Resource tracking by avian frugivores in mountain habitats of northern Spain

Systems comprising avian frugivores and fleshy-fruited plants are commonly used as models to study how animals respond to temporal and spatial variations in food abundance and distribution. Results have been far from conclusive, partly because of methodological constraints. In this study we investigated the fulfilment of a necessary condition for food tracking: correlation over time and space between the abundance of food and the abundance of food trackers. We used a paired-sample design in which fruit sites were compared with fruitless sites on two different spatial scales (landscape and habitat patch) in eight different river basins in the Cantabrian Mountains (NW Spain). Fruit and bird abundances were recorded monthly (October to February) at all sites during three consecutive fruiting seasons. Fruiting seasons (years) could be ranked by fruit abundance as low (2003), intermediate (2002) and high (2004). Since fruit sites were located at considerably higher elevations than fruitless sites, the possible effects of altitude, together with those of other uninvestigated factors, were taken into account by using non-frugivorous birds. Our results apparently suggest a temporal and spatial association between frugivores (avian seed dispersers) and fruit availability at the landscape scale but not at the smaller (habitat patch) scale. This, together with the absence of a similar pattern in non-frugivores, suggests that fruit availability is an important ecological factor affecting the autumn and winter abundance patterns of frugivorous birds and that this factor operates at the landscape scale but not at the habitat patch scale. Moreover, the negative elevational gradient in bird abundance that is typical of temperate mountain ecosystems was clearly reversed for frugivores in the intermediate and high fruiting years (suggesting that the negative effect of higher altitude may have been counterbalanced by the greater availability of food). As in other European upland areas, traditional farming practices in the Cantabrian Mountains are declining and, as a result, the proportion of fruitless habitats is increasing. Our study suggests that in these upland ecosystems frugivorous bird abundances in autumn and winter are associated to fruit supply, thus this habitat change tendency will probably have, in the long-term, a negative effect on bird populations.