5. Ecology  
   


Alpine ibex prefer high altitude meadows alternated with steep rocky slopes (Couturier, 1962; Nievergelt 1966; Hoffman e Nievergelt, 1972; Grignolio et al., 1999). Males and females differ greatly in their habitat choices (Pedrotti, 1995): females use space in a more constant way and are more linked to cliffs, while males move more, selecting different kinds of habitat according to the season and their phisiological needs (Francisci et al., 1985). In spring adult males prefer the more plain meadows (Nievergelt, 1966; Peracino et al., 1989; Pedrotti, 1995), and during the summer, they significantly select alpine meadows (Mascellani, 1997; Grignolio et al., 1999).
Generally, all studies on habitat selection in Alpine ibex, underline how there is a strong preference by this species for environments characterised by rocky steep slopes, which are actively searched for by both sexes especially during the winter season (Nievergelt, 1966; Schaller, 1977; Apollonio & Grimod, 1984; Tosi et al., 1986). This preference may be explained by the high sensibility of this species towards strong snow accumulations (Pedrotti, 1995).

During the summer season, the average altitude of the frequented areas is just below 3000 m a.s.l., with a range from 2300 to 3200-3300 m a.s.l. and, exceptionally, even more. In winter the altitudinal range occupied by Alpine ibex goes from 1800 to 2500-3000 m a.s.l. (Couturier, 1962; Wiersema, 1983). The lowest altitudes are reached in spring, season during which the minimal values are influenced by the altitude of the bottom of the valley and by the presence of wood, as extended forested areas are a limit difficult to cross for Alpine ibex (Nievergelt, 1966).
In areas with a high density of Alpine ibex, the altitude distribution range during summer can be much larger and stable groups of Alpine ibex can be found as low as at 2000 m a.s.l. (Bassano & Peracino, 1987).
The results of numerous studies on Alpine ibex populations recently established following natural migration or reintroductions, appear to dismiss the view of Alpine ibex not being apt to live at low altitudes (von Elsner-Schack, 1982; Choisy, 1983, 1986, 1990 e 1994; Terrier et al., 1994; Raye, 1994). The presence and establishment of stable populations in areas with altitudinal limits even lower than 1000 meters, suggests a high plasticity in habitat selection by the species. It appears however that a necessary condition is the presence of large sectors of steep cliffs, rich in niches, able to create favourable microclimatic conditions (Choisy, 1994).
The subalpine, alpine, and subnival horizons with a mean altitude of 2000 meters, are however the most concerned with the presence of Alpine ibex (Nievergelt, 1966; Pfeffer & Settimo, 1973; Hofmann & Nievergelt, 1972; Peracino et al., 1989; Francisci et al., 1985; Wiersema, 1983).

One of the most important factors influencing habitat selection by Alpine ibex is indeed the gradient and the extension of the slopes. All the authors emphasise that slopes have to be between 30 and 45 degrees, particularly during the winter period. Furthermore, the slopes have to be interrupted by porches and hides in order to favour environmental diversity (Couturier, 1962; Nievergelt, 1966; Hofmann & Nievergelt, 1972; von Elsner Schack, 1982; Wieserma, 1983 e 1984; Tosi et al., 1986; Pedrotti, 1995).

Slopes characterise the different areas chosen by the two sexes: females use steep terrains during the whole year, while males frequent also plain meadows, in accordance with their different seasonal energetic needs (Nievergelt, 1966; Peracino et al.,1989; Pedrotti, 1995).
Slope exposure, for most part of the year, does not influence greatly habitat choice by Alpine ibex: all exposures, even if in a differing extent, are used (Tosi et al., 1986). Detailed analyses of space use by Alpine ibex, in different populations, suggest however that south exposed slopes (south, south-east, south-west) are the most preferred during the whole year and are the only used during winter (Nievergelt, 1966; Peracino et al., 1989). The choice of the kind of exposure appears to be influenced by orography and thus can vary locally (Pedrotti, 1995).

In most Alpine ibex populations, wood does not appear to be important in habitat choice (Bachler, 1935; Rauch, 1937; Baumann, 1949; Kuster, 1961; Nievergelt, 1966; Pedrotti, 1995). Wood appears to be used by Alpine ibex only when it is crossed by ditches or interrupted by cliffs (Couturier, 1962; Hoffmann e Nievergelt, 1972), especially in the case of larch woods (Peracino et al., 1989; Bassano et al., 1992).

During winter seasons with scarce snowfalls, in areas with high density populations, adult and old Alpine ibex males can be found in larch and mixed larch-spruce (Abies alba) woods. Also in this case, however, the presence of small cliffs if of primary importance in the habitat choice (Bassano et al., 1992; Grignolio et al., 1999).

Some populations, more or less recently established in areas with low altitudes and strong vegetal cover , however appear to behave differently. For example, the Alpine ibex population of Hochlantschstock, in eastern Austra, 30 km distance from Graz, lacking of suitable winter terrain, uses coniferous and broadleaf forests for the whole winter, causing a significant damage to the forestry renewal (Kofler, 1981). Recent studies, based on the observation of marked Alpine ibex, have shown that in areas hosting high density populations (Albris-Swiss National Park), Alpine ibex uses forest for a good part of the year and particularly in the months of May, June and September. In these periods the damages to the vegetation, mostly due to Alpine ibex scratching and horning the plants, can be significant (Abderhalden & Buchli, 1998).

Low snow-falls are an important factor for survival in most species of the genus Capra (Schaller, 1977). Areas with precipitation higher than 1500 mm/year appear thus to be unfavourable for the establishment of Alpine ibex populations, while the optimal precipitation range for Alpine ibex goes from 100 to 1000 mm/year (von Elsner Schack, 1982). Villaret (1987), did not find any significant correlation between precipitation and the dynamics of Alpine ibex populations in France.
Intense precipitation can however have an effect on the survival of kids during spring (Nievergelt, 1966). Furthermore, even if the correlation between birth rate and precipitation appears not to be significant in the Gran Paradiso population (rs=-0.33, p= 0.08), the lowest values of birth rate are found in years characterised by heavy precipitation (Bassano et al., 1992).

These contradictory results are almost certainly due to considering snow and rain alike: snow precipitation are important not in terms of centimetres of fallen snow but mostly in terms of height and persistence on the soil of the snow cover (Choisy, 1990).
A significant correlation between mean height of snow cover and Alpine ibex population dynamics in Gran Paradiso National Park has been shown for males (rs= -0.52, p<0.05), females (rs= -0.69, p<0.05) and yearlings (rs= - 0.67, p<0.05)(Bassano et al., 1992).
Places with heavy snow cover are avoided by Alpine ibex (Couturier, 1962; Nievergelt, 1966; Hoffmann & Nievergelt, 1972). This avoidance appears to be due to the alpine ibex being scarcely adapted to snow, as suggested by its particular body mass/body surface ratio and the anatomy of its limbs and hoves (Kofler, 1983).