Questions: 1. Does random colonization predominate in early stages of primary succession? 2. Do pioneer species facilitate the establishment of later arriving species? 3. Does an initially random distribution change to an aggregated pattern with ongoing succession?

Location: Lignite mining region of Lower Lusatia, eastern Germany.

Methods: Individual plants were mapped along a 2 m × 28 m transect during three successive years and classified into two groups (1) the pioneer Corynephorus canescens and (2) ‘all other species’. Using the pair-correlation function, univariate point pattern analysis was carried out by applying a heterogeneous Poisson process as null model. Bivariate analysis and a toroidal shift null model were applied to test for independence between the spatial patterns of the two groups separately for each year, as well by exploring spatiotemporal patterns from different years.

Results: In the first year Corynephorus and ‘all other species’ showed an aggregated pattern on a spatial scale > 40 cm and in the second and third years a significant attraction for distances between 4 and 12 cm, with an increasing radius in the third year. The analyses of interspecific spatiotemporal dynamics revealed a change from independence to attraction between distances of 4 cm and 16 cm when using Corynephorus as focal species. However, applying ‘all other species’ as focal points results in a significant attraction at distances up to 60 cm in the first year and a diminishing attraction in the second and third years with distances ≤ 6 cm.

Conclusions: Facilitative species-species interactions are present in early stages of primary succession, resulting mainly from pioneer species acting as physical barriers and their ability to capture diaspores being drifted by secondary dispersal along the substrate surface. However, due to gradual establishment of perennial species and their ability of lateral extension by vegetative dispersal, facilitation may influence spatial pattern formation predominantly on short temporal and fine spatial scales.

Nomenclature: Wisskirchen & Haeupler (1998).