An emperor penguin population estimate: the first global, synoptic survey of a species from space

Our aim was to estimate the population of emperor penguins (Aptenodytes fosteri) using a single synoptic survey. We examined the whole continental coastline of Antarctica using a combination of medium resolution and Very High Resolution (VHR) satellite imagery to identify emperor penguin colony locations. Where colonies were identified, VHR imagery was obtained in the 2009 breeding season. The remotely-sensed images were then analysed using a supervised classification method to separate penguins from snow, shadow and guano. Actual counts of penguins from eleven ground truthing sites were used to convert these classified areas into numbers of penguins using a robust regression algorithm.We found four new colonies and confirmed the location of three previously suspected sites giving a total number of emperor penguin breeding colonies of 46. We estimated the breeding population of emperor penguins at each colony during 2009 and provide a population estimate of ~238,000 breeding pairs (compared with the last previously published count of 135,000-175,000 pairs). Based on published values of the relationship between breeders and non-breeders, this translates to a total population of ~595,000 adult birds.There is a growing consensus in the literature that global and regional emperor penguin populations will be affected by changing climate, a driver thought to be critical to their future survival. However, a complete understanding is severely limited by the lack of detailed knowledge about much of their ecology, and importantly a poor understanding of their total breeding population. To address the second of these issues, our work now provides a comprehensive estimate of the total breeding population that can be used in future population models and will provide a baseline for long-term research.     

Climate Change Winners: Receding Ice Fields Facilitate Colony Expansion and Altered Dynamics in an Adélie Penguin Metapopulation

There will be winners and losers as climate change alters the habitats of polar organisms. For an Adélie penguin (Pygoscelis adeliae) colony on Beaufort Island (Beaufort), part of a cluster of colonies in the southern Ross Sea, we report a recent population increase in response to increased nesting habitat as glaciers have receded. Emigration rates of birds banded as chicks on Beaufort to colonies on nearby Ross Island decreased after 2005 as available habitat on Beaufort increased, leading to altered dynamics of the metapopulation. Using aerial photography beginning in 1958 and modern satellite imagery, we measured change in area of available nesting habitat and population size of the Beaufort colony. Population size varied with available habitat, and both increased rapidly since the 1990s. In accord with glacial retreat, summer temperatures at nearby McMurdo Station increased by ∼0.50°C per decade since the mid-1980s. Although the Ross Sea is likely to be the last ocean with an intact ecosystem, the recent retreat of ice fields at Beaufort that resulted in increased breeding habitat exemplifies a process that has been underway in the Ross Sea during the entire Holocene. Furthermore, our results are in line with predictions that major ice shelves and glaciers will retreat rapidly elsewhere in the Antarctic, potentially leading to increased breeding habitat for Adélie penguins. Results further indicated that satellite imagery may be used to estimate large changes in Adélie penguin populations, facilitating our understanding of metapopulation dynamics and environmental factors that influence regional populations.     

A method for estimating colony sizes of Adélie penguins using remote sensing imagery

Adélie penguins (Pygoscelis adeliae) are important predators of krill (Euphausia spp.) and Antarctic silverfish (Pleuragramma antarctica) during summer, are a key indicator of the status of the Southern Ocean ecosystem, and are therefore a focal species for the Committee for the Conservation of Antarctic Marine Living Resources (CCAMLR) Ecosystem Monitoring Program. The ability to monitor the population size of species potentially affected by Southern Ocean fisheries, i.e., the Adélie penguin, is critical for effective management of those resources. However, for several reasons, direct estimates of population size are not possible in many locations around Antarctica. In this study, we combine high-resolution (0.6 m) satellite imagery with spectral analysis in a supervised classification to estimate the sizes of Adélie penguin breeding colonies along Victoria Land in the Ross Sea and on the Antarctic Peninsula. Using satellite images paired with concurrent ground counts, we fit a generalized linear mixed model with Poisson errors to predict the abundance of breeding pairs as a function of the area of current-year guano staining identified in the satellite imagery. Guano-covered area proved to be an effective proxy for the number of penguins residing within. Our model provides a robust, quantitative mechanism for estimating the breeding population size of colonies captured in imagery and identifies terrain slope as a significant component influencing apparent nesting density. While our high-resolution satellite imagery technique was developed for the Adélie penguin, these principles are directly transferrable to other colonially nesting seabirds and other species that aggregate in fixed localities.     

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.     

Emigration in emperor penguins: implications for interpretation of long‐term studies

Site fidelity is an important evolutionary trait to understand, as misinterpretation of philopatric behavior could lead to confusion over the key drivers of population dynamics and the environmental or anthropogenic factors influencing populations. Our objective was to explore the hypothesis that emperor penguins are strictly philopatric using satellite imagery, counts from aerial photography, and literature reports on emperor penguin distributions. We found six instances over three years in which emperor penguins did not return to the same location to breed. We also report on one newly‐discovered colony on the Antarctic Peninsula that may represent the relocation of penguins from the Dion Islands, recently confirmed as having been abandoned. Using evidence from aerial surveys and the historical literature, we suggest that emigration may have been partly responsible for the population decline at Pointe Géologie during the 1970s. Our study is the first to use remote sensing imagery to suggest that emperor penguins can and do move between, and establish new, colonies. Metapopulation dynamics of emperor penguins have not been previously considered and represent an exciting, and important, avenue for future research. Life history plasticity is increasingly being recognized as an important aspect of climate change adaptation, and in this regard our study offers new insight for the long‐term future of emperor penguins.     

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.     

Kite aerial photography: a low‐cost method for monitoring seabird colonies

Obtaining aerial high‐resolution images of bird nesting colonies using remote‐sensing technology such as satellite‐based remote sensing, manned aircraft, or Unmanned Aerial Vehicles might not be possible for many researchers due to financial constraints. Kite Aerial Photography (KAP) provides a possible low‐cost alternative. We collected digital images of ground‐nesting seabirds (i.e., cormorants and penguins) in two different ecosystems using a kite‐based platform equipped with consumer‐grade digital cameras with time‐lapse capability to obtain estimates of breeding population size. KAP proved to be an efficient method for acquiring high‐resolution aerial images. We obtained images of colonies of seabirds ranging in size from hundreds to several hundreds of thousands breeding pairs during flights lasting from a few minutes up to three hours, from flat to very steep areas, and in contrasted wind conditions (from 0.5 to 6 Beaufort force). KAP is an efficient low‐cost method for acquiring high‐resolution aerial images and an alternative to ground‐based censuses, especially useful in rugged areas.     

Computer‐automated bird detection and counts in high‐resolution aerial images: a review

Bird surveys conducted using aerial images can be more accurate than those using airborne observers, but can also be more time‐consuming if images must be analyzed manually. Recent advances in digital cameras and image‐analysis software offer unprecedented potential for computer‐automated bird detection and counts in high‐resolution aerial images. We review the literature on this subject and provide an overview of the main image‐analysis techniques. Birds that contrast sharply with image backgrounds (e.g., bright birds on dark ground) are generally the most amenable to automated detection, in some cases requiring only basic image‐analysis software. However, the sophisticated analysis capabilities of modern object‐based image analysis software provide ways to detect birds in more challenging situations based on a variety of attributes including color, size, shape, texture, and spatial context. Some techniques developed to detect mammals may also be applicable to birds, although the prevalent use of aerial thermal‐infrared images for detecting large mammals is of limited applicability to birds because of the low pixel resolution of thermal cameras and the smaller size of birds. However, the increasingly high resolution of true‐color cameras and availability of small unmanned aircraft systems (drones) that can fly at very low altitude now make it feasible to detect even small shorebirds in aerial images. Continued advances in camera and drone technology, in combination with increasingly sophisticated image analysis software, now make it possible for investigators involved in monitoring bird populations to save time and resources by increasing their use of automated bird detection and counts in aerial images. We recommend close collaboration between wildlife‐monitoring practitioners and experts in the fields of remote sensing and computer science to help generate relevant, accessible, and readily applicable computer‐automated aerial photographic census techniques.     

The winter migration of Adelie penguins breeding in the Ross Sea sector of Antarctica

Satellite telemetry was used to monitor the migratory movements of a single Adelie penguin (Pygoscelis adeliae) from Cape Hallett (72.31°S, 170.21°E) following the 1997/1998 breeding season. Locations were obtained using the ARGOS satellite system and compared with the migratory paths taken by two penguins from the Northern Colony at Cape Bird, Ross Island (77.22°S, 166.48°E) following the 1990/1991 breeding season. Although the sample sizes are small, if representative they would indicate that: (1) Adelie penguins breeding in the Ross Sea follow a common migratory path, (2) Adelie penguins breeding in the Ross Sea may travel to a common over-winter feeding ground west and north of the Balleny Islands, and (3) Adelie penguins breeding at 77°S on Ross Island travel nearly twice the distance during their over-winter migration as do those penguins breeding at Cape Hallett and colonies further north. While the Cape Hallett penguin was tracked successfully for 172 days, a record for Adelie penguins, the problem of long-term attachment of transmitters to penguins remains.     

Satellite imagery can be used to detect variation in abundance of Weddell seals (Leptonychotes weddellii) in Erebus Bay, Antarctica

The Weddell seal population in Erebus Bay, Antarctica, represents one of the best-studied marine mammal populations in the world, providing an ideal test for the efficacy of satellite imagery to inform about seal abundance and population trends. Using high-resolution (0.6 m) satellite imagery, we compared counts from imagery to ground counts of adult Weddell seals and determined temporal trends in Erebus Bay during November 2004–2006 and 2009, and December 2007. Seals were counted from QuickBird-2 and WorldView-1 images, and these counts were compared with ground counts at overlapping locations within Erebus Bay during the same time. Counts were compared across years and within individual haul-out locations. We counted a total of 1,000 adult Weddell seals from five images across all years (for a total of 21 satellite-to-ground count comparisons), approximately 72% of the total counted on the ground at overlapping locations. We accurately detected an increase in abundance during 2004–2009. There was a strong, positive correlation (r = 0.98, df = 3, P < 0.003) between ground counts and counts derived from the imagery. The correlation between counts at individual haul-out locations was also strong (r = 0.80, df = 19, P < 0.001). Detection rates ranged from 30 to 88%. Overall, our results showed the utility of high-resolution imagery to provide an accurate way to detect the presence and variation in abundance of Weddell seals. Our methods may be applied to other species in polar regions, such as walruses or polar bears, particularly in areas where little is known about population status.     

The challenges of detecting subtle population structure and its importance for the conservation of emperor penguins

Understanding the boundaries of breeding populations is of great importance for conservation efforts and estimates of extinction risk for threatened species. However, determining these boundaries can be difficult when population structure is subtle. Emperor penguins are highly reliant on sea ice, and some populations may be in jeopardy as climate change alters sea‐ice extent and quality. An understanding of emperor penguin population structure is therefore urgently needed. Two previous studies have differed in their conclusions, particularly whether the Ross Sea, a major stronghold for the species, is isolated or not. We assessed emperor penguin population structure using 4,596 genome‐wide single nucleotide polymorphisms (SNPs), characterized in 110 individuals (10–16 per colony) from eight colonies around Antarctica. In contrast to a previous conclusion that emperor penguins are panmictic around the entire continent, we find that emperor penguins comprise at least four metapopulations, and that the Ross Sea is clearly a distinct metapopulation. Using larger sample sizes and a thorough assessment of the limitations of different analytical methods, we have shown that population structure within emperor penguins does exist and argue that its recognition is vital for the effective conservation of the species. We discuss the many difficulties that molecular ecologists and managers face in the detection and interpretation of subtle population structure using large SNP data sets, and argue that subtle structure should be taken into account when determining management strategies for threatened species, until accurate estimates of demographic connectivity among populations can be made.     

Penguins from space: faecal stains reveal the location of emperor penguin colonies

Aim  To map and assess the breeding distribution of emperor penguins ( Aptenodytes forsteri ) using remote sensing.
Location  Pan-Antarctic.
Methods  Using Landsat ETM satellite images downloaded from the Landsat Image Mosaic of Antarctica (LIMA), we detect faecal staining of ice by emperor penguins associated with their colony locations. Emperor penguins breed on sea ice, and their colonies exist in situ between May and December each year. Faecal staining at these colony locations shows on Landsat imagery as brown patches, the only staining of this colour on sea ice. This staining can therefore be used as an analogue for colony locations. The whole continental coastline has been analysed, and each possible signal has been identified visually and checked by spectral analysis. In areas where LIMA data are unsuitable, freely available Landsat imagery has been supplemented.
Results  We have identified colony locations of emperor penguins at a total of 38 sites. Of these, 10 are new locations, and six previously known colony locations have been repositioned (by over 10 km) due to poor geographical information in old records. Six colony locations, all from old or unconfirmed records, were not found or have disappeared.
Main conclusions  We present a new pan-Antarctic species distribution of emperor penguins mapped from space. In one synoptic survey we locate extant emperor penguin colonies, a species previously poorly mapped due to its unique breeding habits, and provide a vital geographical resource for future studies of an iconic species believed to be vulnerable to future climate change.     

Review of historical population information of emperor penguins

In 1902, the first breeding colony of emperor penguins was discovered. Over the following decades, the number of known emperor penguin colonies increased steadily and new ones are still being discovered. However, rigorous census work has been carried out at only a few colonies and accurate information on trends in breeding populations is limited to a small number of locations. Thus, the total number of breeding pairs is still unknown as is the size of the global population (breeders, non-breeders, juveniles). The International Union for the Conservation of Nature (IUCN) lists the species’ status as ‘least concern’ and states that although the population trend for emperor penguins has not been quantified, the global population appears to be stable. This review summarises the currently available information on the populations of emperor penguins at known colonies in terms of survey methods, count units used and survey frequency. It examines what is known about the state of various colonies and demonstrates that currently available data are inadequate for a trend assessment of the global population.     

Emperor penguins at the West Ice Shelf

In 1956, an emperor penguin (Aptenodytes forsteri) colony had been reported during an aerial survey north of the north-western protrusion of the West Ice Shelf in East Antarctica. About 15,000 birds were estimated to be present. The region often has very heavy pack ice conditions hindering access by vessels. In the summers of 2009–2011, we surveyed the area from the air and sighted two emperor penguin colonies. One was situated on top of the ice shelf and comprised 342 adults and 1,156 chicks. The second colony was seen near the northern edge of the West Ice Shelf on the sea ice about 60?km farther south than in 1956. There were at least 1,498 adults and 3,436 chicks.     

Holocene Adélie penguin diet in Victoria Land,Antarctica

Ornithogenic soils (N = 97) dated up to 7000 Before Present (bp) were sampled in 16 relict and modern breeding colonies of Adélie penguin along the Victoria Land coast (Ross Sea, Antarctica). Taxonomic identification of fish otoliths (N = 677) recovered in these soils allowed to identify the Antarctic silverfish as the most eaten prey (90.1%) throughout the investigated period. A morphometric analysis of the otoliths revealed that the Adélie penguin primarily selected prey averaging 67.23 ± 23 mm of standard length. Temporal distribution of Pleuragramma antarcticum showed a peak between 2,000 and 4,000 years bp, a period corresponding to the maximum spread of Adélie penguin in the Victoria Land. Possible explanations of the variations of the abundance of the fish prey in the diet are discussed in the context of the paleoclimatic events and as possible consequences of dietary shifts due to the temporal variation of prey availability in the Ross Sea ecosystem.     

Diet and foraging effort of Adélie penguins in relation to pack-ice conditions in the southern Ross Sea

We investigated the diet and aspects of foraging effort among Adélie penguins (Pygoscelis adeliae) breeding at three colonies on Ross Island, in the southwestern Ross Sea – Capes Royds, Bird and Crozier – during the chick-provisioning period of three austral summers, 1994–1995, 1995–1996 and 1996–1997. During the study period, pack-ice cover differed in waters offshore of these colonies, by colony, seasons and year. Diet differed among colonies only slightly. The fish Pleuragramma antarcticum was the most important prey, especially during years or periods within years when little pack ice was present. With respect to krill, which composed the remainder of diet, juvenile Euphausia crystallorophias were consumed predominantly in a year of heavy pack-ice cover; more adult krill were consumed in 2 years when pack ice was sparse. Foraging trip duration differed by colony, season and year and was related directly to distance from the colony to the nearest pack ice. The amount of food brought to chicks increased as trip duration increased, to a point (2 days), but then decreased as duration increased further (up to 4 days). On the basis of data on mass of parents and of meal sizes to chicks, it appeared that on the longest trips more of the food gathered by parents was used for self maintenance; on the longest trips, parents lost body mass. Successful foraging during chick rearing, the period when adult foraging is most intense, appears to depend on the proximity of pack ice to nesting colonies for this penguin species. Received: 1 October 1997 / Accepted: 25 April 1998     

Habitat Use by Weddell Seals and Emperor Penguins Foraging in the Ross Sea, Antarctica

The only apex predators that live year-round at high latitudesof the Ross Sea are the Weddell seal and emperor penguin. Theseasonal distribution, foraging depths, and diet of these twospecies appear to overlap. What makes it possible for emperorpenguins and Weddell seals to co-exist at high latitude throughoutthe winter when other marine tetrapods apparently cannot? Bothspecies have similar adaptations for exploitation of the deep-waterhabitat, forage on the same species, and routinely make longand deep dives. Yet, despite these similarities, there is probablylittle trophic overlap between the adults of both species dueto geographical and seasonal differences in habitat use. Forexample, during the winter months while female emperor penguinsare ranging widely in the pack ice, adult seals are foragingand fattening for the upcoming summer fast, literally beneaththe feet of the male penguins. However, there is more extensiveoverlap between juvenile seals and adult penguins, and shiftsin prey abundance and/or distribution would likely affect thesetwo groups similarly. In contrast, juvenile penguins appearto avoid inter- and intra- specific competition by leaving theRoss Sea once they molt.     

Satellite remote sensing in monitoring change of seabirds: use of Spot Image in king penguin population increase at Ile aux Cochons,Crozet Archipelago

The king penguin colony at Ile aux Cochons, Crozet Archipelago (southern Indian Ocean) is the largest in the world. The change in its surface area was investigated by means of satellite remote sensing using spot scene. The satellite picture enabled us to locate the colony and to determine that the surface area occupied by the colony has increased by 56% between 1962 and 1988. The increase in the surface area occupied by the colony was confirmed by a map realised in 1982 (by the Institut Géographique National). The high resolution system available on the spot system which allows at present at 10 m ground resolution, and 5 m in the near future, can thus be a useful tool for monitoring medium to long-term changes in marine bird populations breeding in colonies composed of birds spaced a constant distance from one another. Based on the surface area occupied by the colony and the published data on the population size on Possession Island, as well as information on the breeding frequency of the species, we estimated that the total breeding population is about 1,000,000 pairs for the whole archipelago and that about 70% of birds breed at Ile aux Cochons.     

Corticosterone responses to capture and restraint in emperor and Adelie penguins in Antarctica

Birds respond to capture, handling and restraint with increased secretion of corticosterone, a glucocorticoid hormone that helps birds adjust to stressful situations. Hoods are reported to calm birds, but possible effects of hoods on corticosterone responses have not been reported for any bird. Corticosterone responses to restraint in Adelie penguins held by their legs with their head covered by a hood were markedly lower than responses of penguins restrained in a mesh bag inside a cardboard box (corticosterone at 30 min 15.69+/-1.72 cf. 28.32+/-2.75 ng/ml). The birds restrained by the two methods were sampled at the same location but in different years, so the differences in corticosterone responses cannot unequivocally be ascribed to an effect of hoods to reduce corticosterone responses. Corticosterone responses have been measured in some penguins, but not in the largest, the emperor penguin (Aptenodytes forsteri). The relationship between body mass and corticosterone responses to capture and restraint in penguins was examined in emperor penguins captured on sea ice in McMurdo Sound and Adelie penguins (Pygoscelis adeliae) captured at Cape Bird, Ross Island, Antarctica. Total integrated corticosterone responses were higher in the emperor than the Adelie penguins, but corrected integrated corticosterone responses, which represent the increase in corticosterone from initial concentrations and hence the corticosterone response to restraint, were the same. The results for the emperor and Adelie penguins, together with data from other penguin species, suggest that there is no relationship between the size of corticosterone responses and body mass in penguins.