Spatial pattern of a subalpine forest-alpine pasture ecotone (Las Cutas, Ordesa, Central Pyrenees)

Abstract

We describe the spatial pattern of a subalpine forest-alpine pasture ecotone in the Central Pyrenees, that includes altitudinal timberline and treeline, and it is dominated by Pinus uncinata Ram. A rectangular (30 x 140 m) plot was located crossing the ecotone with its longest side parallel to the slope. We measured for each P. uncinata individual inside the plot: location (coordinates x, y), and structure (e. g. height) and growth form variables (number and type —living or dead, vertical or shrubby— of stems per individual). P. uncinata individuals were classified according to their size (adults, poles, saplings and seedlings) and growth form (krummholz —shrubby and multistemmed individuals— and krummholz with vertical stems). We described quantitatively the type of substrate (bare soil, organic matter, gravel and rock) and cover of herbs, shrubs and P. uncinata using transects parallel to the slope. The ecotone structure was described through: (1) point pattern (Ripley’s K) and (2) surface pattern analyses (spatial correlograms of height); (3) the detection and description of boundaries using density, size or growth form variables; (4) the synthesis of variations of presence and diversity of substrates and herbs and shrubs; and (5) the ordination of quadrats (the plot was previously subdivided into 115 6 x 6 m quadrats), according to their spatial position in the ecotone, the type of substrate, the cover of herbs and shrubs and the number, size and growth form of P. uncinata individuals. Most P. uncinata living individuals were krummholz, located above the timberline. Krummholz individuals showed significant and positive spatial interaction with seedlings. Bigger, vertical and unistemmed individuals predominated in the lower area of the ecotone, in the forest. The change of height with increasing elevation was abrupt and masked an underlying pattern of patches of trees with similar height in the forest. The structure variables were more sensitive because they produced a greater number of boundaries. These boundaries were arranged forming a “diagonal” (in the lower and upper areas of the ecotone for big and small individuals, respectively) because of the sequential location of progressively bigger unistemmed individuals descending across the ecotone. The shrubby individuals were associated with rocky substrates, that decreased in the forest, where organic matter predominated. The snow-wind interaction can explain the location of the studied timberline that could be considered a local phenomenon. Krummholz can buffer seedlings against the harsh climatic conditions of this ecotone (strong wind, reduced snowpack, low temperature). The spatial location of the different classes of individuals, the spatial interaction between seedlings and krummholz individuals, and changes of growth form (from shrubby to vertical growth form or vice versa) can cause some inertia in the response of ecotone P. uncinata populations to environmental changes.