Nest Defence of Nesting Chinstrap Penguins (Pygoscelis antarctica) against Intruders

We tested some predictions of parental investment theory by studying the aggressive behaviour of colonial nesting chinstrap penguins (Pygoscelis antarctica) against human intruders into their nesting territories. We tested for differences in the aggressive behaviour of penguins according to offspring age (eggs vs. chicks), offspring number, nest location in the colonies (central vs. peripheral) and sex. Offspring age was the main factor influencing nest defence, although nest location and sex were also important. Chicks were defended more strongly than eggs, in accordance with changes in the reproductive value of offspring, and this increase in aggressiveness was not related to revisitation of the same individuals. The level of aggression of penguins breeding in central sites was higher than that of peripheral birds, a difference that could be due to the lower residual reproductive value of central-nesting, probably older, birds. The stronger aggressiveness of males could be due to a combination of factors related to sexual selection and life-history traits. Offspring number did not affect the level of nest defence.     

THE LAYING INTERVAL AND INCUBATION PERIOD OF ROCKHOPPER AND MACARONI PENGUINS

Williams, A. J. 1981. The laying interval and incubation period of Rockhopper and Macaroni Penguins. Ostrich 52: 226–229.

The laying interval and incubation period of Rockhopper Penguins Eudyptes chrysocome and Macaroni Penguins E. chrysolophus were studied at Marion Island in 1974–75 and 1976–77. On average, the laying interval was 4,4 and 4,5 days, the incubation period of second-laid eggs was 34,2 and 35,9 days and that of first-laid eggs was 39,1 and 38,0 days in Rock-hopper and Macaroni Penguins respectively. The laying interval in this genus is longer than that in other penguins. The incubation period is similar to that of most other penguins but the second-laid egg normally hatches before the first-laid egg. The long laying interval and the hatching sequence of the eggs both have important affects upon the mortality of eggs in the genus Eudyptes.     

Effects of double bands on Magellanic Penguins

ABSTRACT Banding penguins is controversial because bands can alter the survival, reproduction, and behavior of marked individuals. The effects of bands are not consistent among band types and, although stainless steel is thought to be better than other materials, tests of the long‐term impact of bands on tag‐loss rates and the reproduction and survival of individuals are needed. We tested three types of external tags on Magellanic Penguins (Spheniscus magellanicus) to measure band effects and tag‐loss rates. In 1993, we double‐tagged 300 penguins with aluminum flipper bands, stainless‐steel flipper bands, or small (2 mm × 10 mm) metal tags attached to foot webbing. We searched for double‐tagged birds for 13 of 15 yrs (1994–2008). Aluminum bands deformed, caused feather wear, injured and killed some penguins, and were lost more often than stainless‐steel bands or web tags. During the first 2 yrs of our study, at least nine penguins lost one aluminum band (N= 71 penguins resighted), but no penguins lost a stainless‐steel band (N= 84) or a web tag (N= 88). During the next 13 yrs, five penguins lost one of their two web tags (N= 89), but none lost a stainless‐steel band (N= 84). Females laid eggs of similar size before they carried a band and in the year following tagging (P= 0.09). The type of tags a female carried did not significantly change egg size (P > 0.22). During the first breeding season after tagging, penguins with aluminum bands had lower reproductive success than penguins with stainless‐steel bands or web‐tags (P= 0.04). The annual survival of females with two stainless‐steel bands was lower (0.79) than that of males with two stainless‐steel bands or males and females with two web‐tags (0.87). Aluminum bands injured Magellanic Penguins, were lost at high rates, and should not be used. Double stainless‐steel bands had no apparent effects on adult male Magellanic Penguins, but reduced survival rates of adult females. A single stainless‐steel band would likely have less impact than two bands, and our results suggest that the impact of a single band would be difficult to measure.     

Geographical variation in egg size dimorphism in rockhopper penguins

All crested penguins present a unique reversed hatching asynchrony: the larger second-laid egg (B-egg) hatches before the smaller first-laid egg (A-egg). Although both eggs often hatch, the A-chick generally dies of starvation within days after hatching. However, within rockhopper penguins, the population at the Falkland Islands is unique in that some birds manage to raise both chicks. Although it has been suggested that the egg size dimorphism between A- and B-eggs may explain how long both eggs and chicks survive, this hypothesis has never been explicitly tested. We expect that both eggs are retained longer in the less dimorphic clutches than in the more dimorphic ones. In this paper, we have compiled egg measurements for three rockhopper penguin species (Eudyptes chrysocome, E. filholi and E. moseleyi) in order to compare the intra-clutch egg size dimorphism among these species. Furthermore, we have collected new data to compare egg size dimorphism between two populations of E. chrysocome (Falkland Islands versus Staten Island). A-egg volumes are more variable between species and populations than B-egg volumes. E. chrysocome and especially the population from the Falkland Islands produces the largest A-eggs and the least dimorphic eggs. Nevertheless, as differences in A-egg volumes between species and between the populations of Falkland Islands and Staten Island are stronger and more significant than differences in egg dimorphism, we suggest that A-egg volume, more than egg dimorphism, could be one of the factors influencing the prevalence of twins. A large A-egg and/or reduced egg dimorphism is probably necessary to enable rockhopper penguins to raise two chicks, but other reasons may also be involved which enable them to keep both eggs and chicks.     

Aggressiveness in king penguins in relation to reproductive status and territory location

King penguins, Aptenodytes patagonicus, vigorously defend small territories in very dense colonies. The egg-laying season lasts approximately 4 months, but only pairs that reproduce during the first half of the period succeed in fledging a chick. I examined various factors affecting aggressiveness of king penguins during the breeding season and focused on the differences between central and peripheral territories. Pairs on peripheral territories experienced twice as many encounters with avian predators as did central birds. The vast majority of peripheral birds were late breeders, indicating that reproductive success was very low among penguins defending a territory on the edge of the colony. Time invested in territory defence and rate of agonistic encounters between breeding neighbours increased from the incubation to the brooding period. Parents gave most threat displays to territorial neighbours when the chick was very young and just before crèche formation. Distance to colony edge was not related to aggressiveness in incubating birds, however, the rate of pecking and flipper blows increased from the edge to the centre during brooding. In addition, aggressiveness of breeding penguins towards travelling birds trespassing into their territory increased with distance to edge. Early breeders were not more aggressive than late breeders but the proportion of time spent in territory defence increased with the number of days a bird had spent incubating. As expected, I did not detect any sex difference in aggressive behaviour. Birds occupying territories on the beach were generally more aggressive during the incubation period than those located on the valley sides. Reproductive status (incubating versus brooding) and territory location were the main factors explaining the various levels of aggressiveness observed in breeding king penguins. Copyright 2000 The Association for the Study of Animal Behaviour.     

People in Antarctica — how much do Adélie Penguins Pygoscelis adeliae care?

Summary In the course of physiological field studies, we opportunistically examined the effects of humans and aircraft on breeding Adélie Penguins Pygoscelis adeliae. Proximity to both aircraft and humans caused substantial increases in penguin heart rate even when no external stress was manifest. A solitary human at 20 m distance from commuting penguins on a well-used pathway caused the birds to deviate by 70 m. Birds at nests exposed to a single human fled much more readily when the brood consisted of large chicks (critical distance 6.1 m) rather than small chicks (critical distance 1.3 m) or eggs (critical distance 0.3 m). Aircraft caused birds to panic at distances greater than 1,000 m and 3 days exposure to a helicopter inhibited birds that had been foraging from returning to their nests, caused bird numbers in the colonies to decrease by 15% and produced an active nest mortality of 8%. Based on this data, we make recommendations to minimize stress on Adélie Penguin colonies exposed to man.     

Ontogeny of hepatic microsomal 3-hydroxylation of quinine in Adélie Penguins

Ontogeny of hepatic cytochrome P450 (CYP) content and of quinine 3-hydroxylation, a biomarker of human CYP3A activity, was investigated in Adélie Penguins (Pygoscelis adeliae) by comparing liver microsomes from chicks aged 13-28 days (n=10) with those from adults. The total CYP content in chick microsomes was significantly lower than in adults and correlated with the age of the chicks (r=0.894, P<0.001). Kinetic parameters (mean+/-S.D.) for quinine 3-hydroxylation in chick microsomes were lower than the corresponding values in adult penguins (Km, 122+/-27 vs. 160+/-73 microM; Vmax, 119+/-35 vs. 160+/-72 pmol/min/mg protein) but the differences were not significant, and neither Km nor Vmax correlated with age of the chicks. The pattern of inhibition of quinine 3-hydroxylation by specific human CYP inhibitors suggests that the CYP enzyme mediating quinine 3-hydroxylation in penguin chicks is similar to human CYP3A, but is a different isoform from that in adult penguins. The results imply that Adélie Penguin chicks may have a different susceptibility to environmental pollutants from adult penguins.     

Revisiting the influence of aggressive interactions on the survival of the first‐laid egg in crested penguins (genus Eudyptes)

Crested penguins Eudyptes spp. have evolved a unique form of breeding in which the first of two eggs laid is much smaller than the second and has a higher likelihood of being lost during egg laying and incubation. In this study, we quantified aggressive behaviour in nesting Snares penguins and undertook an egg survival analysis to examine which factors influence egg loss. During 120 h of observation of 50 nests, we recorded a total of 300 aggressive events in which females were repeatedly pecked, bitten and beaten. Aggressive events lasted from less than a minute to up to 55 min (mean 4.6 ± 7.4 min). Single males were the aggressor in 75% of aggressive events and in 50.7% of aggressive events the aggressor was identified as a neighbouring, breeding male. A greater percentage of the small first eggs (34%) were lost than the large second eggs (4%). We found that egg mortality was influenced by 1) whether the other egg within a nest had hatched, 2) who was present at the nest (father, mother or both) and 3) the average duration of aggressive events on the nest. When one egg within a nest had hatched, the other egg had a vastly increased mortality risk irrespective of aggression. However, long, aggressive events directed towards females after their partners had gone foraging, also increased the probability of egg loss. We suggest that the prolonged nest attendance by breeding males well beyond egg laying is in response to the high frequency of aggressive behaviour during this time. Our data show that A‐egg losses occur due to intraspecific aggression in this species. Further research is needed to clarify whether aggressive behaviour in breeding crested penguins is modulated by elevated testosterone levels in the males and whether any reproductive benefits accrue to the aggressors.     

Flipper bands do not affect foraging‐trip duration of Magellanic Penguins

ABSTRACT Flipper bands are used to mark penguins because leg bands can injure their legs. However, concerns remain over the possible effects of flipper bands on penguins. We examined the effects of stainless‐steel flipper bands on the duration of foraging trips by Magellanic Penguins (Spheniscus magellanicus) at Punta Tombo, Argentina, using an automated detection system. We predicted that, if bands were costly and increased drag, flipper‐banded penguins would make longer foraging trips than those with small or no external markings. We tagged 121 penguins with radio‐frequency identification (RFID) tags and an additional external mark. We placed either a stainless‐steel band on the left flipper (N= 62) or a 2×10‐mm small‐animal ear tag in the outside web of the left foot (N= 59). We measured foraging‐trip durations (N= 376 trips) for 68 adult penguins with chicks from 15 December 2007 to 28 February 2008. Contrary to predictions, trip duration was similar for banded and web‐tagged penguins (P= 0.22) and for males and females (P= 0.52), with no interaction between tag type and sex (P= 0.52). No penguins marked in the 2007 breeding season and recaptured between 30 September and 30 November 2008 (N= 113) lost flipper bands or web tags, but three RFID tags failed between March and September 2008. Properly designed and applied flipper bands were a reliable marking method for Magellanic Penguins, had a lower failure rate than RFIDs, and did not affect foraging‐trip duration.     

Finding food in the open ocean: foraging strategies in Humboldt penguins

Penguins are excellent “model” organisms allowing us to study the behaviour of marine homeotherms at sea. Penguins regularly return to their breeding colonies, enabling biologists to equip them with remote sensing devices such as physiological or behavioural data-loggers, radio- or satellite transmitters. Foraging trips at sea can last from days to weeks and after return of the birds to their breeding sites, the devices can easily be removed for analysis of on-board stored data, yielding a wealth of information. Investigation of penguin behaviour at sea becomes particularly revealing when other sources of information can be matched to the data set, such as satellite data on wind, temperature, ice cover, and chlorophyll-a concentrations.

Penguins and other marine homeotherms are true inhabitants of the high seas. Depending on the season, the marine behaviour varies: during reproduction, penguins are central-place foragers, and must return regularly to their nest to feed their chicks. During the remainder of the year, there are no constraints and the birds travel large distances at sea.

Breeding Humboldt penguins react to climatic change by varying their daily foraging range and dive duration. Similar to other representatives of the family Spheniscidae, Humboldt penguins avoid food shortages by migrating into more productive marine areas. Navigational clues such as daylength, sea surface temperature, local wind direction and olfaction might provide important aids in finding patchily distributed prey in the open ocean. DMS, a chemical compound produced by decaying algae, seems to be a further clue that indirectly points the way to feeding areas.     

Foraging behaviour of Magellanic Penguins during the early chick‐rearing period at Isla de los Estados,Argentina

Studies of the at‐sea distribution and trophic ecology of penguins are essential to understand their role in the broader marine food web. Magellanic Penguins Spheniscus magellanicus have a wide distribution and their foraging behaviour varies across breeding sites and between sexes, among others. In this study, we characterized the at‐sea areas, the diving strategies and the relative trophic level of Magellanic Penguins breeding at Isla de los Estados, Argentina, during the early chick‐rearing period. In addition, we quantified the interannual, sexual and individual variability in those parameters during three breeding seasons (2011–2013) using devices recording position and dive depth, and obtained blood samples for stable isotope analysis. During the early chick‐rearing period, Magellanic Penguins showed small differences between the sexes in their foraging behaviour and large overlap in the at‐sea areas used, suggesting no intraspecific variation between sexes. Although there was interannual variability in the foraging behaviour and the trophic level of the penguins, most of the studied nests managed successfully to raise both chicks during the first stage of the breeding cycle (guard stage). The foraging ecology of Magellanic Penguins from this colony was comparable with results of past studies at other breeding colonies. This study contributes to the identification of important at‐sea areas for Magellanic Penguins at the southern edge of their distribution and also to the identification of possible threats in the study area such as interaction with fisheries.     

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.     

BREEDING OF THE ADÉLIE PENGUIN PYGOSCELIS ADELIAE AT CAPE BIRD

Observations were made during four seasons (1967–68, 1968–69, 1969–70, 1970–71) on the breeding of Adelie Penguins at Cape Bird, Ross Island, Antarctica. Breeding data from individuals were related to date of return, laying date, clutch-size, nest location, and change of mate. Some females consistently laid near the mean date of laying, while others were consistently early or late layers. Laying date may be under direct genetic control, or may reflect feeding ability. The mean clutch-size was smaller in peripheral compared to central nests, and smallest of all in isolated nests. Clutches laid late in the season were smaller than those laid near the peak date, and small, late clutches were laid in peripheral rather than central nests. These differences may reflect age and/or feeding ability. Penguins that are better able to find food will return earlier, obtain central sites, and have larger clutches than those with a lesser ability. The two main causes of egg and chick losses were predation and parental failure. Losses were highest in single-egg clutches, at isolated and peripheral nests, and among eggs laid late in the season. These results may be partly related to the age and experience of the penguins. However, regardless of age, peripheral nesting and late laying were always disadvantageous. The sex ratio of adults in the colonies was 117♂:100♀. This may be explained by the higher mortality of females. Some males could not find partners, but females that did not breed had probably been unable to obtain sufficient food for gonad development. The return of penguins, incidence of non-breeding, adult mortality, clutch-size, and breeding success at Cape Bird were all markedly different in 1968–69 compared to the other three seasons studied. This season was marked by the persistence of sea-ice along the northwestern shores of Ross Island. The low reproductive output in 1968–69 was thought to result from a shortage of food for egg-laying and incubation.     

Responses of Emperor Penguins (Aptenodytes forsteri) to encounters with ecotourists while commuting to and from their breeding colony

Emperor penguins (Aptenodytes forsteri) were studied at the Snow Hill breeding colony in November 2006 to determine the effect of people on penguins traveling between the colony and the sea to forage. We tested the null hypothesis that the presence and number of people had no effect on the trajectory of movement or the number and duration of pauses. The distances at which penguins noticed people (mean 35.6 m), changed direction (mean 22.8 m), and the number and duration of pauses increased significantly with increases in the number of tourists in their path, which explained more than 50% of the variance. Undisturbed penguins usually tobogganed on their ventral surface over the ice. When penguins noticed people, they usually stood up and often called. In 10 min observation periods, penguins traveling more than 200 m from people paused an average of <1 min vs. 3.8 min for those passing near people, increasing the energetic cost of commuting. After passing people, penguins rarely stopped. Penguins response to people varied by time of day; later in the day they responded less quickly, changed directions when closer to people, stopped for less time, and passed by people closer than they did earlier in the day. We suggest that the effect of ecotourists on traveling penguins can be partly mitigated by having people walk in small, tight-knit groups, by having people stop moving whenever traveling penguins are within about 25 m to allow the penguins to choose the direction of their passage, and by keeping the visitor pathway separate from the penguin paths insofar as possible.     

Individual repeatability in laying behaviour does not support the migratory carry‐over effect hypothesis of egg‐size dimorphism in Eudyptes penguins

Penguins of the genus Eudyptes are unique among birds in that their first‐laid A‐egg is 54–85% the mass of their second‐laid B‐egg. Although the degree of intra‐clutch egg‐size dimorphism varies greatly among the seven species of the genus, obligate brood reduction is typical of each, with most fledged chicks resulting from the larger B‐egg. Many authors have speculated upon why Eudyptes penguins have evolved and maintained a highly dimorphic 2‐egg clutch, and why it is the first‐laid egg that is so much smaller than the second, but only recently has a testable, proximate mechanism been proposed. In most species of Eudyptes penguins females appear to initiate egg‐formation at sea during return migration to breeding colonies. In macaroni penguins E. chrysolophus, females with a shorter pre‐laying interval ashore (and thus presumably greater overlap between migration and egg‐formation) lay more dimorphic eggs, suggesting a physiological conflict may constrain growth of the earlier‐initiated A‐egg. This migratory carry‐over effect hypothesis (MCEH) was tested in eastern rockhopper penguins E. chrysocome filholi on Campbell Island, New Zealand, by recording the arrival and lay dates, body sizes, and egg masses of transponder‐tagged females over two years. Females with longer pre‐laying intervals laid less dimorphic clutches, as predicted by the MCEH. However, repeated measures of individual females revealed that within‐individual variation in egg‐size dimorphism between years was unrelated to within‐individual variation in pre‐laying interval. Egg masses, and to a lesser extent egg‐size dimorphism, were highly repeatable traits related to body size and body mass. These results and a detailed consideration of the MCEH suggest that egg‐size dimorphism in Eudyptes penguins is unlikely to be caused by a migratory carry‐over effect.     

Type of intruder and reproductive phase influence male territorial defence in wild-caught Siamese fighting fish

This study investigated how parental care increases with offspring age by examining the level of male aggressiveness toward potential intruders during egg guarding in a natural population of Siamese fighting fish (Betta splendens Regan). The degree of aggressiveness was measured at two reproductive phases in response to three types of intruders: male, female and females that have laid eggs. The nest-holding males became more aggressive after their eggs hatched than after the eggs were laid. Nest-holding males displayed gill cover erection, biting, tail beating and attacking at the highest rate towards male intruders, intermediate towards female intruders and the least aggressive towards females, which have laid eggs.     

Evaluation of potential variables contributing to the development and duration of plantar lesions in a population of aquarium-maintained African penguins (Spheniscus demersus)

Bumblefoot (pododermatitis), often described as the most significant environmental disease of captive penguins, is commonly due to excessive pressure or trauma on the plantar surface of the avian foot, resulting in inflammation or necrosis and causing severe swelling, abrasions, or cracks in the skin. Although not formally evaluated in penguins, contributing factors for bumblefoot are thought to be similar to those initiating the condition in raptors and poultry. These factors include substrate, body weight, and lack of exercise. The primary purpose of this retrospective study was to evaluate variables potentially contributing to the development and duration of plantar lesions in aquarium-maintained African penguins (Spheniscus demersus), including sex, weight, age, season, exhibit activity, and territory substrate. Results indicate that males develop significantly more plantar lesions than females. Penguins weighing between 3.51 and 4.0 kg develop plantar lesions significantly more often than penguins weighing between 2.5 and 3.5 kg, and because male African penguins ordinarily weigh significantly more than females, weight is likely a contributing factor in the development of lesions in males compared with females. Significantly more plantar lesions were observed in penguins standing for greater than 50% of their time on exhibit than swimming. Penguins occupying smooth concrete territories developed more plantar lesions compared with penguins occupying grate territories. Recommendations for minimizing bumblefoot in African penguins include training penguins for monthly foot examinations for early detection of plantar lesions predisposing for the disease, encouraging swimming activity, and replacing smooth surfaces on exhibit with surfaces providing variable degrees of pressure and texture on the feet.     

Time‐lapse cameras reveal latitude and season influence breeding phenology durations in penguins

Variation in the phenology of avian taxa has long been studied to understand how a species reacts to environmental changes over both space and time. Penguins (Sphenicidae) serve as an important example of how biotic and abiotic factors influence certain stages of seabird phenology because of their large ranges and the extreme, dynamic conditions present in their Southern Ocean habitats. Here, we examined the phenology of gentoo (Pygoscelis papua) and chinstrap penguins (Pygoscelis antarctica) at 17 sites across the Scotia arc, including the first documented monitoring of phenology on the South Sandwich Islands, to determine which breeding phases are intrinsic, or rather vary across a species range and between years. We used a novel method to measure seabird breeding phenology and egg and chick survival: time‐lapse cameras. Contrary to the long‐standing theory that these phases are consistent between colonies, we found that latitude and season had a predominant influence on the length of the nest establishment, incubation, and guard durations. We observe a trend toward longer incubation times occurring farther south, where ambient temperatures are colder, which may indicate that exposure to cold slows embryo growth. Across species, in colonies located farther south, parents abandoned nests later when eggs were lost or chicks died and the latest record of eggs or chicks in the nest occurred earlier during the breeding period. The variation in both space and time observed in penguin phenology provides evidence that the duration of phases within the annual cycle of birds is not fundamental, or genetic, as previously understood. Additionally, the recorded phenology dates should inform field researchers on the best timing to count colonies at the peak of breeding, which is poorly understood.     

Acoustic recognition in macaroni penguins: an original signature system

Penguins use vocal signatures as cues to identify their kin in dense colonies. Experimental studies of four species in two genera have pointed out that vocal signatures depend on the breeding ecology of the birds. Penguins that have a meeting site for pair members and chicks (genus Pygoscelis), as in many seabirds, have a less complex vocal signature than penguins that have no nests (genus Aptenodytes). To investigate whether this pattern would extend to other nest-building penguins, we studied the vocal signature in a species of the genus Eudyptes, the macaroni penguin, E. chrysolophus. Temporal and spectral features of signatures were analysed to determine which were likely to encode individual identity. Using a methodology derived from the theory of information, we measured and compared the amount of information given by each of these parameters by means of a stereotypy index. We then tested their effective efficiency in playback experiments of modified calls. Like Aptenodytes species, the macaroni penguins used a double coding system that integrated information in both the temporal and spectral domains. The encoding was made through the tempo given by the successive syllables of the call and the harmonic content of the call. However, information was not complementary but was mostly repeated in both domains, and variables were unidimensional as in Pygoscelis signatures. These results point out an original and simple signature system in macaroni penguins. Although they support the hypothesis of simpler systems in nest-building species they also reveal more subtle differences within this category.     

Chick starvation in yellow‐eyed penguins: Evidence for poor diet quality and selective provisioning of chicks from conventional diet analysis and stable isotopes

The link between poor reproductive success and diet was investigated in yellow‐eyed penguins Megadyptes antipodes, by assessing diet at two localities separated by about 30 km: the north coast of Stewart Island where breeding success is low (0.38–0.67 chicks per pair in recent years), and Codfish Island where breeding success is higher (0.96–1.51 chicks per pair), and relating this to published data from South Island localities, where average breeding success was 1.1 chicks per pair. Diet composition, meal sizes and energetic content of meals and prey were determined from stomach contents, and stable isotope analyses of chick down, fledgling feathers and adult blood provided information on diet throughout the fledging period. The high proportion of stomachs that were empty or lacked diagnostic remains reduced sample size considerably, and variability between samples reduced the power to detect significant differences in meal size, proportions of empty stomachs and prey diversity of meals. Energetic content of Stewart Island meals was less than Codfish Island meals, and there was a non‐significant trend for smaller meal sizes and reduced prey diversity among Stewart Island samples. Both localities had lower prey diversity and smaller meals than South Island penguins. Blue cod Parapercis colias accounted for 99% of prey biomass in Stewart Island and 70% in Codfish Island stomach samples, where 27% of prey biomass was opalfish Hemerocoetes monopterygius. Isotopic mixing models carried out on larger sample sizes indicated that opalfish comprised a large proportion of the diet at both locations, with adults selectively provisioning chicks with opalfish while feeding mainly on blue cod themselves. We suggest the large blue cod consumed by Codfish Island and Stewart Island penguins, larger than those consumed by South Island penguins, is difficult to transfer to chicks by regurgitation. Oyster dredging around Stewart Island may have reduced the availability and abundance of alternative prey to Stewart Island penguins.