Population trends and reproductive success at a frequently visited penguin colony on the western Antarctic Peninsula

Petermann Island (65°10′S, 64°10′W), one of the Antarctic Peninsula’s most frequently visited locations, is at the epicenter of a rapid shift in which an Adélie penguin dominated fauna is becoming gentoo penguin dominated. Over the course of five seasons, the breeding productivity of Adélie and gentoo penguins breeding at Petermann Island were monitored to identify drivers of this rapid community change. The impact of tourist visitation on breeding success was also investigated. Consistent with larger trends in this region, the Adélie penguin population decreased by 29% and the gentoo penguin population increased by 27% between the 2003/2004 and 2007/2008 seasons. Reproductive success among Adélie penguins ranged from 1.09 to 1.32 crèched chicks/nest, which was higher than or comparable to other sites and is an unlikely explanation for the precipitous decline of Adélie penguins at Petermann Island. Whereas gentoo penguin reproductive success was lowest in colonies frequently visited by tourists, Adélie penguin colonies frequently visited by tourists had higher reproductive success than those visited only occasionally. These results are placed in the context of other studies on reproductive success and the impact of tourist visitation on breeding colonies of Adélie and gentoo penguins.     

In stark contrast to widespread declines along the Scotia Arc,a survey of the South Sandwich Islands finds a robust seabird community

The South Sandwich Islands, in the South Atlantic Ocean, are a major biological hot spot for penguins and other seabirds, but their remoteness and challenging coastlines preclude regular biological censuses. Here we report on an extensive survey of the South Sandwich Islands, the first since the late 1990s, which was completed through a combination of direct counting, GPS mapping, and interpretation of high-resolution commercial satellite imagery. We find that the South Sandwich Islands host nearly half of the world’s Chinstrap Penguin (Pygoscelis antarctica) population (1.3 million breeding pairs), as well as c. 95,000 breeding pairs of Macaroni Penguins (Eudyptes chrysolophus), and several thousand breeding pairs of Gentoo Penguins (Pygoscelis papua). Despite being at the northern edge of their breeding range, we found an unexpectedly large (≥125,000 breeding pairs) population of Adélie Penguins (Pygoscelis adeliae). Additionally, we report that nearly 1900 pairs of Southern Giant Petrels (Macronectes giganteus) breed in the South Sandwich Islands, 4 % of the global population, almost all of which are found on Candlemas Island. We find that the South Sandwich Islands have not experienced the same changes in penguin abundance and distribution as the rest of the Scotia Arc and associated portions of the western Antarctic Peninsula. This discovery adds important context to the larger conversation regarding changes to penguin populations in the Southern Ocean.     

Plasticity in foraging behaviour and diet buffers effects of inter-annual environmental differences on chick growth and survival in southern rockhopper penguins <Emphasis Type="Italic">Eudyptes chrysocome chrysocome</Emphasis>

In marine ecosystems, primary productivity and consequently food availability for higher trophic levels are often strongly affected by the water temperature. Thus, differences in sea surface temperatures (SST) may lead to differences in the diet composition of predators, but this link is still unknown in many species. By combining GPS tracking and dive analyses on chick-rearing southern rockhopper penguins (Eudyptes chrysocome chrysocome) with stable isotope analyses and monitoring of chick growth rates and chick survival, we here attempted a comprehensive assessment of the effects of inter-annual environmental variability as indicated by SST and chlorophyll a (reflecting primary productivity) data. Inter-annual differences in environmental variables around our study colony on New Island, Falkland/Malvinas Islands, contradicted the general expectation, with higher chlorophyll a concentrations coinciding with higher spring SST in 2010/2011 compared to 2009/2010. Penguins foraged further away from the colony during guard and crèche in 2010/2011 compared to 2009/2010, while performing deeper dives in 2009/2010. Stable isotope mixing models suggested a crustacean-dominated chick diet in 2009/2010, compared to a mixture of squid and fish in 2010/2011. These differences in foraging behaviour and diet, however, had no consequences for chick growth rates or chick survival and thus had no apparent effect on population trajectories. Potentially, environmental conditions in both years could still be seen as favourable compared to other years and breeding sites, enabling the parental birds to buffer the environmental differences by plastic foraging behaviour.     

Kleptoparasitism in foraging gentoo penguins <Emphasis Type="Italic">Pygoscelis papua</Emphasis>

An ongoing challenge in marine top predator research is understanding processes that are responsible for patterns that are often derived from land-based access to individuals. The use of animal-borne camera loggers, however, is proving useful in this regard as researchers can directly observe species and habitat-associated interactions. During an ongoing study investigating the foraging ecology of gentoo penguins (Pygoscelis papua) at the Falkland Islands, we observed the first attempted underwater intraspecific kleptoparasitism event for a penguin. This was revealed through a bird deployed with an animal-borne camera logger. Although unsuccessful in its attempt, the new reported behaviour highlights a novel interaction and further demonstrates the value of cameras in better understanding the ecology of marine vertebrates.     

A comprehensive review of the phenology of <Emphasis Type="Italic">Pygoscelis</Emphasis> penguins

Phenology, the study of stages within the life cycles of plants and animals, has served as a proxy for weather and climate throughout human history, but has only recently become its own field of environmental science. Phenological constraints are particularly demanding in avian species because of the necessity of matching chick provisioning with high food abundance, while allocating time for migration. Within avian species, seabird phenology is of particular interest because many seabird species exhibit colonial breeding behaviour. Penguins, representing roughly 90 % of the biomass in the Southern Ocean, are well studied in the context of population dynamics, prey abundance, and phenology. Here I review the annual cycles of Pygoscelis penguins, a genus including gentoo (Pygoscelis papua), chinstrap (Pygoscelis antarctica), and Adélie (Pygoscelis adeliae) penguins, to better understand what is known about their phenology, what causes known changes, and how their phenology influences fitness. Major differences exist between species, particularly in relation to winter migration, incubation shifts, and the timing of breeding. Even with the numerous studies examining phenology in Pygoscelis penguins, large gaps in our understanding of plasticity in the annual cycle remain. In particular, certain phases are neglected because they are logistically difficult to record or have erroneously been ignored. In addition, temporally, large gaps exist in our understanding of phenology, where studies have not been updated in over 20 years at a particular field site. Because phenology does vary greatly between years, depending on the colony, when possible, researchers should strive to update phenological records by recording the dates of phases each year.     

Size-related dietary changes observed in the squid<Emphasis Type="Italic"> Moroteuthis ingens</Emphasis> at the Falkland Islands: stomach contents and fatty-acid analyses

Dietary composition of the onychoteuthid squid Moroteuthis ingens at the Falkland Islands was related to predator size, as shown by stomach contents and fatty-acid analyses. Comparisons were made between two size classes of squid: those of mantle length <200 mm and those of mantle length >200 mm. Smaller squid had frequently consumed crustaceans and cephalopods; fish were of secondary importance. Larger squid consumed mostly fish and moderate amounts of cephalopods, but had rarely consumed crustaceans. These findings were supported by comparisons drawn between digestive-gland fatty-acid profiles and the fatty-acid profiles of potential prey species. Fatty-acid analyses indicated that the crustaceans Euphausia lucens, Munida gregaria and Themisto gaudichaudii were important prey items of smaller squid, whereas stomach content and fatty-acid analyses indicated that Gymnoscopelus nicholsi of around 100 mm standard length represented much of the fish prey of larger squid.     

Aspects of the breeding biology of the southern rockhopper penguin <Emphasis Type="Italic">Eudyptes c. chrysocome</Emphasis> and new consideration on the intrinsic capacity of the A-egg

The rockhopper penguins Eudyptes chrysocome have recently been split into the northern E. moseleyi and the southern E. chrysocome rockhopper penguin. It is therefore crucial to have a comprehensive understanding of the biology of each species in order to develop appropriate conservation measures. We investigated the breeding biology of the southern rockhopper on New Island, in the western part of the Falklands Islands, by following the breeding attempt of 160 pairs during the 2006/2007 season and examining the effect of lay time and colony habitat on breeding success. Specifically, we compared survival and growth parameters between A- and B-eggs and chicks from non-manipulated and artificially manipulated nests to investigate why southern rockhopper penguins in the Falkland Islands are more able to fledge an A-egg (first laid) than conspecifics elsewhere. Breeding was highly synchronous, with no significant difference in the breeding success between early and late breeders or between pairs breeding in different habitats. We demonstrate for the first time that the A-egg produced by the southern rockhopper penguin has, when alone, the same theoretical intrinsic potential to lead to a fledged chick as the B-egg. In contrast, the hatching success and survival of the B-chick was similar when alone or in a two-egg clutch.     

Spatial and temporal variation in the diet of the gentoo penguin (<Emphasis Type="Italic">Pygoscelis papua</Emphasis>) at Kerguelen Islands

Benthic divers are dependent on local resources and may therefore adopt different foraging strategies to cope with their energetic requirements in varying situations. We investigated the diet of gentoo penguins (Pygoscelis papua) at Kerguelen Islands, comparing its spatial and temporal variations with the general prey distribution. The study was conducted at four sites over 2 years. In total, 212 stomach contents were collected over the entire breeding season. The diet was composed mainly of neritic fish and crustaceans, with important spatial and seasonal variations. Fish dominated the diet at localities facing the open sea (from 38.0% to 94.6% by mass), whereas crustaceans dominated at the more protected site (84.3% by mass). Fish were more abundant in the winter diet and Euphausia vallentini, the major crustacean species, was more abundant in the summer diet. No inter-year variations were detected. These results are consistent with local prey availability, and highlight the large plasticity of the gentoo penguin diet and foraging behaviour.     

Population trends, breeding success and diet composition of gentoo Pygoscelis papua, magellanic Spheniscus magellanicus and rockhopper Eudyptes chrysocome penguins in the Falkland Islands. A review

Data on population size, breeding success and diet composition of gentoo (Pygoscelis papua), magellanic (Spheniscus magellanicus) and rockhopper (Eudyptes chrysocome) penguins, collected as part of the Falkland Island Seabird Monitoring Programme from 1986/1987 to 1998/1999, were analysed with regard to spatial and temporal variation, as well as potential interaction with local commercial fisheries. No significant population trends were detectable, mainly because of the short time-series and large spatial and inter-annual variation in the number of breeding pairs in the colonies monitored. However, the breeding success of all three penguin species has improved slightly over the last few years, indicating a potential for increasing populations in the near future. During the breeding season, all three penguin species preyed opportunistically on a mixture of fish, squid and crustaceans. Diet composition too showed a high degree of spatial and temporal variation. However, in all three penguin species studied, squid gradually disappeared from the diet over successive years, to be replaced by fish. Coincidentally, the commercial catches of the squid species Loligo gahi in Falkland Islands waters decreased and the by-catch of nototheniid fish increased. All three penguin species compete directly with the commercial fishing fleet for L. gahi; however, there may also be competition for Patagonian toothfish (Dissostichus eleginoides), hake (Merluccius sp.) and southern blue whiting (Micromesistius australis), because juveniles of these species were found regularly in penguin diets.     

Winter diet and foraging range of gentoo penguins (<Emphasis Type="Italic">Pygoscelis papua</Emphasis>) from Kidney Cove,Falkland Islands

The winter diet and foraging range of gentoo penguins, Pygoscelis papua, were studied at Kidney Cove, Falkland Islands. The mean wet mass of the 56 stomach-content samples collected from May to October was 32.7ᇃ.4 g. The diet consisted generally of cephalopods, crustaceans and fish, as well as two other molluscs. Among the six cephalopod species identified, the commercially fished Patagonian squid, Loligo gahi, had the highest abundance and was also the main prey by reconstituted mass (53% of the total reconstituted mass). Lobster krill, Munida gregaria, one of five species of crustaceans, was the most abundant prey species by frequency of occurrence and by number (68% and 60%, respectively). Rock cod, Patagonotothen ramsayi, accounted for the majority of the fish diet with 34% of the total reconstituted mass. Most prey species identified in the winter diet were also abundant diet components during the breeding season. However, the known biology of the prey species and their rate of digestion indicated that, in winter, adults may forage further offshore than during the breeding season. This assumption was supported by the results obtained from two birds satellite-tracked during the study period. Both birds remained mainly in inshore waters and returned frequently ashore, but one penguin foraged up to 276 km from the coast. The differences in the foraging behaviour of the two birds were reflected in significant differences with regard to time spent underwater, distance travelled per day and calculated travelling speed. Furthermore, the progress of cohorts of L. gahi over the winter is consistent with results from life-cycle studies in this region and suggests that birds have been foraging in the feeding grounds of L. gahi.     

What is out there: diversity in feeding of gentoo penguins (Pygoscelis papua) around the Falkland Islands (Southwest Atlantic)

During austral spring 2000, the diet of Gentoo penguins, Pygoscelis papua, was studied and compared to prey availability, assessed by trawl and plankton surveys, in waters adjacent to a number of colonies along the coast of West Falkland. There was good agreement in size and abundance of key prey species in the penguin diet and from the plankton and trawl surveys. The diet of birds breeding adjacent to shallow sheltered waters was dominated by demersal species, in particular crustaceans, Munida spp. whilst birds breeding adjacent to deeper exposed waters generally preyed upon pelagic species of fish, Falkland herring Sprattus fuegensis and squid, Moroteuthis ingens. Gentoo penguins generally foraged at depths of up to 30 m, relatively close to the coast, and illustrated a certain degree of selection of prey species where possible, thus providing evidence that they are not fully opportunistic predators.     

Breeding energetics and food requirements of gentoo penguins (Pygoscelis papua) at Heard and Macquarie Islands

The food and energy requirements of breeding gentoo penguins ( Pygoscelis papua ) were studied at Heard and Macquaric Islands by means of isotope turnover techniques.
The food consumption rates of chicks were measured at various stages of growth, providing estimates of the total food provided by adults to rear a chick to fledging.
The energy expenditures of attending adults were also measured at different stages of chick-rearing, allowing the total energy costs associated with breeding to be established for a pair of adults and at the population level on both islands. We estimate the total annual energy budget of a 6·2 kg breeding gentoo penguin to be 1517 MJ which is equivalent to the consumption of 292 kg prey.     

Penguins, fur seals, and fishing: prey requirements and potential competition in the South Shetland Islands, Antarctica

Antarctic and sub-Antarctic seabirds, marine mammals, and human fisheries concentrate their foraging efforts on a single species, Antarctic krill (Euphausiasuperba). Because these predators may have a significant effect on krill abundance, we estimated the energy and prey requirements of Adelie (Pygoscelisadeliae), chinstrap (Pygoscelisantarctica), and gentoo (Pygoscelispapua) penguins and female Antarctic fur seals (Arctocephalusgazella) breeding on the South Shetland Islands, Antarctica and compared these estimates with catch statistics from the Antarctic krill fishery. Published data on field metabolic rate, population size, diet, prey energy content, and metabolic efficiency were used to estimate prey requirements of these breeding, adult, land-based predators and their dependent offspring. Due to their large population size, chinstrap penguins were the most significant krill predators during the period examined, consuming an estimated 7.8 × 10⁸ kg krill, followed by Adelie penguins (3.1 × 10⁷ kg), gentoo penguins (1.2 × 10⁷ kg), and Antarctic fur seals (3.6 × 10⁶ kg). Total consumption of all land-based predators on the South Shetland Islands was estimated at 8.3 × 10⁸ kg krill. The commercial krill fishery harvest in the South Shetland Island region (1.0 × 10⁸ kg) was approximately 12% of this. Commercial harvest coincides seasonally and spatially with peak penguin and fur seal prey demands, and may affect prey availability to penguins and fur seals. This differs from the conclusions of Ichii et al. who asserted that the potential for competition between South Shetland predators and the commercial krill fishery is low. Received: 26 August 1997 / Accepted: 16 December 1997     

The diet of the Antarctic fur seal <Emphasis Type="Italic">Arctocephalus gazella</Emphasis> at the South Orkney Islands in ten consecutive years

To improve the knowledge on the Antarctic fur seal foraging behavior, evaluate whether changes in its diet are associated with changes in prey availability, and evaluate whether fisheries had negative impacts on Antarctic fur seal populations, a total of 1359 scats of non-breeding males were collected in ten consecutive sampling periods between 1994 and 2003 at Laurie Island, South Orkney Islands. Antarctic krill was the most numerous prey throughout the sampling period followed in importance by fish prey. Antarctic krill also predominated by reconstructed mass, except during 1994 and 1998 when penguins were the most important prey, and during 1995 when fish dominated. Among fish, demersal-benthic species (mainly Gobionotothen gibberifrons, Chaenocephalus aceratus and Chionodraco rastrospinosus) dominated the diet except in 1997, 1998, 1999 and 2002 when myctophids (mainly Gymnoscopelus nicholsi and Electrona antarctica) were the most important fish prey. In seasons when fish dominated the diet, the diet was primarily comprised of demersal species. Although our results are in overall agreement with previous studies, the contribution to the diet of the main preys changed throughout the sampling period, probably in parallel with changes in prey abundance. According to historical results on fish consumption, the fisheries carried out in the past have had negative impacts on seal populations from the South Orkney Islands.     

Morphometric and genetic evidence for four species of gentoo penguin

Gentoo penguins (Pygoscelis papua) are found across the Southern Ocean with a circumpolar distribution and notable genetic and morphological variation across their geographic range. Whether this geographic variation represents species‐level diversity has yet to be investigated in an integrative taxonomic framework. Here, we show that four distinct populations of gentoo penguins (Iles Kerguelen, Falkland Islands, South Georgia, and South Shetlands/Western Antarctic Peninsula) are genetically and morphologically distinct from one another. We present here a revised taxonomic treatment including formal nomenclatural changes. We suggest the designation of four species of gentoo penguin: P. papua in the Falkland Islands, P. ellsworthi in the South Shetland Islands/Western Antarctic Peninsula, P. taeniata in Iles Kerguelen, and a new gentoo species P. poncetii, described herein, in South Georgia. These findings of cryptic diversity add to many other such findings across the avian tree of life in recent years. Our results further highlight the importance of reassessing species boundaries as methodological advances are made, particularly for taxa of conservation concern. We recommend reassessment by the IUCN of each species, particularly P. taeniata and P. poncetii, which both show evidence of decline.     

Leukocyte counts in different populations of Antarctic <Emphasis Type="Italic">Pygoscelid</Emphasis> penguins along the Antarctic Peninsula

The Antarctic Peninsula is one of the areas where the climate is changing at the fastest pace, having several effects on the populations of pygoscelid penguins. Few studies have analysed the variation in immune parameters of antarctic birds in a geographical context; thus, analyses of geographical differences in the immune components of wild pygoscelid penguins are still scarce. Leukocyte counts in birds provide information on their immunity and physiological stress. The objective of this study was to analyse the leukocyte counts in penguins of the genus Pygoscelis (gentoo, Adélie and chinstrap penguins), covering sites along the South Shetland Islands and some islands on the west coast of the Antarctic Peninsula. Our results revealed differences in the number of heterophils and eosinophils and in the heterophil/lymphocyte ratio in the northeastern populations of gentoo and Adélie penguins as compared to the rest of the colonies studied. The results contribute to better understanding of the variations in physiological parameters of penguins related to a geographical context.     

Stable isotopes in southern rockhopper penguins: foraging areas and sexual differences in the non-breeding period

Southern rockhopper penguins (Eudyptes chrysocome chrysocome) have experienced severe population declines across their distribution area, potentially in response to bottom-up effects following elevated sea surface temperatures, changes in the food web and prey availability. We conducted stable isotope analysis to compare trophic levels and distribution patterns in the non-breeding period over three consecutive years, and between males and females, using egg membranes, blood cells and feathers of parent birds. Tissues representing the non-breeding season had lower ??¹³C values than prey sampled around the Falklands and red blood cells from breeding rockhopper penguins. In contrast, ??¹⁵N values were higher in red blood cells from the end of winter compared to those from the breeding season and compared to feathers. This indicated that rockhopper penguins left the Falkland Island area in the non-breeding season and foraged either around Burdwood Bank further south, or over the Patagonian Shelf. In winter, only males took more prey of higher trophic level than females. Inter-annual differences in isotopic values partly correlated with sea surface temperatures. However, as prey isotope samples were collected only in 1?year, inter-annual differences in penguin isotopic values may result from different foraging sites, different prey choice or different isotopic baseline values. Our study highlights the potential for stable isotope analyses to detect seasonal and gender-specific differences in foraging areas and trophic levels, while stressing the need for more sampling of isotopic baseline data.     

Gentoo penguin Pygoscelis papua diet as an indicator of planktonic availability in the Kerguelen Islands

Since penguins rely on the main planktonic resources of the Southern Ocean, knowledge of their diet may be used for monitoring these resources During winter and spring of 1987 and 1989, we investigated the composition of the diet of gentoo penguins. Pygoscelis papua, in relation to changes in the availability of two prey species, Euphausia vallentini and Themisto gaudichaudii, sampled during plankton surveys in the Kerguelen Islands. The comparison between plankton surveys and diet analysis was performed on samples taken 2–4 km from the studied colonies. Data on the abundance of Zooplankton derived from penguins' diet matched closely those from net hauls during a year of high plankton availability (1987). On the other hand, a weaker correspondence was found during a year of restricted availability (1989). The mean sizes of amphipods caught by penguins and net hauls were very similar but the size distribution showed comparatively fewer small and large individuals in net-hauls than in penguin stomachs. Detailed studies on the feeding range and foraging effort of penguins are therefore needed in the near future to validate the potential of penguin diet as a measure of plankton abundance.     

Disentangling reproductive biology of the Patagonian toothfish <Emphasis Type="Italic">Dissostichus eleginoides</Emphasis>: skipped vs obligatory annual spawning,foraging migration vs residential life style

Patagonian toothfish Dissostichus eleginoides Smitt 1898, is an important commercially targeted Notothenioid species in South-Atlantic waters. In this study we aimed to clarify several aspects of reproductive biology of Patagonian toothfish in the Falkland Islands waters. Histological examination of female gonads indicates that with the beginning of maturation females maintain at least two populations of oocytes, suggesting that toothfish requires up to two years for oocytes development. Females become mature at an average size of 79.1 cm indicating a decrease of first maturity size if toothfish females in the Falkland Island waters. The majority of females spawn at the size from 101 to 130 cm total length. Distribution of reproductive phases shows an increase of females at developing stage in December and March prior to the spawning peaks in May and August respectively. However, the majority of the toothfish population consist of non-spawning individuals remaining in regressing phase (55.8 to 85.6%) including the spawning period. The skip-spawning for toothfish has been defined as reabsorbing non-reproductive and resting types. The abbreviation of oocytes development in the gonads was observed from 1 to 22.1% of females which omitted the spawning season. Most likely females which remain in the spawning area have the opportunity to spawn more often, whereas females which undergo foraging migration toward Northern parts of the Falklands waters return to the spawning ground less often. Females remain Northern area longer to accumulate necessary amount of energy. This hypnotise is supported by the presence of females in immature, developing and regressing phase throughout surrounding Falkland Islands waters. Presence of post-spawning females in regressing stage throughout the Falklands waters suggest that toothfish may undertake irregular spawning/foraging migration when favourable for spawning condition occur.     

Low MHC variation in the endangered Galápagos penguin (<Emphasis Type="Italic">Spheniscus mendiculus</Emphasis>)

The major histocompatibility complex (MHC) is one of the most polymorphic regions of the genome, likely due to balancing selection acting to maintain alleles over time. Lack of MHC variability has been attributed to factors such as genetic drift in small populations and relaxed selection pressure. The Galápagos penguin (Spheniscus mendiculus), endemic to the Galápagos Islands, is the only penguin that occurs on the equator. It relies upon cold, nutrient-rich upwellings and experiences severe population declines when ocean temperatures rise during El Niño events. These bottlenecks, occurring in an already small population, have likely resulted in reduced genetic diversity in this species. In this study, we used MHC class II exon 2 sequence data from a DRB1-like gene to characterize the amount of genetic variation at the MHC in 30 Galápagos penguins, as well as one Magellanic penguin (S. magellanicus) and two king penguins (Aptenodytes patagonicus), and compared it to that in five other penguin species for which published data exist. We found that the Galápagos penguin had the lowest MHC diversity (as measured by number of polymorphic sites and average divergence among alleles) of the eight penguin species studied. A phylogenetic analysis showed that Galápagos penguin MHC sequences are most closely related to Humboldt penguin (Spheniscus humboldti) sequences, its putative sister species based on other loci. An excess of non-synonymous mutations and a pattern of trans-specific evolution in the neighbor-joining tree suggest that selection is acting on the penguin MHC.