Establishment patterns in a secondary tree line ecotone

On semi-open pre-alpine fen pastures Alder encroachment creates a dynamic mosaic of grassland and woodland, which is rich in ecotones from fen to Carr. The structural diversity in colonisation patterns of Alder on fens suggests a dependency on multiple environmental drivers. Unidirectional progressive ecotone development provides an opportunity to address a current deficit in understanding successional patterns, i.e. process–pattern relationships in a multiple factor regime.We developed an individual-based model of Alder establishment on fen grassland to investigate the dependency of encroachment patterns upon seed production, dispersal distances and safe site availability. The purpose of the model is to provide a causal understanding of establishment patterns of Alder. In the model, all life processes of Alder individuals were parameterised with field data. This allowed us to strictly perform bottom–up simulations and successfully check plausibility by comparing simulated establishment patterns of cohorts with observed ecotone structures.Simulation results show that establishment patterns strongly depend on environmental drivers. Spatial progression of Alder encroachment and width of ecotones, respectively, mainly depend on wind speed during seed dispersal. Dense establishment of Alder leading to community change from fen grassland to Carr, requires windows of opportunity, which are defined by the rare coincidence of widespread dispersal, high seed production and favorable establishment conditions. Life-history traits of Alder (mast year cycle, high seed weight, weak establishment in fen) spatially and temporarily constrain the encroachment process. The structural diversity of long-term encroachment patterns is explained by the event-driven character of encroachment.Modelling individual establishment pathways of seedlings starting from germination revealed an endogenous stochasticity in establishment patterns emerging from low seed densities in the tail of the dispersal function. We conclude an inherent stochastic structure of dispersal-limited tree line ecotones, which limits reconstruction of processes from patterns.In order to describe long-term successional patterns of Alder encroachment at landscape scale, we propose the combination of two concepts: deterministic “patch-movement” of Alder woodland driven by continuous ecotone migration together with rare and stochastic “infiltration” of single Alder trees into open fen grasslands. Conservation management can control predictable “patch-movement” by cutting off maturing saplings around existing Alder woods. But the preservation of the actual large proportion of open grassland in fen pastures from infiltrating Alder seedlings and from the subsequent shift of the pasture to a densely wooded state would require mowing additionally to extensive grazing.

► Alder encroachment on fen grassland is predictable close to seed parents.
► Windows of opportunity exist for stochastic widespread infiltration of fen.
► Constraints are established by life-history traits of Alder.
► Coincidence of environmental drives to open windows is rare.
► Individual life histories of seedlings reveal the stochastic structure of tree lines.