Activity Time Budget during Foraging Trips of Emperor Penguins

We developed an automated method using depth and one axis of body acceleration data recorded by animal-borne data loggers to identify activities of penguins over long-term deployments. Using this technique, we evaluated the activity time budget of emperor penguins (n = 10) both in water and on sea ice during foraging trips in chick-rearing season. During the foraging trips, emperor penguins alternated dive bouts (4.8±4.5 h) and rest periods on sea ice (2.5±2.3 h). After recorder deployment and release near the colony, the birds spent 17.9±8.4% of their time traveling until they reached the ice edge. Once at the ice edge, they stayed there more than 4 hours before the first dive. After the first dive, the mean proportions of time spent on the ice and in water were 30.8±7.4% and 69.2±7.4%, respectively. When in the water, they spent 67.9±3.1% of time making dives deeper than 5 m. Dive activity had no typical diurnal pattern for individual birds. While in the water between dives, the birds had short resting periods (1.2±1.7 min) and periods of swimming at depths shallower than 5 m (0.25±0.38 min). When the birds were on the ice, they primarily used time for resting (90.3±4.1% of time) and spent only 9.7±4.1% of time traveling. Thus, it appears that, during foraging trips at sea, emperor penguins traveled during dives >5 m depth, and that sea ice was primarily used for resting. Sea ice probably provides refuge from natural predators such as leopard seals. We also suggest that 24 hours of sunlight and the cycling of dive bouts with short rest periods on sea ice allow emperor penguins to dive continuously throughout the day during foraging trips to sea.     

The diet of Emperor Penguins Aptenodytes forsteri in Adelie Land, Antarctica

This first quantitative study of the diet of Emperor penguins is based on 29 stomach contents collected with a water off-loading method in Adelie Land. The Emperor is largely ichthyophagous (65% by number and 95% by weight) and feeds extensively on small nototheniids (97% of the fish are 40–125 mm in overall length). These results and data on meal size and feeding frequency of the chick suggest that Emperors are off-shore foraging birds offering little competition for food with other sea-birds or mammals.     

Habitat Use by Weddell Seals and Emperor Penguins Foraging in the Ross Sea, Antarctica

The only apex predators that live year-round at high latitudesof the Ross Sea are the Weddell seal and emperor penguin. Theseasonal distribution, foraging depths, and diet of these twospecies appear to overlap. What makes it possible for emperorpenguins and Weddell seals to co-exist at high latitude throughoutthe winter when other marine tetrapods apparently cannot? Bothspecies have similar adaptations for exploitation of the deep-waterhabitat, forage on the same species, and routinely make longand deep dives. Yet, despite these similarities, there is probablylittle trophic overlap between the adults of both species dueto geographical and seasonal differences in habitat use. Forexample, during the winter months while female emperor penguinsare ranging widely in the pack ice, adult seals are foragingand fattening for the upcoming summer fast, literally beneaththe feet of the male penguins. However, there is more extensiveoverlap between juvenile seals and adult penguins, and shiftsin prey abundance and/or distribution would likely affect thesetwo groups similarly. In contrast, juvenile penguins appearto avoid inter- and intra- specific competition by leaving theRoss Sea once they molt.     

Comparison of Color and Body Condition Between Early and Late Breeding King Penguins

Early breeding is associated with greater reproductive success in many species. In king penguins, Aptenodytes patagonicus , laying extends for 6 mo. Early breeders may fledge a single chick at best, but late breeders virtually never fledge a chick. For early and late breeders, we compared colored ornaments known to be important in mate choice: yellow–orange feathers of the breast and auricular areas, and an ultraviolet and yellow–orange beak spot. Our purpose was to discern differences between males and females in this highly sexually monomorphic species, as well as to discern whether colored ornaments are more important for the more successful early breeders (aspects of color were hue, chroma, and brightness). For this, we weighed and measured 130 penguins. Early males had greater reflectance of ultraviolet color from the beak spot than did early females and late breeders of both sexes, and the early males were heavier and in better condition than late breeding males or females. Late breeding females were the yellowest in breast hue, a trait that has been linked to immunocompetence. Within pairs, males and females were significantly correlated in body mass, but only early in the breeding season. We concluded that early in the breeding season when reproductive success was greatest, potential mates were not only more similar in body mass, but also that females may have chosen males that had brighter beak spots and were in better body condition.     

The post-moult diet of Emperor Penguins (Aptenodytes forsteri) in the eastern Weddell Sea,Antarctica

The diet of emperor penguins Aptenodytes forsteri was studied during late austral summer at Drescher Inlet, eastern Weddell Sea, Antarctica. Antarctic krill Euphausia superba was a major component of the food, accounting for 75% of all prey items. Emperor penguins appear to feed on krill during shallow dives under the fast sea ice. Fish, mainly nototheniids, accounted for less than 20% by number of all prey. An evaluation of the main prey types in terms of mass indicated, however, that fish represented up to 75% approximately of prey mass. Feeding experiments were performed on captive penguins and showed that squid beaks can accumulate for up to 3 weeks within the stomach without any clear signs of erosion. The lack of cephalopod soft parts in the samples makes it likely that all squid beaks were derived from animals captured some time previously. Squid seems to be a very minor dietary component of emperor penguins at the Drescher Inlet.     

Responses of Emperor Penguins (Aptenodytes forsteri) to encounters with ecotourists while commuting to and from their breeding colony

Emperor penguins (Aptenodytes forsteri) were studied at the Snow Hill breeding colony in November 2006 to determine the effect of people on penguins traveling between the colony and the sea to forage. We tested the null hypothesis that the presence and number of people had no effect on the trajectory of movement or the number and duration of pauses. The distances at which penguins noticed people (mean 35.6 m), changed direction (mean 22.8 m), and the number and duration of pauses increased significantly with increases in the number of tourists in their path, which explained more than 50% of the variance. Undisturbed penguins usually tobogganed on their ventral surface over the ice. When penguins noticed people, they usually stood up and often called. In 10 min observation periods, penguins traveling more than 200 m from people paused an average of <1 min vs. 3.8 min for those passing near people, increasing the energetic cost of commuting. After passing people, penguins rarely stopped. Penguins response to people varied by time of day; later in the day they responded less quickly, changed directions when closer to people, stopped for less time, and passed by people closer than they did earlier in the day. We suggest that the effect of ecotourists on traveling penguins can be partly mitigated by having people walk in small, tight-knit groups, by having people stop moving whenever traveling penguins are within about 25 m to allow the penguins to choose the direction of their passage, and by keeping the visitor pathway separate from the penguin paths insofar as possible.     

Penguins bled by vampires

Penguins have a variety of ennemies both on on land and at sea. This is the first account of vampire bats (Desmodus rotundus) preying on juvenile Humboldt Penguins (Spheniscus humboldti). Because of the nightly drain of blood by vampires, and because vampires transmit diseases from one prey species to another, this form of predation may be an important mortality factor.

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Zusammenfassung Pinguine haben eine Vielzahl von Feinden, sowohl im Wasser als auch an Land. Dies ist der erste Hinweis darauf, daß Vampirfledermäuse (Desmodus rotundus) sich vom Blut von Humboldtpinguinküken (Spheniscus humboldti) ernähren. Wegen der nächtlichen Blutverluste durch Vampirfledermäuse, und weil diese Blutsauger Krankheitserreger zwischen ihren Wirten übertragen können, könnte diese Form des Parasitismus einen wichtigen Mortalitätsfaktor darstellen.

     

Homosexual Mating Displays in Penguins

More than 50 yr ago, field studies recorded the same‐sex pairs (and trios) of penguins displaying to each other during the mating season, using behavior patterns typical of heterosexual mating displays. Such observations led to a hypothesis that due to a lack of sex recognition pairing occurs at random with respect to sex, an idea countered by the argument that sex recognition is highly accurate. No quantification of same‐sex mating displays has tested the frequency of such displays in penguins or tested the hypothesis of random display partners with respect to sex. During their mating season, we studied displaying and paired king penguins, Apenodytes patagonicus, at Kerguelen Island and sexed them using a DNA marker, to quantify any occurrence of this behavior. Indeed, same‐sex courtship displays were common (28.3% of 53 displaying pairs), the great majority of which were between males. Some homosexually displaying males eventually paired with females, but such males were significantly slower in heterosexual pairing than males that did not display homosexually. In two extraordinary cases, same‐sex pairs learned each other’s calls, an essential step in the pairing process. The frequency of such pairs was much lower than among displaying couples, significantly so for males. Finally, the frequency of homosexually displaying pairs was significantly lower than expected from random assortment of displaying birds, for both males and females. We examined possible explanations for same‐sex display and its biological significance. A population sex‐ratio bias in favor of males and high concentration of male sex hormones may help to explain non‐reproductive homosexually displaying pairs.     

Fat King Penguins Are Less Steady on Their Feet

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.     

At-Sea Associations in Foraging Little Penguins

Prey distribution, patch size, and the presence of conspecifics are important factors influencing a predator’s feeding tactics, including the decision to feed individually or socially. Little is known about group behaviour in seabirds as they spend most of their lives in the marine environment where it is difficult to observe their foraging activities. In this study, we report on at-sea foraging associations of little penguins (Eudyptula minor) during the breeding season. Individuals could be categorised as (1) not associating; (2) associating when departing from and/or returning to the colony; or (3) at sea when travelling, diving or performing synchronised dives. Out of 84 separate foraging tracks, 58 (69.0%) involved associations with conspecifics. Furthermore, in a total of 39 (46.4%), individuals were found to dive during association and in 32 (38.1%), individuals were found to exhibit synchronous diving. These behaviours suggest little penguins forage in groups, could synchronise their underwater movements and potentially cooperate to concentrate their small schooling prey.     

Sexual Selection on a Coloured Ornament in King Penguins

Species in which the sexes equally exhibit colourful ornaments are an issue for evolutionary theory. Among several hypotheses, sexual selection for mutual mate choice and social selection for signals of behavioural dominance are most commonly supported. We examined the previously documented sex‐similar size of yellow‐orange ear patches in the king penguin, Aptenodytes patagonicus. This species is monogamous and pairs just before reproduction. Raising a chick requires considerable effort by both parents, as they alternate care of their single offspring with foraging at sea. The size of the ear patches appears to signal aggressive territoriality in the breeding colony for both sexes. However, experiments suggest that females prefer large patch size during mate choice, and males do not prefer this trait. We tested whether the size of the coloured ear patch was influenced by sexual selection for couples that had recently paired. We used analyses of covariance to compare the size of the ear patch to a measure of body size and then tested for the difference between males and females. Males were 6.2% larger in ear patch width and 7.7% larger in ear patch area than females, and the distance between the ear patches over the head was 7.5% smaller in males, with all differences highly significant. Consequently, sexual selection appears to favour larger ear patches in males, possibly because of an excess of males that promotes female choice. Social selection also appears to favour the evolutionary maintenance of ear patches of males, and thus both types of selection may contribute to enlarged ear patches.     

Mate switching and copulation behaviour in King Penguins

Extra‐pair paternity (EPP) in monogamous birds may result from either extra‐pair copulations (EPCs) or mate switching. In this study of King Penguins in South Georgia, we observed no EPCs at all, an effect of very efficient mate guarding. Onshore males fast and need not divert attention to foraging or to defending nest or territory, as this species has neither. However, we found that mate switching was common. On average 38% (range: 29%–56%; three years pooled) of the birds established pair bonds with at least one initial partner before switching to the partner they bred with (i.e. the “pair mate”). Of the observed copulations of 44 studied females, 22% were with initial partners and 78% with the pair mate. This and the high proportion of mate switching suggest that roughly 10% of the females could have received sperm from males other than the pair mate. The average copulation frequency was 0.026 h^?1, resulting in an estimated 8.2 copulations per clutch (which consists of one egg). That more copulations than necessary for fertilisation occur suggests that males try to protect paternity by sperm competition, and that this is a result of the potential for EPP due to mate switching in King Penguins. All observed copulations except one took place between days 13 and 5, with the peak 7.5 days prior to egg‐laying. The birds found their pair mates (often not the same as in the previous year) on average about 10 days before egg‐laying, and always established themselves at the outskirts of the colony about 8 days before egg‐laying. Thus, most copulations occurred around the time the birds joined the colony. We suggest that it is adaptive to obtain a breeding spot early, because the colony will grow and pairs joining later will protect the offspring. Additionally, we suggest that early copulation outside the colony is adaptive because of the risk of failing to fertilise the egg when copulating among aggressive neighbours inside the dense colony. Based on these two arguments we suggest a “safe place hypothesis” to explain the early copulation peak in King Penguins.     

Climate Change Increases Reproductive Failure in Magellanic 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.     

Avian pox in Magellanic Penguins (Spheniscus magellanicus)

Avian pox is an enveloped double-stranded DNA virus that is mechanically transmitted via arthropod vectors or mucosal membrane contact with infectious particles or birds. Magellanic Penguins (Spheniscus magellanicus) from two colonies (Punta Tombo and Cabo Dos Bahías) in Argentina showed sporadic, nonepidemic signs of avian pox during five and two of 29 breeding seasons (1982-2010), respectively. In Magellanic Penguins, avian pox expresses externally as wart-like lesions around the beak, flippers, cloaca, feet, and eyes. Fleas (Parapsyllus longicornis) are the most likely arthropod vectors at these colonies. Three chicks with cutaneous pox-like lesions were positive for Avipoxvirus and revealed phylogenetic proximity with an Avipoxvirus found in Black-browed Albatross (Thalassarche melanophrys) from the Falkland Islands in 1987. This proximity suggests a long-term circulation of seabird Avipoxviruses in the southwest Atlantic. Avian pox outbreaks in these colonies primarily affected chicks, often resulted in death, and were not associated with handling, rainfall, or temperature.     

Depth utilization by Penguins and Gentoo Penguin dive patterns

Summary Time-at-depth data, recorded by animal-attached miniature depth gauges, were examined in 8 Gentoo Penguins. Birds showed two depth utiliuation patterns, (1) decreasing time with increasing depth, which we interpret as due to bounce-diving in the pelagic zone and (2) constant time per depth interval down to specific depths where large time peaks were encountered. We interprete this as flat-bottomed diving in benthic-foraging birds. Multiple maximum depth data reported in the literature for various penguin species were analysed to try and reconstruct proportional time-at-depth. These results, together with real time-at-depth data, indicated that penguin depth utilization was strongly mass dependent with larger species spending a greater proportion of time in deeper water.

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Die Tauchmuster des Eselpinguins und die räumliche Ausnutzung der Wassersäule durch Pinguine

Zusammenfassung Miniatur-Tiefenmeßgeräte wurden 8 Eselpinguinen angeheftet und die gewonnenen Daten der Aufenthaltsdauer in verschiedenen Wassertiefen untersucht. Die Vögel zeigten zwei Muster der Tiefenausnutzung, (1) abnehmende Aufenthaltsdauer mit zunehmender Wassertiefe, das wir als Stoßtauchen im Pelagial interpretieren und (2) konstante Aufenthaltsdauer pro Tiefenintervall bis in bestimmte Tiefen, in denen lange Verweildauer registriert wurden. Letzteres interpretieren wir als Grundtauchen bei Vögeln, die am Meeresboden nach Nahrung suchen. Aus der Literatur wurden multiple Tiefenmaxima für verschiedene Pinguinarten entnommen und analysiert, um die Aufenthaltsdauer pro Tiefenintervall zu rekonstruieren. Diese Ergebnisse zeigen zusammen mit tatsächlichen Meßdaten, daß die Tiefenausnutzung der Pinguine stark massenabhängig ist, wobei größere Arten eine größeren Zeitanteil in tieferem Wasser verbringen.

     

Inter-Annual Variability of Fledgling Sex Ratio in King Penguins

As the number of breeding pairs depends on the adult sex ratio in a monogamous species with biparental care, investigating sex-ratio variability in natural populations is essential to understand population dynamics. Using 10 years of data (2000–2009) in a seasonally monogamous seabird, the king penguin (Aptenodytes patagonicus), we investigated the annual sex ratio at fledging, and the potential environmental causes for its variation. Over more than 4000 birds, the annual sex ratio at fledging was highly variable (ranging from 44.4% to 58.3% of males), and on average slightly biased towards males (51.6%). Yearly variation in sex-ratio bias was neither related to density within the colony, nor to global or local oceanographic conditions known to affect both the productivity and accessibility of penguin foraging areas. However, rising sea surface temperature coincided with an increase in fledging sex-ratio variability. Fledging sex ratio was also correlated with difference in body condition between male and female fledglings. When more males were produced in a given year, their body condition was higher (and reciprocally), suggesting that parents might adopt a sex-biased allocation strategy depending on yearly environmental conditions and/or that the effect of environmental parameters on chick condition and survival may be sex-dependent. The initial bias in sex ratio observed at the juvenile stage tended to return to 1∶1 equilibrium upon first breeding attempts, as would be expected from Fisher’s classic theory of offspring sex-ratio variation.     

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.