Properties, goods and services
Mountain regions (circa 20-24% of all land, scattered throughout the globe) exhibit many climate types corresponding to widely separated latitudinal belts within short horizontal distances. Consequently, although species richness decreases with elevation, mountain regions support many different ecosystems and have among the highest species richness globally (e.g., Väre et al., 2003; Moser et al., 2005; Spehn and Körner, 2005). Mountain ecosystems have a significant role in biospheric carbon storage and carbon sequestration, particularly in semi-arid and arid areas (e.g., the western U.S., – Schimel et al., 2002; Tibetan plateau – Piao et al., 2006). Mountain ecosystem services such as water purification and climate regulation extend beyond their geographical boundaries and affect all continental mainlands (e.g., Woodwell, 2004). Local key services allow habitability of mountain areas, e.g. through slope stabilisation and protection from natural disasters such as avalanches and rockfall. Mountains increasingly serve as refuges from direct human impacts for many endemic species. They provide many goods for subsistence livelihoods, are home to many indigenous peoples, and are attractive for recreational activities and tourism. Critically, mountains harbour a significant fraction of biospheric carbon (28% of forests are in mountains).
The TAR identified mountain regions as having experienced above-average warming in the 20th century, a trend likely to continue (Beniston et al., 1997; Liu and Chen, 2000). Related impacts included an earlier and shortened snow-melt period, with rapid water release and downstream floods which, in combination with reduced glacier extent, could cause water shortage during the growing season. The TAR suggested that these impacts may be exacerbated by ecosystem degradation pressures such as land-use changes, over-grazing, trampling, pollution, vegetation destabilisation and soil losses, in particular in highly diverse regions such as the Caucasus and Himalayas (Gitay et al., 2001). While adaptive capacities were generally considered limited, high vulnerability was attributed to the many highly endemic alpine biota (Pauli et al., 2003). Since the TAR, the literature has confirmed a disproportionately high risk of extinction for many endemic species in various mountain ecosystems, such as tropical montane cloud forests or forests in other tropical regions on several continents (Williams et al., 2003; Pounds and Puschendorf, 2004; Andreone et al., 2005; Pounds et al., 2006), and globally where habitat loss due to warming threatens endemic species (Pauli et al., 2003; Thuiller et al., 2005b).
Because temperature decreases with altitude by 5-10°C/km, relatively short-distanced upward migration is required for persistence (e.g., MacArthur, 1972; Beniston, 2000; Theurillat and Guisan, 2001). However, this is only possible for the warmer climatic and ecological zones below mountain peaks (Gitay et al., 2001; Peñuelas and Boada, 2003). Mountain ridges, by contrast, represent considerable obstacles to dispersal for many species which tends to constrain movements to slope upward migration (e.g., Foster, 2001; Lischke et al., 2002; Neilson et al., 2005; Pounds et al., 2006). The latter necessarily reduces a species’ geographical range (mountain tops are smaller than their bases). This is expected to reduce genetic diversity within species and to increase the risk of stochastic extinction due to ancillary stresses (Peters and Darling, 1985; Gottfried et al., 1999), a hypothesis confirmed by recent genetic analysis showing gene drift effects from past climate changes (e.g., Alsos et al., 2005; Bonin et al., 2006). A reshuffling of species on altitude gradients is to be expected as a consequence of individualistic species responses that are mediated by varying longevities and survival rates. These in turn are the result of a high degree of evolutionary specialisation to harsh mountain climates (e.g., Theurillat et al., 1998; Gottfried et al., 1999; Theurillat and Guisan, 2001; Dullinger et al., 2005; Klanderud, 2005; Klanderud and Totland, 2005; Huelber et al., 2006), and in some cases they include effects induced by invading alien species (e.g., Dukes and Mooney, 1999; Mack et al., 2000). Genetic evidence for Fagus sylvatica, e.g., suggests that populations may show some capacity for an in situ adaptive response to climate change (Jump et al., 2006). However, ongoing distributional changes (Peñuelas and Boada, 2003) show that this response will not necessarily allow this species to persist throughout its range.
Upper tree lines, which represent the interface between sub-alpine forests and low-stature alpine meadows, have long been thought to be partly controlled by carbon balance (Stevens and Fox, 1991). This hypothesis has been challenged (Hoch and Körner, 2003; Körner, 2003a). Worldwide, cold tree lines appear to be characterised by seasonal mean air temperatures of circa 6°C (Körner, 1998; Körner, 2003a; Grace et al., 2002; Körner and Paulsen, 2004; Millar et al., 2004; Lara et al., 2005; Zha et al., 2005). In many mountains, the upper tree line is located below its potential climatic position because of grazing, or disturbances such as wind or fire. In other regions such as the Himalaya, deforestation of past decades has transformed much of the environment and has led to fragmented ecosystems (Becker and Bugmann, 2001). Although temperature control may be a dominant determinant of geographical range, tree species may be unable to migrate and keep pace with changing temperature zones (Shiyatov, 2003; Dullinger et al., 2004; Wilmking et al., 2004).
Where warmer and drier conditions are projected, mountain vegetation is expected to be subject to increased evapotranspiration (Ogaya et al., 2003; Jasper et al., 2004; Rebetez and Dobbertin, 2004; Stampfli and Zeiter, 2004; Jolly et al., 2005; Zierl and Bugmann, 2005; Pederson et al., 2006). This leads to increased drought, which has been projected to induce forest dieback in continental climates, particularly in the interior of mountain ranges (e.g., Fischlin and Gyalistras, 1997; Lischke et al., 1998; Lexer et al., 2000; Bugmann et al., 2005), and mediterranean areas. Even in humid tropical regions, plants and animals have been shown to be sensitive to water stress on mountains (e.g., Borneo – Kitayama, 1996; Costa Rica – Still et al., 1999). There is very high confidence that warming is a driver of amphibian mass extinctions at many highland localities, by creating increasingly favourable conditions for the pathogenic Batrachochytrium fungus (Pounds et al., 2006).
The duration and depth of snow cover, often correlated with mean temperature and precipitation (Keller et al., 2005; Monson et al., 2006), is a key factor in many alpine ecosystems (Körner, 2003c; Daimaru and Taoda, 2004). A lack of snow cover exposes plants and animals to frost and influences water supply in spring (Keller et al., 2005). If animal movements are disrupted by changing snow patterns, as has been found in Colorado (Inouye et al., 2000), increased wildlife mortality may result. At higher altitudes, the increased winter precipitation likely to accompany warming leads to greater snowfall, so that earlier arriving altitudinal migrants are confronted with delayed snowmelt (Inouye et al., 2000).
Disturbances such as avalanches, rockfall, fire, wind and herbivore damage interact and are strongly dependent on climate (e.g., Peñuelas and Boada, 2003; Whitlock et al., 2003; Beniston and Stephenson, 2004; Cairns and Moen, 2004; Carroll et al., 2004; Hodar and Zamora, 2004; Kajimoto et al., 2004; Pierce et al., 2004; Schoennagel et al., 2004; Schumacher et al., 2004). These effects may prevent recruitment and thus limit adaptive migration responses of species, and are exacerbated by human land use and other anthropogenic pressures (e.g., Lawton et al., 2001; Dirnböck et al., 2003; Huber et al., 2005).
Ecotonal (see Glossary) sensitivity to climate change, such as upper tree lines in mountains (e.g., Camarero et al., 2000; Walther et al., 2001; Diaz, 2003; Sanz-Elorza et al., 2003), has shown that populations of several mountain-restricted species are likely to decline (e.g., Beever et al., 2003; Florenzano, 2004). The most vulnerable ecotone species are those that are genetically poorly adapted to rapid environmental change, reproduce slowly, disperse poorly, and are isolated or highly specialised, because of their high sensitivity to environmental stresses (McNeely, 1990). Recent findings for Europe, despite a spatially coarse analysis, indicate that mountain species are disproportionately sensitive to climate change (about 60% species loss – Thuiller et al., 2005b). Substantial biodiversity losses are likely if human pressures on mountain biota occur in addition to climate change impacts (Pounds et al., 1999, 2006; Lawton et al., 2001; Pounds, 2001; Halloy and Mark, 2003; Peterson, 2003; Solorzano et al., 2003; Pounds and Puschendorf, 2004).