Differences in colonization strategies of three common pioneer woody species in post mining heaps.

Willow (Salix caprea), birch (Betula pendula) and aspen (Populus tremula) are common pioneer woody species, however little is known about colonization strategies in large-scale disturbances. Here we have compared the strategies of establishment of these pioneer woody species in unreclaimed sites on a large (1957 ha) spoil heap in Czechia. For all species, seedlings numbers peaked in the 17 year old (successional age - time since overburden heaping) plot, suggesting that initial soil development promotes seedling establishment while covering of the surface by litter and organic layers reduces the establishment of pioneer species. The proportion of willow decreased from the edge of the heap and analysis of the age structure suggests that willow establishment was correlated with the presence of older willows in the vicinity of willows of certain ages (13 and 23 years being particularly important). The proportion of birch increased with its distance from the heap edge, and it is correlated with suitable weather conditions in the year of establishment, mainly July rainfall. Aspen proportion does not change significantly with its distance from the heap edge and year of establishment. It correlates with the number of trees in both surrounding and climatic conditions. Detailed analysis of young trees shows that vegetative propagation by root suckers (offspring) is rare in birch. In willow they represent about half of the trees while in aspen all of the young trees were root suckers derived from older aspen trees. This indicates a different colonization strategy of individual species. Birch is capable of long-distance seed transfer, which establishes most of the population, and its establishment is highly influenced by climatic conditions. Willow spreads massively over a short distance, and its establishment is highly influenced by the presence of 21-23 year old willow individuals in the vicinity. Only few scattered trees are able to establish at longer distances, where they can later spread locally by seeds. Aspen spreads over a long distance in low numbers but when some trees are established it spreads massively locally by clones.