The diet of the King Penguin Aptenodytes patagonicus at Macquarie Island

The diet of King Penguins Aptenodytes patagonicus at Macquarie Island was studied between November 1984 and November 1985 based on stomach flushed samples (obtaining 93% of the total stomach content) from ten birds each month. The mean stomach content mass of the 118 samples was 923 0 g. Percentage by number, percentage by weight and dietary coefficient analysis all showed the main prey of the penguins to be myctophid lantern fish of the species Electrona carlsbergi and Krefftichthys anderssoni. Juvenile fish of both species were eaten from December to July, and adults in August and September. Cephalopods were relatively unimportant in contrast to previous indications. The amount of food brought ashore and the composition of the diet varied over the year, with K. anderssoni the dominant food in all but the winter months when E. carlsbergi replaced it as the principal food item.     

Diet of the King Penguin Aptenodytes patagonicus during three summers at South Georgia

King Penguins Aptenodytes patagonicus which were rearing chicks were studied during three summers from November 1991 to March 1994 at South Georgia. Stomach samples ( n =115) collected by flushing had a mean mass of 1308 g. Fish mass was allocated to each species based on the relationship between fish mass and otolith length. Three mesopelagic lanternfishes (Myctophidae), Krefftichthys anderssoni, Electrona carlsbergi and Protomyctophum choriodon , dominated the diet both by numbers and mass. They were small fish with mean mass of 3–7 g. Overall, K. anderssoni dominated the diet in terms of numbers and mass. Although Barracudina Notolepis coatsi occurred in <3% of the diet by numbers, it was large (106 g) and was second most important in terms of mass. Squid represented <3% of the diet by mass. Although the chick-rearing success was poor in the 1993–1994 summer, meal size was not reduced but foraging trips were longer. In the 1993–1994 summer, a larger proportion of the otoliths were not identifiable because they were more completely digested. Fewer otoliths were identified as being those of K. anderssoni , but we argue that about 90% of the unidentified otoliths were K. anderssoni. There was also more squid and N. coatsi in the diet during the poor summer. A consistent trend was that P. choriodon was rare or absent in early summer but more important later in the year, and at the end of 1992–1993, it was the dominant prey. We conclude that myctophid fish, especially K. anderssoni , are the main summer prey of King Penguins rearing chicks at South Georgia, as found in other recent studies in the Southern Ocean.     

Individual breeding decisions and long-term reproductive strategy in the King Penguin Aptenodytes patagonicus

Although studied for 35 years, knowledge of the reproductive biology of the King Penguin Aptenodytes patagonicus remains incomplete. The chick requires more than 12 months of care, which extends the breeding cycle, including moult, to more than one year, i.e. the King Penguin is neither annual nor biennial. In an attempt to resolve ambiguities in the literature and to elucidate the long-term breeding strategy of the species, we studied breeciing frequency at the individual level, considering the decision to breed in relation to breeding history over the previous few years. Although adult birds attempted to breed annually (0.83 breeding attempts per year), successful rearing occurred, at best, every two years only (maximum of 0.41 fledged chick per pair). Comparing successive years, the number of breeding birds in the colony was stable but the number of fledged chicks varied from 29 to 278 over eight years. These results suggest that King Penguins adopt (as individuals) an opportunistic reproductive strategy, in that they usually lay an egg every year, even when failure is certain. Nevertheless, the decision to breed was not entirely blind, and we identified groups of birds that invested differentially in breeding attempts. The decision to breed was related to the previous breeding frequency, i.e. 81% of the birds that had bred continuously in the past started a new breeding attempt, but only 67% of birds that had missed a year did so. In intermittent breeders, birds that had bred frequently, more often started a new breeding attempt than birds that had largely missed years (71% versus 57%, respectively). Classes of breeders could correspond to age classes, to birds of different breeding quality or to alternative breeding strategies coexisting in the species. Testing the hypothesis that reproductive effort increases with age should be possible in future.     

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.     

Prey species and nutritive value of food fed during summer to King Penguin Aptenodytes patagonica chicks at Possession Island, Crozet Archipelago

The diet of King Penguins Aptenodytes patagonica at Possession Island was studied in February 1989 by analysis of 20 stomach contents collected from adult birds, just before they fed their single chicks. The mean mass of the samples was 1.84 kg, equivalent to 15% of the unladen adult body-mass. Fish accounted for 99.8% and squids for 0.2% by mass. The main prey were mesopelagic myctophid fish which live in dense shoals and perform a daily vertical migration. Subadults/adults of Electrona carlsbergi and juveniles and subadults/adults of Krefftichthys anderssoni represented 73.7 and 13.4% of the diet by mass, respectively. No difference was found in the diet of male and female King Penguins. Comparison of individual samples suggests that these birds catch a large quantity of fish from only a limited number of shoals. The mean caloric content of the food was 7 kJ/g wet mass. The total energy requirement of each chick during its initial period of growth was estimated to amount to 328,000 kJ, equivalent to 55 kg of food. The rate at which energy was delivered to the chick was calculated to be 50 W during this period.     

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.     

Eye structure and foraging in King Penguins Aptenodytes patagonicus

Anterior eye structure and retinal visual fields were determined in King Penguins Aptenodytes patagonicus using keratometry and an ophthalmoscopic reflex technique. The cornea is relatively flat (radius 32.9 mm) and hence of low refractive power (10.2 dioptres in air) and this may be correlated with the amphibious nature of penguin vision. The large size of the eye and of the fully dilated pupil may be correlated with activity at low light levels. In air, the binocular field is long (vertical extent 180̀) and narrow (maximum width 29̀), with the bill placed approximately centrally—a topography found in a range of bird species which employ visual guidance of bill position when foraging. Upon immersion in water, the optical power of the cornea is abolished, with the effect that the monocular fields decrease and binocularity is lost. King Penguins have a pupil type which has not hitherto been recorded in birds. In daylight it contracts to a square-shaped pinhole but dilates to a large circular aperture in darkness. This change alters retinal illumination by 300-fold (2.5 log₁₀ units). When diving, this permits the retina to be pre-adapted to the low ambient light levels that the birds encounter upon reaching mesopelagic depths. These penguins also forage at depths where ambient light levels, even during the day, can fall below the equivalent of terrestrial starlight. Under these conditions, the birds must rely upon the detection of light from the photophores of their prey. In this they are aided by their absolutely large pupil size and broad cyclopean visual field.     

Finding your mate in a seabird colony: contrasting strategies of the Guillemot Uria aalge and King Penguin Aptenodytes patagonicus

Capsule King Penguins recognize their mates by voice, but Guillemots do not need acoustic cues even though their calls show individual variation.

Aims To determine whether the structure of Guillemot calls could allow individual recognition, as with King Penguin, and whether acoustic cues are used to locate mates among a dense mass of conspecifics at a colony.

Methods Observations were made on breeding Guillemots and King Penguins. Calls made by birds returning to their mates were recorded, the signals digitized and the calls analysed. Calls were later played back to the mates of the birds concerned and the effects noted on both them and their neighbours.

Results Both Guillemots and King Penguins emitted calls on return to the breeding site which contained individual signatures and were therefore potentially usable for mate recognition. In King Penguins, auditory recognition was essential for finding a mate, whereas in Guillemots most of the arriving birds located their mate in a dense crowd of conspecifics without the help of acoustic signals. Guillemots could differentiate neighbours from strangers without auditory cues.

Conclusion Calls are essential for the successful identification of mates by King Penguins but not by Guillemots.     

Foraging range of king penguins Aptenodytes patagonicm during summer at Marion Island

Foraging ranges of king penguins Aptenodytes patagonicus were estimated by combining information on the feeding rates to chicks and brood shift lengths of adults (assessed by daily weighings of large chicks and daily checks of marked birds brooding small chicks) with measurements of travelling speeds and activity budgets at sea (assessed using remote recording devices). Adults brooding small chicks were relieved on average every 13 days and large chicks were fed every four days. Adults with large chicks spent 36% of their time, between attachment of the device and recapture, travelling at an average speed of 8.7 km.h^-1. This gives an estimated mean maximum foraging range of about 300 km. Adults attending small chicks spent 19% of their time away swimming, giving an estimated mean maximum foraging range of 225 km. Extreme foraging ranges for all birds were 75 and 902 km for penguins returning between two and 24 days at sea, respectively. Total distance travelled was highly correlated with time away from the colony.     

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.     

Diet composition of Emperor Penguin chicks Aptenodytes forsten' at two Mawson Coast colonies, Antarctica

The diet composition of Emperor Penguin Aptenodytes forsteri chicks was examined at Auster and Taylor Glacier colonies, near Australia's Mawson station, Antarctica, between hatching in mid-winter and fledging in mid-summer by "water-offloading" adults. Chicks at both colonies were fed a similar suite of prey species. Crustaceans occurred in 82% of stomach samples at Auster and 87% of stomachs at Taylor Glacier and were heavily digested: their contribution to food mass could not be quantified. Fish, primarily bentho-pelagic species, accounted for 52% by number and 55% by mass of chick diet at Auster, and squid formed the remainder. At Taylor Glacier the corresponding values were 27% by number and 31% by mass of fish and 73% by number and 69% by mass of squid. Of the 33 species or taxa identified, the fish Trematomus eulepidotus and the squid Psychroteuthis glacialis and Allu-roteuthis antarcticus accounted for 64% and 74% of the diets by mass at Auster and Taylor Glacier, respectively. The sizes of fish varied temporally but not in a linear manner from winter to summer. Adult penguins captured fish ranging in length from 60 mm ( Pfeura-gramma antarcticum ) to 250 mm ( T. eulepidotus ) and squid (P. glacialis ) from 19 to 280 mm in mantle length. The length-frequency distribution of P. glacialis showed seasonal variation, with the size of squid increasing from winter to summer. The energy density of chick diet mix increased significantly prior to "fledging".     

Sex identification in King Penguins Aptenodytes patagonicus through morphological and acoustic cues

In the context of sexual selection, animals have developed a variety of cues conveying information about the sex of an individual to conspecifics. In many colonial seabird species, where females and males are monomorphic and do not show obvious differences in external morphology, acoustic cues are an important signal for individual and sex recognition. Here, we study the vocal and morphological sex dimorphism in the King Penguin Aptenodytes patagonicus, a colonial, monomorphic seabird for which our knowledge about the role of vocalizations and morphology in mate choice is very limited. Data were collected at Possession Island, Crozet Archipelago, in a breeding colony consisting of about 16 000 breeding pairs. Using measurements of six morphological features and analysing acoustic parameters of call recordings of adult individuals, we show that King Penguins can be sexed based on a single morphological measurement of the beak with an accuracy of 79%. We found a sex‐specific syntax in adult King Penguin calls that provided a 100% accurate method to distinguish between the sexes in our study population. To confirm the method at the species level, we analysed calls recorded from King Penguin adults in Kerguelen Island, 1300 km away from our study population and found the same accuracy of the sex‐specific syntax. This sex‐specific syllable arrangement is rare in non‐passerines and is a first step in understanding the mate choice process in this species. Furthermore, it offers a cost‐effective, non‐invasive technique for researchers to sex King Penguins in the field.     

Responses of king penguin Aptenodytes patagonicus adults and chicks to two food‐related odours

Increasing evidence suggests that penguins are sensitive to dimethyl sulphide (DMS), a scented airborne compound that a variety of marine animals use to find productive areas of the ocean where prey is likely to be found. Here we present data showing that king penguins Aptenodytes patagonicus are also sensitive to DMS. We deployed DMS on a lake near a king penguin colony at Ratmanoff beach in the Kerguelen archipelago. We also presented DMS to ‘sleeping’ adults on the beach. On the lake, penguins responded to the DMS deployments by swimming more, while on the beach, penguins twitched their heads and woke up more for the DMS than for the control presentations. Interestingly, penguins did not respond to cod liver oil deployments on the lake; mirroring at‐sea studies of other penguins. Although at‐sea studies are needed to confirm that king penguins use DMS as a surface cue that informs them of productivity under the water, this study is an important first step in understanding how these birds locate prey over significant distances.     

Spheniscins, avian beta-defensins in preserved stomach contents of the king penguin, Aptenodytes patagonicus

During the last part of egg incubation in king penguins, the male can preserve undigested food in the stomach for several weeks. This ensures survival of the newly hatched chick, in cases where the return of the foraging female from the sea is delayed. In accordance with the characterization of stress-induced bacteria, we demonstrate the occurrence of strong antimicrobial activities in preserved stomach contents. We isolated and fully characterized two isoforms of a novel 38-residue antimicrobial peptide (AMP), spheniscin, belonging to the beta-defensin subfamily. Spheniscin concentration was found to strongly increase during the period of food storage. Using a synthetic version of one of two spheniscin isoforms, we established that this peptide has a broad activity spectrum, affecting the growth of both pathogenic bacteria and fungi. Altogether, our data suggest that spheniscins and other, not yet identified, antimicrobial substances may play a role in the long term preservation of stored food in the stomach of king penguins.     

Post-Fledging Dispersal of King Penguins (Aptenodytes patagonicus) from Two Breeding Sites in the South Atlantic

Most studies concerning the foraging ecology of marine vertebrates are limited to breeding adults, although other life history stages might comprise half the total population. For penguins, little is known about juvenile dispersal, a period when individuals may be susceptible to increased mortality given their naïve foraging behaviour. Therefore, we used satellite telemetry to study king penguin fledglings (n = 18) from two sites in the Southwest Atlantic in December 2007. The two sites differed with respect to climate and proximity to the Antarctic Polar Front (APF), a key oceanographic feature generally thought to be important for king penguin foraging success. Accordingly, birds from both sites foraged predominantly in the vicinity of the APF. Eight king penguins were tracked for periods greater than 120 days; seven of these (three from the Falkland Islands and four from South Georgia) migrated into the Pacific. Only one bird from the Falkland Islands moved into the Indian Ocean, visiting the northern limit of the winter pack-ice. Three others from the Falkland Islands migrated to the eastern coast of Tierra del Fuego before travelling south. Derived tracking parameters describing their migratory behaviour showed no significant differences between sites. Nevertheless, generalized linear habitat modelling revealed that juveniles from the Falkland Islands spent more time in comparatively shallow waters with low sea surface temperature, sea surface height and chlorophyll variability. Birds from South Georgia spent more time in deeper waters with low sea surface temperature and sea surface height, but high concentrations of chlorophyll. Our results indicate that inexperienced king penguins, irrespective of the location of their natal site in relation to the position of the APF, develop their foraging skills progressively over time, including specific adaptations to the environment around their prospective breeding site.     

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.     

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.     

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.