FACTORS GOVERNING FEEDING RATE, FOOD REQUIREMENT AND BROOD SIZE OF NESTLING GREAT TITS PARUS MAJOR
Authors: Royama T.R 
Published: April 2008

SUMMARYObservations were made on feeding rates and food‐consumption of nestling Great Tits Parus major mainly in Larch plantations at lake Yamanaka, Japan.Feeding frequencies were recorded by an automatic recorder. There were marked differences between early and late broods; the feeding frequencies were twice as great in early than in late broods of the same size. No clear tendency was observed in the variations of feeding frequencies in relation to brood size. There was, however, a clear inverse relationship between the frequencies and the average size of food brought to the nests. The males' share in terms of feeding frequencies is described. These figures, however, did not follow the males' contribution in terms of weight of food, which was nearly always higher than the females'. It is pointed out that feeding frequencies are far too variable to be used as a true index of food consumption by nestlings, and are not reliable.Attempts were made to measure the weight of food; the method is described. The average weight of food brought by males was lighter in early than in later broods. The total weight of food was estimated. The trend of daily food consumption per chick was similar to that of the chick's growth curve. It was found that up to about the tenth day of the nestling period daily food‐intake per chick increased linearly as body weight increased.At some nests, rate of defaecation was observed. This was at first low, but it increased steeply on the third day, with a steady increase thereafter.By comparing the rates of food intake, faeces output, and weight increment of a chick, it was found that only 20–30% of digested matter (the difference between food‐intake and faeces‐output was used up daily (for body temperature regulation various external effort, etc.). The factors responsible for this high efficiency of growth in nestlings are discussed.There was a clear inverse relationship between the total weight of food brought per chick per day and the brood size. This is largely because the heat‐loss is greater in small than in large broods, so that a chick from a small brood in fact needs more energy to maintain its body temperature after a certain age than one from a large brood. This is discussed in detail.Factors which caused variations in size of food are discussed in relation to feeding frequencies.It is pointed out that, because of the inverse relationship between energy requirement by each chick and brood size, the total food requirement by a brood as a whole did not vary directly in proportion to the brood size. An estimation showed that a b/3 still required about 75% of the total food required by a b/8. A smaller brood is less advantageous than expected to parents feeding nestlings when they encounter adverse conditions, e.g. food shortage in the habitat, or a lack of help by their mates, etc. On the other hand, it is suggested that once they have left the nest, the food‐demand by a brood of fledglings the parents have to feed, so that, in the fledging period, in times of food shortage it would certainly be advantageous to have fewer young. It is suggested that, although fledglings may consume three to four times as much food as nestlings, the parents, in providing this food, would not work proportionately harder, since the parents' efficiency of providing food could be higher in feeding the fledglings, which always follow the parents as they are hunting, than in feeding the nestlings to which food has to be brought.On this basis, the adaptive significance of the length of the nestling period in nidicolous species is discussed in relation to clutch size, brood size and food requirement.