High resilience of bryophyte assemblages in streamside compared to upland forests.

Landscape heterogeneity causes spatial variation in disturbance regimes and resilience. We asked whether the resilience of bryophyte (liverwort and moss) assemblages to clear-cutting differs between streamside and upland boreal forests in northern Sweden. We hypothesized that bryophyte survival and recolonization rates are higher in streamside areas, thus raising resilience. Conversely, disturbance-intolerant but also invading species should be more frequent here, potentially reducing resilience. In each of 18 sites, we compared two 0.1-ha plots (one streamside and one upland) located in old forest that had never been clear-cut with two matching plots in young stands established after clear-cutting of old forests 30-50 years earlier. We used the magnitude of the difference in assemblages between old and young stands as a measure of change and, therefore, resilience (large difference implying low resilience). Species assemblages were more resilient in streamside than in upland forests. Species composition changed significantly in upland but not in streamside forests. Reductions in species richness were more pronounced in upland forests for total richness and for eight subgroups of species. Two results indicated lower survival/recolonization in upland forests: (1) species had a stronger association with old stands in upland areas, and (2) among species present in both the old streamside and old upland plot in a site, fewer appeared in the young upland than in the corresponding streamside plot. Simultaneously, a higher proportion of species invaded streamside areas; 40 of the 262 species encountered in streamside forests increased their occupancy by two or more sites compared to only two of 134 species in uplands. We suggest that in boreal forests spatial variation in resilience of assemblages of forest organisms intolerant of canopy removal is related to factors governed mainly by topography. More generally, we argue that landscape-scale variation in resilience of assemblages is influenced by spatial variation in (1) stress and resource availability, (2) number of disturbance-intolerant species, and (3) magnitude of environmental changes brought about by a disturbance with a specific intensity. We also suggest that rapid recovery in the short term does not necessarily imply higher long-term ability to return to the pre-disturbance state.