THE ROLE OF NEST SITES IN LIMITING THE NUMBERS OF HOLE-NESTING BIRDS - A REVIEW

The breeding densities of many bird-species which nest in tree cavities are in some areas limited by shortage of sites. This is evident from circumstantial evidence in which the numbers of breeding pairs in different areas correlate with the numbers of local nest sites, or where changes in the numbers of nest sites resulting from natural processes or human action are followed by changes in the numbers of pairs. It is also shown experimentally, where nest site provision or removal has been followed by a corresponding change in breeding density. When nest boxes were provided, they were often occupied in the same year, leading to an immediate rise in breeding density. This implied that surplus birds were available in the vicinity and able to take them up. In future years, pair numbers often increased further but eventually they levelled off. This implied that, at this higher level, other limiting factors came into play. Experiments on nest site provision thus provide examples of different factors acting successively to limit breeding density at different levels. By constraining breeding density, shortage of nest sites not only prevents some birds from breeding, but must also limit the total numbers of breeders and non-breeders, because no population can increase beyond a certain point, when the production of young is limited. Once all available nest sites are occupied, reproduction is inevitably density-dependent with respect to the total population of breeders and non-breeders. Because several bird species can use the same type of site, competition is frequent, and the numbers of dominant species can affect the numbers and distribution of others. In extreme situations, a species may be totally absent from areas where all suitable nest sites are occupied by dominant competitors. In less extreme situations, the numbers of a subordinate species may vary from year to year or from place to place in inverse relation to the numbers of a dominant species. Both these situations can be changed by the provision of extra sites. In managed woods and modern tree plantations, most hole-nesting bird species are excluded or kept at extremely low levels by shortage of nest sites. This is a direct result of forestry practice, which usually entails removal of the old and dying trees that normally provide the sites. The problem can be rectified by allowing a number of old and dead trees to remain in managed forests, or (on a more local scale) by the provision of nest boxes. By appropriate design and positioning of the boxes, the most desired species can be favoured.