Breeding energetics and food requirements of gentoo penguins (Pygoscelis papua) at Heard and Macquarie Islands

The food and energy requirements of breeding gentoo penguins ( Pygoscelis papua ) were studied at Heard and Macquaric Islands by means of isotope turnover techniques.
The food consumption rates of chicks were measured at various stages of growth, providing estimates of the total food provided by adults to rear a chick to fledging.
The energy expenditures of attending adults were also measured at different stages of chick-rearing, allowing the total energy costs associated with breeding to be established for a pair of adults and at the population level on both islands. We estimate the total annual energy budget of a 6·2 kg breeding gentoo penguin to be 1517 MJ which is equivalent to the consumption of 292 kg prey.     

Foraging strategies of incubating and brooding king penguins Aptenodytes patagonicus

For oceanic birds like king penguins, a major constraint is the separation of foraging areas from the breeding colony, largely because swimming increases foraging costs. However, the relationship between foraging strategy and breeding stage has been poorly investigated. Using time-depth recorders, we studied the diving behaviour of two groups of king penguins that were either incubating or brooding chicks at Crozet Islands (Southern Indian Ocean) at the same period of the year. Although birds with chicks had the highest predicted energy demand, they made foraging trips half as long as incubating birds (6 vs. 14 days) and modified their time and depth utilisation. Birds with chicks dived deeper during daylight (mean maximum depth of 280 m vs. 205 m for those incubating). At night, birds with chicks spent twice as much time diving as those incubating, but birds at both stages never dived beyond 30 m. Movements to greater depths by brooding birds are consistent with the vertical distribution of myctophid fish which are the main prey. As chick provisioning limits trip duration, it is suggested that it is more efficient for parents to change their diving patterns rather than to restrict their foraging range. Received: 23 June 1997 / Accepted: 3 November 1997     

Contrasting bottom-up effects of warming ocean on two king penguin populations

Breeding success is often correlated with climate, but the underlying bottom-up mechanisms remain elusive—particularly in marine environments. Consequently, conservation plans of many species often consider climate change as a unilateral threat, ignoring that even nearby populations can show contradicting trends with climate. Better understanding the relationship between climate and environment at different scales can help us interpret local differences in population trends, ultimately providing better tools to evaluate the global response of a species to threats such as global warming. We studied a growing king penguin population nesting at Kerguelen island (Southern Indian Ocean), hosting one of the largest colonies in the world. We used a unique dataset of foraging, breeding success, and climate data spanning over 25 years to examine the links between climate, marine environment, and breeding success at this colony. The results were then compared to the neighboring population of Crozet, which experienced the steepest decline for this species over the past few decades. At Crozet, penguins experienced lower breeding success in warmer years due to productive currents shifting away from the colony, affecting foraging behavior during chick rearing. At Kerguelen, while chick mass and survival experienced extreme variation from year to year, the annual variation was not associated with the position of the currents, which varied very little compared to the situation in Crozet. Rather than being affected by prey distribution shifts, we found evidence that chick provisioning in Kerguelen might be influenced by prey abundance, which seem to rather increase in warmer conditions. Furthermore, warmer air temperature in winter increased chick survival rate, likely due to reduced thermoregulation cost. Investigating the mechanisms between climate and fitness allowed us to predict two different fates for these populations regarding ongoing global warming.     

Food requirements of breeding king penguins at Heard Island and potential overlap with commercial fisheries

The diet composition of king penguins (Aptenodytes patagonicus) at Heard Island (53°05′S; 73°30′E) was determined from stomach contents of 98 adults captured as they returned to the island throughout 1992. During the two growth seasons, the diet was dominated by the myctophid fish Krefftichthys anderssoni (94% by number, 48% by mass). The paralepidid fish Magnisudis prionosa contributed <1% by numbers but 17% by mass. Mackerel icefish (Champsocephalus gunnari) accounted for 17% by mass of chick diet in late winter, when chicks were malnourished and prone to starvation, although its annual contribution to the penguins' diet was only 3%. Squid was consumed only between April and August; Martialia hyadesi was the commonest squid taken, comprising 40–48% of the winter diet. The remainder of the diet consisted of the squid Moroteuthis ingens and fish other than K. anderssoni. The energy content of the diet mix fed to the chicks varied seasonally being highest during the growth seasons (7.83 ± 0.25 kJ g⁻¹) and lowest in winter (6.58 ± 0.19 kJ g⁻¹). From energetic experiments we estimated that an adult penguin consumed 300 kg of food each, of which its chick received 55 kg during the 1992 season. The chicks received large meals at the beginning of winter (1.2 ± 0.3 kg) and during the middle of the second growth season (1.2 ± 0.3 kg), and their smallest meals in late winter (0.4 ± 0.1 kg). The gross energy required to rear a king penguin chick was estimated to be 724 MJ. The potential impact of commercial fisheries on the breeding activities of king penguins is discussed. Received: 20 October 1997 / Accepted: 27 April 1998     

Satellite data identify decadal trends in the quality of Pygoscelis penguin chick‐rearing habitat

Pygoscelis penguins are experiencing general population declines in their northernmost range whereas there are reported increases in their southernmost range. These changes are coincident with decadal‐scale trends in remote sensed observations of sea ice concentrations (SIC) and sea surface temperatures (SST) during the chick‐rearing season (austral summer). Using SIC, SST, and bathymetry, we identified separate chick‐rearing niche spaces for the three Pygoscelis penguin species and used a maximum entropy approach (MaxEnt) to spatially and temporally model suitable chick‐rearing habitats in the Southern Ocean. For all Pygoscelis penguin species, the MaxEnt models predict significant changes in the locations of suitable chick‐rearing habitats over the period of 1982–2010. In general, chick‐rearing habitat suitability at specific colony locations agreed with the corresponding increases or decreases in documented population trends over the same time period. These changes were the most pronounced along the West Antarctic Peninsula where there has been a rapid warming event during at least the last 50 years.     

Relevance of forage fish in the diet of Magellanic penguins breeding in northern Patagonia,Argentina

We quantified the trophic niche of Magellanic penguins (Spheniscus magellanicus) breeding and moulting in Golfo San Jorge, Argentina, through conventional stomach content and stable isotope analysis. A total of 112 adults were flushed during the early and late chick stages of 2011 and 2012 at Isla Vernacci Norte, and at least 15 prey taxa were found, including fishes, cephalopods, crustaceans and polychaetes. Overall, Argentine anchovy (Engraulis anchoita) showed the highest contribution in terms of importance by mass (68.1–85.3%, depending on chick stage and year), except for the old chick stage in 2011 when the shortfin squid (Illex argentinus) was the main prey consumed (56.0%). Based on carbon and nitrogen isotopic values from a total of 256 blood samples, corresponding to young and old chicks and to adults of both sexes sampled throughout the incubation, chick and moult stages at the above mentioned colony and years, Bayesian mixing model outputs showed that Argentine anchovy was always the main prey (48–86%). Bayesian mixing model outputs obtained from adults of both sexes and their chicks during the late chick stage of 2013 at Isla Vernacci Norte, Isla Tova and Isla Leones also showed that Argentine anchovy was the main prey consumed. This is the first comprehensive assessment of Magellanic penguin diet composition in northern Patagonia, quantifying the relative contribution of prey in the diet of adults and chicks at different stages of the annual cycle and years, and confirms the relevance of a forage fish such as the Argentine anchovy in its trophic ecology.     

Evaluating the prudence of parents: daily energy expenditure throughout the annual cycle of a free-ranging bird, the macaroni penguin Eudyptes chrysolophus

We measured daily energy expenditure (DEE) continuously for a whole year in a free ranging bird, the macaroni penguin Eudyptes chrysolophus . We combined these measurements with concurrently recorded foraging behaviour, and literature information on body mass and dietary factors to estimate prey consumption rates and foraging success. DEE was at a maximum during late chick-rearing but was equally high during all other active phases of the breeding season. DEE was approximately 4×resting metabolic rate, which accords with established theory and suggests a common 'energetic ceiling' throughout the summer period. However, whether this represents a maximum in physiological capacity, or a rate which optimises fitness is still unclear. Rates of prey consumption and foraging success followed different patterns from daily energy expenditure. Daily prey consumption was high as the penguins prepared for long fasts associated with moulting and incubation but relatively low during chick-rearing, when foraging areas were restricted and foraging success lower. It appears that the energy intake of macaroni penguins is subject to extrinisic or environmental constraints rather than to intrinsic physiological limits.     

Penguins use the two-voice system to recognize each other

The sound-producing structure in birds is the syrinx, which is usually a two-part organ located at the junction of the bronchi. As each branch of the syrinx produces sound independently, many birds have two acoustic sources. Thirty years ago, we had anatomical, physiological and acoustical evidence of this two-voice phenomenon but no function was known. In songbirds, often these two voices with their respective harmonics are not activated simultaneously but they are obvious in large penguins and generate a beat pattern which varies between individuals. The emperor penguin breeds during the Antarctic winter, incubating and carrying its egg on its feet. Without the topographical cue of a nest, birds identify each other only by vocal means when switching duties during incubation or chick rearing. To test whether the two-voice system contains the identity code, we played back the modified call of their mate to both adults and also the modified call of their parents to chicks. Both the adults and the chicks replied to controls (two voices) but not to modified signals (one voice being experimentally suppressed). Our experiments demonstrate that the beat generated by the interaction of these two fundamental frequencies conveys information about individual identity and also propagates well through obstacles, being robust to sound degradation through the medium of bodies in a penguin colony. The two-voice structure is also clear in the call of other birds such as the king penguin, another non-nesting species, but not in the 14 other nesting penguins. We concluded that the two-voice phenomenon functions as an individual recognition system in species using few if any landmarks to meet. In penguins, this coding process, increasing the call complexity and resisting sound degradation, has evolved in parallel with the loss of territoriality.     

Nutrient reserve storage,energetics, and food consumption during the prebreeding and premoulting foraging periods of king penguins

Nutrient reserve storage during the prefasting foraging trips of king penguins Aptenodytes patagonicus was investigated by measuring body composition at the beginning and the end of the prebreeding and premoulting periods at sea. Both periods were marked by a 45–47% increase in body mass (4.6 and 5.6 kg during the prebreeding and premoulting trips, respectively) including body water (31% and 55% of the total increase in body mass), protein (13% and 16%) and lipid (46% and 23%), but not ash. Fat accretion accounted for most of the energy stored (86% and 71% versus 14% and 29% for protein) and it was mainly located in the subdermal depot, the larger increase in body protein occurring in pectoral muscles. Daily mass gain was lower during the prebreeding foraging trip than during the premoulting one (195 versus 328 g/day), while there was no difference in the rate of energy storage (4,097 and 4,155 kJ/day). The total energy costs of the foraging periods were calculated to be 381 and 315 MJ during the prebreeding and premoulting trips respectively, which corresponded to a 70-kg and a 58-kg food intake. The consumption of marine resources by the king penguin population from Crozet Islands was estimated to be 166,000 tons in spring/summer during the two foraging periods preceding long-term fasts.     

Impact of predation on king penguin Aptenodytes patagonicus in Crozet Archipelago

Predation can have major effects on population dynamics, but predator–prey interactions in marine ecosystems have rarely been studied. While the king penguin is one of the most studied seabirds, little is known about the impact of predation on its population dynamics. Here, we determine the impact of the main predators (giant petrels and skuas) on king penguin breeding success taking into account the nocturnal predation of petrels. We found that predation is the most important source of breeding failure for king penguins. The smallest chicks within crèches are the most hunted. The periphery of the colony suffers the highest risk of predation during summer. Our study shows the unequal quality of some areas inside the colony in terms of predation risk and breeding success, and points out the importance of timing in successful breeding.     

Nocturnal predation of king penguins by giant petrels on the Crozet Islands

Dietary segregation of sympatric seabirds in the Southern Ocean is partly linked to differences in their foraging techniques. We have investigated the activity of giant petrels (Macronectes spp.) in a king penguin (Aptenodytes patagonicus) colony day and night during the austral winter of 2001 on the Crozet Islands. Using an automatic identification system and an infrared video camera, we followed 15 petrels tagged with micro transponders. Our data show that giant petrels predate king penguin chicks during the night. The activity of giant petrels is even slightly higher during nighttime than during the day. In addition, our data show a higher nocturnal activity by northern giant petrels (M. halli) than by southern giant petrels (M. giganteus). These unexpected results raise questions concerning visual adaptations to nocturnal foraging in giant petrels and their potential impact on the sleep, vigilance and crèching behavior of penguin chicks.     

Ontogeny of thermoregulatory mechanisms in king penguin chicks (Aptenodytes patagonicus)

The rapid maturation of thermoregulatory mechanisms may be of critical importance for optimising chick growth and survival and parental energy investment under harsh climatic conditions. The ontogeny of thermoregulatory mechanisms was studied in growing king penguin chicks from hatching to the full emancipation observed at 1 month of age in the sub-Antarctic area (Crozet Archipelago). Newly hatched chicks showed small, but significant regulatory thermogenesis (21% rise in heat production assessed by indirect calorimetry), but rapidly became hypothermic. Within a few days, both resting (+32%) and peak (+52%) metabolic rates increased. The first week of life was characterised by a two-fold rise in thermogenic capacity in the cold, while thermal insulation was not improved. During the second and third weeks of age, thermal insulation markedly rose (two-fold drop in thermal conductance) in relation to down growth, while resting heat production was slightly reduced (-13%). Shivering (assessed by electromyography) was visible right after hatching, although its efficiency was limited. Thermogenic efficiency of shivering increased five-fold with age during the first weeks of life, but there was no sign of non-shivering thermogenesis. We conclude that thermal emancipation of king penguin chicks may be primarily determined by improvement of thermal insulation after thermogenic processes have become sufficiently matured. Both insulative and metabolic adaptations are required for the rapid ontogeny of thermoregulation and thermal emancipation in growing king penguin chicks.     

Behavioural time budget of breeding king penguins (Aptenodytes patagonica)

As do so many other seabirds, penguins fast when ashore for breeding. For penguins in dense colonies, territory defence seems to imply conflicting energetic requirements because of its assumed high energy cost, when the birds need to limit energy expenditure to cope with their fast. In this context, behavioural time budget over 24 h was investigated during breeding in the king penguin, Aptenodytes putugonicu , by using a remote-controlled videocamera. The comparison of day-night activity was performed in relation to breeding status (incubation vs. brooding) and duration of fasting (beginning vs. end of incubation shift). Five categories of behaviours were quantified: territoly defence, comfort, resting, sleeping and chick-feeding. Breeding king penguins remain active by day as well as by night. Between incubation and brooding we found a three-fold increase in the energy consuming temtory defence, together with a drastic decrease in that posture which corresponds to deep sleep, is. when most energy is saved. These increases in aggressiveness and vigilance may be related to protection of the newly hatched chick. Between the onset and the end of an incubation shift, the time spent in sleep increases three-fold, whereas territory defence remains unchanged. These data for penguins under natural conditions accord with previous studies on captive birds which have shown that an increasing proportion of sleep during the course of fasting may contribute to energy saving. On the other hand, both resting (which is the main component of penguins'time budget; about 65%) and comfort (about 16% of time) show no change either between incubation and brooding or during the course of fasting.     

Diet and foraging effort of Adélie penguins in relation to pack-ice conditions in the southern Ross Sea

We investigated the diet and aspects of foraging effort among Adélie penguins (Pygoscelis adeliae) breeding at three colonies on Ross Island, in the southwestern Ross Sea – Capes Royds, Bird and Crozier – during the chick-provisioning period of three austral summers, 1994–1995, 1995–1996 and 1996–1997. During the study period, pack-ice cover differed in waters offshore of these colonies, by colony, seasons and year. Diet differed among colonies only slightly. The fish Pleuragramma antarcticum was the most important prey, especially during years or periods within years when little pack ice was present. With respect to krill, which composed the remainder of diet, juvenile Euphausia crystallorophias were consumed predominantly in a year of heavy pack-ice cover; more adult krill were consumed in 2 years when pack ice was sparse. Foraging trip duration differed by colony, season and year and was related directly to distance from the colony to the nearest pack ice. The amount of food brought to chicks increased as trip duration increased, to a point (2 days), but then decreased as duration increased further (up to 4 days). On the basis of data on mass of parents and of meal sizes to chicks, it appeared that on the longest trips more of the food gathered by parents was used for self maintenance; on the longest trips, parents lost body mass. Successful foraging during chick rearing, the period when adult foraging is most intense, appears to depend on the proximity of pack ice to nesting colonies for this penguin species. Received: 1 October 1997 / Accepted: 25 April 1998     

Temporal variation in the diet of gentoo penguins at the Falkland Islands

Knowledge of diet is critical in interpreting the ecological roles of marine top predators and provides information towards their conservation and management. The Falkland Islands hold the largest number of breeding gentoo penguins. Yet knowledge of gentoo penguin diet at the Falklands is limited to either broad taxonomic divisions of prey items or dietary samples collected only on a single day. This study is the first to investigate gentoo penguin diet at Cow Bay, Falklands, to the species level, over repeated sampling intervals during the breeding period. Through stomach content analysis, we determined diet over a large temporal scale (2002/2003/2004–2011/2012/2013) and between the guard and crèche periods of chick rearing. The principle prey item by reconstituted mass was rock cod fish Patagonotothen spp., for all periods (47–78 %) except that of the 2012/2013 crèche period (19 %) when Falkland herring Sprattus fugensis made up the bulk of the diet (52 %). Of the cephalopods recovered, Patagonian squid Doryteuthis gahi was prominent (1–24 %), while crustaceans contributed negligibly to gentoo penguin diet. Our findings revealed that gentoo penguins breeding at the Falkland Islands were primarily demersal foragers with an ability for pelagic feeding. Diet choice appears to reflect prey availability.     

Stomach stones in king penguin chicks

Many animals that possess a gizzard swallow stones or sandy grit, supposedly to aid in the mechanical breakdown of food. While this has been well documented in the literature, our study is the first to report the presence of stones in the gizzard of king penguin chicks. We found stones, so called ‘gastroliths’, in the pyloric region of the gizzard, the part of the digestive tract that is specialised for the mechanical breakdown of food. Stones were already present in the gizzard of chicks and, hence, during the first year of the life of king penguins, which is spent on land. Some chicks were found to have more than 130 stones (0.5–22 mm in size) in their gizzard. The gastroliths we found in king penguins are of the same geological origin as rocks present at the colony, which suggests that birds swallowed them there. The functional role of gastroliths in penguin chicks and adults is still unknown. We discuss the potential roles that these gastroliths might play in king penguins (i.e. aid in digestion, buoyancy control during foraging at sea, adaptation to fasting).     

The post-moult diet of Emperor Penguins (Aptenodytes forsteri) in the eastern Weddell Sea,Antarctica

The diet of emperor penguins Aptenodytes forsteri was studied during late austral summer at Drescher Inlet, eastern Weddell Sea, Antarctica. Antarctic krill Euphausia superba was a major component of the food, accounting for 75% of all prey items. Emperor penguins appear to feed on krill during shallow dives under the fast sea ice. Fish, mainly nototheniids, accounted for less than 20% by number of all prey. An evaluation of the main prey types in terms of mass indicated, however, that fish represented up to 75% approximately of prey mass. Feeding experiments were performed on captive penguins and showed that squid beaks can accumulate for up to 3 weeks within the stomach without any clear signs of erosion. The lack of cephalopod soft parts in the samples makes it likely that all squid beaks were derived from animals captured some time previously. Squid seems to be a very minor dietary component of emperor penguins at the Drescher Inlet.     

Ticks Ixodes uriae and the breeding performance of a colonial seabird, king penguin Aptenodytes patagonicus

High densities of penguins in their colonies and the continuous use of these sites over consecutive reproductive periods increase the risk of development of tick populations. We have studied the effects of tick parasitism by Ixodes uriae in a colony of king penguins Aptenodytes patagonicus at Possession Island during three breeding seasons. We investigated the prevalence and periods of tick infestation during the one-year breeding cycle of penguins. The effects of tick parasitism on penguin breeding performance were assessed from photographs of the colony and with an automatic penguin identification system. We compared two groups of penguins carrying individual subcutaneous electronic tags, one group breeding in an infested area and the other in a non-infested area. Tick feeding activity was coincident with the periods when adult penguins stayed ashore for six days or more, i.e. during the incubating period. This duration corresponds to the duration of a tick meal on the host. The level of infestation varied between years. Penguins showed a lower incubating success in infested areas during a year of high infestation. In an infested area, individuals seen with ticks had a lower breeding success in rearing a one-year old chick than those seen without ticks.     

The food and feeding ecology of the giant petrels Macronectes halli and M. giganteus at South Georgia

Giant petrels are the chief scavenging seabirds in the Southern Ocean. Quantitatively analysed regurgitations from chicks of both Macronectes halli and M. giganteus at Bird Island, South Georgia throughout the chick rearing period consisted predominantly of adult Macaroni penguins Eudyptes chrysolophus. Significant quantities of burrowing petrels and krill were taken but the amount varied in different years. Only male M. halli fed regularly at fur seal carcases. Although 10 species were found in regurgitations, squid were unimportant in the diet. Fish are possibly more important during the winter.
Intersexual differences in diet were of greater significance than interspecific ones with males taking more carrion and females more crustaceans. Chicks were fed during both day and night and in both species males received significantly more food than females during the fledging period. M. giganteus chicks received more than M. halli at all ages.
Despite their extensive reliance on carrion both species take a variety of prey and this may be especially important during the winter when penguin and seal carrion is scarce.     

Diurnal sleep depth changes in the king penguin (<Emphasis Type="Italic">Aptenodytes patagonicus</Emphasis>)

Avian sleep quality depends on its depth (deeper sleep being of better quality). In king penguins (Aptenodytes patagonica), sleep may be disturbed by congeners passing in the sleeper's vicinity. As king penguin activity is increased in the morning, sleep disturbances are more likely to occur during this time period. One might therefore assume that afternoon sleepers (AS) sleep more profoundly than morning sleepers (MS). To test this hypothesis, we examined the diurnal variations in sleep depth of king penguins sleeping in resting sites adjacent to the colony of 'La Baie du Marin' (Crozet Archipelago). We measured the bodily tactile arousal threshold at the upper back level. The arousal threshold in AS was twice as high as in MS. This study demonstrates for the first time that sleep depth changes according to time of day in a diurnal wild bird. We postulate that diurnal sleep depth is increased due to decreased congener movements close to the sleeping penguin.