Positive selection over the mitochondrial genome and its role in the diversification of gentoo penguins in response to adaptation in isolation.

Although mitochondrial DNA has been widely used in phylogeography, evidence has emerged that factors such as climate, food availability, and environmental pressures that produce high levels of stress can exert a strong influence on mitochondrial genomes, to the point of promoting the persistence of certain genotypes in order to compensate for the metabolic requirements of the local environment. As recently discovered, the gentoo penguins (Pygoscelis papua) comprise four highly divergent lineages across their distribution spanning the Antarctic and sub-Antarctic regions. Gentoo penguins therefore represent a suitable animal model to study adaptive processes across divergent environments. Based on 62 mitogenomes that we obtained from nine locations spanning all four gentoo penguin lineages, we demonstrated lineage-specific nucleotide substitutions for various genes, but only lineage-specific amino acid replacements for the ND1 and ND5 protein-coding genes. Purifying selection (dN/dS < 1) is the main driving force in the protein-coding genes that shape the diversity of mitogenomes in gentoo penguins. Positive selection (dN/dS > 1) was mostly present in codons of the Complex I (NADH genes), supported by two different codon-based methods at the ND1 and ND4 in the most divergent lineages, the eastern gentoo penguin from Crozet and Marion Islands and the southern gentoo penguin from Antarctica respectively. Additionally, ND5 and ATP6 were under selection in the branches of the phylogeny involving all gentoo penguins except the eastern lineage. Our study suggests that local adaptation of gentoo penguins has emerged as a response to environmental variability promoting the fixation of mitochondrial haplotypes in a non-random manner. Mitogenome adaptation is thus likely to have been associated with gentoo penguin diversification across the Southern Ocean and to have promoted their survival in extreme environments such as Antarctica. Such selective processes on the mitochondrial genome may also be responsible for the discordance detected between nuclear- and mitochondrial-based phylogenies of gentoo penguin lineages.

Phase transitions in huddling emperor penguins.

Emperor penguins (Aptenodytes forsteri) are highly adapted to the harsh conditions of the Antarctic winter: they are able to fast for up to 134 days during breeding. To conserve energy, emperor penguins form tight groups (huddles), which is key for their reproductive success. The effect of different meteorological factors on the huddling behaviour, however, is not well understood. Using time-lapse image recordings of an emperor penguin colony, we show that huddling can be described as a phase transition from a fluid to a solid state. We use the colony density as order parameter, and an apparent temperature that is perceived by the penguins as the thermodynamic variable. We approximate the apparent temperature as a linear combination of four meteorological parameters: ambient temperature, wind speed, global radiation and relative humidity. We find a wind chill factor of -2.9 °C/(ms -1), a humidity chill factor of -0.5°C/% rel. humidity, and a solar radiation heating factor of 0.3 °C//(Wm 2). In the absence of wind, humidity and solar radiation, the phase transition temperature (50% huddling probability) is -48.2°C for the investigated time period (May 2014). We propose that higher phase transition temperatures indicate a shrinking thermal insulation and thus can serve as a proxy for lower energy reserves of the colony, integrating pre-breeding foraging success at sea and energy expenditure at land due to environmental conditions. As current global change is predicted to have strong detrimental effects on emperor penguins within the next decades, our approach may thus contribute towards an urgently needed long-term monitoring system for assessing colony health.

Adjustments of gastric pH, motility and temperature during long-term preservation of stomach contents in free-ranging incubating king penguins.

Male king penguins are able to store undigested food in their stomach for up to 3 weeks during their incubation fast, which evidently implies some modification of their digestive process. Using small electronic recorders, we studied the change in gastric pH, motility and temperature during the first week of food storage. The pH could be maintained at values as high as 6 throughout the incubation fast, a pH that is unfavourable for avian gastric proteinase activity. Gastric motility was never completely inhibited but could be markedly reduced. Stomach temperature was maintained at around 38 degrees C. The fact that stomach temperature of incubating birds did not show a daily rhythmic fluctuation as seen in non-breeding birds could be due to temperature constraints on embryo development. Thus the present study demonstrates substantial adjustments of pH and gastric motility in incubating king penguins, which may contribute to the inhibition of digestive gastric processes.

Long-term effects of flipper bands on penguins.

Changes in seabird populations, and particularly of penguins, offer a unique opportunity for investigating the impact of fisheries and climatic variations on marine resources. Such investigations often require large-scale banding to identify individual birds, but the significance of the data relies on the assumption that no bias is introduced in this type of long-term monitoring. After 5 years of using an automated system of identification of king penguins implanted with electronic tags (100 adult king penguins were implanted with a transponder tag, 50 of which were also flipper banded), we can report that banding results in later arrival at the colony for courtship in some years, lower breeding probability and lower chick production. We also found that the survival rate of unbanded, electronically tagged king penguin chicks after 2-3 years is approximately twice as large as that reported in the literature for banded chicks.