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iForest -Biogeosciences and Forestry
IntroductionFuture impacts of global climate change such as predicted increases in annual air and surface temperatures and variations in pre cipitation will cause significant alterations in forest ecosystems (Eastaugh 2008). Impacts of these changes will be proportionally more perceptible at high elevations (Bensiton et al. 1997).In timberline ecotones near the upper limit of closed forests tree growth, forest structure and forest dynamics are mainly temperaturedriven (Tranquillini 1979, Innes 1991, Körner 1998. The sensitivity of these biomes to climate variability is high and thus of special interest for understanding the ef fects of global change.Swiss stone pine (Pinus cembra L.) is dis tributed in timberline ecotones across Europe from the Carpathian Mountains to the French Alps (Polunin & Walters 1986, Ulber et al. 2004. During several hundred years of hu man activities such as alpine farming or tim ber extraction Swiss stone pine was often eliminated and therefore restricted to stands on inaccessible slopes exposed to the North (Holtmeier 1966, Motta & Nola 2001, Höhn et al. 2009). In recent decades, socio-eco nomic and silvicultural changes have fa voured the establishment of Swiss stone pine (Motta et al. 2006). This five-needled conifer tree is well adapted to the harsh subalpine climate conditions in the Central European Alps (Ulber et al. 2004) and is often asso ciated with mountain pine (Pinus montana Miller), Scots pine (Pinus sylvestris L.), European larch (Larix decidua Miller) and Norway spruce (Picea abies L. Karst). In the continental subalpine forests of the Central Alps with relatively low rainfall and mean annual temperatures below 1.5 °C (Ellenberg 1996), stands develop from early-successio nal stages dominated by mountain pine to a late-successional stage dominated by Swiss stone pine and European Larch (Risch et al. 2003, Höhn et al. 2009).This review will summarize the evidence of Swiss stone pine responses to climate change at the timberline ecotone. The review will consider all life stages, and possible dis tribution shifts of Swiss stone pine popula tions in the future will be discussed.
Seedling establishmentSwiss stone pine is a monoecious, wind pollinated species which reaches reproduc tive maturity at 40-60 years of age (Ulber et al. 2004) with good seed production years occurring on average twice in ten years (Mattes 1982). Seed production is especially sensitive to climate because important deve lopmental processes such as the initiation of flower and cone primordia, meiosis and the release of pollen depend to a large degree on climatic variables (Pigott 1992). The tempo ral dynamics of seed production and the in fluence of climate change on seed produc tion of Swiss stone pine have however not been comprehensively investigated to date.The wingless Swiss stone pine seeds are mainly dispersed by the European nutcracker bird (Nucifraga caryocatactes L.). Swiss stone pine seeds are the main food source for European nutcrackers, and those bird...