Chappell MA, Janes DN, Shoemaker VH, Bucher TL, Maloney SK (1993). Reproductive effort in Adélie Penguins. Behavioral Ecology and Sociobiology 33:173-182.

ABSTRACT -- We estimated reproductive effort (energy expenditures for reproduction, as opposed to maintenance) in Adélie penguins breeding at Palmer Station, Antarctica. Data on body composition changes and metabolic rate were obtained using isotopic methods. Adélie breeding behavior consists of an initial courtship stage (during which both sexes fast), incubation, the 'guard' stage (when chicks are 1 to 18-28 days old), and the 'creche' stage (from the end of guarding until chicks are 38-45 days old). Both males and females lost considerable mass during the initial stages of the reproductive season, but males fasted longer and lost more mass. Mass losses of both sexes were comprised of 66% depot fat and 34% lean tissue. Mass and body composition remained constant once birds resumed feeding. The metabolic expenditure for the foraging necessary to accumulate the mass lost while fasting -- one component of reproductive effort -- was about 63 MJ in males and 39 MJ in females. Field metabolic rates (FMR) were low during courtship and while incubating, increasing more than 2-fold when birds resumed foraging. Although mean FMR increased between incubation and the creche stage, differences between stages were small and not significant. We used FMR data and an energy balance model to estimate the cost of feeding chicks. Results suggest a maintenance FMR of about 2.7 X basal metabolism (BMR), increasing to 3.4-3.6 X BMR during the creche stage. The reproductive effort (as metabolic expenditures) associated with feeding chicks is 31 MJ (males) to 36 MJ (females). Cumulative reproductive effort is 94 MJ in males and 75 MJ in females, or 5.3-6.2% of the annual energy budget. The reproductive effort devoted to chick care does not appear to be constrained by physiological or time limitations. Instead, selection to reduce the risk of predation may prevent the evolution of increased parental care.