• facilitation;
  • geographical range;
  • microenvironment;
  • spatial heterogeneity;
  • tree recruitment


  • 1
    We analyse the factors controlling seedling establishment of Scots pine at its southernmost geographical limit (southern Spain), by monitoring emergence, survival and growth for up to 4 years in the microhabitats to which seeds are dispersed. Naturally established seedlings were monitored in two mountain ranges, and experimental sowings were performed both in woodlands and in adjacent successional shrublands into which the forest could expand.
  • 2
    Emergence was high in all microhabitats, although it was highest under the canopy of shrubs. Overall survival was low, with c. 90% of seedlings dying in the first growing season (c. 98% after several growing seasons). Survival differed among microhabitats, being highest under shrubs and extremely low (or zero) under pines or in bare soil.
  • 3
    Seedling growth was the highest in areas of bare soil, intermediate under shrubs, and very low under pines.
  • 4
    Establishment under pines was prevented by both mortality and poor performance, and good performance cannot counteract high mortality in the open. Shrubs, however, acted as nurse plants, buffering summer drought without reducing radiation to levels critical for growth, and protecting seedlings from ungulate trampling, hail and frost heave.
  • 5
    Patterns of recruitment were similar for woodland stands and successional shrublands. In addition, patterns of survival for naturally established seedlings were similar to those of seedlings originating from experimental sowings.
  • 6
    Juveniles were positively associated with shrubs but negatively with bare soil or areas below pine canopies. The facilitative effect of shrubs on seedling survival therefore changes the spatial pattern of recruitment from that determined by germination.
  • 7
    Overall, processes controlling seedling establishment in these southern Scots pine forests differ sharply from those operating in its main distribution area. The comparison among contrasting geographical ranges may contribute to an understanding of the role of environmental conditions in the balance between competition and facilitation, and assist in forecasting plant regeneration responses to global climate change.