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     

Foraging parameters of gentoo penguins Pygoscelis papua at Marion Island

Summary We measured the foraging parameters of breeding gentoo penguins Pygoscelis papua at sub-Antarctic Marion Island. Mean swimming speed was 7.9 km h^-1. Penguins spent on average 8.1 h away from the colony if they returned on the same day they left and 23.7 h away if they remained at sea overnight. Sixteen percent of the total time away at sea was spent swimming. Time spent swimming, and consequently distance travelled, during a foraging trip were highly variable and showed significant intercolony differences. However, 80% of all foraging trips totalled less than 40 km. Meal sizes were small and there was no correlation between meal size and distance travelled, suggesting a low availability of food. Prey items in the diet consisted mainly of the benthic shrimp Nauticaris marionis and the demersal fish Notothenia squamifrons. On evidence from stomach contents and the distances the penguins travelled, we suggest that Nauticaris marionis has a more restricted distribution around Marion Island than does Notothenia squamifrons. The concentration of gentoo penguin breeding colonies along the east coast of Marion Island and the south-east coast of Prince Edward Island may be attributable to favourable feeding conditions between the two islands.     

Seasonal variation in the diet of the king penguin (Aptenodytes patagonicus) at sub-Antarctic Marion Island

King penguins are important consumers of marine resources, throughout the year, at the Prince Edward Islands. Meal size varied from 8.5–12.6% of adult mass, depending on the method of determination. In spite of the biases in the analysis favouring the overestimation of squid, fish and, in particular, myctophid fish accounted for the largest proportion of the stomach samples, 87% by wet mass, 75% by numbers and 69% by reconstituted mass. The relative abundance of fish in the diet dropped markedly in winter followed by a subsequent rise to nearly 100% in summer. This rise coincided with an increase in the chick growth rate and the king penguin population at the island and suggests the rise in relative abundance offish reflects a real increase in the availability of fish around the islands.
Juvenile and adult Krefftichthys anderssoni/Protomyctophum tenisoni and adult Electrona carlsbergi were the most common fish consumed. There was an increase in the modal size of K. anderssoni/P. tenisoni throughout the year which we interpret as growth of a single fish population. Juvenile Kondakoviu longimana was the important squid species taken by king penguins. Crustaceans were only rarely recorded in the diet and may have come from digestion of fish and squid stomachs.
This is the first study of the diet of a Southern Ocean pelagic predator that has identified myctophid fish as a major component of its diet. All three important fish species taken by king penguins at Marion Island have a wide distribution throughout the Southern Ocean and consequently may prove to be important dietary components of other Southern Ocean pelagic predators.     

Short-term responses of king penguins Aptenodytes patagonicus to helicopter disturbance at South Georgia

The short-term behavioural effects of helicopter overflights on breeding king penguins Aptenodytes patagonicus at South Georgia were examined. Seventeen helicopter overflights were made at altitudes between 230 and 1,768 m (750–5,800 ft) above ground level. Noise from the aircraft engines and helicopter blades increased sound levels in the colony from a background level of 65–69 dB(A) to a maximum mean peak level of 80 dB(A) during overflights. Penguin behaviour changed significantly during all overflights at all altitudes compared to the pre- and post-flight periods. Pre-overflight behaviour resumed within 15 min of the aircraft passing overhead and no chicks or eggs were observed to be taken by predators during overflights. Non-incubating birds showed an increased response with reduced overflight altitude, but this was not observed in incubating birds. Variability in overflight noise levels did not affect significantly the behaviour of incubating or non-incubating birds. Penguins exhibited a reduced response to overflights as the study progressed (despite later flights generally being flown at lower altitudes) suggesting some degree of habituation to aircraft. To minimise disturbance to king penguins we recommend a precautionary approach such that overflights are undertaken at the maximum altitude that is operationally practical, or preferably are avoided altogether.     

At‐sea distribution and habitat use in king penguins at sub‐Antarctic Marion Island

King penguins make up the bulk of avian biomass on a number of sub‐Antarctic islands where they have a large functional effect on terrestrial and marine ecosystems. The same applies at Marion Island where a substantial proportion of the world population breeds. In spite of their obvious ecological importance, the at‐sea distribution and behavior of this population has until recently remained entirely unknown. In addressing this information deficiency, we deployed satellite‐linked tracking instruments on 15 adult king penguins over 2 years, April 2008 and 2013, to study their post‐guard foraging distribution and habitat preferences. Uniquely among adult king penguins, individuals by and large headed out against the prevailing Antarctic Circumpolar Current, foraging to the west and southwest of the island. On average, individuals ventured a maximum distance of 1,600 km from the colony, with three individuals foraging close to, or beyond, 3,500 km west of the colony. Birds were mostly foraging south of the Antarctic Polar Front and north of the southern boundary of the Antarctic Circumpolar Current. Habitat preferences were assessed using boosted regression tree models which indicated sea surface temperate, depth, and chorophyll a concentration to be the most important predictors of habitat selection. Interestingly, king penguins rapidly transited the eddy‐rich area to the west of Marion Island, associated with the Southwest Indian Ocean Ridge, which has been shown to be important for foraging in other marine top predators. In accordance with this, the king penguins generally avoided areas with high eddy kinetic energy. The results from this first study into the behavioral ecology and at‐sea distribution of king penguins at Marion Island contribute to our broader understanding of this species.     

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.     

New southerly breeding location of king penguins (Aptenodytes patagonicus) on Elephant Island (Maritime Antarctic)

In the 2009–2010 austral summer, two breeding pairs of king penguins were recorded at Stinker Point, Elephant Island, Maritime Antarctic. This is the first record of king penguins breeding south of 60°S. The finding suggests a possible range extension of this species and increases the number of breeding bird species at Stinker Point, which was recently appointed as an Important Bird Area in Antarctica.     

Metabolism and thermoregulation during fasting in king penguins, Aptenodytes patagonicus, in air and water

We measured oxygen consumption rate (Vo(2)) and body temperatures in 10 king penguins in air and water. Vo(2) was measured during rest and at submaximal and maximal exercise before (fed) and after (fasted) an average fasting duration of 14.4 +/- 2.3 days (mean +/- 1 SD, range 10-19 days) in air and water. Concurrently, we measured subcutaneous temperature and temperature of the upper (heart and liver), middle (stomach) and lower (intestine) abdomen. The mean body mass (M(b)) was 13.8 +/- 1.2 kg in fed and 11.0 +/- 0.6 kg in fasted birds. After fasting, resting Vo(2) was 93% higher in water than in air (air: 86.9 +/- 8.8 ml/min; water: 167.3 +/- 36.7 ml/min, P < 0.01), while there was no difference in resting Vo(2) between air and water in fed animals (air: 117.1 +/- 20.0 ml O(2)/min; water: 114.8 +/- 32.7 ml O(2)/min, P > 0.6). In air, Vo(2) decreased with M(b), while it increased with M(b) in water. Body temperature did not change with fasting in air, whereas in water, there were complex changes in the peripheral body temperatures. These latter changes may, therefore, be indicative of a loss in body insulation and of variations in peripheral perfusion. Four animals were given a single meal after fasting and the temperature changes were partly reversed 24 h after refeeding in all body regions except the subcutaneous, indicating a rapid reversal to a prefasting state where body heat loss is minimal. The data emphasize the importance in considering nutritional status when studying king penguins and that the fasting-related physiological changes diverge in air and water.     

The impact of avian predator-scavengers on King Penguin Aptenodytes patagonicus chicks at Marion Island

While ashore King Penguins Aptenodytes patagonicus are fed upon by a guild of five predator-scavenger seabirds. During the winter (April-October) male Southern Giant Petrels Macronectes giganteus killed an estimated 6430 (11.2%) of Marion Island's King Penguin chicks, although most birds only scavenged in the colonies. The rate of predation varied, with peaks in April-May and in September. The proportion of successful attacks was 22.7%. There was a strong correlation between colony size and the rate of accumulation of chick corpses. Kelp Gulls Larus dominicanus and Lesser Sheathbills Chionis minor also scavenged penguin corpses but Northern Giant Petrels M. halli and female Southern Giant Petrels rarely entered the colonies. During the summer predation was mainly by Sub-Antarctic Skuas Catharacta lonnbergi which took eggs and small chicks.     

Summer diet of king penguins (Aptenodytes patagonicus) at the Falkland Islands, southern Atlantic Ocean

The diet of king penguins (Aptenodytes patagonicus) brooding chicks was investigated during February 2001 at the Falkland Islands, where a small but increasing population is located at the limit of the breeding range of this species. Fish was the most important food source by number (98.0%) and reconstituted mass (97.8%), squids accounting for the remainder. Myctophid fishes represented the main part of the diet (97.7% by number and 96.6% by reconstituted mass), Protomyctophum choriodon being by far the main prey item (84.2% and 88.1%, respectively). Four other myctophids and one squid species each contributed to more than 1% of the diet by number: Krefftichthys anderssoni (4.8%), Electrona carlsbergi (4.6%), Gymnoscopelus nicholsi (2.2%) and Protomyctophum tenisoni (1.8%), together with small juveniles of Gonatus antarcticus (1.8%). Twelve squid species were identified from accumulated lower beaks, including the ommastrephid Martialia hyadesi (48.3% by number), the onychoteuthids Moroteuthis ingens (15.6%), Kondakovia longimana (10.5%) and Moroteuthis knipovitchi (7.3%), and Gonatus antarcticus (9.2%). The stable-carbon and stable-nitrogen isotopic composition of chick food and adult blood differed in a way that suggests that, during the same trip, adult birds fed for themselves in distant foraging grounds, and fed for their chicks on their way back to the colony. The study emphasizes that king penguins are specialist myctophid eaters throughout their breeding range in summer, and highlights the importance of Protomyctophum choriodon as a link between zooplankton and top predators in the pelagic ecosystem of the southwestern Atlantic Ocean.     

Survival in king penguins Aptenodytes patagonicus: temporal and sex-specific effects of environmental variability

We investigated annual adult survival rates of king penguins Aptenodytes patagonicus breeding at South Georgia during 6 years in relation to age/breeding experience, sex, and food availability. During the first 3 years of the study, when food availability was good, survival was 97.7% for experienced breeders, which confirmed the very high survival rates observed in penguins in general. In these years survival did not differ between the sexes, presumably because parental investment is shared equally between the sexes, and the sexual dimorphism is small in king penguins. Survival was lower for young, first-time breeders (83.0%). In experienced birds the annual survival rate decreased to 68-82% following a catastrophic year when food availability was extremely low. We address the question how parents balance their current investment in offspring against their chances to reproduce in the future. We argue that the high mortality rate among breeding individuals after the year of food stress provides support for previous suggestions that the response to increased costs in seabirds might be complex to predict and does not always follow intuitive expectations according to general life-history theory. We also found that females survived significantly less well than males following the bad year. We explain this result as follows: the male-biased sex ratio (56:44) that we observed in our study colony clearly does not result from lower female survival during normal conditions. An already existing skewed sex ratio forces males to delay the onset of breeding because of a lack of breeding partners. This in turn causes breeding females to be, on average, younger and less experienced than males and to have lower survival following a year of food shortage. In this study survival was linked with food availability and we suggest that this was connected to climatic/oceanographic features, such as the position of the Antarctic Polar Front Zone. We could, however, not verify this by anomalies in sea surface temperature data.     

Comparison of foraging strategies of incubating king penguins Aptenodytes patagonicus from Macquarie and Heard islands

The foraging strategies of king penguins from Heard and Macquarie islands were compared using satellite telemetry, time-depth recorders and diet samples. Trip durations were 16.8±3.6 days and 14.8±4.1 days at Macquarie and Heard islands, respectively. At Macquarie Island, total distances travelled were 1281±203 km compared to 1425±516 km at Heard Island. The total time the penguins spent at sea was 393±66 h at Macquarie Island and 369±108 h at Heard Island. The penguins from Macquarie Island performed more deep dives than those from Heard Island. King penguins from Macquarie Island travelled 1.5±0.2 km h⁻¹ day⁻¹ compared to 1.3±0.1 km h⁻¹ day⁻¹. At Macquarie Island, 19% of dives were upto 70–90 m depth compared to 35% at Heard Island. The main dietary prey species were the fish Krefftychthis anderssoni and the squid Moroteuthis ingens in both groups. The differences in the at-sea distribution and the foraging behaviour of the two groups of penguins were possibly related to differences in oceanography and bathymetric conditions around the two islands. Dietary differences may be due to interannual variability in prey availability since the two colonies were studied during incubation but in different years.     

Effect of fasting on the VO2-fh relationship in king penguins, Aptenodytes patagonicus

King penguins (Aptenodytes patagonicus) may fast for up to 30 days during their breeding period. As such extended fasting may affect the relationship between the rate of O(2) consumption (Vo(2)) and heart rate (f(H)), five male king penguins were exercised at various speeds on repeated occasions during a fasting period of 24-31 days. In addition, Vo(2) and f(H) were measured in the same animals during rest in cold air and water (4 degrees C). Vo(2) and f(H) at rest and Vo(2) during exercise decreased with fasting. There was a significant relation between Vo(2) and f(H) (r(2) = 0.56) that was improved by including speed, body mass (M(b)), number of days fasting (t), and a cross term between f(H) and t (r(2) = 0.92). It was concluded that there was a significant change in the Vo(2)-f(H) relationship with fasting during exercise. As t is measurable in the field and was shown to be significant and, therefore, a practical covariate, a regression equation for use when birds are ashore was obtained by removing speed and M(b). When this equation was used, predicted Vo(2) was in good agreement with the observed data, with an overall error of 3.0%. There was no change in the Vo(2)-f(H) relationship in penguins at rest in water.     

Effects of weather on activity and sleep in brooding king penguins (Aptenodytes patagonicus)

Avian sleep is sensitive to thermal challenges. Brooding king penguins (Aptenodytes patagonicus) defending a territory are exposed to wide daily fluctuations in ambient temperature and wind conditions. We studied the daytime behavioural time budget of 89 groups of territorial adults brooding a 1- to 5-week-old chick on Crozet Island during summer 1998. Scans were performed every 10 min and each bird was categorized as either active, resting or sleeping. Three ranges of ambient temperatures (T1=5-9°C, T2=10-13°C, T3=14-19°C) and wind conditions (calm-light, moderate, strong) were distinguished. Wind conditions did not affect the behavioural time budget of king penguins during summer. Resting, which represented about half of the daytime behavioural time budget, increased by 17% when ambient temperature increased from T1 to T3, mostly at the expense of active behaviours. The percentage of time spent sleeping was low, but was reduced by 66% when ambient temperature increased over 10°C. Thus, behavioural sleep was mainly observed in a range of temperatures within which resting metabolic rate of adult king penguins is minimal, i.e. between -1 and 11°C. It is also interesting to note that the range of ambient temperatures in which sleeping was high coincides with the most common microclimatology prevailing at the colony during summer, i.e. ambient temperatures between 5 and 10°C.     

Core temperature variability in diving king penguins (Aptenodytes patagonicus): a preliminary analysis

Core temperature was determined in two king penguins living in the wild at Ile de la Possession, Crozel Archipelago, using implantable four-channel temperature loggers. Core temperatures derived from bird no. 1 (sensor placed under the sternum, in the vicinity of the liver and upper stomach) were closely correlated with diving activity (as determined by an external light recorder), and ranged from 38.3°C, (on land) to a minimum of 37.2°C during a dive. Core temperatures measured in bird no. 2 showed that temperatures near the heart were generally 1°C lower than those under the sternum or in the lower abdomen. Core temperatures declined continuously during dives (by 0.8, 1.2 and 2.7°C in the lower abdomen, under the sternum and near the heart, respectively) and showed precipitous drops to 35°C, probably associated with ingestion of food. Temperatures measured near the heart fluctuated over a period of 288 s, corresponding to the duration (from the literature) of the surface/dive cycle. The relevance of these findings with respect to diving physiology, blood perfusion of tissues, tissue metabolism and aerobic dive limits is discussed.     

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     

Seasonal change in foraging areas and dive depths of breeding king penguins at Heard Island

The seasonal variation in the foraging behaviour of king penguins (Aptenodytes patagonicus) was studied at Heard Island (53°05′S, 73°30′E) during 1992/1993. On seven occasions throughout the breeding cycle, time-depth-light recorders were deployed on breeding adults to record the dive activities and foraging. Foraging locations changed with season: in autumn and spring 1992, adults foraged between 48–52°S and 74–78°E, about 370 km NNE of Heard Island close to the Polar Front. Two penguins tracked in winter travelled 2220 km east of Heard Island (95°E) along the northern ice limit, and 1220 km south of Heard Island to approximately 65°S, respectively. In spring (October), the penguins again foraged further north than during winter. The foraging area utilised in October overlapped the area where the penguins foraged in March/April. The penguins' diving behaviour also varied seasonally: the modal depth of deep dives (>50 m) increased from about 100 m in February to 220 m in October. Mean dive depths increased from 70 ± 52 m in March 1992 to 160 ± 68 m in August 1992. Penguins dived deep (>50 m) only during daylight hours (16 h in February, 9 h in July). Mean dive durations ranged from 2.9 ± 1.1 min in March 1992 to 5.1 ± 1.2 min in August 1992. Associated with changes in foraging location and dive behaviour was a change in diet composition: during summer the penguins ingested mainly myctophid fish (>90%) while in winter the most important diet item was squid. Accepted: 19 October 1998     

Diving patterns of female macaroni penguins breeding on Marion Island, South Africa

Despite the large biomass of macaroni penguins Eudyptes chrysolophus in the Southern Ocean, their feeding ecology is poorly known at some important breeding localities. We investigated the diving behaviour and diet of female macaroni penguins feeding small chicks on Marion Island (46o52′S, 37o5′E), South Africa, one of the species’ most northerly breeding sites, supporting 4% of their global population. We then compared our results with similar studies from other localities. In December 2008, we collected information on 12 foraging trips from 6 individuals using time-depth recorders, as well as diet from 42 individuals. Median trip duration was 22.8 h (5.6–80.8 h). Penguins performed 42.8 ± 15.9 dives per hour at sea, with dive depths averaging 24.6 ± 8.6 m and lasting 40.8 ± 12.1 s, although 74.3% of dives were <10 m. Euphasids dominated their diet (86% by mass), mainly Thysanoessa vicina. A second peak in dive depths at 55–80 m might reflect the 12% of fish in their diet. The substantial proportion of shallow night dives (30% of total dives) suggests some foraging occurs at night. Differences in diving patterns of individual macaroni penguins in this study confirmed the behavioural flexibility of these birds reported from other breeding localities. However, most other studies assumed that dives <3–5 m were commuting dives whereas our study suggests that at least some prey are caught during shallow dives. We highlight how different analytical methods can change the outcome of studies. Despite macaroni penguins’ apparent flexibility in foraging behaviour during the breeding season, their numbers are decreasing globally. Further investigations of their foraging behaviour are needed to assess potential competition with other predators and krill fisheries.     

Accounting for body condition improves allometric estimates of resting metabolic rates in fasting king penguins, Aptenodytes patagonicus

We describe a method that allows prediction of resting metabolic rate (RMR, ml O₂ · min⁻¹) in adult male and female king penguins on shore by measuring body mass (M _b) and the length of the foot, flipper and beak. This method is accurate, underestimating measured RMR (n=114) by 4% in a data set consisting of 44 birds (33 males and 11 females). Measurement error was unbiased with respect to fasting duration and can therefore estimate RMR during any stage of fasting. This new method provides significant cost and logistical savings when estimating RMR during fieldwork, allowing RMR of a large number of birds to be measured quickly. These findings suggest the possibility that the use of M _b and morphometrics will allow development of general and specific equations to estimate RMR in other species.     

Gastrointestinal helminths of King penguins (Aptenodytes patagonicus) at Crozet Archipelago

Communities of helminths are known to be related to feeding behaviors of hosts. While climate change and overfishing can impact food availability for Antarctic piscivorous predators, knowledge about infectious and parasitic diseases among Antarctic species is scarce or fragmentary. We studied the helminth community of King penguins (Aptenodytes patagonicus) from the Crozet Archipelago, the main breeding area of the species. Based on a sample of 41 individuals found freshly dead from predation or starvation, the gastrointestinal helminth community in King penguins was composed of 1 species of cestode (Tetrabothrius wrighti) and 2 species of nematodes (Tetrameres wetzeli and Contracaecum heardi). Cestodes formed the core of the helminth community (97.5% of worms collected) with a prevalence of infestation of 100% and a mean intensity of 178.6 worms per host. Sources of infestation and pathologies caused by these worms are also discussed.