Worldwide fisheries generate large volumes of fishery waste and it is often assumed that this additional food is beneficial to populations of marine top-predators. We challenge this concept via a detailed study of foraging Cape gannets Morus capensis and of their feeding environment in the Benguela upwelling zone. The natural prey of Cape gannets (pelagic fishes) is depleted and birds now feed extensively on fishery wastes. These are beneficial to non-breeding birds, which show reduced feeding effort and high survival. By contrast, breeding gannets double their diving effort in an attempt to provision their chicks predominantly with high-quality, live pelagic fishes. Owing to a scarcity of this resource, they fail and most chicks die. Our study supports the junk-food hypothesis for Cape gannets since it shows that non-breeding birds can survive when complementing their diet with fishery wastes, but that they struggle to reproduce if live prey is scarce. This is due to the negative impact of low-quality fishery wastes on the growth patterns of gannet chicks. Marine management policies should not assume that fishery waste is generally beneficial to scavenging seabirds and that an abundance of this artificial resource will automatically inflate their populations.

SUMMARY The main objective of this study was to determine heart rate(fh) and the energetic costs of specific behaviours of king penguins while ashore and while foraging at sea during their breeding period. In particular, an estimate was made of the energetic cost of diving in order to determine the proportion of dives that may exceed the calculated aerobic dive limit (cADL; estimated usable O2 stores/estimated rate of oxygen consumption during diving). An implanted data logger enabled fh and diving behaviour to be monitored from 10 free-ranging king penguins during their breeding period. Using previously determined calibration equations, it was possible to estimate rate of oxygen consumption(V̇O2) when the birds were ashore and during various phases of their foraging trips. Diving behaviour showed a clear diurnal pattern, with a mixture of deep (>40 m),long (>3 min) and shallow (<40 m), short (<3 min) dives from dawn to dusk and shallow, short dives at night. Heart rate during dive bouts and dive cycles (dive + post-dive interval) was 42% greater than that when the birds were ashore. During diving, fh was similar to the `ashore'value (87±4 beats min–1), but it did decline to 76% of the value recorded from king penguins resting in water. During the first hour after a diving bout, fh was significantly higher than the average value during diving (101±4 beats min–1) and for the remainder of the dive bout. Rates of oxygen consumption estimated from these (and other) values of fh indicate that when at sea, metabolic rate (MR) was 83%greater than that when the birds were ashore [3.15 W kg–1(–0.71, +0.93), where the values in parentheses are the computed standard errors of the estimate], while during diving bouts and dive cycles,it was 73% greater than the `ashore' value. Although estimated MR during the total period between dive bouts was not significantly different from that during dive bouts [5.44 W kg–1 (–0.30, +0.32)], MR during the first hour following a dive bout was 52% greater than that during a diving bout. It is suggested that this large increase following diving(foraging) activity is, at least in part, the result of rewarming the body,which occurs at the end of a diving bout. From the measured behaviour and estimated values of V̇O2, it was evident that approximately 35% of the dives were in excess of the cADL. Even if V̇O2 during diving was assumed to be the same as when the birds were resting on water,approximately 20% of dives would exceed the cADL. As V̇O2 during diving is, in fact, that estimated for a complete dive cycle, it is quite feasible that V̇O2 during diving itself is less than that measured for birds resting in water. It is suggested that the regional hypothermia that has been recorded in this species during diving bouts may be at least a contributing factor to such hypometabolism.

SUMMARY Warm-blooded diving animals wintering in polar regions are expected to show a high degree of morphological adaptation allowing efficient thermal insulation. In stark contrast to other marine mammals and seabirds living at high latitudes, Arctic great cormorants Phalacrocorax carbo have very limited thermal insulation because of their partly permeable plumage. They nonetheless winter in Greenland, where they are exposed to very low air and water temperatures. To understand how poorly insulated diving endotherms survive the Arctic winter, we performed year-round recordings of heart rate,dive depth and abdominal temperature in male great cormorants using miniature data loggers. We also examined the body composition of individuals in the spring. Abdominal temperatures and heart rates of birds resting on land and diving showed substantial variability. However, neither hypothermia nor significantly lower heart rate levels were recorded during the winter months. Thus our data show no indication of general metabolic depression in great cormorants wintering in Greenland. Furthermore, great cormorants did not reduce their daily swimming time during the coldest months of the year to save energy; they continued to forage in sub-zero waters for over an hour every day. As birds spent extended periods in cold water and showed no signs of metabolic depression during the Arctic winter, their theoretical energy requirements were substantial. Using our field data and a published algorithm we estimated the daily food requirement of great cormorants wintering in Greenland to be 1170±110 g day-1. This is twice the estimated food requirement of great cormorants wintering in Europe. Great cormorants survive the Arctic winter but we also show that they come close to starvation during the spring, with body reserves sufficient to fast for less than 3 days. Lack of body fuels was associated with drastically reduced body temperatures and heart rates in April and May. Concurrent, intense feeding activity probably allowed birds to restore body reserves. Our study is the first to record ecophysiological parameters in a polar animal on a year-round basis. It challenges the paradigm that efficient thermal insulation is a prerequisite to the colonization of polar habitats by endotherms.

Summary 1.Knowledge of the functional response of predators to prey densities conditions our understanding of food webs. Such links are still poorly understood within the higher trophic levels of marine ecosystems. 2. We present the first field study recording the foraging effort and foraging yield of a seabird (the Great Cormorant, Phalacrocorax carbo ) as well as the abundance and quality of prey within its foraging area. 3. We confirm that Great Cormorants foraging off West-Greenland show the highest foraging performance recorded for a marine predator (between 17 and 41 g fish caught per minute underwater). Former work suggests that such high foraging yield should be based upon the exploitation of extremely profitable prey patches. 4. Contrary to this hypothesis, average prey abundances estimated within the foraging areas of the cormorants were low (0·03-0·09 prey m − 2 , depending on methods), as was the average calorific value of the prey items (4·2 kJ g − 1 ). 5. Our study suggests that Great Cormorants remain highly successful predators even when exploiting modest prey resources. These findings have implications for our understanding of predator-prey relationships, and for the management of Great Cormorant populations.