of competitors is higher. Second broods tend to have smaller clutches. Insularity also affects clutch size, with great tits on offshore islands laying smaller clutches with larger eggs than mainland birds. The eggs are white with red spots. The female undertakes all incubation duties, and is fed by the male during incubation. The bird is a close sitter, hissing when disturbed. The timing of hatching, which is best synchronised with peak availability of prey, can be manipulated when environmental conditions change after the laying of the first egg by delaying the beginning of incubation, laying more eggs or pausing during incubation. The incubation period is between 12 and 15 days. The chicks, like those of all tits, hatch unfeathered and blind. Once feathers begin to erupt, the nestlings are unusual for altricial birds in having plumage coloured with carotenoids similar to their parents (in most species it is dun-coloured to avoid predation). The nape is yellow and attracts the attention of the parents by its ultraviolet reflectance. This may be to make them easier to find in low light, or be a signal of fitness to win the parents' attention. This patch turns white after the first moult at age two months, and diminishes in size as the bird grows. Chicks are fed by both parents, usually receiving of food a day. Both parents provision the chicks with food and aid in nest sanitation by removing faecal packets, with no difference in the feeding effort between the sexes. The nestling period is between 16 and 22 days, with chicks being independent of the parents eight days after fledging. Feeding of the fledgeling may continue after independence, lasting up to 25 days in chicks from the first brood, but as long as 50 days in the second brood. Nestlings from second broods have weaker immune systems and body condition than those from first broods, and hence have a lower juvenile survival rate. Inbreeding depression occurs when the offspring produced as a result of a mating between close relatives show reduced fitness. The reduced fitness is generally considered to be a consequence of the increased expression of deleterious recessive alleles in these offspring. In natural populations of P. major, inbreeding is avoided by dispersal of individuals from their birthplace, which reduces the chance of mating with a close relative. Ecology The Eurasian sparrowhawk is a predator of great tits, with the young