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A scanning electron microscope study of the morphological changes which occur in shells of hen and quail eggs during incubation is described. The results are compared with observations on the shells of hatched eggs taken from a range of species. It was found that less change occurred in the shells of altricial species than precocial ones, the difference being associated presumably with a smaller calcium requirement for developing altricial embryos. The cores rather than the sides of mammillae appeared to be the major sites of erosion.

Summary 1. Three basic hatching methods are described together with one subsidiary method. The symmetrical method is characterized by rotation of the chick in the egg during hatching climax; a line of damage around the circumference of the egg is evident at the beginning of a pushing phase which causes a fairly symmetrical cap to be broken from the shell. The asymmetrical method is used by a few long-billed species; it involves little or no rotation of the chick in the egg, and produces asymmetrical shell remains. The megapodes have developed a unique hatching method in response to their unusual incubation conditions. Parental assistance has been observed in some species, but only as an auxiliary to either the symmetrical or asymmetrical method. Approximately 150 species have been categorized according to the hatching method(s) they use. 2. Among those species adopting the symmetrical method, there is considerable variability as to how far the chick turns in the egg during hatching climax. On this basis, a spectrum of behaviour, expressed in terms of angle of rotation (θ), has been proposed. At one end lie species such as the bobwhite quail and little owl (θ≥ 360°), and at the other, the ostrich and rhea (θ≤ 90°). 3. The theory that, with the exception of the megapodes, there is only one basic hatching method is examined. The tenets of this theory are found to be inconsistent with recent observations of species differences in hatching behaviour. It is concluded that hatching behaviour is governed by an intrinsic species-specific programme, in turn influenced by mechanical or other external factors. 4. The literature contains several suggestions as to the external factors that might influence hatching technique, but only one detailed investigation. It is proposed that interspecific differences in the mechanical properties of the egg integument (the shell and its underlying membranes) can be regarded as forming a spectrum from very brittle to comparatively tough. The amount of climax rotation by the chick is seen as an adaptive response to brittleness or toughness of the egg integument. The hatching technique of the chick is further modified as a response to the effect of moisture content on the integument. 5. The selective pressures leading to the evolution of the symmetrical and asymmetrical hatching methods are discussed. The previous model for the development of the asymmetrical method is amended to account for those species of gull which may adopt either method.

Mechanical tests, including experiments designed to simulate the pipping and hatching processes, were combined with acoustic-emission analysis to show that the eggshells of some avian species (e.g. domestic hen and domestic duck) may be classified as hard and brittle, and others (e.g. Japanese quail and pigeon) as tougher and more flexible. Laboratory and field studies of hatching, and observations of cine films, demonstrated that these differences in the mechanical properties of the shell were related to hatching technique. Whilst chicks in hard brittle Shells made relatively few holes before pushing away a cap, the hatching of chicks contained in tougher, more flexible Shells was characterized by extensive Perforation along a latitude of the shell. Indeed, the latter often continued to perforate the shell for more than a complete revolution before pushing away a cap. The results are then discussed in relation to avian nest environment.