Østnes, J.E., B.M. Jenssen and C. Bech (2001). Growth and development of homeothermy in nestling European Shags (Phalacrocorax aristotelis). The Auk 118: 983-995.

European Shag (Phalacrocorax aristotelis) nestlings were studied on a small island off the coast of central Norway. Increase in body mass (BM) with age (t, days) was described by the logistic equation: BM = 1,622 g / [1 + e0.172(t 19.9)]. All growth parameters measured (body mass, and length of tarsus, wing and head) showed the highest relative growth rate when the nestlings were 5-10 days old, that is, before the nestlings had achieved homeothermy. An incipient endothermic response was noted when the nestlings were 9 days old, and they became homeothermic at ages of 15-18 days. Respective mass-specific resting metabolic rates for nestlings 0, 15 and 45 days old were 47, 261 and 147% of the predicted value for adult nonpasserine birds of similar body masses. Mass-specific minimal thermal conductance decreased from 366% of predicted adult value at hatching, to 220% of that predicted when nestlings were 21 days old. For nestlings 15 days old the factorial metabolic scope (resting metabolic rate/peak metabolic rate) was only 1.5, but that increased rapidly thereafter. Rapid increase in the mass-specific RMR and decrease in the minimal thermal conductance is suggested to contribute importantly to improve the homeothermic ability during the first two weeks of the developmental period. At hatching, leg and pectoral muscles constituted 5.3 and 2.2%, respectively, of total wet body mass. Relative leg muscle mass increased rapidly and had almost reached adult proportions when the nestlings were 25-30 days old. In contrast, pectoral-muscle mass increased in an almost direct proportion to the body mass during the first 30 days of the growth period, and increased rapidly thereafter. At hatching, the water fraction (water content/lipid-free wet mass) was significantly lower in the leg than in the pectoral muscles (0.920 vs. 0.931). The water fraction of the leg muscles also remained lower during the entire growth period. Judging from the proportionately greater mass and higher degree of maturity of the leg compared to the pectoral muscles, the former would seem to be the main site of cold-induced heat production during early development of homeothermy in European Shag nestlings.