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Concept: Penguins


One of the notable features of penguin evolution is the occurrence of very large species in the early Cenozoic, whose body size greatly exceeded that of the largest extant penguins. Here we describe a new giant species from the late Paleocene of New Zealand that documents the very early evolution of large body size in penguins. Kumimanu biceae, n. gen. et sp. is larger than all other fossil penguins that have substantial skeletal portions preserved. Several plesiomorphic features place the new species outside a clade including all post-Paleocene giant penguins. It is phylogenetically separated from giant Eocene and Oligocene penguin species by various smaller taxa, which indicates multiple origins of giant size in penguin evolution. That a penguin rivaling the largest previously known species existed in the Paleocene suggests that gigantism in penguins arose shortly after these birds became flightless divers. Our study therefore strengthens previous suggestions that the absence of very large penguins today is likely due to the Oligo-Miocene radiation of marine mammals.

Concepts: Evolution, Bird, Mammal, Phylogenetics, Penguin, Dinosaur, Penguins, Flightless bird


Returning to the shore after a feeding sojourn at sea, king penguins often undertake a relatively long terrestrial journey to the breeding colony carrying a heavy, mostly frontal, accumulation of fat along with food in the stomach for chick-provisioning. There they must survive a fasting period of up to a month in duration, during which their complete reliance on endogenous energy stores results in a dramatic loss in body mass. Our aim was to determine if the king penguin’s walking gait changes with variations in body mass. We investigated this by walking king penguins on a treadmill while instrumented with an acceleration data logger. The stride frequency, dynamic body acceleration (DBA) and posture of fat (pre-fasting; 13.2 kg) and slim (post fasting; 11 kg) king penguins were assessed while they walked at the same speed (1.4km/h) on a treadmill. Paired statistical tests indicated no evidence for a difference in dynamic body acceleration or stride frequency between the two body masses however there was substantially less variability in both leaning angle and the leaning amplitude of the body when the birds were slimmer. Furthermore, there was some evidence that the slimmer birds exhibited a decrease in waddling amplitude. We suggest the increase in variability of both leaning angle and amplitude, as well as a possibly greater variability in the waddling amplitude, is likely to result from the frontal fat accumulation when the birds are heavier, which may move the centre of mass anteriorly, resulting in a less stable upright posture. This study is the first to use accelerometry to better understand the gait of a species within a specific ecological context: the considerable body mass change exhibited by king penguins.

Concepts: Mass, Bird, Classical mechanics, Penguin, Emperor Penguin, King Penguin, Walking, Penguins


Crested penguins (genus Eudyptes) have a peculiar hatching pattern, with the first-laid egg (A-egg) hatching after the second-laid egg (B-egg) and chicks from A-eggs typically having a much lower survival probability. Maternal yolk androgens have been suggested to contribute to the competitive superiority of the B-chick in southern rockhopper penguins Eudyptes chrysocome, given their important role in mediating sibling competition in other species. We therefore increased the yolk androgen levels in freshly-laid eggs and examined the consequences for sibling competition–via effects on embryonic developmental times, chick growth and early survival. We placed one androgen-treated egg and one control egg into each foster nest, matching them for mass, laying date and laying order. The androgen treatment did not significantly affect embryonic developmental times or chick measurements at hatching. However, elevated yolk androgen levels benefitted chick growth in interaction with the number of siblings in a brood. Chicks from androgen-treated eggs had faster growth in the presence of a sibling than chicks from control eggs. Under these circumstances they also had a higher survival probability. Thus maternal androgens appear to reinforce the observed hatching pattern, facilitating brood reduction. This contrasts to most previous studies in other species where yolk androgens have been shown to compensate for the negative consequences of delayed hatching within the brood hierarchy.

Concepts: Egg, Androgen, Rockhopper penguin, Western Rockhopper Penguin, Penguins


Climate change is causing more frequent and intense storms, and climate models predict this trend will continue, potentially affecting wildlife populations. Since 1960 the number of days with >20 mm of rain increased near Punta Tombo, Argentina. Between 1983 and 2010 we followed 3496 known-age Magellanic penguin (Spheniscus magellanicus) chicks at Punta Tombo to determine how weather impacted their survival. In two years, rain was the most common cause of death killing 50% and 43% of chicks. In 26 years starvation killed the most chicks. Starvation and predation were present in all years. Chicks died in storms in 13 of 28 years and in 16 of 233 storms. Storm mortality was additive; there was no relationship between the number of chicks killed in storms and the numbers that starved (P = 0.75) or that were eaten (P = 0.39). However, when more chicks died in storms, fewer chicks fledged (P = 0.05, R (2) = 0.14). More chicks died when rainfall was higher and air temperature lower. Most chicks died from storms when they were 9-23 days old; the oldest chick killed in a storm was 41 days old. Storms with heavier rainfall killed older chicks as well as more chicks. Chicks up to 70 days old were killed by heat. Burrow nests mitigated storm mortality (N = 1063). The age span of chicks in the colony at any given time increased because the synchrony of egg laying decreased since 1983, lengthening the time when chicks are vulnerable to storms. Climate change that increases the frequency and intensity of storms results in more reproductive failure of Magellanic penguins, a pattern likely to apply to many species breeding in the region. Climate variability has already lowered reproductive success of Magellanic penguins and is likely undermining the resilience of many other species.

Concepts: Precipitation, Climate, Weather, Climate change, Penguin, Penguins, Magellanic Penguin, Punta Tombo


Climate change is a major threat to global biodiversity. Antarctic ecosystems are no exception. Investigating past species responses to climatic events can distinguish natural from anthropogenic impacts. Climate change produces ‘winners’, species that benefit from these events and ‘losers’, species that decline or become extinct. Using molecular techniques, we assess the demographic history and population structure of Pygoscelis penguins in the Scotia Arc related to climate warming after the Last Glacial Maximum (LGM). All three pygoscelid penguins responded positively to post-LGM warming by expanding from glacial refugia, with those breeding at higher latitudes expanding most. Northern (Pygoscelis papua papua) and Southern (Pygoscelis papua ellsworthii) gentoo sub-species likely diverged during the LGM. Comparing historical responses with the literature on current trends, we see Southern gentoo penguins are responding to current warming as they did during post-LGM warming, expanding their range southwards. Conversely, Adélie and chinstrap penguins are experiencing a ‘reversal of fortunes’ as they are now declining in the Antarctic Peninsula, the opposite of their response to post-LGM warming. This suggests current climate warming has decoupled historic population responses in the Antarctic Peninsula, favoring generalist gentoo penguins as climate change ‘winners’, while Adélie and chinstrap penguins have become climate change ‘losers’.

Concepts: Climate, Climate change, Penguin, Antarctica, Global warming, Penguins, Pygoscelis, Gentoo Penguin


Group foraging provides predators with advantages in over-powering prey larger than themselves or in aggregating small prey for efficient exploitation. For group-living predatory species, cooperative hunting strategies provide inclusive fitness benefits. However, for colonial-breeding predators, the benefit pay-offs of group foraging are less clear due to the potential for intra-specific competition. We used animal-borne cameras to determine the prey types, hunting strategies, and success of little penguins (Eudyptula minor), a small, colonial breeding air-breathing marine predator that has recently been shown to display extensive at-sea foraging associations with conspecifics. Regardless of prey type, little penguins had a higher probability of associating with conspecifics when hunting prey that were aggregated than when prey were solitary. In addition, success was greater when individuals hunted schooling rather than solitary prey. Surprisingly, however, success on schooling prey was similar or greater when individuals hunted on their own than when with conspecifics. These findings suggest individuals may be trading-off the energetic gains of solitary hunting for an increased probability of detecting prey within a spatially and temporally variable prey field by associating with conspecifics.

Concepts: Predation, Hunting, Penguin, Optimal foraging theory, Penguins, Little Penguin, Eudyptula, White-flippered Penguin


In contrast to their terrestrial call, the offshore call of penguins during their foraging trips has been poorly studied due to the inaccessibility of the foraging site-the open ocean-to researchers. Here, we present the first description of the vocal behaviour of penguins in the open ocean and discuss the function of their vocal communication. We deployed an animal-borne camera on gentoo penguins (Pygoscelis papua) and recorded their foraging behaviour during chick guarding. From the video recordings, we collected 598 offshore calls from 10 individuals in two breeding seasons (2014-2015 and 2015-2016), and we analysed the acoustic characteristics and behavioural contexts of these calls, including diving patterns, group association events, and foraging behaviour. The offshore calls varied in their dominant frequency and length, and penguins produced calls of different lengths in succession. Group associations were observed within one minute following an offshore call in almost half of the instances (43.18%). Penguins undertook dives of shallower depths and shorter durations after producing an offshore call than those before producing an offshore call. Our findings show that penguins may use vocal communication in the ocean related with group association during foraging trips.

Concepts: Psychology, Penguin, Ocean, Penguins, Pygoscelis, Gentoo Penguin


The African Penguin (Spheniscus demersus) is a highly social and vocal seabird. However, currently available descriptions of the vocal repertoire of African Penguin are mostly limited to basic descriptions of calls. Here we provide, for the first time, a detailed description of the vocal behaviour of this species by collecting audio and video recordings from a large captive colony. We combine visual examinations of spectrograms with spectral and temporal acoustic analyses to determine vocal categories. Moreover, we used a principal component analysis, followed by signal classification with a discriminant function analysis, for statistical validation of the vocalisation types. In addition, we identified the behavioural contexts in which calls were uttered. The results show that four basic vocalisations can be found in the vocal repertoire of adult African Penguin, namely a contact call emitted by isolated birds, an agonistic call used in aggressive interactions, an ecstatic display song uttered by single birds, and a mutual display song vocalised by pairs, at their nests. Moreover, we identified two distinct vocalisations interpreted as begging calls by nesting chicks (begging peep) and unweaned juveniles (begging moan). Finally, we discussed the importance of specific acoustic parameters in classifying calls and the possible use of the source-filter theory of vocal production to study penguin vocalisations.

Concepts: Bird, Principal component analysis, Seabird, Penguins, Flightless birds, Magellanic Penguin, Humboldt Penguin, Spheniscus


Climate-driven sympatry may lead to competition for food resources between species. Rapid warming in the West Antarctic Peninsula (WAP) is coincident with increasing gentoo penguin and decreasing Adélie penguin populations, suggesting that competition for food may exacerbate the Adélie penguin decline. On fine scales, we tested for foraging competition between these species during the chick-rearing period by comparing their foraging behaviors with the distribution of their prey, Antarctic krill. We detected krill aggregations within the horizontal and vertical foraging ranges of Adélie and gentoo penguins, and found that krill selected for habitats that balance the need to consume food and avoid predation. In overlapping Adélie and gentoo penguin foraging areas, four gentoo penguins switched foraging behavior by foraging at deeper depths, a strategy which limits competition with Adélie penguins. This suggests that climate-driven sympatry does not necessarily result in competitive exclusion of Adélie penguins by gentoo penguins. Contrary to a recent theory, which suggests that increased competition for krill is one of the major drivers of Adélie penguin population declines, we suggest that declines in Adélie penguins along the WAP are more likely due to direct and indirect climate impacts on their life histories.

Concepts: Penguin, Antarctica, Penguins, Pygoscelis, Gentoo Penguin


The aim of this work is to contribute to the knowledge of gastrointestinal parasites of the Gentoo penguin (Pygoscelis papua) from 25 de Mayo/King George Island (South Shetlands, Antarctica). Gastrointestinal tracts of 37 fresh dead individuals (21 chicks, 10 juveniles, and 6 adults) were collected from December 2006 to February 2012 and examined for macroparasites. Four adult parasite species were found: one Cestoda species (Parorchites zederi), two Nematoda species (Stegophorus macronectes and Tetrameres wetzeli), and one Acanthocephalan (Corynosoma shackletoni). Two species of immature acanthocephalans, Corynosoma hamanni and Corynosoma bullosum, were found in a single host. This is the first record of Tetrameres wetzeli in Gentoo penguins. The low parasite richness observed could be related to the stenophagic and pelagic diet of this host species which feeds almost exclusively on krill.

Concepts: Penguin, Penguins, Pygoscelis, Gentoo Penguin