Managing heterogeneity: the importance of grazing and environmental variation on post-fire succession in heathlands.
Semi-natural habitats have been shaped by human disturbance regimes for centuries. Spatially and temporally heterogeneous land use practices, such as cutting, burning, grazing and turf-cutting, have resulted in complex mosaic landscapes that are of high priority for conservation in Europe. Contemporary conservation subjects these systems to management regimes that are generally less diverse, in terms of disturbances and fine-scale temporal and spatial variability, than traditional land use, but the ecological consequences of these simplifications are unclear. We investigated the interactive effects of fire and grazing on plant species composition and diversity along local environmental (moisture) gradients in coastal Calluna vulgaris heathlands in western Norway. A replicated series of post-fire successions (n=12) was initiated in three heathland habitats and the areas subjected to two grazing regimes. Floristic and environmental data were recorded in permanent plots over a 5-year period. Community data were analysed using multivariate ordination techniques (principal components analysis, partial redundancy analyses, and principal response curves) and generalized linear models. Fire induced strong successional trends in the species composition of the heathlands. These trends differed among heathland habitats, and with grazing. Strong interactions between fire, habitat and grazing implied that the effect of grazing on the successional dynamics differed among habitats. Species diversity decreased in the first year after fire but increased beyond the pre-fire levels during succession. This trend was not affected by local environment or grazing, although there were main effect differences in diversity between environments and grazing regimes. Our results demonstrate that the two management practices do not have simple additive effects within the semi-natural system studied, as grazing created ecological opportunities for additional sets of species, increased variability among habitats, and added complexity to the post-fire successional dynamics. In order to preserve diversity, conservation management should thus aim to preserve the level of complexity of the traditional management regimes, both in terms of the actual disturbances (e.g. fire and grazing) as well as the spatial scales at which they are applied. Further, the considerable change in these effects along the local environmental gradient brings into question the efficiency of general management prescriptions, and indicates that local environmental variability should be taken into account in the conservation of semi-natural habitats.