Comparative fuel metabolism in Gentoo and King Penguins: adaptation to brief versus prolonged fasting

To compare fuel utilization in large birds adapted to brief or prolonged fasting, protein and lipid utilization were quantified in the Gentoo Penguin (Pygoscelis papua) and the King Penguin (Aptenodytes patagonica). The inshore feeder Gentoo Penguin fasts for only a few days in its colony, while King Penguin chicks starve for several months in the subantarctic winter and male King Penguins starve for 5–6 weeks at the beginning of their breeding cycle. After an initial decrease in both daily body mass loss and nitrogen excretion during the first days of food deprivation, these two parameters thereafter stabilized at low values. At that time, protein utilization accounted for 15% of total energy expenditure in Gentoo Penguins and only 6% in King Penguin chicks during winter, the remainder (85% and 94%, respectively) being provided by fat oxidation. Similar percentages in fuel metabolism as seen in chicks during winter were reached in fasting adult King Penguins and spring chicks. However, a seasonal adaptation occurs in fasting chicks because energy expenditure is lower during winter. As previously described in starved mammals, the effectiveness in protein sparing could be related to the initial adiposity of the birds: the larger the fat stores (about 9% and 30% in Gentoo Penguins and winter chicks of King Penguins, respectively), the longer the fast duration and the better the level of protein conservation.     

Breeding sites and population of seabirds on Admiralty Bay,King George Island,Antarctica

The monitoring of the status and distribution of seabird populations is necessary to understand their spatial and temporal responses to rapid climate changes occurring in the Western Antarctic Peninsula area. We surveyed and mapped Admiralty Bay bird communities and related them to climate variables—temperature, temperature anomaly, Antarctic Oscillation Index and El-Niño Southern Oscillation Index. We recorded 13 breeding seabird species over three seasons (2009/2010, 2010/2011 and 2011/2012) and mapped 10 of them over an area of 149.5 ha. The ice-free areas with the greatest number of species were Point Thomas, Keller Peninsula and Hennequin Point. The most abundant species was the Adelie Penguin (Pygoscelis adeliae) followed by the Chinstrap Penguin (P. antarcticus). We observed that the number of breeding pairs of Gentoo Penguins (P. papua), Chinstrap Penguins and skuas (Catharacta maccormicki and C. antarctica) are related to temperature, temperature anomaly and El-Niño Southern Oscillation Index. The size of breeding populations and their distributions have been fluctuating over the last 30 years in ice-free areas of Admiralty Bay. Most species showed a decreasing trend from 1978 to 2012, with the exception of Chinstrap Penguins, Southern Giant Petrels (Macronectes giganteus) and skuas, which seem to be stable in numbers in the last two decades. Decreases in seabird populations from the Antarctic Peninsula are widely recognized as a response to environmental change and anthropogenic influences such as tourism and building activities, thus highlighting the importance of monitoring to support mitigation measures.     

In stark contrast to widespread declines along the Scotia Arc,a survey of the South Sandwich Islands finds a robust seabird community

The South Sandwich Islands, in the South Atlantic Ocean, are a major biological hot spot for penguins and other seabirds, but their remoteness and challenging coastlines preclude regular biological censuses. Here we report on an extensive survey of the South Sandwich Islands, the first since the late 1990s, which was completed through a combination of direct counting, GPS mapping, and interpretation of high-resolution commercial satellite imagery. We find that the South Sandwich Islands host nearly half of the world’s Chinstrap Penguin (Pygoscelis antarctica) population (1.3 million breeding pairs), as well as c. 95,000 breeding pairs of Macaroni Penguins (Eudyptes chrysolophus), and several thousand breeding pairs of Gentoo Penguins (Pygoscelis papua). Despite being at the northern edge of their breeding range, we found an unexpectedly large (≥125,000 breeding pairs) population of Adélie Penguins (Pygoscelis adeliae). Additionally, we report that nearly 1900 pairs of Southern Giant Petrels (Macronectes giganteus) breed in the South Sandwich Islands, 4 % of the global population, almost all of which are found on Candlemas Island. We find that the South Sandwich Islands have not experienced the same changes in penguin abundance and distribution as the rest of the Scotia Arc and associated portions of the western Antarctic Peninsula. This discovery adds important context to the larger conversation regarding changes to penguin populations in the Southern Ocean.     

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.     

Cycles of <Emphasis Type="Italic">Euphausia superba</Emphasis> recruitment evident in the diet of Pygoscelid penguins and net trawls in the South Shetland Islands,Antarctica

Size and sex of Antarctic krill taken from chinstrap and gentoo penguin diet were compared to those from scientific net surveys in the South Shetland Islands from 1998 to 2006 in order to evaluate penguin diet as a sampling mechanism and to look at trends in krill populations. Both penguin diet and net samples revealed a 4–5 year cycle in krill recruitment with one or two strong cohorts sustaining the population during each cycle. Penguin diet samples contained adult krill of similar lengths to those caught in nets; however, penguins rarely took juvenile krill. Penguin diet samples contained proportionately more females when the krill population was dominated by large adults at the end of the cycles; net samples showed greater proportions of males in these years. These patterns are comparable to those reported elsewhere in the region and are likely driven by the availability of different sizes and sexes of krill in relation to the colony.     

Pinguine – Überlebenskünstler in der Antarktis. Klimawandel

The future of penguin population development in the Western Antarctic Peninsula (= WAP) is largely depending on ecological factors like food availability (mostly krill) due to primary production of algae which itself depends on sea ice conditions, water‐ and air temperature and salinity. The extraordinary rise in temperature in the WAP area seems to cause a change in population numbers of Adelie and Gentoo Penguins: Adelies are declining in the north and Gentoos were occupying these sites instead. Gentoos have already reached the southern polar circle. These trends occurred in parallel with regional long‐term warming and significant reduction in sea ice extent. There is a lack of available information for penguin populations breeding possibly more south in the WAP area. We still have large gaps in our present knowledge in Adelies and Emperor Penguins southernmost breeding distribution.     

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 ecosystem evolution of penguin colonies in the past 8,500 years on Vestfold Hills, East Antarctica

Penguin colony is one of the Earth’s simplest ecosystems. As the seabird with the largest population in Antarctica, penguin is a unique indicator of Antarctic environment and climate changes. In this study, we collected an ornithogenic sediment core from Gardner Island in Vestfold Hills, East Antarctica, reconstructed an 8,500 years variation history of penguin population and vegetation abundance on this island, and examined the evolution of the penguin colony. We used the levels of two molecular markers cholesterol and cholestanol as the proxy indicators of penguin population size. Other molecular markers, including C_24:0 alkenoic acid, C₁₈ n-alkanol and phytol were used as the proxy indicators of aquatic moss, algae, and general vegetation, respectively. It is shown that the growth of algae was mainly affected by the nutritional supply from penguin droppings, so their abundance was positively linked with penguin population. The growth of aquatic moss, however, was controlled more by the degree of water body transparency than by nutrient availability. Because the pollution of water body increased as penguin population grew, aquatic moss abundance showed a seesaw-like relationship with penguin population. These results suggested that penguins played a dominant role in this simple ecosystem in the Antarctic environment. The reconstructed relationship between penguin population and vegetation abundance may offer new insights to understand ancient Antarctic environment and ecology.     

Population structure and phylogeography of the Gentoo Penguin (Pygoscelis papua) across the Scotia Arc

Climate change, fisheries' pressure on penguin prey, and direct human disturbance of wildlife have all been implicated in causing large shifts in the abundance and distribution of penguins in the Southern Ocean. Without mark‐recapture studies, understanding how colonies form and, by extension, how ranges shift is challenging. Genetic studies, particularly focused on newly established colonies, provide a snapshot of colonization and can reveal the extent to which shifts in abundance and occupancy result from changes in demographic rates (e.g., reproduction and survival) or migration among suitable patches of habitat. Here, we describe the population structure of a colonial seabird breeding across a large latitudinal range in the Southern Ocean. Using multilocus microsatellite genotype data from 510 Gentoo penguin (Pygoscelis papua) individuals from 14 colonies along the Scotia Arc and Antarctic Peninsula, together with mitochondrial DNA data, we find strong genetic differentiation between colonies north and south of the Polar Front, that coincides geographically with the taxonomic boundary separating the subspecies P. p. papua and P. p. ellsworthii. Using a discrete Bayesian phylogeographic approach, we show that southern Gentoos expanded from a possible glacial refuge in the center of their current range, colonizing regions to the north and south through rare, long‐distance dispersal. Our findings show that this dispersal is important for new colony foundation and range expansion in a seabird species that ordinarily exhibits high levels of natal philopatry, though persistent oceanographic features serve as barriers to movement.     

Short-term responses of king penguins Aptenodytes patagonicus to helicopter disturbance at South Georgia

The short-term behavioural effects of helicopter overflights on breeding king penguins Aptenodytes patagonicus at South Georgia were examined. Seventeen helicopter overflights were made at altitudes between 230 and 1,768 m (750–5,800 ft) above ground level. Noise from the aircraft engines and helicopter blades increased sound levels in the colony from a background level of 65–69 dB(A) to a maximum mean peak level of 80 dB(A) during overflights. Penguin behaviour changed significantly during all overflights at all altitudes compared to the pre- and post-flight periods. Pre-overflight behaviour resumed within 15 min of the aircraft passing overhead and no chicks or eggs were observed to be taken by predators during overflights. Non-incubating birds showed an increased response with reduced overflight altitude, but this was not observed in incubating birds. Variability in overflight noise levels did not affect significantly the behaviour of incubating or non-incubating birds. Penguins exhibited a reduced response to overflights as the study progressed (despite later flights generally being flown at lower altitudes) suggesting some degree of habituation to aircraft. To minimise disturbance to king penguins we recommend a precautionary approach such that overflights are undertaken at the maximum altitude that is operationally practical, or preferably are avoided altogether.     

Climate as a driver of population variability in breeding Gentoo Penguins Pygoscelis papua at the Falkland Islands

Detecting and predicting how populations respond to environmental variability are eminent challenges in conservation research and management. This is particularly true for wildlife populations at high latitudes, many of which demonstrate changes in population dynamics associated with global warming. The Falkland Islands (Southwest Atlantic) hold one of the largest Gentoo Penguin Pygoscelis papua populations in the world, representing c. 34% of the global population. The numbers of breeding Gentoo Penguins at the Falkland Islands have shown a high degree of inter‐annual variability since monitoring commenced in 1990. However, proximate causes of annual variability in breeding numbers have not been explored. Here we examine 21 years of Gentoo Penguin breeding surveys from the Falkland Islands and assess whether inter‐annual variability in the number of breeding pairs were correlated with proxies of environmental variability. There was a positive correlation between the number of breeding pairs and a broad‐scale climatic variation index, the Southern Oscillation Index (SOI). In turn, the SOI was significantly correlated with spring sea surface temperature anomalies, indicating a more immediate atmospherically forced response to El Niño Southern Oscillation variability in the Southwest Atlantic than previously reported. However, we also describe a non‐linear response to environmental variability that may highlight foraging plasticity and/or the complexity of regional ecosystem interactions that operate across a range of different scales.     

Fluctuations in Adélie penguin prey size in the mid to late Holocene, northern Marguerite Bay, Antarctic Peninsula

We investigated temporal changes in Adélie penguin prey size in northern Marguerite Bay, Antarctic Peninsula, through excavations of three abandoned and one active colony at Lagoon (67°35'S, 68°16'W) and Ginger Islands (67°45'S, 68°41'W), respectively, in austral summer 1999/2000. Radiocarbon dates on penguin bones and eggshell fragments collected at each site indicate that Lagoon Island was first occupied after 6000 BP and Ginger Island near 2275 BP. Identifiable non-krill prey remains (otoliths and squid beaks) were recovered from ornithogenic soils at all sites, with Antarctic silverfish (Pleuragramma antarcticum) and squid (Psychroteuthis glacialis) being the most abundant species represented in the deposits. Estimated mean standard lengths and mantle lengths of these two prey taxa, based on regressions with otolith and beak measurements respectively, indicate that Adélie penguins primarily select these prey within a mean size range of 95-117 mm. Prey size also varied significantly across seven occupation periods from 6000 BP to the present, but did not correlate with climate change.     

Ornithogenic soils of the cape bird adelie penguin rookeries,Antarctica

Summary The activity of soil-forming processes in the ornithogenic soils under the adelie penguin rookeries at Cape Bird, Ross Island, Antarctica, is limited by low temperatures and aridity, and is dominated by the influence of organic input from the penguins. A study of soil chemistry, biochemistry and biology of ornithogenic soils was carried out during the austral summers of 1981/1982 and 1983/1984. For the study, five sites were used to represent a relative age sequence in biological activity, from a presently occupied colony, a recently (previous season) abandoned colony, a re-worked guano site, and a long abandoned (many years) colony, to compare with a site that apparently had never been occupied by penguins. The sites were selected by the apparent age of the guano based on its morphology. Characteristics of the soils and guanos within the sequence are now described and illustrated, to complement the chemistry, biochemistry and biology of these sites, results of which have been published previously in this journal. A conventional radiocarbon age of 8 080±160 years BP (14C No. 5990) was obtained from a penguin bone in the long abandoned colony guano. A few temperature records indicated that soil temperatures were higher from soils with guano cover than from adjacent soils without guano cover. This difference in temperature should contribute to the increased biological and biochemical activity in the guano soils.     

Population variability in heat shock proteins among three Antarctic penguin species

Heat shock proteins (HSPs) are synthesised under stressful conditions such as exposure to elevated temperatures, contamination, free radicals, UV light or pathophysiological states resulting from parasites and/or pathogens. HSPs function to protect cells by means of modulation of protein folding. In Antarctica, these proteins have been studied in such organisms as protozoa and fishes, without attention to geographical variation. We studied the variation of HSP70 and HSP60 levels in Gentoo, Adelie and Chinstrap penguins among different populations along the Antarctic Peninsula from King George Island (62°15′S) to Avian Island (67°46′S). Our results show that the northern population of Gentoo penguin showed higher levels of HSP70 and HSP60 than the southern population. High temperature, human impact and immunity as a proxy for parasites and diseases in northern locations could explain such variation. Adelie penguin only showed significant geographical variation in HSP70, increasing north to south, a pattern perhaps related to increased UV radiation and decreased temperatures from north to south. Chinstrap penguin shows no population differences in the variation in neither HSP70 nor HSP60, although HSP70 showed marginally significant differences. Sexual differences in the level of these proteins are also discussed.     

High juvenile annual survival probabilities in Southern Rockhopper Penguins Eudyptes chrysocome are independent of individual fledging traits

Juvenile survival is an important demographic parameter. Southern Rockhopper Penguins Eudyptes chrysocome have undergone a dramatic population decline in the past century across their distribution, but the demographic processes are poorly understood. To estimate juvenile annual survival probabilities, Rockhopper Penguin chicks from two cohorts on New Island, Falkland Islands, were marked with transponders and recorded in subsequent years using an automated gateway. We first estimated annual survival and detection probabilities using a Cormack‐Jolly‐Seber (CJS) model, and found that both probabilities were extremely high (81% in the first and 98% in the second, third and fourth years of life), even in comparison with adult birds. Because detection probability after 3 years was effectively 1, and our sample size (n = 114) was too small to explore the effects of individual traits on survival in a CJS model, we assessed whether sex, cohort, body mass and laying sequence affected whether juveniles returned to the colony during their first 3 years of life using a simple generalized linear model that assumed perfect detection. Juveniles from the first cohort and males showed a higher return probability than juveniles from the second cohort and females. There was no clear effect of fledging body mass on return rate, probably related to the favourable environmental conditions during the study period. The laying sequence did not markedly affect the return probability of chicks, indicating that, once fledged, first‐laid A‐chicks have the same probability to return as second‐laid B‐chicks despite a much larger initial maternal investment in B‐eggs in this species. This study demonstrates extraordinarily high juvenile survival probabilities and will help to understand the recent changes in the population dynamics of the Falkland Islands Southern Rockhopper Penguins.     

Phosphorus limitation for the colony formation, growth and photosynthesis of an edible cyanobacterium, Nostoc sphaeroides

The percentage of spherical colonies from the trichomes of Nostoc sphaeroides reached 62–73% after 16 days with 50 and 250 μM P, but only10–15% at 0.5 and 5 μM P. During colony formation from microcolonies to macrocolonies, the growth rates were 95, 206 and 244% higher, respectively at 5, 50 and 250 μM P than that at 0.5 μM P. The light-saturated photosynthetic rate, maximum electron transport rate and light-limited photosynthetic efficiency at 0.5 μM P decreased, respectively, by 45, 51 and 32% than those at 250 μM P. These indicated that the colony development, growth and photosynthetic capacities were restricted at low P level, suggesting that P might be an important factor limiting the productivity and distribution of N. sphaeroides in the field.     

Testing methods to estimate abundance of Magellanic Penguins Spheniscus magellanicus

Capsule Using simulations in a geographical information system our results showed that systematic sampling with quadrats was the most accurate, precise and cost-effective method to survey Magellanic Penguin colonies.

Aims To determine which sampling method gives better estimations of penguin abundance.

Methods A virtual colony was generated deriving spatial parameters from a real survey and applying Kriging interpolation. Three sampling methods were then applied on this virtual colony: random sampling with quadrats; systematic sampling with quadrats; systematic sampling with fixed-width transects. The estimated abundance for each trial was compared to the abundance of the virtual colony to have a measure of accuracy and precision.

Results Systematic sampling with quadrats estimated penguin abundance better than random or systematic sampling with transects since it achieved 100% accuracy and great precision after sampling only 2.1% of the virtual colony.

Conclusion The use of a simulated colony allowed the comparison of several sampling methods traditionally used in Magellanic Penguin surveys. The results of this study are important in order to standardize sampling protocols for Magellanic Penguins and to have more comparable estimations to detect trends over time. Also, the methodological approach used here could be used to assess sampling methods for other colonial bird species.     

Foraging ecology of Gentoo Penguins Pygoscelis papua at Macquarie Island during the period of chick care

The diet, diving behaviour, swimming velocity and foraging range of Gentoo Penguins Pygoscelis papua were studied at Macquarie Island during the breeding season in the 1993–1994 austral summer. Gentoo Penguins are considered to be inshore feeders, and at Macquarie Island the diet and estimated foraging ranges supported this. The diet consisted of 91.6% fish and 8.3% squid, by mass. The dominant prey taxa were the fish Gymnoscopelus sp. and Paranotothenia magellanica. A mixture of pelagic and benthic prey was consumed, with a greater proportion of benthic species occurring later in the season. The penguins exhibited a strong diurnal pattern in their diving behaviour. Deep diving (≥30 m) began near sunrise (03.00 h) and finished close to sunset (21.00 h). Diving at night was less common and very shallow (<10 m). Early in the breeding season, dive profiles indicated that birds were probably following vertically migrating pelagic prey through the water column and were foraging in waters over 100 m deep. Later in the season, more uniform, shallower depths were used, suggesting an increase in benthic foraging activity. These changes in dive pattern and depth were consistent with the habitat preferences of prey species found in the diet. Gentoo Penguins swam at 1.04 m per s and had a maximum potential foraging range of about 26 km for single-day trips. They tended to forage within 14 km of the colony, with a mean range of 5.4 km. This range encompassed the deep ocean habitat to the west and east of the island and a shallow area to the north.     

The end of the Kachemak tradition on the Kenai Peninsula, southcentral Alaska

The Kachemak tradition was established by ca. 3000 B.P. in Kachemak Bay. Probably somewhat later a variant termed Riverine Kachemak, with a population adapted to salmon and terrestrial resources, appeared on the northern Kenai Peninsula. The Kachemak tradition people seem to have abandoned Kachemak Bay by ca. 1400 B.P. Seven of 12 available Kachemak tradition dates predate 1400 B.P. even at two sigma. Scattered younger dates are thus suspect outliers. The end of Riverine Kachemak tradition has been placed at ca. 1000 B.P., at which time the population was supposedly replaced by in-migrating groups ancestral to the Dena'ina Athapaskans. Close examination of the numerous available radiocarbon dates shows that most Riverine Kachemak dates cluster in the early centuries of the First Millennium A.D. and most Dena'ina dates substantially postdate 1000 A.D. Probably the Riverine Kachemak and Dena'ina peoples never met on the Kenai River. However, the correspondence in date ranges between Kachemak Bay and Riverine Kachemak is striking, suggesting their fates were linked. Both traditions collapsed by 1400-1500 B.P. The causes are probably multiple but do not include cultural replacement.