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.     

Penguin responses to climate change in the Southern Ocean

Penguins are adapted to live in extreme environments, but they can be highly sensitive to climate change, which disrupts penguin life history strategies when it alters the weather, oceanography and critical habitats. For example, in the southwest Atlantic, the distributional range of the ice‐obligate emperor and Adélie penguins has shifted poleward and contracted, while the ice‐intolerant gentoo and chinstrap penguins have expanded their range southward. In the Southern Ocean, the El Niño‐Southern Oscillation and the Southern Annular Mode are the main modes of climate variability that drive changes in the marine ecosystem, ultimately affecting penguins. The interaction between these modes is complex and changes over time, so that penguin responses to climate change are expected to vary accordingly, complicating our understanding of their future population processes. Penguins have long life spans, which slow microevolution, and which is unlikely to increase their tolerance to rapid warming. Therefore, in order that penguins may continue to exploit their transformed ecological niche and maintain their current distributional ranges, they must possess adequate phenotypic plasticity. However, past species‐specific adaptations also constrain potential changes in phenology, and are unlikely to be adaptive for altered climatic conditions. Thus, the paleoecological record suggests that penguins are more likely to respond by dispersal rather than adaptation. Ecosystem changes are potentially most important at the borders of current geographic distributions, where penguins operate at the limits of their tolerance; species with low adaptability, particularly the ice‐obligates, may therefore be more affected by their need to disperse in response to climate and may struggle to colonize new habitats. While future sea‐ice contraction around Antarctica is likely to continue affecting the ice‐obligate penguins, understanding the responses of the ice‐intolerant penguins also depends on changes in climate mode periodicities and interactions, which to date remain difficult to reproduce in general circulation models.     

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.     

Sea ice predicts long‐term trends in Adélie penguin population growth,but not annual fluctuations: Results from a range‐wide multiscale analysis

Understanding the scales at which environmental variability affects populations is critical for projecting population dynamics and species distributions in rapidly changing environments. Here we used a multilevel Bayesian analysis of range‐wide survey data for Adélie penguins to characterize multidecadal and annual effects of sea ice on population growth. We found that mean sea ice concentration at breeding colonies (i.e., “prevailing” environmental conditions) had robust nonlinear effects on multidecadal population trends and explained over 85% of the variance in mean population growth rates among sites. In contrast, despite considerable year‐to‐year fluctuations in abundance at most breeding colonies, annual sea ice fluctuations often explained less than 10% of the temporal variance in population growth rates. Our study provides an understanding of the spatially and temporally dynamic environmental factors that define the range limits of Adélie penguins, further establishing this iconic marine predator as a true sea ice obligate and providing a firm basis for projection under scenarios of future climate change. Yet, given the weak effects of annual sea ice relative to the large unexplained variance in year‐to‐year growth rates, the ability to generate useful short‐term forecasts of Adélie penguin breeding abundance will be extremely limited. Our approach provides a powerful framework for linking short‐ and longer term population processes to environmental conditions that can be applied to any species, facilitating a richer understanding of ecological predictability and sensitivity to global change.     

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.     

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.     

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.     

The influence of historical climate changes on Southern Ocean marine predator populations: a comparative analysis

The Southern Ocean ecosystem is undergoing rapid physical and biological changes that are likely to have profound implications for higher‐order predators. Here, we compare the long‐term, historical responses of Southern Ocean predators to climate change. We examine palaeoecological evidence for changes in the abundance and distribution of seabirds and marine mammals, and place these into context with palaeoclimate records in order to identify key environmental drivers associated with population changes. Our synthesis revealed two key factors underlying Southern Ocean predator population changes; (i) the availability of ice‐free ground for breeding and (ii) access to productive foraging grounds. The processes of glaciation and sea ice fluctuation were key; the distributions and abundances of elephant seals, snow petrels, gentoo, chinstrap and Adélie penguins all responded strongly to the emergence of new breeding habitat coincident with deglaciation and reductions in sea ice. Access to productive foraging grounds was another limiting factor, with snow petrels, king and emperor penguins all affected by reduced prey availability in the past. Several species were isolated in glacial refugia and there is evidence that refuge populations were supported by polynyas. While the underlying drivers of population change were similar across most Southern Ocean predators, the individual responses of species to environmental change varied because of species specific factors such as dispersal ability and environmental sensitivity. Such interspecific differences are likely to affect the future climate change responses of Southern Ocean marine predators and should be considered in conservation plans. Comparative palaeoecological studies are a valuable source of long‐term data on species’ responses to environmental change that can provide important insights into future climate change responses. This synthesis highlights the importance of protecting productive foraging grounds proximate to breeding locations, as well as the potential role of polynyas as future Southern Ocean refugia.     

Effects of climate change on an emperor penguin population: analysis of coupled demographic and climate models

Sea ice conditions in the Antarctic affect the life cycle of the emperor penguin (Aptenodytes forsteri). We present a population projection for the emperor penguin population of Terre Adélie, Antarctica, by linking demographic models (stage‐structured, seasonal, nonlinear, two‐sex matrix population models) to sea ice forecasts from an ensemble of IPCC climate models. Based on maximum likelihood capture‐mark‐recapture analysis, we find that seasonal sea ice concentration anomalies (SIC_a) affect adult survival and breeding success. Demographic models show that both deterministic and stochastic population growth rates are maximized at intermediate values of annual SIC_a, because neither the complete absence of sea ice, nor heavy and persistent sea ice, would provide satisfactory conditions for the emperor penguin. We show that under some conditions the stochastic growth rate is positively affected by the variance in SIC_a. We identify an ensemble of five general circulation climate models whose output closely matches the historical record of sea ice concentration in Terre Adélie. The output of this ensemble is used to produce stochastic forecasts of SIC_a, which in turn drive the population model. Uncertainty is included by incorporating multiple climate models and by a parametric bootstrap procedure that includes parameter uncertainty due to both model selection and estimation error. The median of these simulations predicts a decline of the Terre Adélie emperor penguin population of 81% by the year 2100. We find a 43% chance of an even greater decline, of 90% or more. The uncertainty in population projections reflects large differences among climate models in their forecasts of future sea ice conditions. One such model predicts population increases over much of the century, but overall, the ensemble of models predicts that population declines are far more likely than population increases. We conclude that climate change is a significant risk for the emperor penguin. Our analytical approach, in which demographic models are linked to IPCC climate models, is powerful and generally applicable to other species and systems.     

Leukocyte counts in different populations of Antarctic <Emphasis Type="Italic">Pygoscelid</Emphasis> penguins along the Antarctic Peninsula

The Antarctic Peninsula is one of the areas where the climate is changing at the fastest pace, having several effects on the populations of pygoscelid penguins. Few studies have analysed the variation in immune parameters of antarctic birds in a geographical context; thus, analyses of geographical differences in the immune components of wild pygoscelid penguins are still scarce. Leukocyte counts in birds provide information on their immunity and physiological stress. The objective of this study was to analyse the leukocyte counts in penguins of the genus Pygoscelis (gentoo, Adélie and chinstrap penguins), covering sites along the South Shetland Islands and some islands on the west coast of the Antarctic Peninsula. Our results revealed differences in the number of heterophils and eosinophils and in the heterophil/lymphocyte ratio in the northeastern populations of gentoo and Adélie penguins as compared to the rest of the colonies studied. The results contribute to better understanding of the variations in physiological parameters of penguins related to a geographical context.     

Warming effects in the western Antarctic Peninsula ecosystem: the role of population dynamic models for explaining and predicting penguin trends

The western Antarctica Peninsula and Scotia Sea ecosystems appear to be driven by complex links between climatic variables, primary productivity, krill and Avian predators. There are several studies reporting statistical relationships between climate, krill and Penguin population size. The Adélie (Pygoscelis adeliae), Chinstrap (P. antarctica) and Gentoo (P. papua) penguins appear to be influenced by interannual variability in sea-ice extent and krill biomass. In this paper we developed simple conceptual models to decipher the role of climate and krill fluctuations on the population dynamics of these three Pygoscelis penguin species inhabiting the Antarctic Peninsula region. Our results suggest that the relevant processes underlying the population dynamics of these penguin species at King George Island (South Shetland Islands) are intra-specific competition and the combined effects of krill abundance and sea-ice cover. Our results using population theoretical models appear to support that climate change, specifically regional warming on the western Antarctic Peninsula, represents a major driver. At our study site, penguins showed species-specific responses to climate change. While Chinstrap penguins were only influenced by krill abundance, the contrasting population trends of Adélie and Gentoo penguins appear to be better explained by the “sea-ice hypothesis”. We think that proper population dynamic modeling and theory are essential for deciphering and proposing the ecological mechanisms underlying dynamics of these penguin populations.     

The interaction between island geomorphology and environmental parameters drives Adélie penguin breeding phenology on neighboring islands near Palmer Station,Antarctica

Despite many studies on Adélie penguin breeding phenology, understanding the drivers of clutch initiation dates (CIDs, egg 1 lay date) is limited or lacks consensus. Here, we investigated Adélie penguin CIDs over 25 years (1991–2016) on two neighboring islands, Torgersen and Humble (<1 km apart), in a rapidly warming region near Palmer Station, Antarctica. We found that sea ice was the primary large‐scale driver of CIDs and precipitation was a secondary small‐scale driver that fine‐tunes CID to island‐specific nesting habitat geomorphology. In general, CIDs were earlier (later) when the spring sea ice retreat was earlier (later) and when the preceding annual ice season was shorter (longer). Island‐specific effects related to precipitation and island geomorphology caused greater snow accumulation and delayed CIDs by ~2 days on Torgersen compared to Humble Island. When CIDs on the islands were similar, conditions were mild with less snow across breeding sites. At Torgersen Island, the negative relationship between CID and breeding success highlights detrimental effects of delayed breeding and/or snow on penguin fitness. Past phenological studies reported a relationship between air temperature and CID, assumed to be related to precipitation, but we found air temperature was more highly correlated to sea ice, revealing a misinterpretation of temperature effects. Finally, contrasting trends in CIDs based on temporal shifts in regional sea ice patterns revealed trends toward earlier CIDs (4–6 day advance) from 1979 to 2009 as the annual ice season shortened, and later CIDs (7–10 day delay) from 2010 to 2016 as the annual ice season lengthened. Adélie penguins tracked environmental conditions with flexible breeding phenology, but their life history remains vulnerable to subpolar weather conditions that can delay CIDs and decrease breeding success, especially on landscapes where geomorphology facilitates snow accumulation.     

Morphological aspects of the heart of the northern rockhopper penguin (Eudyptes chrysocome moseleyi ): possible implication in diving behavior and ecology?

We compared the heart morphology of the small, deep-diving northern rockhopper penguin to the hearts of small, shallow-diving and large, deep-diving penguin species. The rockhopper penguin had a heart larger than expected for its body mass, and its heart weight/body weight was significantly greater than in the larger Adélie penguin. We found the rockhopper's right ventricle weight/heart weight to be significantly greater than this relationship in both the larger chinstrap and Adélie penguins. The relationship of the right to left ventricular weights in the rockhopper heart is not different to that of the large, deepest-diving emperor penguin. A larger heart in the rockhopper penguin might be related to its diving behavior and ecology if it contributes to diving efficiency during foraging by increasing lung perfusion during surface recovery. This would lead to decreased surface time. Accepted: 20 May 2000     

Marine pelagic ecosystems: the west Antarctic Peninsula

The marine ecosystem of the West Antarctic Peninsula (WAP) extends from the Bellingshausen Sea to the northern tip of the peninsula and from the mostly glaciated coast across the continental shelf to the shelf break in the west. The glacially sculpted coastline along the peninsula is highly convoluted and characterized by deep embayments that are often interconnected by channels that facilitate transport of heat and nutrients into the shelf domain. The ecosystem is divided into three subregions, the continental slope, shelf and coastal regions, each with unique ocean dynamics, water mass and biological distributions. The WAP shelf lies within the Antarctic Sea Ice Zone (SIZ) and like other SIZs, the WAP system is very productive, supporting large stocks of marine mammals, birds and the Antarctic krill, Euphausia superba. Ecosystem dynamics is dominated by the seasonal and interannual variation in sea ice extent and retreat. The Antarctic Peninsula is one among the most rapidly warming regions on Earth, having experienced a 2 degrees C increase in the annual mean temperature and a 6 degrees C rise in the mean winter temperature since 1950. Delivery of heat from the Antarctic Circumpolar Current has increased significantly in the past decade, sufficient to drive to a 0.6 degrees C warming of the upper 300 m of shelf water. In the past 50 years and continuing in the twenty-first century, the warm, moist maritime climate of the northern WAP has been migrating south, displacing the once dominant cold, dry continental Antarctic climate and causing multi-level responses in the marine ecosystem. Ecosystem responses to the regional warming include increased heat transport, decreased sea ice extent and duration, local declines in icedependent Adélie penguins, increase in ice-tolerant gentoo and chinstrap penguins, alterations in phytoplankton and zooplankton community composition and changes in krill recruitment, abundance and availability to predators. The climate/ecological gradients extending along the WAP and the presence of monitoring systems, field stations and long-term research programmes make the region an invaluable observatory of climate change and marine ecosystem response.     

Sea ice affects the population dynamics of Adélie penguins in Terre Adélie

Overall Adélie penguin population size in Pointe Géologie Archipelago increased between 1984 and 2003 at a rate of 1.77% per year, and averaged 33,726±5,867 pairs. As predicted by the optimum model proposed by Smith et al. (Bioscience 49:393–404, 1999). Adélie penguin population size increased when sea ice extent and concentration (SIE and SIC) decreased six years earlier, indicating that the conditions around reproduction or first years at sea, were determinant. The breeding success averaged 85.2±35.45% and was not related to environmental variables. Adult survival probability varied between years from 0.64 to 0.82. Southern oscillation index (SOI) had a strong negative effect on adult annual survival. Adult survival of Adélie penguins increased during warmer events, especially during winter and spring at the beginning of reproduction. Therefore, we speculate that the rapid decreases in 1988–1991 and 1996 of the breeding population size were related to a decrease in adult mortality. However, adult survival varied little, and could not explain the strong increasing population trend. The sea ice conditions during breeding or during the first year at sea appeared determinant and influenced the population dynamics through cohort effects, probably related to the availability of productive feeding habitats.     

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.     

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.     

Long-term population trends in seven Antarctic seabirds at Pointe Géologie (Terre Adélie) Human impact compared with environmental change

Antarctic seabird populations have been much studied over the last decades as bioindicators of the nature of variability in the Southern Ocean marine ecosystem, and most attention has been focused on the role of food supply and the extent of sea ice. In addition, the rapid spread of tourism and the activities of researchers since the early 1960s have raised questions related to their real and potential impact on bird populations. Our data sets start in 1952 for several species of Antarctic seabirds and this study documents the trends over a 14-year period (1985–1999) in seven species breeding on Pointe Géologie archipelago (Terre Adélie, Antarctica). This is the first study where the direct impact of destruction of breeding sites (for building of an airstrip) is examined and where such long-term populations trends have been assessed in such a number of Antarctic species at one site. Trends from 1985 show that for the whole archipelago and when excluding islands destroyed, Adélie penguins and south polar skuas were the only species to show a significant increase (>3.5% annual change). The others species showed opposite trends, three increasing slightly (southern fulmars +0.4%, cape petrels +2.3%, snow petrels +0.9%) and two decreasing (emperor penguin −0.9%, southern giant petrel −3.9%). Three species particularly affected by the destruction of their breeding habitat (Adélie penguin, cape petrel, snow petrel) showed the capability to restore their populations. The availability of food and nesting sites is discussed in relation to environmental change. Species feeding on krill (Adélie penguins and cape petrels) increased more than other species; however, decrease of ice cover can increase availability of nesting sites. The importance of long-term studies is shown when assessing the role of human activities in Antarctica compared to larger-scale changes. Accepted: 18 September 2000     

Too much of a good thing: sea ice extent may have forced emperor penguins into refugia during the last glacial maximum

The relationship between population structure and demographic history is critical to understanding microevolution and for predicting the resilience of species to environmental change. Using mitochondrial DNA from extant colonies and radiocarbon‐dated subfossils, we present the first microevolutionary analysis of emperor penguins (Aptenodytes forsteri) and show their population trends throughout the last glacial maximum (LGM, 19.5–16 kya) and during the subsequent period of warming and sea ice retreat. We found evidence for three mitochondrial clades within emperor penguins, suggesting that they were isolated within three glacial refugia during the LGM. One of these clades has remained largely isolated within the Ross Sea, while the two other clades have intermixed around the coast of Antarctica from Adélie Land to the Weddell Sea. The differentiation of the Ross Sea population has been preserved despite rapid population growth and opportunities for migration. Low effective population sizes during the LGM, followed by a rapid expansion around the beginning of the Holocene, suggest that an optimum set of sea ice conditions exist for emperor penguins, corresponding to available foraging area.     

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.