A multivariate approach to the study of the spatial structure of treeline ecotones

Abstract

We study the structure of two contrasting alpine forest.pasture ecotones located in the Central Pyrenees (sites Ordesa and Tessó). We define ecotone structure as the spatial distribution of trees of different size classes and growth-forms, and the relationship between these aspects and the spatial distribution of understory vegetation and substrate. The studied ecotones are dominated by Pinus uncinata and have been little affected by recent anthropogenic disturbances (logging, grazing). One rectangular plot (30 x 140 m2) was located within each site encompassing treeline and timberline with its longest side parallel to the slope. The distribution of size and growth-form classes at Ordesa followed a clear sequence of increasing size downslope, from shrubby krummholz individuals to bigger arborescent trees. At Ordesa, regeneration was concentrated near the krummholz area and over rocky substrates. At Tessó, regeneration was abundant above the treeline, where the cover of the dominant understory shrub (Rhododendron ferrugineum) decreased. Detrended canonical correspondence analysis of tree and plant cover data, with respect to spatial location in the ecotone and substrate cover, demonstrated that elevation was an important factor controlling the distribution of trees and understory plants in both ecotones. Finally, k-means clustering with spatial constraint revealed abrupt spatial clusters along the slope at Ordesa. However, the ecotone at site Tessó was composed of elongated downslope spatial clusters suggesting greater spatial heterogeneity and subtle gradual changes due to other factors in addition to the altitudinal gradient (snow avalanches). These contrasting structures correspond well with the ecotone (sharp boundary, Ordesa) and ecocline (gradual transition, Tessó) concepts. This suggests the dominant role of different local environmental factors: wind at site Ordesa and avalanches at Tessó. Positive feedbacks, like facilitation among P. uncinata individuals (nurse effect), may maintain and intensify the sharpness of the ecotone at Ordesa.