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.     

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.     

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.     

Fish prey of southern elephant seals, <Emphasis Type="Italic">Mirounga leonina</Emphasis>, at King George Island

Between November and February in the years 1993/1994 to 1999/2000, 153 southern elephant seals, Mirounga leonina, were stomach lavaged at King George Island. Fish occurred in 16 of 108 stomachs containing food items. Myctophids were the dominant fish prey, contributing 76.6% of the 145 sagittal otoliths found. The most important prey species was Gymnoscopelus nicholsi, which constituted 69% of the otoliths and occurred in 75% of stomach samples. Next in importance was the nototheniid Pleuragramma antarcticum, which occurred in 31.3% of samples and represented 11.7% in numbers. We suggest that while myctophids may be the dominant fish prey of elephant seals in areas close to King George Island, they are probably replaced by P. antarcticum as seals migrate towards higher latitudes where this fish is extremely abundant.     

The fish diet of black-browed albatrossDiomedea melanophris and grey-headed albatrossD. Chrysostoma at South Georgia

 The fish component of the diet of black-browed and grey-headed albatrosses at South Georgia was investigated by intercepting 155 meals from adults arriving to feed chicks during February 1986 and 1994. Fish represented 30% and 72% by mass of the diet of black-browed albatrosses and 14% and 60% by mass of the diet of grey-headed albatrosses in 1986 and 1994 respectively. We determined the identity and quantified the contribution (by numbers, size and mass) of fish species mainly by using otoliths (54 representing 9 taxa and 57 representing 17 taxa in black-browed and grey-headed albatross samples respectively). For black-browed albatrosses in 1986 the main fish prey was Patagonotothen guntheri (77% of otoliths, 51% of estimated fish biomass) and a single large specimen of Icichthys australis (40% estimated biomass), whereas in 1994 Pseudochaenichthys georgianus was the main fish prey (57% of estimated biomass) with Magnisudis prionosa (30%) and Champsocephalus gunnari (12%) also making substantial contributions. Grey-headed albatross samples from 1986 were dominated by southern lampreys (40% by number, 79% of estimated biomass), lanternfish (32% of numbers, 9% by mass) and Patagonotothen guntheri (11% by mass); in 1994 Champsocephalus gunnari (42% by numbers, 24% by mass), Magnisudis prionosa (13% by number, 36% by mass), Muraenolepis microps (90% by number), Pseudochaenichthys georgianus (15% by mass) and lanternfish (18% by number but only 1% by mass) were the main prey. The importance of Patagonotothen guntheri to both species in 1986 and its absence in 1994 probably reflect albatrosses obtaining it from the commercial fishery, which was active in 1986 but closed in 1994. Otherwise the fish diet of black-browed albatrosses is dominated by krill-feeding fish, characteristic of the waters of the South Georgia shelf. In contrast, the grey-headed albatross diet comprises deeper water mesopelagic species, especially lanternfish, which reflect its affinity for the Antarctic Polar Frontal Zone and associated oceanic upwellings. Received: 28 June 1995 / Accepted: 8 October 1995     

Food and feeding ecology of Wilson's storm petrel Oceanites oceanicus at South Georgia

At South Georgia, the diet of Wilson's storm petrel was studied using the regurgitates of adults arriving to feed chicks. Feeding frequency and meal size were estimated by weighing chicks twice daily, or in some cases every 3 h during daytime. Crustaceans contributed 98% of the total number of individual items and 68% of the total weight consumed; fish, 1% of the number of items and 28% of the weight. The most abundant crustacean was the amphipod Themisto gaudichaudii, which accounted for 90% of the total number but only 44% of the total weight of crustaceans eaten. Most (79%) of the Themisto were juveniles. Euphausiids were much less numerous in the diet (5% by number) but were the main group by weight (55%); most (52%) were Antarctic krill, with juveniles and sub-adults (25–50 mm long) predominating. Mysids, copepods and barnacle larvae were also present. Fish were all myctophids (lanternfish), Protomyctophum normani and P. bolini being identified; specimens were 50–85 mm long and weighed 1–4 g. Meal sizes averaged 6-5-7-5 g (14–22% adult body mass); about 75% of chicks were fed each day (mainly at night), about 10% probably receiving meals from both parents. A review of storm petrel diets emphasizes the importance of fish to Oceanodroma species and of crustaceans to Oceanites, Garrodia and Pelagodroma. Euphausiids and amphipods (chiefly Themisto and Hyperia) are the main crustacean prey and range from 5–50 mm and 0-005-0-7 g. Myctophids are the main fish prey and range from 20–100 mm and 1–5 g. Meal size ranges from 15–25% adult body mass and chicks are fed on 50–85% of days. This low delivery rate is mainly responsible for the disproportionately slow growth and long fledging period of storm petrel chicks.     

Fish prey of Antarctic fur seals Arctocephalus gazella at Ile de Croy,Kerguelen

 The composition of Antarctic fur seal prey was assessed through analysis of scats collected in March 1994 on Ile de Croy, Iles Nuageuses. Fish remains predominated in samples, occurring in 95% of droppings. A total of 968 otoliths allowed the identification of 16 fish species. Myctophid fish (12 species) dominated the diet both by number (94% of the otoliths) and by fish reconstituted mass (76%). Three fish species constituted together 87% of the reconstituted mass: the myctophids Gymnoscopelus nicholsi (52%) and G. piabilis (12%), and the channichthyid Champsocephalus gunnari (23%). Prey distribution suggests that during late summer seals forage in upper slope waters in the northeast of the Kerguelen Archipelago. Received:1 March 1996/Accepted: 20 May 1996     

Shared dietary niche between sexes in Magellanic Penguins

Prey use by male and female Magellanic Penguins (Spheniscus magellanicus) was quantified during chick rearing in northern Golfo San Jorge, Argentina and the dietary niches and diet composition of each sex were assessed and compared. A total of 67 males and 45 females were flushed during the chick stages of 2011 and 2012. Argentine Anchovy (Engraulis anchoita) was the main prey in terms of importance by mass in males and females in all stages and years (52.4–86.6% and 83.8–88.7%, respectively), except during the old chick stage in 2011 when the main prey in males and females was the Shortfin Squid (Illex argentinus) (63.1% and 44.4%, respectively). Analyses of similarity showed that diet composition in terms of contribution by mass of the main prey species was similar between sexes. Based on the isotopic values of δ¹³C and δ¹⁵N corresponding to 38 males and 21 females, the Bayesian mixing model outputs showed that Argentine Anchovy was also the main prey. In both years and chick stages, the isotopic niche of males and females was similar. The similarity in the dietary niche between male and female Magellanic Penguins raising chicks may be due to the high availability of Argentine Anchovy during the study period and the consequent dominance of this forage species in the diet of both sexes.     

Sexual differences in the diet of king cormorants at Macquarie Island

Sexual differences in the diet of king cormorants,Phalacrocorax albiventer, were investigated during the 1993/1994 austral summer at Macquarie Island. The major food items, identified by otoliths in regurgitations, were demersal fish; fish mass consumed could be estimated using a wet mass-otolith length relationship. Two fish species,Paranotothenia magellanica andHarpagifer georgianus, constituted 98% of the wet mass (male and female cormorants combined). Estimated individual fish mass ofParanotothenia magellanica (19.6±11.6 g) was greater than that ofH. georgianus (2.8±1.3 g). Total wet mass of food and number of fish in regurgitations did not differ statistically between the sexes of cormorants. However, males tended to feed on larger fish than did females.     

Comparative fuel metabolism in Gentoo and King Penguins: adaptation to brief versus prolonged fasting

To compare fuel utilization in large birds adapted to brief or prolonged fasting, protein and lipid utilization were quantified in the Gentoo Penguin (Pygoscelis papua) and the King Penguin (Aptenodytes patagonica). The inshore feeder Gentoo Penguin fasts for only a few days in its colony, while King Penguin chicks starve for several months in the subantarctic winter and male King Penguins starve for 5–6 weeks at the beginning of their breeding cycle. After an initial decrease in both daily body mass loss and nitrogen excretion during the first days of food deprivation, these two parameters thereafter stabilized at low values. At that time, protein utilization accounted for 15% of total energy expenditure in Gentoo Penguins and only 6% in King Penguin chicks during winter, the remainder (85% and 94%, respectively) being provided by fat oxidation. Similar percentages in fuel metabolism as seen in chicks during winter were reached in fasting adult King Penguins and spring chicks. However, a seasonal adaptation occurs in fasting chicks because energy expenditure is lower during winter. As previously described in starved mammals, the effectiveness in protein sparing could be related to the initial adiposity of the birds: the larger the fat stores (about 9% and 30% in Gentoo Penguins and winter chicks of King Penguins, respectively), the longer the fast duration and the better the level of protein conservation.     

Fish diet of the Antarctic fur seal,Arctocephalus gazella,in summer,at Stranger Point,King George Island,South Shetland Islands

 In February 1992, 34 faecal samples from non-breeding male Antarctic fur seals, Arctocephalus gazella, were collected at Stranger Point, King George Island, South Shetlands. Fish constituted an important part of the diet, occurring in 90% of those scats containing prey remains. From 1162 otoliths found in the faeces, 1086 were identified to at least family level. Myctophids and nototheniids represented together almost 90% of the fish eaten. The dominant species were Gymnoscopelus nicholsi, Pleuragramma antarcticum and Electrona antarctica, contributing 33.3%, 30.8% and 12.0% of the otoliths respectively. The standard length of these three species was estimated from otoliths with little or no signs of erosion. This study showed that fur seals fed mainly on pelagic fish species that are often associated with krill. These findings are corroborated by fur seal diving patterns. Received: 7 October 1994/Accepted: 5 October 1995     

Chick provisioning rates and growth in Blacklbrowed Albatross Diomedea melanophris and Grey-headed Albatross D. chrysostoma at Bird Island, South Georgia

We compared the parental division of labour and the pattern and rate of parental provisioning by two sympatric species of albatross of similar mass and breeding timetable but differing in diet and in the duration of chick‐rearing. Using electronic weighing platforms inside artificial nests, we recorded chick mass of Black‐browed Albatross and Grey‐headed Albatross at Bird Island, South Georgia every 10 minutes for both species in 1993 and 1994 and for each species in two other years between 1990 and 1996. The chick mass data (nearly one million weighings) were used to calculate meal mass (over 5000 meals) and intervals between meals. Adult birds were fitted with radio‐transmitters which allowed each meal to be allocated to the appropriate parent. The combination of meal mass and foraging trip duration were used to calculate provisioning rates for chicks and individual adults. Overall, Black‐browed Albatrosses delivered significantly lighter meals (569 g) than Grey‐headed Albatrosses (616 g) but more frequently (every 2.07 days and 2.50 days respectively). Thus combining foraging trip data for both parents, Black‐browed Albatross chicks received a meal every 1.22 days compared with 1.26 days for Greyheaded Albatross. These rates did not differ significantly. The contribution of each sex of each species in chick provisioning fluctuated between years, being similar in some years or biased towards males in others. Chicks of both species that failed to fledge received smaller, less frequent meals than successful chicks. In 1990 and 1994, Black‐browed Albatross chick provisioning rates were lower than in 1992 and 1993. In 1990, both meal mass and trip duration were affected, but only in 1994 was trip duration longer. Greyheaded Albatross chick provisioning rate was lower in 1994 than in other years but trip duration was longer. In each species, significant changes in meal mass and trip duration occurred within the chick‐rearing period. Chick provisioning rates invariably declined before chicks attained their peak mass. For both species, chick growth rates and peak and fledging mass, but not fledging age, were affected by differences in provisioning rate.     

Effect of diet on otolith composition in Pomatomus saltatrix, an estuarine piscivore

To test the hypothesis that elemental composition of otoliths (sagittae) could be influenced by differences in natural prey type, young‐of‐the‐year bluefish Pomatomus saltatrix were captured immediately after their migration from oceanic waters into mid‐Atlantic Bight estuaries and fed either shrimp, Crangon septemspinosa and Palaemonetes spp. or fish Menidia menidia under similar temperature and salinity regimes in two separate 60 day experiments. Unlimited rations of fish and shrimp prey were provided in the first experiment which led to differences in bluefish growth rate between the two prey treatments; fish prey was limited in the second experiment to ensure that growth rates of bluefish in the two prey treatments were similar. Concentrations of seven elements in bluefish otoliths were determined using solution‐based inductively coupled plasma mass spectrometry (ICPMS). There was no significant effect of diet on five of the seven elements examined (Na, Mg, K, Ca and Mn). The levels of Sr and Ba in the otoliths of shrimp‐fed bluefish, however, were significantly higher than fish‐fed bluefish in both experiments. Concentrations of Ba in shrimp‐fed bluefish otoliths were double that found in fish‐fed bluefish. The results suggest that diet can explain some of the variation in otolith chemistry previously attributed to physical and chemical properties of the water.     

Foraging ecology of Gentoo Penguins Pygoscelis papua at Macquarie Island during the period of chick care

The diet, diving behaviour, swimming velocity and foraging range of Gentoo Penguins Pygoscelis papua were studied at Macquarie Island during the breeding season in the 1993–1994 austral summer. Gentoo Penguins are considered to be inshore feeders, and at Macquarie Island the diet and estimated foraging ranges supported this. The diet consisted of 91.6% fish and 8.3% squid, by mass. The dominant prey taxa were the fish Gymnoscopelus sp. and Paranotothenia magellanica. A mixture of pelagic and benthic prey was consumed, with a greater proportion of benthic species occurring later in the season. The penguins exhibited a strong diurnal pattern in their diving behaviour. Deep diving (≥30 m) began near sunrise (03.00 h) and finished close to sunset (21.00 h). Diving at night was less common and very shallow (<10 m). Early in the breeding season, dive profiles indicated that birds were probably following vertically migrating pelagic prey through the water column and were foraging in waters over 100 m deep. Later in the season, more uniform, shallower depths were used, suggesting an increase in benthic foraging activity. These changes in dive pattern and depth were consistent with the habitat preferences of prey species found in the diet. Gentoo Penguins swam at 1.04 m per s and had a maximum potential foraging range of about 26 km for single-day trips. They tended to forage within 14 km of the colony, with a mean range of 5.4 km. This range encompassed the deep ocean habitat to the west and east of the island and a shallow area to the north.     

Impact of food abundance, diet and food quality on the breeding of the fruit pigeon, Parea Hemiphaga novaeseelandiae chathamensis, on Chatham Island, New Zealand

Breeding and diet of Parea Hemiphaga novaeseelandiae chathamensis were studied in relation to food abundance and quality on Chatham Island from 1991 to 1994. Although pairs were found breeding in all months, they nested predominantly during winter and spring (June-November). The timing of the nesting season, the proportion of pairs that bred and the number of chicks reared per pair varied between nesting seasons. During the 1992–1993 and 1994–1995 nesting seasons, all pairs nested, and many pairs reared two chicks, often involving clutch overlap (58% of 12 cases in 1992–1993, 37% of eight cases in 1994–1995). In 1993–1994, when fruit was scarce, nesting began 2 months later, only 44% of pairs nested and no pairs attempted to rear a second chick. Prior to and during the productive nesting seasons (1992–1993, 1994–1995), the diet of Parea consisted mainly of fruit, particularly that of Matipo Myrsine chathamica in autumn (March-May) and Hoho Pseudopanax chathamicus in winter and spring. Nutrient analyses of the main Parea foods showed that the pulp of these fruit were rich in lipids and available carbohydrates compared with those in foliage foods. Heavy fruiting of Matipo and Hoho promoted early nesting and prolonged nesting for 6 months, including nesting during winter.     

From mass of body elements to fish biomass: a direct method to quantify food intake of fish eating birds

The winter diet of great crested grebe (Podiceps cristatus) was studied in the major lakes of Insubria region, N Italy, in 2000–2003 by analysing stomach and oesophagus contents of birds found dead. Winter diet of cormorant (Phalacrocorax carbo) was studied using pellet analysis. Diet was expressed in terms of numerical abundance, frequency and biomass of prey. A detailed methodology is described of how to prepare a reference collection of otoliths, pharyngeal bones and chewing pads (key-elements), essential for species-specific identification of fish remains in stomachs and pellets, and to estimate ingested fish biomass. Correlations between mass (mg) of key-elements and mass of corresponding fish were significant, and exponential regression models were used to calculate biomass of each single prey item directly from the mass of its key-elements. Advantages of this method are compared with the more traditional two step approach in which size of elements is used to estimate fish length, followed by calculating fresh mass from fish length. Cyprinids formed the largest part of the diet for both predators. Great crested grebes selected small prey, feeding mainly on bleak (Alburnus alburnus alborella), while diet composition of cormorants varied significantly in relation to prey availability. Handling editor: K. Martens     

Fish prey of the Wandering Albatross Diomedea exulans at South Georgia

Summary The fish diet (45% of total diet by weight) of Wandering Albatrosses rearing chicks at South Georgia during the austral winters of 1983 and 1984 was investigated using otoliths retrieved from regurgitations. These provide the first quantitative data for this species and for any albatross. By number of identified otoliths (32% could be identified only as ?Macrouridae and ?Moridae), Pseudochaenichthys georgianus (35%), Muraenolepis microps (33%) and Chaenocephalus aceratus (20%) predominated, with Notothenia gibberifrons, Pagothenia hansoni and Champsocephalus gunnari (together 12%) also present. Composition by weight (estimated from otolith length) of the main species was Pseudochaenichthys 51%, Muraenolepis 14%, Chaenocephalus 27%; if digestion and wear had reduced otoliths by 10% the values would be Pseudochaenichthys 54%, Chaenocephalus 25%, Muraenolepis 13%. Composition by weight (actual or corrected values) was almost identical between years but epipelagic fish were significantly more abundant in 1983 than 1984. All identified fish eaten by Wandering Albatrosses are common on the South Georgia continental shelf and most of them are caught in the commercial fishery there. However, two of the three main target species of this fishery in 1983–1984, Notothenia rossii and Champsocephalus were not, or rarely, caught by Wandering Albatrosses. It seems unlikely, therefore, that the albatrosses depend greatly on the fishery for acquisition of fish prey but how they catch several species, including Muraenolepis, which are mainly benthic in habit is unknown.     

Modeling the marine resources consumed in raising a king penguin chick: an energetics approach

Accurate estimates of penguin energetics would represent an important contribution to our understanding of the trophodynamics of the Southern Ocean ecosystem and our ability to predict effects of environmental change on these species. We used the heart rate-rate of oxygen consumption technique to estimate rate of energy expenditure in adult king penguins raising a chick, in combination with data from the literature on changes in adult mass, chick energy requirements, and prey energy density. Our model estimated a variety of energetic costs and quantities of prey consumption related to raising a king penguin chick during the austral summer. The total energy requirements of a king penguin chick at the Crozet Archipelago from hatching until reaching a mass of 8 kg 90 d later is 271 MJ, representing the consumption of 38.4 kg of myctophid fish. A successfully breeding male requires 0.78 kg d(-1) of fish during the entirety of the incubation period and 1.14 kg d(-1) during the subsequent 90 d of chick rearing. Assuming the same energy requirements for females, the estimated 580,000 pairs of king penguins that breed successfully at Crozet each year, together with their chicks, consume a total of around 190,000 tons of fish during the incubation and summer rearing periods combined. If, due to depletion of fish stocks, the diet of breeders and chicks during the summer becomes identical to the typical diet of adults during the austral winter, the mass of prey required by both adults and chicks combined (where the chick still reaches 8 kg after 90 d) would increase by more than 25%.     

The use of otoliths as indicators of Little Penguin Eudyptula minor diet

The validity of using otoliths from stomach contents quantitatively to determine the number and size of fish consumed was tested on Little Penguins. They were fed different meal sizes of known number and size of fish and the stomach contents were recovered after various time intervals. There were no differences in estimates of original fish size when calculated from otolith length or weight. Rate of digestion of otoliths tended to decrease with increased meal size but increased with time after ingestion. Digestion of otoliths proceeds rapidly and, if ignored, estimates of numbers of fish consumed and of original fish size can be significantly underestimated. This problem can be partially solved by inspection of otolith condition and restricting calculations of fish size to otoliths unaffected by digestion. Many factors introduce variations into rate of otolith degradation and further species-specific studies are required before appropriate correction factors can be applied.     

The use of otoliths as indicators of Little Penguin Eudyptula minor diet

The validity of using otoliths from stomach contents quantitatively to determine the number and size of fish consumed was tested on Little Penguins. They were fed different meal sizes of known number and size of fish and the stomach contents were recovered after various time intervals. There were no differences in estimates of original fish size when calculated from otolith length or weight. Rate of digestion of otoliths tended to decrease with increased meal size but increased with time after ingestion. Digestion of otoliths proceeds rapidly and, if ignored, estimates of numbers of fish consumed and of original fish size can be significantly underestimated. This problem can be partially solved by inspection of otolith condition and restricting calculations of fish size to otoliths unaffected by digestion. Many factors introduce variations into rate of otolith degradation and further species-specific studies are required before appropriate correction factors can be applied.