Ontogeny of thermoregulation and energy metabolism in pygoscelid penguin chicks

The ontogeny of thermoregulation and energy metabolism of chinstrap (Pygoscelis antarctica) and gentoo (P. papua) penguins was studied on King George Island, South Shetland Island, Antarctica. The major findings of this study are: Chinstrap and gentoo penguin chicks hatched completely poikilothermic, due to their poor heat-production ability at low ambient temperatures. They were able to maintain high body temperatures and metabolic rates only by being brooded by adults. Newly hatched chinstrap penguin chicks had, at a specified ambient temperature, significantly higher metabolic rates than newly hatched gentoos. Moreover, chinstrap chicks maintained a significantly higher body temperature. It is suggested that this is a non-acclimatory metabolic adaptation of chinstrap penguin chicks to the lower mean temperatures of their breeding areas. On the 15th day after hatching, chinstrap chicks were completely, and gentoo chicks almost completely, homeothermic. In spite of their high thermogenic capacity from about day 10, chicks were not at that time capable of controlling heat dissipation, and were still dependent on their parents. In older downy chicks and fledglings, heat loss at low temperatures, expressed as heat conductance (CA), was similar to that found for the adults of other penguin species. Just before moulting the CA of chicks was lower than after moulting. Moulting alone did not cause a clear increase in CA. Towards the end of their stay on land the CA of pre-fledged gentoos decreased by 31%. This decrease was not connected with the development of feathers or growth in the chicks' weight. The combination of the low CA and high SMR of chicks gave very low lower critical temperatures, near -15 degrees C. The wide thermoneutral zones of the chicks covered the whole range of air temperature variations in the breeding colonies of both species studied on King George Island. The CA values of homeothermic chinstrap chicks were not lower than those of gentoos, despite the more southern breeding range of the former species. The older chicks of both species are well protected against cold. Any further increase in insulation in chinstrap chicks would be of no adaptative importance.     

Blood glucose partition and levels of glycolytic enzymes in erythrocytes and somatic tissues of penguins

1. A comparative study was carried out on blood glucose partition and glucose metabolism of penguin erythrocytes and somatic tissues. Pygoscelidae penguins (Pygoscelis antarctica and P. papua) were used in these experiments. 2. Blood glucose partition was established by assaying whole blood and plasma glucose in several individuals of the gentoo and chinstrap penguins. 3. It was found that almost all the whole blood sugar is compartmentalized at the plasma site, the red blood cells being ineffective in regard to glucose metabolism. 4. Levels of hexokinase, phosphoglucose isomerase, phosphofructokinase, fructose bisphosphate aldolase, glyceraldehyde phosphate dehydrogenase, phosphoglycerate kinase, phosphopyruvate hydratase (enolase), pyruvate kinase, alpha-glycerolphosphate dehydrogenase and fructose bisphosphate phosphatase were estimated in the erythrocytes of both gentoo and chinstrap penguins, the same determinations being carried out also on the somatic tissues (leg muscle, breast muscle, heart muscle, liver and brain) of the gentoo.     

Assessing penguin colony size and distribution using digital mapping and satellite remote sensing

Changes in penguin abundance and distribution can be used to understand the response of species to climate change and fisheries pressures, and as a gauge of ecosystem health. Traditionally, population estimates have involved direct counts, but remote sensing and digital mapping methodologies can provide us with alternative techniques for assessing the size and distribution of penguin populations. Here, we demonstrate the use of a field-based digital mapping system (DMS), combining a handheld geographic information system with integrated geographical positioning system as a method for: (a) assessing penguin colony area and (b) ground-truthing colony area as derived from satellite imagery. Work took place at Signy Island, South Orkneys, where colonies of the three congeneric pygoscelid penguins: Adélie Pygoscelis adeliae, chinstrap P. antarctica and gentoo P. papua were surveyed. Colony areas were derived by mapping colony boundaries using the DMS with visual counts of the number of nesting birds made concurrently. Area was found to be a good predictor for number of nests for all three species of penguin. Using a maximum likelihood multivariate classification of remotely sensed satellite imagery (QuickBird2, 18 January 2010; Digital Globe ID: 01001000B90AD00), we were able to identify penguin colonies from the spectral signature of guano and differentiate between colonies of Adélie and chinstrap penguins. The area classified (all species combined) from satellite imagery versus area from DMS data was closely related (R ² = 0.88). Combining these techniques gives a simple and transferrable methodology for examining penguin distribution and abundance at local and regional scales.     

Chinstrap penguins (Pygoscelis antarctica) nesting on Sabrina Islet, Balleny Islands, Antarctica

Although the majority of chinstrap penguins (Pygoscelis antarctica) inhabit the Scotia Arc, a minuscule population of ten breeding pairs has been reported several times from Chinstrap Islet, in the Balleny Islands, 5,000 km distant on the opposite side of Antarctica. An aerial photographic census (December 2000) reveals that the overall penguin population of the Balleny Islands has declined by 8% since the last census 16 years ago, while the combined populations of the two largest colonies, Chinstrap Islet and Sabrina Islet, have increased by 11%. A ground visit confirmed for the first time that chinstrap penguins also breed on Sabrina Islet, occupying 20-24 nests on the margins of the Adélie colony (3,790 pairs in all). The small size and anomalous location of the Balleny Islands chinstrap population clearly warrant further study and the strongest protective measures.     

Divergent responses of Pygoscelis penguins reveal a common environmental driver

The responses of predators to environmental variability in the Antarctic Peninsula region have exhibited divergent patterns owing to variation in the geographic settings of colonies and predator life-history strategies. Five breeding colonies of Pygoscelis penguins from King George Island and Livingston Island, South Shetland Islands, Antarctica, were examined to (1) compare the responses of sympatric congeners to recent changes in their Antarctic ecosystem and (2) assess underlying causes for such responses. We used linear regression and correlation analyses to compare indices of abundance, recruitment, and summer breeding performance of the Adélie (P. adeliae), gentoo (P. papua), and chinstrap penguins (P. antarctica). Breeding colonies of Adélie and chinstrap penguins have declined by roughly 50% since the mid-1970s, and recruitment indices of Adélie penguins have declined by roughly 80%, but no such patterns are evident for gentoo penguins. Fledging success, however, has remained stable at all breeding colonies. The different trends in abundance and recruitment indices for each species, despite generally similar indices of summer performance, suggest that winter conditions contribute to the divergent responses among the penguins. In particular, strong correlations between indices of penguin and krill recruitment suggest that penguins in the South Shetland Islands may live under an increasingly krill-limited system that has disproportionate effects on the survival of juvenile birds.     

Geographic morphological variation of Gentoo penguin (Pygoscelis papua) and sex identification: using morphometric characters and molecular markers

Gentoo penguins Pygoscelis papua have sexual dimorphism, which females have smaller morphological measurements than males. P. papua also shows a geographic morphological variation, decreasing in size toward southern latitudes. Therefore, morphological measurements may vary in space and overlap in the distribution of traits between females and males, which could lead to sex misidentification. A total of 300 blood samples and eight measurements and weight were obtained for Gentoo penguins from three localities in the South Shetland Islands and the Antarctic Peninsula. The molecular sex was identified for all individuals using two primer pairs (P2/P8 and PL/PR). We obtained higher success amplification for P2/P8 compared to PL/PR. We used 172 adults to create a morphological discriminant function using the backward stepwise method. We established a discriminant function for sex identification of each locality due to significant morphological differences found between sexes and between the three studied localities for adult Gentoo penguins. Using these functions, the percentages of individuals correctly assigned was 93.87 % in Ardley Island, 88.09 % in Bernardo O’Higgins, and 83.95 % in Gabriel Gonzalez Videla. The most southern locality studied was the most morphologically divergent from the others and with low divergence between sexes. Moreover, we also established a general function for the Gentoo penguin, which can be used to identify the sex of penguins from a broad number of localities. Therefore, molecular and biometric are useful techniques for sexing Gentoo penguins.     

Differential leucocyte cell counts from the pygoscelid penguins of Antarctica

Differential leucocyte counts were obtained for three cogeneric species of wild antarctic penguins, Pygoscelis adelie (adelie), Pygoscelis papua (gentoo), and Pygoscelis antarctica (chinstrap). Significant differences between the differential leucocyte counts of the three species were not observed.     

Detection, differentiation, and abundance estimation of penguin species by high-resolution satellite imagery

Due to its high spatial resolution, broad spatial coverage, and cost-effectiveness, commercial satellite imagery is rapidly becoming a key component of biological monitoring in the Antarctic. While considerable success in surveying emperor penguins (Aptenodytes forsteri) has been facilitated by their large size and the visual simplicity of their habitat, there has been considerably less progress in mapping colonies on the Antarctic Peninsula and associated sub-Antarctic islands where smaller penguin species breed on topographically complex terrain composed of mixed substrates. Here, we demonstrate that Adélie penguin (Pygoscelis adeliae), chinstrap penguin (P. antarcticus), gentoo penguin (P. papua), and macaroni penguin (Eudyptes chrysolophus) colonies can be detected by high-resolution (2-m multispectral, 40–50-cm panchromatic) satellite imagery and that under ideal conditions, such imagery is capable of distinguishing among groups of species where they breed contiguously. To demonstrate the potential for satellite imagery to estimate penguin population abundance, we use satellite imagery of Paulet Island (63°35′S, 55°47′W) to estimate a site-wide population of 115,673 (99,222–127,203) breeding pairs of Adélie penguins.     

Geographic variation in the immunoglobulin levels in pygoscelid penguins

Antarctic organisms, including penguins, are susceptible to parasites and pathogens. Effects of infestation could differ in different locations along a geographical gradient from north to south consistent with conditions that affect the prevalence and virulence of parasites and pathogens. The immune system, including immunoglobulins as the main component of the humoral immune response, is the major way by which organisms confront infestation. We investigated the variation in immunoglobulin levels in three species of antarctic penguins (Pygoscelis antarctica, Pygoscelis papua, and Pygoscelis adeliae) along a geographical gradient from King George Island (62°15′S) to Avian Island (67°46′S). We found that immunoglobulin levels increased northwards in all the three species. This could indicate a higher impact of parasites and/or pathogens relative to the existing gradient in temperatures along this coast. Changing temperatures, consistent with global climate change, could be altering the ecology of parasite or pathogen infestation within the biota of northern Antarctica. We have also found marginal differences in immunoglobulin levels between sexes in both chinstrap and gentoo penguins.     

Isolation and characterization of macaroni penguin (Eudyptes chrysolophus) microsatellite loci and their utility in other penguin species (Spheniscidae, AVES)

We report the characterization of 25 microsatellite loci isolated from the macaroni penguin (Eudyptes chrysolophus). Thirteen loci were arranged into four multiplex sets for future genetic studies of macaroni penguin populations. All 25 loci were tested separately in each of four other penguin species [Adélie penguin (Pygoscelis adeliae), chinstrap penguin (Pygoscelis antarctica), gentoo penguin (Pygoscelis papua) and king penguin (Aptenodytes patagonicus)]. Between eight and 12 loci were polymorphic per species. These loci are expected to be useful for studies of population genetic structure in a range of penguin species.     

Killer whale predation on penguins in Antarctica

We report here the first published observations of killer whales (Orcinus orca) feeding on penguins in Antarctica. The sightings took place in the Gerlache Strait off the western Antarctic Peninsula during February 2010. Two species of pygoscelid penguins were taken—gentoo (Pygoscelis papua, at least four individuals) and chinstrap (P. antarctica, 2). From remains left at the surface, it was clear that the killer whales fed mainly on the breast muscles, although some penguins may have been swallowed whole. The killer whales were ecotype B, which are purported seal specialists, but we also saw ecotype A, prey specialists on Antarctic minke whales Balaenoptera bonaerensis, chase, but not catch penguins. Because of their small relative size, if penguins are regularly targeted by killer whales in Antarctica, the impact on their populations could be significant.     

First direct, site-wide penguin survey at Deception Island, Antarctica, suggests significant declines in breeding chinstrap penguins

Deception Island (62°57′S, 60°38′W) is one of the most frequently visited locations in Antarctica, prompting speculation that tourism may have a negative impact on the island’s breeding chinstrap penguins (Pygoscelis antarctica). Discussions regarding appropriate management of Deception Island and its largest penguin colony at Baily Head have thus far operated in the absence of concrete information regarding the current size of the penguin population at Deception Island or long-term changes in abundance. In the first ever field census of individual penguin nests at Deception Island (December 2–14, 2011), we find 79,849 breeding pairs of chinstrap penguins, including 50,408 breeding pairs at Baily Head and 19,177 breeding pairs at Vapour Col. Our field census, combined with a simulation designed to capture uncertainty in an earlier population estimate by Shuford and Spear (Br Antarct Surv Bull 81:19–30, 1988), suggests a significant (>50?%) decline in the abundance of chinstraps breeding at Baily Head since 1986/1987. A comparative analysis of high-resolution satellite imagery for the 2002/2003 and the 2009/2010 seasons suggests a 39?% (95th percentile CI?=?6–71?%) decline (from 85,473?±?23,352 to 52,372?±?14,309 breeding pairs) over that 7-year period and provides independent confirmation of population decline in the abundance of breeding chinstrap penguins at Baily Head. The decline in chinstrap penguins at Baily Head is consistent with declines in this species throughout the region, including sites that receive little or no tourism; as a consequence of regional environmental changes that currently represent the dominant influence on penguin dynamics, we cannot ascribe any direct link between chinstrap declines and tourism from this study.     

Population trends and reproductive success at a frequently visited penguin colony on the western Antarctic Peninsula

Petermann Island (65°10′S, 64°10′W), one of the Antarctic Peninsula’s most frequently visited locations, is at the epicenter of a rapid shift in which an Adélie penguin dominated fauna is becoming gentoo penguin dominated. Over the course of five seasons, the breeding productivity of Adélie and gentoo penguins breeding at Petermann Island were monitored to identify drivers of this rapid community change. The impact of tourist visitation on breeding success was also investigated. Consistent with larger trends in this region, the Adélie penguin population decreased by 29% and the gentoo penguin population increased by 27% between the 2003/2004 and 2007/2008 seasons. Reproductive success among Adélie penguins ranged from 1.09 to 1.32 crèched chicks/nest, which was higher than or comparable to other sites and is an unlikely explanation for the precipitous decline of Adélie penguins at Petermann Island. Whereas gentoo penguin reproductive success was lowest in colonies frequently visited by tourists, Adélie penguin colonies frequently visited by tourists had higher reproductive success than those visited only occasionally. These results are placed in the context of other studies on reproductive success and the impact of tourist visitation on breeding colonies of Adélie and gentoo penguins.     

Contrasting population changes in sympatric penguin species in association with climate warming

Climate warming and associated sea ice reductions in Antarctica have modified habitat conditions for some species. These include the congeneric Adélie, chinstrap and gentoo penguins, which now demonstrate remarkable population responses to regional warming. However, inconsistencies in the direction of population changes between species at different study sites complicate the understanding of causal processes. Here, we show that at the South Orkney Islands where the three species breed sympatrically, the less ice‐adapted gentoo penguins increased significantly in numbers over the last 26 years, whereas chinstrap and Adélie penguins both declined. These trends occurred in parallel with regional long‐term warming and significant reduction in sea ice extent. Periodical warm events, with teleconnections to the tropical Pacific, caused cycles in sea ice leading to reduced prey biomass, and simultaneous interannual population decreases in the three penguin species. With the loss of sea ice, Adélie penguins were less buffered against the environment, their numbers fluctuated greatly and their population response was strong and linear. Chinstrap penguins, considered to be better adapted to ice‐free conditions, were affected by discrete events of locally increased ice cover, but showed less variable, nonlinear responses to sea ice loss. Gentoo penguins were temporarily affected by negative anomalies in regional sea ice, but persistent sea ice reductions were likely to increase their available niche, which is likely to be substantially segregated from that of their more abundant congeners. Thus, the regional consequences of global climate perturbations on the sea ice phenology affect the marine ecosystem, with repercussions for penguin food supply and competition for resources. Ultimately, variability in penguin populations with warming reflects the local balance between penguin adaptation to ice conditions and trophic‐mediated changes cascading from global climate forcing.     

Dynamics of antarctic penguin populations in relation to inter-annual variability in sea ice distribution

To investigate the role of sea ice cover on penguin populations we used principal component analysis to compare population variables of Adélie (Pygoscelis adeliae) and chinstrap (Pygoscelis antarctica) penguins breeding on Signy Island, South Orkney Islands with local (from direct observations) and regional (from remote sensing data) sea ice variables. Throughout the study period, the Adélie penguin population size remained stable, whereas that of chinstrap penguins decreased slightly. For neither species were there significant relationships between population size and breeding success, except for an apparent inverse density-dependent relationship between the number of Adélie breeding pairs and the number of eggs hatching. For both species, no general relationship was found between either population size or breeding success and the local sea ice conditions. However, the regional sea ice extent at a particular time prior to the start of the breeding season was related to the number of birds that arrived to breed. For both species, this period occurred before the sea ice reached its maximum extent and was slightly earlier for Adélie than for chinstrap penguins. These results suggest that sea ice conditions outside the breeding season may play an important role in penguin population processes.     

An observation of between-mates feeding behaviour in chick-guarding chinstrap penguins

Courtship feeding has not been reported for chinstrap penguins (Pygoscelis antarctica). However, we observed a male chinstrap penguin fed to his mate during the guard stage of chick. After being fed, the female regurgitated the food to her chick. Our observation suggests that the chicks’ behaviour of begging for food may be retained in adults, but it is usually restrained.     

Absence of haematozoa in a wild chinstrap penguin Pygoscelis antarctica population

No haematoza were detected in 40 adult and 58 nestling wild chinstrap penguins (Pygoscelis antarctica) from Antarctica examined by blood smear. Received: 27 November 1996 / Accepted: 24 March 1997     

Breeding ecology of the gentoo penguin <Emphasis Type="Italic">Pygoscelis papua</Emphasis> at Kerguelen Archipelago

The gentoo penguin (Pygoscelis papua) is one of the most widespread penguin species and has been proven to be highly plastic in many aspects of its ecology. However, data from their sub-Antarctic range suggest an unexplained decline of their populations over the last 10–20 years, stressing the need for additional knowledge on their breeding ecology and demography. The present study provides insights into the breeding ecology of the gentoo penguin at a major breeding site, Kerguelen Archipelago, over three breeding seasons (1987, 2002 and 2003). Similarly to other northern populations, gentoo penguins breeding at Kerguelen exhibited winter laying, slow provisioning rate, slow growth rate associated with an extended rearing period and relatively low breeding success compared to southern populations. Our study also revealed interannual differences in the timing of laying and growth parameters as well as unusual sex differences in parental investment. Despite their high plasticity, there are indications that gentoo penguins at the northern edge of their range might work at the upper limit of their capacities. Sub-Antarctic populations would, therefore, be more sensitive to environmental changes than more southerly ones and need to be closely monitored.     

Gentoo penguin Pygoscelis papua diet as an indicator of planktonic availability in the Kerguelen Islands

Since penguins rely on the main planktonic resources of the Southern Ocean, knowledge of their diet may be used for monitoring these resources During winter and spring of 1987 and 1989, we investigated the composition of the diet of gentoo penguins. Pygoscelis papua, in relation to changes in the availability of two prey species, Euphausia vallentini and Themisto gaudichaudii, sampled during plankton surveys in the Kerguelen Islands. The comparison between plankton surveys and diet analysis was performed on samples taken 2–4 km from the studied colonies. Data on the abundance of Zooplankton derived from penguins' diet matched closely those from net hauls during a year of high plankton availability (1987). On the other hand, a weaker correspondence was found during a year of restricted availability (1989). The mean sizes of amphipods caught by penguins and net hauls were very similar but the size distribution showed comparatively fewer small and large individuals in net-hauls than in penguin stomachs. Detailed studies on the feeding range and foraging effort of penguins are therefore needed in the near future to validate the potential of penguin diet as a measure of plankton abundance.     

Genetic structure of declining chinstrap penguin (Pygoscelis antarcticus) populations from South Shetland Islands (Antarctica)

Seabirds and their response to climate perturbations are important bioindicators of changes in Antarctic ecosystems. During 30?years of observations of two chinstrap penguin (Pygoscelis antarcticus) colonies, one on King George Island and the other on Penguin Island (South Shetland Islands, Antarctica), the size of the breeding populations decreased by 84 and 41?%, respectively. We applied analyses of amplified fragment length polymorphisms to study the genetic structure of the two populations and to evaluate the influence of the sudden population decrease. Our data indicate that there were only weak genetic differences between the populations, which were not strong enough to support the hypothesis of population differentiation. Weak genetic differences observed between the two populations seem not to be determined by selection processes. We hypothesize that the very low level of between-population genetic structure can be explained by some extent of genetic drift, which is largely compensated by gene flow. Moreover, the two populations seem to remain in a stationary state. Our results support the hypothesis of limited natal philopatry in chinstrap penguins. The observed decrease in population size is probably caused by emigration or a rise in juvenile mortality due to the increasing krill limitation of the marine food web. However, detailed research is required to address this issue.