The spatiotemporal dynamics of a primary succession


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  • 1Conceptual models of ecosystem development commonly predict a phase of initial colonization characterized by the nucleation, growth and coalescence of discrete patches of pioneer plants. Spatiotemporal dynamics during subsequent development may follow one of three different models: the classical model, in which initially discrete patches of competitive dominant (secondary) colonists coalesce to form a homogeneous cover; the patch dynamics model, in which renewal mechanisms such as disturbance create a shifting mosaic of patches at different stages; and the geoecological model, in which the vegetation gradually differentiates along edaphic gradients related to the underlying physical template.
  • 2These models of spatiotemporal dynamics were tested using vegetation and soil data from an 850-year chronosequence, comprised of seven lava flows on Mt Hekla, Iceland. The scale and intensity of spatial pattern were quantified on each flow using spatial analyses (mean-variance ratios, quadrat variance techniques and indices of autocorrelation). Changes in spatial pattern with increasing terrain age were compared with predicted trajectories, in order to identify which of the models of spatiotemporal dynamics was most consistent with the observations.
  • 3The early stages of ecosystem development were characterized by colonization of the pioneer species, especially Racomitrium mosses, in discrete patches (‘Pioneer colonization stage’, < 20 years), which then grew laterally and coalesced to form a continuous, homogeneous carpet (‘Pioneer expansion stage’, 20–100 years). Later in the sequence, higher plants established in discrete patches within this pioneer matrix (‘Higher plant colonization stage’, 100–600 years). Over time, heterogeneity re-emerged at a larger spatial scale as the vegetation differentiated according to topographic variations in the underlying substrate (‘Differentiation stage’, > 600 years).
  • 4Synthesis. The spatiotemporal dynamics observed in the early stages of this succession were consistent with a model of pioneer nucleation in micro-scale safe sites, followed by growth, coalescence and eventual fragmentation of pioneer patches. The spatial patterns which emerged later in development support the geoecological model, with spatial differentiation of vegetation related to meso-scale substrate topography. The findings provide insight on how vegetation patterns emerge at different stages of ecosystem development in response to differing scales of heterogeneity in the underlying physical environment.