In his seminal paper about using seabirds as indicators of marine food supplies, Cairns (1987, Biol Oceanogr 5:261-271) predicted that (1) parameters of seabird biology and behavior would vary in curvilinear fashion with changes in food supply, (2) the threshold of prey density over which birds responded would be different for each parameter, and (3) different seabird species would respond differently to variation in food availability depending on foraging behavior and ability to adjust time budgets. We tested these predictions using data collected at colonies of common murre Uria aalge and black-legged kittiwake Rissa tridactyla in Cook Inlet, Alaska. (1) Of 22 seabird responses fitted with linear and non-linear functions, 16 responses exhibited significant curvilinear shapes, and Akaike's information criterion (AIC) analysis indicated that curvilinear functions provided the best-fitting model for 12 of those. (2) However, there were few differences among parameters in their threshold to prey density, presumably because most responses ultimately depend upon a single threshold for prey acquisition at sea. (3) There were similarities and some differences in how species responded to variability in prey density. Both murres and kittiwakes minimized variability (CV < 15%) in their own body condition and growth of chicks in the face of high annual variability (CV = 69%) in local prey density. Whereas kittiwake breeding success (CV = 63%, r 2 = 0.89) reflected prey variability, murre breeding success did not (CV = 29%, r 2 < 0.00). It appears that murres were able to buffer breeding success by reallocating discretionary 'loafing' time to foraging effort in response (r 2 = 0.64) to declining prey density. Kittiwakes had little or no discretionary time, so fledging success was a more direct function of local prey density. Implications of these results for using 'seabirds as indicators' are discussed.

Summary1. Life-history theory predicts a trade-off between costs of current reproduction and future survival of individuals. Studies of short-lived animals in general support this prediction. However, the effect of nutritional stress during reproduction on survival of long-lived animals is poorly understood. 2. We examined the link between nutritional stress, fecundity and return to a breeding colony (hereafter 'survival') of black-legged kittiwakes (Rissa tridactyla) at two colonies with contrasting patterns in adult survival, fecundity, and numerical trends. 3. We tested the observational (at Duck and Gull Is., Cook Inlet, Northern Gulf of Alaska) and experimental (at Middleton I., Gulf of Alaska) relationships between variations in the secretion of the stress hormone corticosterone (CORT) and food abundance. Then, we examined the relationships between nutritional stress (as reflected in CORT), reproduction, and survival of individuals. 4. On average, CORT was higher in kittiwakes breeding on Duck I. (declining, low fecundity, high survival) compared to those breeding on Gull I. (increasing, high fecundity, low survival). 5. At both colonies, CORT was directly negatively correlated with food abundance quantified at sea. Experimental feeding of individuals ad libitum resulted in a reduction of CORT in birds breeding on Middleton I. These results suggest that CORT is a reliable measure of food availability and defines nutritional stress (stress) in kittiwakes. 6. On Gull I., where survival is low (86%), production of young declined as stress increased. On Duck I., where survival is high (93%), parents always failed in raising young, though they experienced a wide range of stress levels. 7. Survival of individuals is linked to their CORT levels during reproduction. High levels of CORT predicted disappearance of individuals from both colonies. 8. The results support the hypothesis that nutritional stress during reproduction affects both survival and reproduction in long-lived animals. However, even within a species the ways in which survival and reproduction trade-off against each other may vary among populations. Results suggest that reproductive consequences of nutritional stress might differ between declining and increasing populations, which should be tested. We conclude that severity of nutritional stress during reproduction is one of the major factors defining population processes in kittiwakes.

Flexible time budgets allow individual animals to buffer the effects of variable food availability by allocating more time to foraging when food density decreases. This trait should be especially important for marine predators that forage on patchy and ephemeral food resources. We examined flexible time allocation by a long-lived marine predator, the Common Murre (Uria aalge), using data collected in a five-year study at three colonies in Alaska (USA) with contrasting environmental conditions. Annual hydroacoustic surveys revealed an order-of-magnitude variation in food density among the 15 colony-years of study. We used data on parental time budgets and local prey density to test predictions from two hypotheses: Hypothesis A, the colony attendance of seabirds varies nonlinearly with food density; and Hypothesis B, flexible time allocation of parent murres buffers chicks against variable food availability. Hypothesis A was supported; colony attendance by murres was positively correlated with food over a limited range of poor-to-moderate food densities, but independent of food over a broader range of higher densities. This is the first empirical evidence for a nonlinear response of a marine predator's time budget to changes in prey density. Predictions from Hypothesis B were largely supported: (1) chick-feeding rates were fairly constant over a wide range of densities and only dropped below 3.5 meals per day at the low end of prey density, and (2) there was a nonlinear relationship between chick-feeding rates and time spent at the colony, with chick-feeding rates only declining after time at the colony by the nonbrooding parent was reduced to a minimum. The ability of parents to adjust their foraging time by more than 2 h/d explains why they were able to maintain chick-feeding rates of more than 3.5 meals/d across a 10-fold range in local food density.

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