Coordinated movements prevent jamming in an Emperor penguin huddle

For Emperor penguins (Aptenodytes forsteri), huddling is the key to survival during the Antarctic winter. Penguins in a huddle are packed so tightly that individual movements become impossible, reminiscent of a jamming transition in compacted colloids. It is crucial, however, that the huddle structure is continuously reorganized to give each penguin a chance to spend sufficient time inside the huddle, compared with time spent on the periphery. Here we show that Emperor penguins move collectively in a highly coordinated manner to ensure mobility while at the same time keeping the huddle packed. Every 30-60 seconds, all penguins make small steps that travel as a wave through the entire huddle. Over time, these small movements lead to large-scale reorganization of the huddle. Our data show that the dynamics of penguin huddling is governed by intermittency and approach to kinetic arrest in striking analogy with inert non-equilibrium systems, including soft glasses and colloids.     

Penguins use the two-voice system to recognize each other

The sound-producing structure in birds is the syrinx, which is usually a two-part organ located at the junction of the bronchi. As each branch of the syrinx produces sound independently, many birds have two acoustic sources. Thirty years ago, we had anatomical, physiological and acoustical evidence of this two-voice phenomenon but no function was known. In songbirds, often these two voices with their respective harmonics are not activated simultaneously but they are obvious in large penguins and generate a beat pattern which varies between individuals. The emperor penguin breeds during the Antarctic winter, incubating and carrying its egg on its feet. Without the topographical cue of a nest, birds identify each other only by vocal means when switching duties during incubation or chick rearing. To test whether the two-voice system contains the identity code, we played back the modified call of their mate to both adults and also the modified call of their parents to chicks. Both the adults and the chicks replied to controls (two voices) but not to modified signals (one voice being experimentally suppressed). Our experiments demonstrate that the beat generated by the interaction of these two fundamental frequencies conveys information about individual identity and also propagates well through obstacles, being robust to sound degradation through the medium of bodies in a penguin colony. The two-voice structure is also clear in the call of other birds such as the king penguin, another non-nesting species, but not in the 14 other nesting penguins. We concluded that the two-voice phenomenon functions as an individual recognition system in species using few if any landmarks to meet. In penguins, this coding process, increasing the call complexity and resisting sound degradation, has evolved in parallel with the loss of territoriality.     

How do weather conditions affect the huddling behaviour of emperor penguins?

Huddling allows emperor penguins to conserve energy and survive their long winter fast while facing harsh climatic conditions. Here we report the first investigation into the effects of changes in wind speed and ambient temperature on different components of penguin huddling behaviour. We attached light and temperature recorders to male emperor penguins at the Pointe Géologie colony, Antarctica, which recorded huddling events. We then compared the frequency, duration, occurrence and intensity of huddling bouts, with ambient air temperatures and wind speeds. Huddling occurrence increased with lower ambient temperatures and higher wind speeds, whereas huddling intensity was mainly enhanced by lower ambient temperatures. Moreover, huddling group movements were linked to wind direction and its global density to lower ambient temperatures. Hence, emperor penguins complex huddling behaviour was modulated differently depending on these two parameters. Weather conditions may then affect emperor penguins ability to save energy and survive their winter fast.     

Hidden keys to survival: the type,density, pattern and functional role of emperor penguin body feathers

Antarctic penguins survive some of the harshest conditions on the planet. Emperor penguins breed on the sea ice where temperatures drop below −40°C and forage in −1.8°C waters. Their ability to maintain 38°C body temperature in these conditions is due in large part to their feathered coat. Penguins have been reported to have the highest contour feather density of any bird, and both filoplumes and plumules (downy feathers) are reported absent in penguins. In studies modelling the heat transfer properties and the potential biomimetic applications of penguin plumage design, the insulative properties of penguin plumage have been attributed to the single afterfeather attached to contour feathers. This attribution of the afterfeather as the sole insulation component has been repeated in subsequent studies. Our results demonstrate the presence of both plumules and filoplumes in the penguin body plumage. The downy plumules are four times denser than afterfeathers and play a key, previously overlooked role in penguin survival. Our study also does not support the report that emperor penguins have the highest contour feather density.     

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

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

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

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

Responses of king penguin Aptenodytes patagonicus adults and chicks to two food‐related odours

Increasing evidence suggests that penguins are sensitive to dimethyl sulphide (DMS), a scented airborne compound that a variety of marine animals use to find productive areas of the ocean where prey is likely to be found. Here we present data showing that king penguins Aptenodytes patagonicus are also sensitive to DMS. We deployed DMS on a lake near a king penguin colony at Ratmanoff beach in the Kerguelen archipelago. We also presented DMS to ‘sleeping’ adults on the beach. On the lake, penguins responded to the DMS deployments by swimming more, while on the beach, penguins twitched their heads and woke up more for the DMS than for the control presentations. Interestingly, penguins did not respond to cod liver oil deployments on the lake; mirroring at‐sea studies of other penguins. Although at‐sea studies are needed to confirm that king penguins use DMS as a surface cue that informs them of productivity under the water, this study is an important first step in understanding how these birds locate prey over significant distances.     

Penguin heat-retention structures evolved in a greenhouse Earth

Penguins (Sphenisciformes) inhabit some of the most extreme environments on Earth. The 60+ Myr fossil record of penguins spans an interval that witnessed dramatic shifts in Cenozoic ocean temperatures and currents, indicating a long interplay between penguin evolution and environmental change. Perhaps the most celebrated example is the successful Late Cenozoic invasion of glacial environments by crown clade penguins. A major adaptation that allows penguins to forage in cold water is the humeral arterial plexus, a vascular counter-current heat exchanger (CCHE) that limits heat loss through the flipper. Fossil evidence reveals that the humeral plexus arose at least 49 Ma during a 'Greenhouse Earth' interval. The evolution of the CCHE is therefore unrelated to global cooling or development of polar ice sheets, but probably represents an adaptation to foraging in subsurface waters at temperate latitudes. As global climate cooled, the CCHE was key to invasion of thermally more demanding environments associated with Antarctic ice sheets.     

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.     

Fibre types in primary ‘flight’ muscles of the African Penguin (Spheniscus demersus)

The African penguin (Spheniscus demersus) is an endangered seabird that resides on the temperate southern coast of Africa. Like all penguins it is flightless, instead using its specialized wings for underwater locomotion termed ‘aquatic flight’. While musculature and locomotion of the large Antarctic penguins have been well studied, smaller penguins show different biochemical and behavioural adaptations to their habitats. We used histochemical and immunohistochemical methods to characterize fibre type composition of the African penguin primary flight muscles, the pectoralis and supracoracoideus. We hypothesized the pectoralis would contain predominantly fast oxidative–glycolytic (FOG) fibres, with mainly aerobic subtypes. As the supracoracoideus and pectoralis both power thrust, we further hypothesized these muscles would have a similar fibre type complement. Our results supported these hypotheses, also showing an unexpected slow fibre population in the deep parts of pectoralis and supracoracoideus. The latissimus dorsi was also examined as it may contribute to thrust generation during aquatic flight, and in other avian species typically contains definitive fibre types. Unique among birds studied to date, the African penguin anterior latissimus dorsi was found to consist mainly of fast fibres. This study shows the African penguin has specialized flight musculature distinct from other birds, including large Antarctic penguins.     

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.     

Ticks Ixodes uriae and the breeding performance of a colonial seabird, king penguin Aptenodytes patagonicus

High densities of penguins in their colonies and the continuous use of these sites over consecutive reproductive periods increase the risk of development of tick populations. We have studied the effects of tick parasitism by Ixodes uriae in a colony of king penguins Aptenodytes patagonicus at Possession Island during three breeding seasons. We investigated the prevalence and periods of tick infestation during the one-year breeding cycle of penguins. The effects of tick parasitism on penguin breeding performance were assessed from photographs of the colony and with an automatic penguin identification system. We compared two groups of penguins carrying individual subcutaneous electronic tags, one group breeding in an infested area and the other in a non-infested area. Tick feeding activity was coincident with the periods when adult penguins stayed ashore for six days or more, i.e. during the incubating period. This duration corresponds to the duration of a tick meal on the host. The level of infestation varied between years. Penguins showed a lower incubating success in infested areas during a year of high infestation. In an infested area, individuals seen with ticks had a lower breeding success in rearing a one-year old chick than those seen without ticks.     

Fossil penguin (Aves: Sphenisciformes) cranial material from the Eocene of Seymour Island (Antarctica)

Our knowledge of the cranial morphology of early penguins remains poor, particularly for Paleogene taxa. This paper describes a partial penguin skull and additional isolated cranial elements from the Eocene La Meseta Formation of Seymour Island, Antarctica. These specimens cannot be assigned to named taxa at present, but there is a strong possibility they belong to La Meseta penguins known only from postcranial elements. The skull shares extensive dorsal development of the temporal fossae with extant and fossil Spheniscus and the fossil penguins Paraptenodytes and Marplesornis, indicating the adductor complex was powerful in early penguins. Partial mandibles belonging to a much larger penguin are similar to Paraptenodytes and differ from all living penguins in the lack of a hooked medial process of the articular and the presence of a foramen anterior to the mandibular cotyles. Given the rarity of penguin cranial remains, these specimens provide important new insight into early penguin evolution.     

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.     

Genetic evidence for three species of rockhopper penguins, Eudyptes chrysocome

The taxonomy of rockhopper penguins, Eudyptes chrysocome (Forster 1781), is contentious. Some authorities recognise three subspecies based on morphological differences and geographical separation of breeding populations while others suggest that morphological differences support classifying rockhopper penguins as two distinct species. The taxonomy of rockhopper penguins is of more than academic interest as breeding colonies worldwide have declined markedly in size since the 1930s and rockhopper penguins are currently listed as vulnerable by the IUCN. We compared the genetic distances between three mitochondrial gene regions from the three putative rockhopper penguin subspecies with the distances between various penguin sister species to clarify the taxonomy and systematics of rockhopper penguins. Genetic distances between the rockhopper penguin taxa, relative to other closely related penguin species, support reclassifying the three rockhopper penguin subspecies as species. Reclassification of rockhopper penguins as three species could result in their conservation status being upgraded from vulnerable to endangered.     

Sex Identification of Four Penguin Species Using Locus‐Specific PCR

Traditional methods for sex identification are not applicable to sexually monomorphic species, leading to difficulties in the management of their breeding programs. To identify sex in sexually monomorphic birds, molecular methods have been established. Two established primer pairs (2550F/2718R and p8/p2) amplify the CHD1 gene region from both the Z and W chromosomes. Here, we evaluated the use of these primers for sex identification in four sexually monomorphic penguin species: king penguins (Aptenodytes patagonicus), rockhopper penguins (Eudyptes chrysocome), gentoo penguins (Pygoscelis papua), and Magellanic penguins (Spheniscus magellanicus). For all species except rockhopper penguins, primer pair 2550F/2718R resulted in two distinct CHD1Z and CHD1W PCR bands, allowing for sex identification. For rockhopper penguins, only primer pair p8/p2 yielded different CHD1Z and CHD1W bands, which were faint and similar in size making them difficult to distinguish. As a result, we designed a new primer pair (PL/PR) that efficiently determined the gender of individuals from all four penguin species. Sequencing of the PCR products confirmed that they were from the CHD1 gene region. Primer pair PL/PR can be evaluated for use in sexing other penguin species, which will be crucial for the management of new penguin breeding programs. Zoo Biol 32:257–261, 2013. © 2012 Wiley Periodicals, Inc.     

Heat loss and site-dependent fecundity in chinstrap penguins (Pygoscelis antarctica)

We study the effects of heat loss and nest site quality on fecundity in a chinstrap penguin (Pygoscelis antarctica) colony on Deception Island, Antarctica. During the austral summers of 1990–1991, 1993–1994, and 1995–1996, 441 randomly selected nests were analyzed. Penguins breeding in the center of subcolonies hatched significantly earlier and had larger broods and than those nesting near or at the edge of the subcolonies. These differences, however, were significantly affected by interaction between year and nest location, being highly significant in colder years, when peripheral nests produce fewer young. Analysis of the rate of heat loss showed that penguins breeding at the edge of subcolonies lost heat twice as rapidly as those breeding in the interior of the subcolony. In a re-occupation experiment, evacuated center nests were re-occupied almost ten times as rapidly as edge nests. An analysis of mean fecundity in the period 1991–1996 and mean wind chill suggested that most of the variability in fecundity among years was related to differences in the rate of heat loss. Subcolonies tend to be as circular as possible, thereby decreasing the proportion of edge nests as the size of subcolony increases. Our results support the site-dependent fecundity hypothesis.     

Vision of the Humboldt penguin (Spheniscus humboldti) in air and water

Refractive states measured by retinoscopy and photorefraction indicate that the eyes of the Humboldt penguin, Spheniscus humboldti, are approximately emmetropic in air and water. Extensive myopia in air, as predicted by earlier authors and by a recent anatomical study, is non-existent. Photorefractive measurements of the refractive state, in water, of the Humboldt penguin indicate that it can accommodate sufficiently to make up the loss of the refractive power of the cornea. The cornea of the Humboldt penguin is flattened relative to the overall size of the eye. In all these respects (corneal flattening, and accommodation in air and water) the eyes of Humboldt penguins are like those of gentoo, (Pygoscelis papua), rockhopper (Eudyptes crestatus), Magellanic (Spheniscus magellanicus), and king penguins (Aptenodytes patagonica).     

Foraging behaviour of the little penguin, Eudyptula minor: initial results and assessment of instrument effect

We investigated the foraging behaviour of little penguins using a new archival electronic activity recorder which simultaneously measures speed and depth against time. We present the first integrated data of foraging behaviour of two little penguins, from which we were able to distinguish between several types of travelling and foraging behaviours. The little penguins foraged mainly within 15 m of the surface and travelled at speeds between 8 and 9 km h^-1. Using attachments ranging between 1.4 and 11.8% penguin cross-sectional area (0.1 and 6% penguin mass), and isotopic water, we also assessed the effects of carrying devices while foraging. Both water influx and metabolic rates were significantly lower in penguins carrying devices, compared to penguins foraging without devices attached. Even the relatively small attachments resulted in a decreased foraging efficiency and we suggest that there is no simple or fixed relationship between size of device and the effect on the bearer.