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).     

Eye structure and foraging in King Penguins Aptenodytes patagonicus

Anterior eye structure and retinal visual fields were determined in King Penguins Aptenodytes patagonicus using keratometry and an ophthalmoscopic reflex technique. The cornea is relatively flat (radius 32.9 mm) and hence of low refractive power (10.2 dioptres in air) and this may be correlated with the amphibious nature of penguin vision. The large size of the eye and of the fully dilated pupil may be correlated with activity at low light levels. In air, the binocular field is long (vertical extent 180̀) and narrow (maximum width 29̀), with the bill placed approximately centrally—a topography found in a range of bird species which employ visual guidance of bill position when foraging. Upon immersion in water, the optical power of the cornea is abolished, with the effect that the monocular fields decrease and binocularity is lost. King Penguins have a pupil type which has not hitherto been recorded in birds. In daylight it contracts to a square-shaped pinhole but dilates to a large circular aperture in darkness. This change alters retinal illumination by 300-fold (2.5 log₁₀ units). When diving, this permits the retina to be pre-adapted to the low ambient light levels that the birds encounter upon reaching mesopelagic depths. These penguins also forage at depths where ambient light levels, even during the day, can fall below the equivalent of terrestrial starlight. Under these conditions, the birds must rely upon the detection of light from the photophores of their prey. In this they are aided by their absolutely large pupil size and broad cyclopean visual field.     

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.     

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.     

Rush and grab strategies in foraging marine endotherms: the case for haste in penguins

The speed at which air-breathing marine predators that forage by diving should swim is likely to depend on a variety of factors that differ substantially from those relevant in animals for which access to oxygen is unlimited. We used loggers attached to free-living penguins to examine the speed at which three species swam during periods searching for prey and compared this to their speeds during actual prey pursuit. All penguin species appeared to travel at similar speeds around 2 m/s during normal commuting between the surface and feeding depths, which accords closely with minimum costs of transport. However, Adélie penguins, Pygoscelis adeliae, slowed down to feed, Magellanic penguins, Spheniscus magellanicus, speeded up and king penguins, Aptenodytes patagonicus, travelled at a variety of speeds, although mean speed did not change from normal commuting. Since energy expenditure, and therefore oxygen usage, in swimming animals increases with the cube of the speed, we hypothesized that prey escape speed (a function of prey size) and prey density would prove critical in determining optimum pursuit speeds in predators. Simple models of this type help explain why it is that some penguin species apparently benefit by increasing speed to capture prey while others benefit by decreasing speed.     

Energetic cost of foraging in free-diving emperor penguins.

Hypothesizing that emperor penguins (Aptenodytes forsteri) would have higher daily energy expenditures when foraging for their food than when being hand-fed and that the increased expenditure could represent their foraging cost, we measured field metabolic rates (FMR; using doubly labeled water) over 4-d periods when 10 penguins either foraged under sea ice or were not allowed to dive but were fed fish by hand. Surprisingly, penguins did not have higher rates of energy expenditure when they dove and captured their own food than when they did not forage but were given food. Analysis of time-activity and energy budgets indicated that FMR was about 1.7 x BMR (basal metabolic rate) during the 12 h d(-1) that penguins were lying on sea ice. During the remaining 12 h d(-1), which we termed their "foraging period" of the day, the birds were alert and active (standing, preening, walking, and either free diving or being hand-fed), and their FMR was about 4.1 x BMR. This is the lowest cost of foraging estimated to date among the eight penguin species studied. The calculated aerobic diving limit (ADL(C)), determined with the foraging period metabolic rate of 4.1 x BMR and known O(2) stores, was only 2.6 min, which is far less than the 6-min ADL previously measured with postdive lactate analyses in emperors diving under similar conditions. This indicates that calculating ADL(C) from an at-sea or foraging-period metabolic rate in penguins is not appropriate. The relatively low foraging cost for emperor penguins contributes to their relatively low total daily FMR (2.9 x BMR). The allometric relationship for FMR in eight penguin species, including the smallest and largest living representatives, is kJ d(-1)=1,185 kg(0.705).     

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.     

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.     

Eye structure and amphibious foraging in albatrosses

Anterior eye structure and retinal visual fields were determined in grey-headed and black-browed albatrosses, Diomedea melanophris and D. chrysostoma (Procellariiformes, Diomedeidae), using keratometry and an ophthalmoscopic reflex technique. Results for the two species were very similar and indicate that the eyes are of an amphibious optical design suggesting that albatross vision is well suited to the visual pursuit of active prey both on and below the ocean surface. The corneas are relatively flat (radius ca. 14.5 mm) and hence of low absolute refractive power (ca. 23 dioptres). In air the binocular fields are relatively long (vertical extent ca. 70 degrees) and narrow (maximum width in the plane of the optic axes 26–32 degrees), a topography found in a range of bird species that employ visual guidance of bill position when foraging. The cyclopean fields measure approximately 270 degrees in the horizontal plane, but there is a 60 degrees blind sector above the head owing to the positioning of the eyes below the protruding supraorbital ridges. Upon immersion the monocular fields decrease in width such that the binocular fields are abolished. Anterior eye structure, and visual field topography in both air and water, show marked similarity with those of the Humboldt penguin.     

Genetic diversity in populations of Gentoo penguins (Pygoscelis papua)

RAPD analysis was used to examine the extent of genetic polymorphism in two populations of Gentoo penguin (Pygoscelis papua) from Antarctic Islands (Petermann and Livingston). The chosen two of three 10 mer oligonucleotide primers accordingly to preliminary results showed different levels of polymorphism in Gentoo penguins at Petermann Island (from 23.53 to 42.86%) and Livingston Island (from 52.94 to 57.14%). Nei's similarity coefficients were in range from 0.5606 (when Gentoo genome profiles were compared with RAPD profiles of two related penguin species: Pygoscelis adeliae (Adelie) and Pygoscelis antarctica (Chinstrep)) to 0.9281 among observed Gentoo penguin populations. Nei's distances values ranged from 0.0746 to 0.5787 among the populations and species. The obtained results will be used for further estimation of genetic diversity of Gentoo penguins and determination of their taxonomic status.     

The refractive significance of the nictitating membrane of the bird eye

Summary Previous authors have suggested that in certain aquatic birds a transparent nictitating membrane of high refractive index could compensate for the refractive loss of the cornea in water. Measurements of refractive indices, refractive state and curvature carried out on a number of aquatic birds indicate that the nictitating membrane does not have a refractive function; the refractive indices of the cornea and nictitating membrane are very similar (approximately 1.37), the nictitating membrane does not alter the refractive state of the eye, and its curvature, when it is in place in front of the globe, is virtually the same as that of the cornea.This research was supported by grants from the National Research Council of Canada and the Canadian National Sportsmen's Fund. The assistance of Mote Marine Laboratory, The Mount Desert Island Biological Laboratory, the Niska Wildlife Foundation and the Felicadades Wildlife Foundation is gratefully acknowledged. The herring gulls and black guillemots were provided by Dr. W.B. Kinter. In addition, the authors are grateful for the assistance provided by Drs. J.L. Lincer and D. Peakall and Mr. R. Orleib.     

A history of shifting fortunes for African penguins

Africa is home today to only a single breeding species of penguin, Spheniscus demersus (black‐footed penguin), which is endangered with extinction. Spheniscus demersus has been the only breeding species of penguin to share African coastlines with humans over the last 400 000 years. Interestingly, African penguin diversity was substantially higher before the evolution of archaic humans. The fossil record indicates that a diverse assemblage of penguin species inhabited the southern African coasts for much of the Neogene. Previous excavations have identified four distinct species in Early Pliocene coastal marine deposits. Here we extend this pattern of high diversity and report the oldest record of penguins from Africa. Seventeen penguin specimens were identified from the Saldanha Steel locality, revealing the presence of at least four distinct species in South Africa during the Miocene. The largest of these species reached the size of the extant Aptenodytes patagonicus (king penguin), whereas the smallest was approximately the size of the smallest extant penguin Eudyptula minor (little blue penguin). Recovery of Miocene penguin remains is in accordance with earlier predictions of multiple pre‐Pliocene colonizations of Africa and supports a higher level of ecological diversity amongst African penguins in the past. © 2013 The Linnean Society of London     

Modeling the marine resources consumed in raising a king penguin chick: an energetics approach

Accurate estimates of penguin energetics would represent an important contribution to our understanding of the trophodynamics of the Southern Ocean ecosystem and our ability to predict effects of environmental change on these species. We used the heart rate-rate of oxygen consumption technique to estimate rate of energy expenditure in adult king penguins raising a chick, in combination with data from the literature on changes in adult mass, chick energy requirements, and prey energy density. Our model estimated a variety of energetic costs and quantities of prey consumption related to raising a king penguin chick during the austral summer. The total energy requirements of a king penguin chick at the Crozet Archipelago from hatching until reaching a mass of 8 kg 90 d later is 271 MJ, representing the consumption of 38.4 kg of myctophid fish. A successfully breeding male requires 0.78 kg d(-1) of fish during the entirety of the incubation period and 1.14 kg d(-1) during the subsequent 90 d of chick rearing. Assuming the same energy requirements for females, the estimated 580,000 pairs of king penguins that breed successfully at Crozet each year, together with their chicks, consume a total of around 190,000 tons of fish during the incubation and summer rearing periods combined. If, due to depletion of fish stocks, the diet of breeders and chicks during the summer becomes identical to the typical diet of adults during the austral winter, the mass of prey required by both adults and chicks combined (where the chick still reaches 8 kg after 90 d) would increase by more than 25%.     

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.     

Concentration of trace metals in excrements of two species of penguins from different locations of the Antarctic Peninsula

The presence of metals in Antarctica is becoming an issue that needs to be more investigated as human presence is increasing in the region, especially in the Antarctic Peninsula, where most of the polar animals, scientific bases and tourists are concentrated. Penguins are endemic species of Antarctica situated at the top of food web and are useful sentinels of pollution. To improve data of trace metal contamination in the Antarctic Peninsula region, concentrations of Cd, Pb, As, Cu and Zn in fresh excrement of penguins were determined by atomic absorption. Samples were collected in several locations during the 2011/2012 austral summer: eight sites (O’Higgins Station, Yankee Harbour, Mikkelsen Harbor, Danco Island, Gonzalez Videla Base, Yelcho Station and Brown Station) from gentoo penguin (Pygoscelis papua) colonies and four locations (Hydrurga Rocks, Cape Shirreff, Narebski Point and Kopaitic Island) from chinstrap penguin (Pygoscelis antarctica) colonies. Data showed that gentoo penguin colonies had higher levels of metals (μg g^?1 dw) like Pb, Cu and Zn at locations such as O’Higgins (2.89 for Pb, 199.95 for Cu and 379.99 for Zn) and Gonzalez Videla (2.74 for Pb and 222.51 for Cu). Levels of Cd in excreta of chinstrap penguins were higher at Narebski Point (3.13 μg g^?1). The levels of Pb, As, Cd, Cu and Zn were similar or even higher to those reported in excreta of the same species by previous studies at Antarctica. Data suggest that metals ingested by these two penguin species feeding in the sea end up in terrestrial ecosystems.     

Morphological aspects of the heart of the northern rockhopper penguin (Eudyptes chrysocome moseleyi ): possible implication in diving behavior and ecology?

We compared the heart morphology of the small, deep-diving northern rockhopper penguin to the hearts of small, shallow-diving and large, deep-diving penguin species. The rockhopper penguin had a heart larger than expected for its body mass, and its heart weight/body weight was significantly greater than in the larger Adélie penguin. We found the rockhopper's right ventricle weight/heart weight to be significantly greater than this relationship in both the larger chinstrap and Adélie penguins. The relationship of the right to left ventricular weights in the rockhopper heart is not different to that of the large, deepest-diving emperor penguin. A larger heart in the rockhopper penguin might be related to its diving behavior and ecology if it contributes to diving efficiency during foraging by increasing lung perfusion during surface recovery. This would lead to decreased surface time. Accepted: 20 May 2000     

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.     

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.     

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.