In this study, we combined the analysis of a coenocline with Huisman–Olff–Fresco (HOF) modelling of species behaviour to describe colonization patterns of woody species on reclaimed coal wastes in northern Spain. We hypothesize that fine‐scale variation in abiotic factors along the forest‐grassland gradient that appears after mining restoration affects colonization pattern of woody species. To understand these factors, we analysed changes in floristic composition and abundance of the main woody colonizers according to the spatial distance from forest limit, as well as the spatial variability of some edaphic and microclimatic parameters. Our results showed how the primary coenocline extracted from DCA1 reflected a gradient in environmental conditions influencing the species composition of the woody plant communities, from the more shade, moist and dense forest on acid and rich soils to the open, dry and oligotrophic grassland. The colonization pattern of woody species is affected by fine‐scale variations in abiotic factors, such as the increase in pH, P, light, and the decrease in K, N, C/N, organic matter and soil moisture, from the forest to the restored mine. Most species HOF modelled showed monotone responses with decreasing trends along the environmental gradient from forest to mine, being Quercus petraea the species clearly dominant in the forest. On the contrary, Cytisus scoparius and Genista florida colonize intensely in the mine area, reaching the maximum abundance around 15 and 10 m distance from the forest limit, respectively. We conclude that the composition of woody species in the forest is not a good predictor of the colonizing intensity, not even of the species composition in the mine area. Copyright © 2011 John Wiley & Sons, Ltd.
We evaluated the ecological significance of the boundary form between two patches with contrasting vegetation (mine grassland and adjacent forest) on woody colonization and forest expansion in open-cast coal mines in Northern Spain. Woody colonization and browsing traces were measured on three mine sites, along 24 transects that were laid out perpendicular to the forest-mine boundary and classified according to their shape (concave, convex, straight). Mine sites were colonized from the close forest by woody species, whose colonization intensity depends on the boundary form. The overall colonization intensity decreased with increasing distance to the forest and differed depending on the boundary form. The more intense colonization was found in concave boundaries and the strongest decrease in convex boundaries close to the forest, whereas straight boundaries showed an intermediate colonization pattern. Concave boundaries reached higher woody cover in the basal strata of the mines than convex (up to 2 m) or straight boundaries (up to 1 m) from 11 m to the forest edge, mainly by the presence of dense patches of Cytisus scoparius (L.) Link, with a scattered overstory of Genista florida L. These shrubs might reduce the browsing intensity and act as nurse plants facilitating the establishment of Quercus petraea (Matt.) Liebl. in mine areas at greater distances from the forest edge. The forest-mine boundary form does not affect the forest vertical structure that is homogenous and does not help explain the woody colonization pattern in the mines. We conclude that edge characteristics have a strong potential to be used in the restoration of native forests based on natural processes. The implications of our results for sessile oak (Quercus petraea (Matt.) Liebl.) forest expansion along edges in fragmented Mediterranean forest landscapes were discussed.
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