Energy requirements of Adélie penguin (Pygoscelis adeliae) chicks

Summary The energy requirements of Adélie penguin (Pygoscelis adeliae) chicks were analysed with respect to body mass (W, 0.145–3.35 kg, n=36) and various forms of activity (lying, standing, minor activity, locomotion, walking on a treadmill). Direct respirometry was used to measure O₂ consumption ( ) and CO₂ production. Heart rate (HR, bpm) was recorded from the ECG obtained by both externally attached electrodes and implantable HR-transmitters. The parameters measured were not affected by hand-rearing of the chicks or by implanting transmitters. HR measured in the laboratory and in the field were comparable. Oxygen uptake ranged from in lying chicks to at maximal activity, RQ=0.76. Metabolic rate in small wild chicks (0.14–0.38 kg) was not affected by time of day, nor was their feeding frequency in the colony (Dec 20–21). Regressions of HR on were highly significant (p< 0.0001) in transmitter implanted chicks (n=4), and two relationships are proposed for the pooled data, one for minor activities ( ), and one for walking ( ). Oxygen consumption, mass of the chick (2–3 kg), and duration of walking (T, s) were related as , whereas mass-specific O₂ consumption was related to walking speed (S, m·s^-1) as .Abbreviations bpm beats per minute - D distance walked (m) - ECG electrocardiogram - HR heart rate (bpm) - ns number of steps - RQ respiratory quotient - S walking speed (m·s^-1) - T time walked (s) - W body mass (kg)     

Adverse effects of instrumentation in incubating Adélie penguins (Pygoscelis adeliae)

The use of data-loggers has permitted to explore the biology of free-ranging animals. However, this method has also been reported to reduce reproductive success while the reasons of this deleterious effect remain poorly documented. In this study, we aimed to identify critical periods of the breeding cycle of Adélie penguins (Pygoscelis adeliae) when the reproductive success may decrease because of instrumentation. For this purpose, we monitored 40 pairs, where one parent was instrumented before egg laying and 30 pairs without devices (controls). These pairs were followed at least during the incubation period but the majority was monitored during the entire breeding season. Reproductive success was affected in pairs where males were instrumented. This was not due to extra chick mortality during chick rearing but to a significantly lower hatching success. Moreover, the use of artificial eggs recording incubation temperatures and egg rotation indicated that in instrumented incubating males, eggs spent as much time at optimal incubation temperatures as control eggs but were rotated at a higher frequency. In Adélie penguins, males initiate incubation and it has been established that the early stage of incubation is one of the most critical periods for embryonic development. The low hatching rate observed in instrumented males was associated with a higher egg rotation rate, perhaps as a stress response to the presence of the instrument. Even though the causal effects remain unclear, instrumentation severely affected hatching success. For these reasons, we recommend equipping birds after the early incubation.     

Satellite telemetry of the winter migration of Adélie penguins (Pygoscelis adeliae)

Adélie penguins (Pygoscelis adeliae), after breeding in Antarctica during the austral summer, undergo a winter migration before returning to the breeding grounds 8 months later. It is the major source of adult mortality, with about a quarter of them not returning. Here we describe the first attempt to track the winter migration of Adélie penguins using satellite telemetry. Transmitters were attached to two penguins on 16 February 1991 after their post-breeding moult at Cape Bird, Ross Island, Antarctica. Transmissions were received from one penguin (bird #1) for 4.4 months, during which time it travelled 2792.6 km from the rookery (nearly 1500 km straight-line distance). Transmissions were received from the other penguin (bird #2) for 2.5 months during which time it followed a path remarkably similar to that of bird #1. The penguins travelled northwards up the coast of Victoria Land, keeping within 100 km of the coast, rounding Cape Adare soon after 29 March and were midway between the Balleny Islands and the Antarctic coast on 3 May. Thereafter, the record from bird #1 shows that it travelled further westwards until, when opposite the Mastusevich Glacier Tongue of the Mastusevich Glacier, it turned due north and moved away from the coast. By 29 June, when transmissions ended, its progression had slowed and it was northwest of the Balleny Islands near a zone where pack ice covered 75% of the surface of the sea. Two novel points that arise from this study are: (1) that Adélie penguins from Cape Bird undergo winter migrations of not less than 5000 km, and (2) that they may be travelling to common overwinter feeding grounds.     

Human-induced behaviour in Adélie penguins Pygoscelis adeliae

Summary Adélie penguins Pygoscelis adeliae appear to be little perturbed by man. We examined the incidence of nest desertion and duration of foraging trip in Adélie penguins when manipulated and fitted with devices of differing sizes. Birds with ca. 1 cm clipped from their tail feathers stayed at sea 50% longer than unmarked controls. The length of foraging trip and incidence of nest desertion increased with increasing device volume. Penguins fitted with devices did not reduce foraging trip length to that of unpackaged birds for at least 19 days. The susceptibility of Adélie penguins to disturbance should be carefully considered when activity patterns are being studied.     

In situ heart rate and activity of incubating Adélie penguins (Pygoscelis adeliae)

Summary Heart rates and activity were monitored over 24 h in unrestrained, incubating Adélie penguins (Pygoscelis adeliae) exposed to natural conditions in the colony. Heart rate (HR in bpm) increased linearly with wind speed (w; range 0–19 m/s): HR = 85.8+1.35 w, but was unrelated (P>0.05) to temperature (-2.5°–6°C), humidity (37%–100%) cloud cover (0–8/8) and estimated solar radiation (0–12). Wind-induced heat loss was apparently compensated to a large degree by increased metabolic activity. Activity (A) measured as frequency of standing per hour, decreased linearly with temperature (t) and wind speed (w): A = 1.651–0.033w–0.090t. After correcting for meteorological influences, heart rate and bird activity showed no diurnal periodicity. When incubating, metabolism and activity of Adélie penguins appear to be mainly governed by climatic variations.     

Delayed laying and prolonged fasting in Adélie Penguins Pygoscelis adeliae

Observations of nesting Adélie Penguins (Pygoscelis adeliae) were made at Ardley Island during spring 1990 when snow cover was unusually thick at some subcolony sites. Adélie Penguins at these sites had to delay egg laying until the snow melted. Maximum length of fasting periods comprising pre-breeding and incubation was 50 days. Long fasting seemed to have no detrimental effect on breeding. Furthermore, there was no relationship between penguin arrival mass and duration of fast. Even birds with small mass had sufficient reserves to undergo long fasting periods.In spring 1990, when we started with a monitoring study for CEMP (CCAMLR 1990) at Ardley Island, there were still high quantities of snow at the subcolony sites. Adélie Penguins at Ardley Island inhabit both small rocky outcrops and flat, stony hillocks (storm bars). The latter had a distinctly thicker snow cover at this time so that the pebbles necessary for nest building were unattainable. Consequently, we observed the behaviour of the penguins in this situation, recorded the laying dates and lengths of fasting periods.     

Observations on the thermal conductance of Adélie (Pygoscelis adeliae) and Humboldt (Spheniscus humboldti) penguins

 Analyses of cooling rates in one Adélie and one Humboldt penguin yielded calculated thermal conductance values of 0.1040 and 0.1672 W(kg ^°C)^-1, respectively. We review the methods used to calculate penguin surface area, an important component in calculating conductance values, and suggest that, in comparative studies of thermal balance, the use of body mass is a better estimator of body size than surface area. Using previously published data on penguin species, we found a significant model to predict thermal conductance from body mass according to: log C= log 0.1083−0.474 log M, where C is minimal specific thermal conductance in W(kg ^°C)^-1 and M is body mass in kilograms. Received: 13 June 1995/Accepted: 11 March 1996     

Adélie penguin (Pygoscelis adeliae) survival rates and their relationship to environmental indices in the South Shetland Islands,Antarctica

Seabird life history is typified by low fecundity, high adult survival rates, and relatively long lives. Such traits act as buffers, enabling persistence of populations under variable environmental conditions. Numerous studies, however, have suggested strong sensitivity of seabirds to environmental variability. In the Antarctic Peninsula region, for example, Adélie penguin (Pygoscelis adeliae) populations have declined during the last three decades, attributed largely to rapid changes in environmental conditions and food availability. We use 30 years of mark-recapture data from known-age individuals in the South Shetland Islands and capture-mark-recapture models to estimate survival rates with respect to such environmental variation. We investigated specifically whether negative trends in survival rates were evident and whether indices of global, regional, and local environmental conditions considered important for Adélie penguin survival explained the variability in survival rates. Overall, negative trends in juvenile survival were evident, but adult survival rates exhibited high interannual variability. Indices of sea ice extent had the strongest correlations with survival rates, particularly Weddell Sea ice extent during spring among adults (r = 0.62) and during winter for juveniles (r = 0.46). An analysis of deviance, however, suggested that single environmental covariates explained <30 % of the observed variation in the full mark-recapture models. Despite positive effects of sea ice extent on survival rates of Adélie penguins, limited explanatory power of several environmental conditions previously identified as important for Adélie penguin survival underscores the difficulty of predicting future population responses in this region of rapid environmental change.     

Sex-Based Differences in Adélie Penguin (Pygoscelis adeliae) Chick Growth Rates and Diet

Sexually size-dimorphic species must show some difference between the sexes in growth rate and/or length of growing period. Such differences in growth parameters can cause the sexes to be impacted by environmental variability in different ways, and understanding these differences allows a better understanding of patterns in productivity between individuals and populations. We investigated differences in growth rate and diet between male and female Adélie Penguin (Pygoscelis adeliae) chicks during two breeding seasons at Cape Crozier, Ross Island, Antarctica. Adélie Penguins are a slightly dimorphic species, with adult males averaging larger than adult females in mass (~11%) as well as bill (~8%) and flipper length (~3%). We measured mass and length of flipper, bill, tibiotarsus, and foot at 5-day intervals for 45 male and 40 female individually-marked chicks. Chick sex was molecularly determined from feathers. We used linear mixed effects models to estimate daily growth rate as a function of chick sex, while controlling for hatching order, brood size, year, and potential variation in breeding quality between pairs of parents. Accounting for season and hatching order, male chicks gained mass an average of 15.6 g d^-1 faster than females. Similarly, growth in bill length was faster for males, and the calculated bill size difference at fledging was similar to that observed in adults. There was no evidence for sex-based differences in growth of other morphological features. Adélie diet at Ross Island is composed almost entirely of two species—one krill (Euphausia crystallorophias) and one fish (Pleuragramma antarctica), with fish having a higher caloric value. Using isotopic analyses of feather samples, we also determined that male chicks were fed a higher proportion of fish than female chicks. The related differences in provisioning and growth rates of male and female offspring provides a greater understanding of the ways in which ecological factors may impact the two sexes differently.     

Feeding strategies of free-ranging Adélie penguins Pygoscelis adeliae analysed by multiple data recording

The fine-scale feeding behaviour of free-ranging Adélie penguins (Pygoscelis adeliae) during a single foraging trip was investigated by monitoring three parameters simultaneously at a frequency of 1 Hz, these being depth, swim speed and oesophagus temperature. Ingestion events were detected as abrupt drops in the oesophageal temperature and related to the birds' foraging behaviour. Although a high percentage of oesophageal temperature loggers were rejected, 1 complete foraging trip was recorded for all the 3 parameters from 1 bird while 92% and 67% of the foraging trip was recorded for 2 other birds; 12.3% of the temperature drops occurred at the surface but they were mainly small, except 61 of them probably representing snow ingestion while the birds were on land. All other drops were observed during dives, 88% of them during the undulatory (and occasionally the ascent) phase of dives deeper than 40 m. The mean swim speed during non-feeding shallow and exploratory dives was relatively constant throughout the dive, around 2.1 m s^-1, whereas during feeding deep dives, swim speed during the undulatory phase was lower (1.71 m s^-1) than during the descent and ascent and was characterised by a series of rapid accelerations and decelerations; 42.6% of these accelerations were followed by one or more ingestion events and birds swam upward in 60% of the accelerations. Such multiple data recording opens new paths for the examination of the decision-making processes in foraging penguins.     

Using logistic regression models to predict breeding success in male Adélie penguins (Pygoscelis adeliae)

Measures of breeding success are traditionally derived using the proportion of total nests that successfully reach strategic stages across a breeding season, such as pair formation, egg laying or fledging chicks. The use of logistic regression has recently become a popular tool in avian literature for identifying influential factors that predict nest and/or individual breeding success. In this study, we use logistic regression models to assess the importance of a range of factors affecting male Adélie penguin (Pygoscelis adeliae) breeding success during the 2002–2003 austral summer, when the presence of icebergs and extensive sea-ice meant overall breeding success of Adélie penguins was low (16/51 focal males had chicks reach fledging point). Logistic regression models for the early breeding season showed that good/average nest quality and central/middle nest location within the sub-colony were the best predictors of successful pair formation. Later, during incubation, the most successful males were those that not only returned earlier but were also heavier upon arrival and built nests of at least average quality. During the final stage when chicks had begun to fledge, the combined parameters of heavier male weight, early arrival time and good nest quality were the best predictors of breeding success. The logistic regression approach used here showed that the predictive ability of these parameters varied as the season progressed. However, most importantly, our logistic models fit the data well across all breeding stages.     

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.     

Foraging rhythms in Adélie penguins (Pygoscelis adeliae) at hope bay,Antarctica; determination and control

Summary We studied diel periodicity in activity of Adélie penguins (Pygoscelis adeliae) during the austral summer solstice and one month later by counting, hourly, numbers of birds leaving colonies to forage, numbers arriving back after foraging, numbers in a colony and numbers resting outside the colonies. During the solstice large numbers of birds were arriving at and departing from the colony at all times of the day although there was a tendency for more birds to be at sea when light intensity was highest at mid-day. Generally, birds not brooding chicks did not rest on land. A month later, when visibility was poor at mid-night, the percentage of birds at sea was highly positively correlated with light intensity. Birds returning from foraging in the evening fed chicks immediately and then either rested in the colonies or on snow patches between the colonies and the sea until the following morning.     

Fluoride turnover in adélie penguins (Pygoscelis adeliae) and other brid species

Summary Fluoride concentrations found in soft tissues of free ranging birds are below 10 mg/kg d.w. In ducks (Anas platyrhynchos) 99.5% of all fluoride is bound in the skeleton. Krill-eating adélie penguins (Pygoscelis adeliae) reached equilibrium concentrations in the order of 10000 mg F/kg d.w. in femur. Fluoride uptake with the diet is buffered by the skeleton where it has a half-life of 3–4 weeks. Fluoride from fresh krill does not reach the organs of adélie penguins to the same extent as NaF. Most fluoride remains bound to the krill cuticle and is not absorbed by the penguin gut during digestion. The availability of krill fluoride was in the range of 21%–30% for muscle and 48%–68% for femur of adélie penguins as compared to NaF (100%). The role of the salt glands in fluoride excretion is neglegible. The uptake of approx. 240 mg F daily with the krill diet is balanced by excretion via the cloaca.This work was supported by a grant from the DFG Nr. MZ-AD 24/9 and by a stipend from the Studienstiftung des deutschen Volkes     

Post-breeding dispersal of Ad��lie penguins (Pygoscelis adeliae) nesting at Signy Island, South Orkney Islands

ARGOS satellite telemetry and Global Location Sensors (geolocators) were used to identify the moult locations and the winter foraging dispersal of Adélie penguins after they left their breeding colonies on Signy Island in the South Orkney Islands. Animals were tracked during the period December 2004 to October 2005. All birds displayed a similar pattern of migratory behaviour, remaining away from colonies for approximately 9 months, at distances of up to 2,235 km. Moult locations were within the pack ice. Mean daily travel speeds to the moult locations were significantly faster when moving through open water than through pack ice. Moult occurred during February/March within a narrow latitudinal range (65–71°S), at a mean distance of 126 km from the ice edge; the mean duration of individual moult was c. 18.6 days. After moult, penguins spent the subsequent winter months moving north or north-eastward within the expanding winter pack ice, at a mean distance of 216 km from the ice edge, and in areas with ice cover >80%. The penguins returned to the vicinity of their colony between September 26 and October 22, 2005. This dependence of Adélie penguins on sea ice habitat suggests that any further reductions in sea ice extent in the Weddell Sea region would potentially have important impacts on the population processes of this pagophilic species.     

Fluoride excretion in adélie penguins (Pygoscelis adeliae) and mallard ducks (Anas platyrynchos)

• 1.1. Chronic F⁻ doses of NaF (40 and 80mg F⁻/kg per day) given to adélie penguins for 6–7 weeks led to a significant increase in F⁻ concentrations of nasal salt secretions with 1.16 and 2.22μg F⁻/g (wet wt) respectively as opposed to 0.66 μg F⁻/g in controls.
• 2.2. Single oral administration of 6 mg fluoride as NaF to mallard ducks led to increased F⁻ concentrations in the nasal salt secretions, with a mean of 0.67μg F⁻/g as opposed to 0.11 μg F⁻/g in controls.
• 3.3. In mallard ducks as well as in adélie penguins the amount of F⁻ excreted via the cloaca was 10³-10⁴ times that excreted via the nasal salt glands.

     

Characterization of CYP1A enzyme in Adélie penguin liver

As CYP1A enzymes are induced by certain contaminants, their induction pattern has been used as a biomarker for exposure of certain pollutants. Ethoxyresorufin O-deethylase (EROD) activities are widely used in environmental assessments of polychlorinated biphenyls in many wildlife species. The EROD activity, a typical probe for CYP1A enzyme was studied in liver microsomes prepared from Adélie penguins (Pygoscelis adeliae) (n=10). Penguin liver microsomes (0.5 mg/mL) were incubated with the substrate ethoxyresorufin and NADPH at 37 degrees C for 10 min, and the reaction was terminated by addition of methanol. The formation of the metabolite resorufin was assayed by an HPLC method. EROD activity was present in all liver samples studied. Penguin liver microsomal fraction exhibits typical Michaelis-Menten kinetics in the O-deethylation of ethoxyresorufin. The data were best described by a biphasic kinetic model, which could be interpreted in terms of two populations of CYP enzyme. Mean (+/-S.D.) K(m) values for high- and low-affinity components of EROD were 51+/-109 (range: 0.16 to 358) and 872+/-703 (range: 303 to 2450) nM, respectively. The corresponding mean V(max) values for the high- and low-affinity enzyme activities were 1.8+/-1.4 (range: 0.21 to 5.1) and 9.6+/-3.7 (range: 6.0 to 18.3) pmol/min/mg. The EROD activity in penguin liver microsomes was inhibited by CYP1A inhibitors (phenacetin, 7-ethoxycoumarin and proportional variant-naphthoflavone), whereas other CYP inhibitors for CYP2C9 (tolbutamide), 2C19 (mephenytoin), 2D6 (debrisoquin) and 2E1 (diethyldithiocarbamate) had no effect. These results suggest that CYP1A-like enzymes are present in penguin livers. The activity of this enzyme may be a useful biomarker for assessing the environmental impact of pollutants on Antarctic wildlife.     

Diet of Adélie penguins Pygoscelis adeliae during the post-hatching period at Esperanza Bay,Antarctica, 1987/88

The diet of the Adélie penguin Pygoscelis adeliae was studied at Esperanza Bay, Antarctic Peninsula, during the post-hatching period by quantitative analysis of adult stomach contents. Euphausiids constituted on average 96% by mass, while fish contributed 4% by mass. Amphipods were present in small amounts. Antarctic krill Euphausia superba was the predominant component throughout the sampling period. In contrast, Euphausia crystallorophias occurred rarely. Differences in the size of krill taken by Adélie penguins appear to reflect either local changes in the availability of certain age classes at various times in the breed ing season, or differences in foraging areas, or are due to year-to-year differences in prey availability and abundance, possibly caused by variations in seasonal ice cover. Pleuragramma antarcticum constituted the bulk of the fish portion, particularly during the guard period.     

Adélie penguin survival: age structure,temporal variability and environmental influences

The driving factors of survival, a key demographic process, have been particularly challenging to study, especially for winter migratory species such as the Adélie penguin (Pygoscelis adeliae). While winter environmental conditions clearly influence Antarctic seabird survival, it has been unclear to which environmental features they are most likely to respond. Here, we examine the influence of environmental fluctuations, broad climatic conditions and the success of the breeding season prior to winter on annual survival of an Adélie penguin population using mark–recapture models based on penguin tag and resight data over a 16-year period. This analysis required an extension to the basic Cormack–Jolly–Seber model by incorporating age structure in recapture and survival sub-models. By including model covariates, we show that survival of older penguins is primarily related to the amount and concentration of ice present in their winter foraging grounds. In contrast, fledgling and yearling survival depended on other factors in addition to the physical marine environment and outcomes of the previous breeding season, but we were unable to determine what these were. The relationship between sea-ice and survival differed with penguin age: extensive ice during the return journey to breeding colonies was detrimental to survival for the younger penguins, whereas either too little or too much ice (between 15 and 80% cover) in the winter foraging grounds was detrimental for adults. Our results demonstrate that predictions of Adélie penguin survival can be improved by taking into account penguin age, prior breeding conditions and environmental features.