphytes and terricolous species near tree trunks, few studies exist, and still fewer compare mixed stands with each corresponding pure tree species stand. We evaluated and quantified the influence of forest composition on tree-associated bryophyte diversity (species richness, abundance, composition) in mixed and pure oak-pine stands in a French lowland forest. The main explanatory variables for bryophyte diversity at tree-level were the identity of the phorophyte tree species and the mixture type (pure versus mixed). At the plot level, the main explanatory variable was the stand type (pure oak, pure pine and mixed). We also explored the role of other variables including the chemistry of the bryophyte substrates (soil, bark) and water supply (stemflow, throughfall), as well as stand abundance variables (basal area, interfering plant cover). We analyzed data with Generalized Linear Models under Bayesian statistics, to take into account the spatial autocorrelation between plots and any under- or over-dispersion of our data. At the tree-level, bryophyte richness and abundance were higher on oak than on pine. Pine bryophyte richness was higher in mixed compared to pure stands, whereas for oak, mixed stands did not enhance bryophyte richness. At the plot level, mixed stands hosted bryophyte communities of similar richness to those in pure oak stands, whereas pure pine stands were clearly poorer. Our exploratory models suggested strong effects of water supply chemistry (stemflow and throughfall pH or conductivity) and basal area; the latter had a strong quadratic effect on epiphytic richness at the plot level. In terms of composition, three species were more likely to be found on pine phorophytes, seven species clearly occurred more frequently on oaks. Some species were more likely to be associated to pine in mixed than in pure stands, and one species was found more often under pines in pure stands. Therefore, bryophyte diversity at the landscape level should benefit from the simultaneous presence of the three stand composition types – pure oak, pure pine and mixed stands.

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