One of the longest continuing data sets involving a marine organism in the Antarctic is that of annual estimates of breeding population size of Adélie penguins Pygoscelis adeliae at colonies on Ross Island, Ross Sea, 1959 to 1997. The sizes of these colonies have displayed significant interannual variability during the 29-yr period. We hypothesized that changes are related to natural environmental factors; and used path analysis to analyze annual variation in population growth in relation to physical environmental factors during that part of the record with comparable sea-ice satellite imagery from 1973 to 1997. The Ross Sea sector of the Southern Ocean lying north of Ross Island, from 150°E to 130°W, comprised our study area. Annual population growth measured during summer was explained best, and inversely, by the extent of sea-ice in the study area 5 winters earlier, and in some way related to the Southern Oscillation. Analysis of a subset of the sea-ice data from 1979 to 1997 indicated strong correlations to ice conditions in the eastern portion of the study area (174 to 130°W), and virtually no correlations to the western half (150°E to 175°W). This result supported other indirect evidence that the Ross Island penguins winter in the eastern Ross Sea/western Amundsen Sea. A demographic model indicated that variation in survival of juveniles and subadults might account for the observed population variation, and would also explain the 5-yr lag as 5 yr is the average age of recruitment to the summer breeding population. Extensive sea-ice during winter appears to reduce subadult survival, expressed subsequently when these cohorts reach maturation. We hypothesize that extensive (more northerly) sea-ice limits access of penguins to productive waters known to occur south of the southern boundary of the Antarctic Circumpolar Current, with starvation or increased predation disproportionately affecting less-experienced birds. The observed patterns of penguin population change, including those preceding the satellite era, imply that sea-ice extent has changed significantly over recent decades.

In an investigation of the factors leading to geographic structuring among Adélie Penguin (Pygoscelis adeliae) populations, we studied the size and overlap of colony‐ specific foraging areas within an isolated cluster of colonies. The study area, in the southwestern Ross Sea, included one large and three smaller colonies, ranging in size from 3900 to 135 000 nesting pairs, clustered on Ross and Beaufort Islands. We used triangulation of radio signals from transmitters attached to breeding penguins to determine foraging locations and to define colony‐specific foraging areas during the chick‐provisioning period of four breeding seasons, 1997–2000. Colony populations (nesting pairs) were determined using aerial photography just after egg‐laying; reproductive success was estimated by comparing ground counts of chicks fledged to the number of breeding pairs apparent in aerial photos. Foraging‐trip duration, meal size, and adult body mass were estimated using RFID (radio frequency identification) tags and an automated reader and weighbridge. Chick growth was assessed by weekly weighing. We related the following variables to colony size: foraging distance, area, and duration; reproductive success; chick meal size and growth rate; and seasonal variation in adult body mass. We found that penguins foraged closest to their respective colonies, particularly at the smaller colonies. However, as the season progressed, foraging distance, duration, and area increased noticeably, especially at the largest colony. The foraging areas of the smaller colonies overlapped broadly, but very little foraging area overlap existed between the large colony and the smaller colonies, even though the foraging area of the large colony was well within range of the smaller colonies. Instead, the foraging areas of the smaller colonies shifted as that of the large colony grew. Colony size was not related to chick meal size, chick growth, or parental body mass. This differed from the year previous to the study, when foraging trips of the large colony were very long, parents lost mass, and chick meals were smaller. In light of existing data on prey abundance in neritic waters in Antarctica suggesting that krill are relatively evenly distributed and in high abundance in the Southern Ross Sea, we conclude that penguins depleted or changed the availability of their prey, that the degree of alteration was a function of colony size, and that the large colony affected the location (and perhaps ultimately the size) of foraging areas for the smaller colonies. It appears, therefore, that foraging dynamics play a role in the geographic structuring of colonies in this species.

Abstract:Simultaneous, but contrary, decadal-scale changes in population trajectories of two penguin species in the western Pacific and Ross Sea sectors of the Southern Ocean, during the early/mid-1970s and again during 1988-89, correspond to changes in weather and sea ice patterns. These in turn are related to shifts in the semi-annual and Antarctic oscillations. Populations of the two ecologically dissimilar penguin species -Adélie Pygoscelis adeliae and emperor Aptenodytes forsteri -have been tallied annually since the 1950s making these the longest biological datasets for the Antarctic. Both species are obligates of sea ice and, therefore, allowing for the demographic lags inherent in the response of long-lived species to habitat or environmental variation, the proximate mechanisms responsible for the shifts involved changes in coastal wind strength and air and sea temperatures, which in turn affected the seasonal formation and decay of sea ice and polynyas. The latter probably affected such rates as the proportion of adults breeding and ultimately the reproductive output of populations in ways consistent with the two species' opposing sea ice needs. Corresponding patterns for the mid-1970s shift were reflected also in ice-obligate Weddell seal Leptonychotes weddelli populations and the structure of shallow-water sponge communities in the Ross Sea. The 1988-89 shift, by which time many more datasets had become available, was reflected among several ice-frequenting vertebrate species from all Southern Ocean sectors. Therefore, the patterns most clearly identified in the Pacific Sector were apparently spread throughout the high latitudes of the Southern Ocean.

In animal populations, a minority of individuals consistently achieves the highest breeding success and therefore contributes the most recruits to future generations. On average, foraging performance is important in determining breeding success at the population level, but evidence is scarce to show that more successful breeders (better breeders) forage differently than less successful ones (poorer breeders). To test this hypothesis, we used a 10-year, three-colony, individual-based longitudinal data set on breeding success and foraging parameters of a long-lived bird, the Adélie Penguin, Pygoscelis adeliae. Better breeders foraged more efficiently than poorer breeders under harsh environmental conditions and when offspring needs were higher, therefore gaining higher net energy profit to be allocated to reproduction and survival. These results imply that adverse "extrinsic" conditions might select breeding individuals on the basis of their foraging ability. Adélie Penguins show sufficient phenotypic plasticity that at least a portion of the population is capable of surviving and successfully reproducing despite extreme variability in their physical and biological environment, variability that is likely to be associated with climate change and, ultimately, with the species' evolution. This study is the first to demonstrate the importance of "extrinsic" conditions (in terms of environmental conditions and offspring needs) on the relationship between foraging behavior and individual quality.

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High survival and breeding philopatry was previously confirmed for the Adélie penguin (Pygoscelis adeliae) during a period of stable environmental conditions. However, movements of breeding adults as a result of an unplanned natural experiment within a fourcolony meta-population provided interesting insights into this species' population dynamics. We used multistate mark-recapture models to investigate apparent survival and dispersal of breeding birds in the southwestern Ross Sea during 12 breeding seasons (1996)(1997)(1998)(1999)(2000)(2001)(2002)(2003)(2004)(2005)(2006)(2007). The natural experiment was facilitated by the temporary grounding of two immense icebergs that (i) erected a veritable fence separating colonies and altering migration routes and (ii) added additional stress by trapping extensive sea ice in the region during 5 of 12 y. Colony size varied by orders of magnitude, allowing investigation of apparent survival and dispersal rates in relation to both environmental conditions and colony size within this meta-population. Apparent survival was lowest for the smallest colony (4,000 pairs) and similar for the medium (45,000 pairs) and large colonies (155,000 pairs), despite increased foraging effort expended by breeders at the largest colony. Dispersal of breeding birds was low (<1%), except during years of difficult environmental conditions when movements increased, especially away from the smallest colony (3.5%). Decreased apparent survival at the smallest colony could reflect differences in migration chronology and winter habitat use compared with the other colonies, or it may reflect increased permanent emigration to colonies outside this meta-population. Contrary to current thought, breeding penguins are not always philopatric. Rather, stressful conditions can significantly increase dispersal rates.Antarctica | avian demography | climate change | philopatry | population dynamics

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Measurements of the size of Adélie penguin (Pygoscelis adeliae) colonies of the southern Ross Sea are among the longest biologic time series in the Antarctic. We present an assessment of recent annual variation and trends in abundance and growth rates of these colonies, adding to the published record not updated for more than two decades. High angle oblique aerial photographic surveys of colonies were acquired and penguins counted for the breeding seasons 1981–2012. In the last four years the numbers of Adélie penguins in the Ross and Beaufort Island colonies (southern Ross Sea metapopulation) reached their highest levels since aerial counts began in 1981. Results indicated that 855,625 pairs of Adélie penguins established breeding territories in the western Ross Sea, with just over a quarter (28%) of those in the southern portion, constituting a semi-isolated metapopulation (three colonies on Ross Island, one on nearby Beaufort Island). The southern population had a negative per capita growth rate of −0.019 during 1981–2000, followed by a positive per capita growth rate of 0.067 for 2001–2012. Colony growth rates for this metapopulation showed striking synchrony through time, indicating that large-scale factors influenced their annual growth. In contrast to the increased colony sizes in the southern population, the patterns of change among colonies of the northern Ross Sea were difficult to characterize. Trends were similar to southern colonies until the mid-1990s, after which the signal was lost owing to significantly reduced frequency of surveys. Both climate factors and recovery of whale populations likely played roles in the trends among southern colonies until 2000, after which depletion of another trophic competitor, the Antarctic toothfish (Dissostichus mawsoni), may explain the sharp increasing trend evident since then.