Krill-feeding behaviour of gentoo penguins as shown by animal-borne camera loggers

Animal-borne camera loggers were used to examine the patterns of prey encounter and feeding behaviour of gentoo penguins at King George Island, Antarctica. The still images from the camera loggers showed that the penguins encountered the swarms of krill for 25.5% (range: 8–38%) of their dives (>5 m) on average, during their foraging trips (mean duration of 5.4 h, n = 7 trips). They encountered krill swarms during the dives to 10–70 m depth, in pelagic as well as benthic habitats. In the benthic habitat, the penguins swam just above the sea floor and headed downward over a krill swarm, probably using the sea floor to assist them to feed on mobile swarms. The shallow coastal waters would be the important foraging habitat of gentoo penguins breeding in King George Island.     

Antarctic krill (Euphausia superba Dana) eat salps

 Feeding behaviour of Antarctic krill (Euphausia superba) on salps was observed in shipboard experiments during the 1994/1995 Kaiyo Maru Antarctic Ocean research cruise. The feeding rate was more than 0.5 salp/krill per day. When offered ethanol extracts of four prey types, salps, phytoplankton, krill and polychaetes, krill preferred the salp extracts. This evidence implies that the substances extracted from salps were most attractive to krill. These results might indicate a tight ecological relationship between krill and salps. Received: 24 May 1995/Accepted: 8 October 1995     

Food and feeding ecology of emperor penguins in the eastern Weddell Sea

Summary The diet of the emperor penguin Aptenodytes forsteri in the eastern Weddell Sea, Antarctica was studied during October and November 1986 by stomach content analysis. Emperor penguins fed mainly on Antarctic krill Euphausia superba, Antarctic silverfish Pleuragramma antarcticum and squid Psychroteuthis glacialis. Benthic prey was not found. The prey composition suggests two different feeding strategies, shallow dives exploring the rugged underside of sea ice where krill is taken, and deep dives when mesopelagic fish and squid are consumed. Chicks were fed on average every 1.44 days.     

Size and stage composition of age class 0 Antarctic krill (<Emphasis Type="Italic">Euphausia superba</Emphasis>) in the ice–water interface layer during winter/early spring

The condition and survival of Antarctic krill (Euphausia superba) strongly depends on sea ice conditions during winter. How krill utilize sea ice depends on several factors such as region and developmental stage. A comprehensive understanding of sea ice habitat use by krill, however, remains largely unknown. The aim of this study was to improve the understanding of the krill’s interaction with the sea ice habitat during winter/early spring by conducting large-scale sampling of the ice–water interface (0–2 m) and comparing the size and developmental stage composition of krill with the pelagic population (0–500 m). Results show that the population in the northern Weddell Sea consisted mainly of krill that were <1 year old (age class 0; AC0), and that it was comprised of multiple cohorts. Size per developmental stage differed spatially, indicating that the krill likely were advected from various origins. The size distribution of krill differed between the two depth strata sampled. Larval stages with a relatively small size (mean 7–8 mm) dominated the upper two metre layer of the water column, while larger larvae and AC0 juveniles (mean 14–15 mm) were proportionally more abundant in the 0- to 500-m stratum. Our results show that, as krill mature, their vertical distribution and utilization of the sea ice appear to change gradually. This could be the result of changes in physiology and/or behaviour, as, e.g., the krill’s energy demand and swimming capacity increase with size and age. The degree of sea ice association will have an effect on large-scale spatial distribution patterns of AC0 krill and on predictions of the consequences of sea ice decline on their survival over winter.     

Diatom-Specific Highly Branched Isoprenoids as Biomarkers in Antarctic Consumers

The structure, functioning and dynamics of polar marine ecosystems are strongly influenced by the extent of sea ice. Ice algae and pelagic phytoplankton represent the primary sources of nutrition for higher trophic-level organisms in seasonally ice-covered areas, but their relative contributions to polar marine consumers remain largely unexplored. Here, we investigated the potential of diatom-specific lipid markers and highly branched isoprenoids (HBIs) for estimating the importance of these two carbon pools in an Antarctic pelagic ecosystem. Using GC-MS analysis, we studied HBI biomarkers in key marine species over three years in Adélie Land, Antarctica: euphausiids (ice krill Euphausia crystallorophias and Antarctic krill E. superba), fish (bald notothens Pagothenia borchgrevinki and Antarctic silverfish Pleuragramma antarcticum) and seabirds (Adélie penguins Pygoscelis adeliae, snow petrels Pagodroma nivea and cape petrels Daption capense). This study provides the first evidence of the incorporation of HBI lipids in Antarctic pelagic consumers. Specifically, a di-unsaturated HBI (diene) of sea ice origin was more abundant in ice-associated species than in pelagic species, whereas a tri-unsaturated HBI (triene) of phytoplanktonic origin was more abundant in pelagic species than in ice-associated species. Moreover, the relative abundances of diene and triene in seabird tissues and eggs were higher during a year of good sea ice conditions than in a year of poor ice conditions. In turn, the higher contribution of ice algal derived organic matter to the diet of seabirds was related to earlier breeding and higher breeding success. HBI biomarkers are a promising tool for estimating the contribution of organic matter derived from ice algae in pelagic consumers from Antarctica.     

Seabird and fur seal responses to vertically migrating winter krill swarms in Antarctica

Summary The foraging behaviour of fur seals and two species of surface feeding seabirds was observed over swarms of vertically migrating krill along the Antarctic Peninsula in July 1987. Fur Seal haul out patterns were correlated with krill in the upper 30 m of the water column. Krill moved to the surface at night; seals subsequently foraged from 1400-0700 hours before returning to floes. Foraging was continuous through the night. Dive duration decreased as krill moved up to the surface; shorter dives may have been more successful than longer ones. It is possible that very deep dives, which occur early in a foraging bout, represent more of an attempt to assess krill depth and distribution rather than being a genuine foraging effort. Seabirds responded to the presence of a surface krill swarm by circling over it and foraging; krill at depths greater than 30 m elicited directional flight and low frequencies of prey capture attempts. Both Snow Petrels and Antarctic Terns preyed on krill, but each species approached the swarms from different habitats. Snow Petrels primarily overflew areas covered by ice; terns preferred open water. This suggested that prey encounters are essentially opportunistic, although the search for prey is limited to rather specific marine habitats. This feature may be important to our understanding of the factors that determine the pelagic distribution of seabirds.     

Potential Climate Change Effects on the Habitat of Antarctic Krill in the Weddell Quadrant of the Southern Ocean

Antarctic krill is a cold water species, an increasingly important fishery resource and a major prey item for many fish, birds and mammals in the Southern Ocean. The fishery and the summer foraging sites of many of these predators are concentrated between 0° and 90°W. Parts of this quadrant have experienced recent localised sea surface warming of up to 0.2°C per decade, and projections suggest that further widespread warming of 0.27° to 1.08°C will occur by the late 21^st century. We assessed the potential influence of this projected warming on Antarctic krill habitat with a statistical model that links growth to temperature and chlorophyll concentration. The results divide the quadrant into two zones: a band around the Antarctic Circumpolar Current in which habitat quality is particularly vulnerable to warming, and a southern area which is relatively insensitive. Our analysis suggests that the direct effects of warming could reduce the area of growth habitat by up to 20%. The reduction in growth habitat within the range of predators, such as Antarctic fur seals, that forage from breeding sites on South Georgia could be up to 55%, and the habitat’s ability to support Antarctic krill biomass production within this range could be reduced by up to 68%. Sensitivity analysis suggests that the effects of a 50% change in summer chlorophyll concentration could be more significant than the direct effects of warming. A reduction in primary production could lead to further habitat degradation but, even if chlorophyll increased by 50%, projected warming would still cause some degradation of the habitat accessible to predators. While there is considerable uncertainty in these projections, they suggest that future climate change could have a significant negative effect on Antarctic krill growth habitat and, consequently, on Southern Ocean biodiversity and ecosystem services.     

Ecological drivers of change at South Georgia: the krill surplus,or climate variability

Macaroni penguins Eudyptes chrysolophus are thought to be one of the most important mesopredators in the Southern Ocean having a greater impact on prey availability and abundance than any other seabird species. Their population centre has long been held to be South Georgia where populations were thought to comprise many million animals. Here we report the results of a recent census of the macaroni population at South Georgia undertaken using aerial survey methods. We report dramatic declines in numbers (～1.0 million breeding pairs) compared to numbers observed in the late 1970s (～5.4 million pairs), but show that these reductions have occurred principally at sites where numbers had previously been very large. During the breeding season, the main foraging grounds of birds from these sites overlap with the foraging grounds of Antarctic fur seals Arctocephalus gazella, a major competitor for their principal prey, Antarctic krill Euphausia superba. We suggest that the redistribution of the macaroni penguin population at South Georgia reflects the recent recovery of fur seal populations and thus the ongoing consequences of human intervention at South Georgia, a process which started more than 2 centuries previously. The implied resource competition and the observed population changes may also be exacerbated by recent reductions in Antarctic krill abundance which have been linked with reductions in seasonal sea ice following recent, rapid, regional warming in the Antarctic; however, the recovery of fur seal populations, and the ongoing recovery of krill‐eating whale populations argues that tropho‐dynamic interactions may be sufficient to explain the observed changes.     

Ecosystem Oceanography of Seabird Hotspots: Environmental Determinants and Relationship with Antarctic Krill Within an Important Fishing Ground

The discipline of ecosystem oceanography provides a framework for assessing the role of mesoscale physical processes on the formation and occurrence of biological hotspots. We used shipboard surveys over nine years to investigate environmental determinants of seabird hotspots near the Antarctic Peninsula, a region experiencing rapid climate change and an expanding krill fishery. We hypothesize that seabird hotspots are structured by mesoscale ocean conditions that reflect differences in prey distribution within oceanic and coastal waters. We used generalized additive models to quantify functional relationships of seabird hotspots with krill biomass, and a suite of remotely sensed environmental variables, such as eddy kinetic energy. The spatial organization, changes in intensity, and distribution shifts of seabird hotspots indicate different environmental drivers within coastal and oceanic domains and reflect the seasonal variability of the ecosystem. Our results indicate at least eight mesoscale hotspot zones that represent ecologically important areas where significant krill and predator biomass may be concentrated. Our ecosystem assessment of seabird hotspots identified critical foraging habitat and provided reference points to benefit research on estimating their trophic impacts on Antarctic ecosystems and potential effects from the krill fishery. Our approach is generally applicable to other pelagic ecosystems that are structured by hydrographic fronts and eddies, and containing schooling forage species shared by multiple wide-ranging predators. Furthermore, identification of biological hotspots is useful for the designation of marine protected areas most critical to potentially endangered wildlife and fisheries resources.     

A method for estimating colony sizes of Adélie penguins using remote sensing imagery

Adélie penguins (Pygoscelis adeliae) are important predators of krill (Euphausia spp.) and Antarctic silverfish (Pleuragramma antarctica) during summer, are a key indicator of the status of the Southern Ocean ecosystem, and are therefore a focal species for the Committee for the Conservation of Antarctic Marine Living Resources (CCAMLR) Ecosystem Monitoring Program. The ability to monitor the population size of species potentially affected by Southern Ocean fisheries, i.e., the Adélie penguin, is critical for effective management of those resources. However, for several reasons, direct estimates of population size are not possible in many locations around Antarctica. In this study, we combine high-resolution (0.6 m) satellite imagery with spectral analysis in a supervised classification to estimate the sizes of Adélie penguin breeding colonies along Victoria Land in the Ross Sea and on the Antarctic Peninsula. Using satellite images paired with concurrent ground counts, we fit a generalized linear mixed model with Poisson errors to predict the abundance of breeding pairs as a function of the area of current-year guano staining identified in the satellite imagery. Guano-covered area proved to be an effective proxy for the number of penguins residing within. Our model provides a robust, quantitative mechanism for estimating the breeding population size of colonies captured in imagery and identifies terrain slope as a significant component influencing apparent nesting density. While our high-resolution satellite imagery technique was developed for the Adélie penguin, these principles are directly transferrable to other colonially nesting seabirds and other species that aggregate in fixed localities.     

Shearwater Foraging in the Southern Ocean: The Roles of Prey Availability and Winds

Background

Sooty (Puffinus griseus) and short-tailed (P. tenuirostris) shearwaters are abundant seabirds that range widely across global oceans. Understanding the foraging ecology of these species in the Southern Ocean is important for monitoring and ecosystem conservation and management.

Methodology/Principal Findings

Tracking data from sooty and short-tailed shearwaters from three regions of New Zealand and Australia were combined with at-sea observations of shearwaters in the Southern Ocean, physical oceanography, near-surface copepod distributions, pelagic trawl data, and synoptic near-surface winds. Shearwaters from all three regions foraged in the Polar Front zone, and showed particular overlap in the region around 140°E. Short-tailed shearwaters from South Australia also foraged in Antarctic waters south of the Polar Front. The spatial distribution of shearwater foraging effort in the Polar Front zone was matched by patterns in large-scale upwelling, primary production, and abundances of copepods and myctophid fish. Oceanic winds were found to be broad determinants of foraging distribution, and of the flight paths taken by the birds on long foraging trips to Antarctic waters.

Conclusions/Significance

The shearwaters displayed foraging site fidelity and overlap of foraging habitat between species and populations that may enhance their utility as indicators of Southern Ocean ecosystems. The results highlight the importance of upwellings due to interactions of the Antarctic Circumpolar Current with large-scale bottom topography, and the corresponding localised increases in the productivity of the Polar Front ecosystem.     

The effects of prey demography on humpback whale (Megaptera novaeangliae) abundance around Anvers Island, Antarctica

Baleen whales and Adelie penguins in the near-shore waters around the Antarctic Peninsula forage principally on Antarctic krill. Given the spatial overlap in the distribution of these krill predators (particularly humpback whales) and their dependence on krill, the goals of this paper are to determine if the inter-annual community structure and relative abundance of baleen whales around Anvers Island is related to krill demography and abundance, and if the potential exists for inter-specific interactions between Adelie penguins and baleen. We use whale sightings and prey data from both net tows and Adelie penguin stomach samples to correlate the abundance of humpback whales with krill demography and abundance from 1993 to 2001. We find significant relationships between whale abundance and the size–frequency distribution of krill targeted by Adelie penguins, as well as the foraging success of Adelie penguins. These findings suggest both krill predators share common prey preferences in the upper portions of the water column around Anvers Island. These findings highlight the need for better knowledge of baleen whale foraging ecology and inter-specific interactions with penguins, as sea ice and krill populations around the Antarctic Peninsula are affected by rapid changes in climate.     

Diet and diving behaviour of European Storm Petrels Hydrobates pelagicus in the Mediterranean (ssp. melitensis)

Capsule Unlike Atlantic populations, which feed on krill, Mediterranean populations feed mainly on pelagic fish Gymnammodites cicerellus.

Aims To determine the diet and dive depth of the Mediterranean subspecies of European Storm Petrels Hydrobates pelagicus melitensis.

Methods Analysis of regurgitates of adults arriving at the colony for chick feeding and by determination of dives depth using the capillary tube method.

Results The main prey is Gymnammodites cicerellus, a pelagic fish. Storm Petrels dive for their prey and can reach up to 5 m in depth. They also make short foraging trips just outside the colony where they capture Opossum Shrimps Misydacea.

Conclusions European Storm Petrels in the Mediterranean exploit pelagic fish which are taken by diving. This contrasts with the Atlantic populations which feed mainly on krill. Mediterranean birds also feed on Opossum Shrimps Mysidacea during short foraging trips made at night just outside the colony. Differences in diet between long and short foraging trips may be because adults have to forage for both themselves and their chicks.     

南极Gerlache海峡海鸟的取食集群及食性

1997年12月至1998年2月,我们对南极半岛席尔瓦角善于飞翔海鸟与企鹅的取食关联性进行了研究,同时调查了取食集团中主要鸟种的食性.发现每个取食集团中有35.6-37.0只善于飞翔的海鸟,其中几乎都有纹颊企鹅群 (Pygoscelis antarctica),黑背鸥(Larus dominicanus)、灰贼鸥(Catharacta maccormicki)、花斑鹱(Daption capensis)和巨鹱(Macronectes giganteus)是各集团中最常见的鸟类.各取样单元内有相关性的种数随季节变化而减少,一些种类的减少与特定的物候期有关.南极磷虾(Euphausia superba)是绝大部分飞翔海鸟的主要食物,研究发现黑背鸥与纹颊企鹅所捕食的南极磷虾的大小最为接近.飞翔海鸟的觅食行为表明:在海面上短时停留的飞翔海鸟也能够成功捕捉到磷虾,这可能与磷虾躲避企鹅的捕食有关[动物学报 53(3):425-430,2007].     

Harp seal foraging behaviour during summer around Svalbard in the northern Barents Sea: diet composition and the selection of prey

The harp seal Pagophilus groenlandicus is a major high trophic level predator in the Barents Sea, and to better understand their function in the Barents Sea ecosystem, we need to understand their foraging behaviour during their most intensive feeding period. We analysed the diet composition and prey preference of 184 harp seals and 94 faeces samples, sampled in the northern Barents Sea (around Svalbard) during the period May–August in 1996, 1997, and 2004–2006. Concurrent with the sampling of seals, prey availability was assessed in one area in 1996 and 1997 and in two areas in 2006 using standard acoustic methods. Our study showed that harp seal diet composition varied significantly both in time (year) and space, and that their diets appeared to be size dependent. Both subadult (<150 cm) and adult seals were associated with pelagic crustaceans (particularly krill), whereas primarily adult seals were associated with fish (capelin, gadoids and flatfish). Krill was the most important prey group (63 %) followed by polar cod (16 %) and other fish species (10 %). The prey preference of harp seals varied in time and space; polar cod was often preferred by the seals whereas krill was commonly consumed in lower proportion than observed in the survey area. Gadoids and capelin had either been exploited in the same or less proportion as observed in the survey sea. This study emphasises the ecological significance of krill as prime food for harp seals during their intensive feeding period in summer.     

Spatial distributions and population dynamics of two salp species, Ihlea racovitzai and Salpa thompsoni, in the waters north of Lützow-Holm Bay (East Antarctica) during austral summers of 2005 and 2006

Information on the sequences of generations and reproductive states of salps in the Southern Ocean is essential for an improved understanding of salp population growth, although changes in distribution patterns of two species of salps, Salpa thompsoni and Ihlea racovitzai, in the Southern Ocean have the potential to alter the Southern Ocean ecosystem. We used stratified, quantitative sampling from the surface to 2,000 m depth with an RMT 8 net in January of 2005 and 2006 to determine the distribution and population structure of salps in the north of Lützow-Holm Bay, East Antarctica. Ihlea racovitzai occurred in both 2005 and 2006, but S. thompsoni was found only in 2005. Ihlea racovitzai occurred abundantly along the ice edge where Antarctic Winter Water was well developed, whereas S. thompsoni was more abundant at northern stations affected by warm Modified Circumpolar Deep Water. Small solitary stages of I. racovitzai dominated in 2005, but they had declined significantly by 2006. The S. thompsoni population was composed of small immature aggregates and mature solitary stages, suggesting that the solitary stages were reproducing. We did not find mature aggregate and immature solitary stages in the present study and thus suggested that S. thompsoni was unable to complete its life cycle in the north of Lützow-Holm Bay because of failure of sexual reproduction in the aggregate stage. The S. thompsoni population was therefore probably transported to our study area by advection.     

Life history buffering in Antarctic mammals and birds against changing patterns of climate and environmental variation

The consequences of warming for Antarctic long‐lived organisms depend on their ability to survive changing patterns of climate and environmental variation. Among birds and mammals of different Antarctic regions, including emperor penguins, snow petrels, southern fulmars, Antarctic fur seals and Weddell seals, we found strong support for selection of life history traits that reduce interannual variation in fitness. These species maximize fitness by keeping a low interannual variance in the survival of adults and in their propensity to breed annually, which are the vital rates that influence most the variability in population growth rate (λ). All these species have been able to buffer these rates against the effects of recent climate‐driven habitat changes except for Antarctic fur seals, in the Southwest Atlantic. In this region of the Southern Ocean, the rapid increase in ecosystem fluctuation, associated with increasing climate variability observed since 1990, has limited and rendered less predictable the main fur seal food supply, Antarctic krill. This has increased the fitness costs of breeding for females, causing significant short‐term changes in population structure through mortality and low breeding output. Changes occur now with a frequency higher than the mean female fur seal generation time, and therefore are likely to limit their adaptive response. Fur seals are more likely to rely on phenotypic plasticity to cope with short‐term changes in order to maximize individual fitness. With more frequent extreme climatic events driving more frequent ecosystem fluctuation, the repercussions for life histories in many Antarctic birds and mammals are likely to increase, particularly at regional scales. In species with less flexible life histories that are more constrained by fluctuation in their critical habitats, like sea‐ice, this may cause demographic changes, population compensation and changes in distribution, as already observed in penguin species living in the Antarctic Peninsula and adjacent islands.     

Intra-seasonal variation in foraging behavior among Ad��lie penguins (Pygocelis adeliae) breeding at Cape Hallett, Ross Sea, Antarctica

We investigated intra-seasonal variation in foraging behavior of chick-rearing Adélie penguins, Pygoscelis adeliae, during two consecutive summers at Cape Hallett, northwestern Ross Sea. Although foraging behavior of this species has been extensively studied throughout the broad continental shelf region of the Ross Sea, this is the first study to report foraging behaviors and habitat affiliations among birds occupying continental slope waters. Continental slope habitat supports the greatest abundances of this species throughout its range, but we lack information about how intra-specific competition for prey might affect foraging and at-sea distribution and how these attributes compare with previous Ross Sea studies. Foraging trips increased in both distance and duration as breeding advanced from guard to crèche stage, but foraging dive depth, dive rates, and vertical dive distances travelled per hour decreased. Consistent with previous studies within slope habitats elsewhere in Antarctic waters, Antarctic krill (Euphausia superba) dominated chick meal composition, but fish increased four-fold from guard to crèche stages. Foraging-, focal-, and core areas all doubled during the crèche stage as individuals shifted distribution in a southeasterly direction away from the coast while simultaneously becoming more widely dispersed (i.e., less spatial overlap among individuals). Intra-specific competition for prey among Adélie penguins appears to influence foraging behavior of this species, even in food webs dominated by Antarctic krill.     

Stable isotopes document the winter foraging ecology of king penguins and highlight connectivity between subantarctic and Antarctic ecosystems

The poorly known winter foraging ecology of the king penguin, a major Southern Ocean consumer, was investigated at the subantarctic Crozet Islands where the largest global population breeds. Blood δ¹³C and δ¹⁵N values were used as proxies of the birds’ foraging habitat and diet, respectively, and circulating prolactin levels helped in determining the birds’ reproductive status. Plasma prolactin concentrations showed that king penguin adults of unknown breeding status (n = 52) that were present at the colony in winter were in fact breeders and failed breeders, but were not non ‐breeders. Circulating prolactin was neither related to δ¹³C nor δ¹⁵N values, thus suggesting that both breeders and failed breeders used the same foraging habitats and fed on the same prey. Plasma and blood cell isotopic values depicted four new relevant biological features on the feeding strategies of king penguins during the critical winter period: (1) 42% of the birds foraged in the distant Antarctic Zone, but 58% fed primarily in subantarctic waters (δ¹³C), (2) they preyed upon myctophids in both zones (δ¹⁵N), (3) individuals were consistent in their foraging strategies over the winter months (δ¹³C and δ¹⁵N), and (4) a higher proportion of females (77%–80%) than males (27%–31%) favored feeding in distant Antarctic waters (δ¹³C). This study highlights trophic connectivity between subantarctic and Antarctic ecosystems and hence the key role of energy export from Antarctic waters to sustain breeding populations of subantarctic predators, including during the Austral winter.     

The overwintering strategy of Antarctic krill under the pack-ice of the Weddell Sea

Summary Antarctic krill (Euphausia superba Dana) occurs in enormous swarms in Antarctic waters during the ice-free summer months. The winter whereabouts of this stock were hitherto unknown. Evidence collected during the Winter Weddell Sea Project 1986 (WWSP'86, G. Hempel 1988) covering a large area of the eastern and southern Weddell Sea indicates that the seasonal sea ice cover sustains the bulk of the krill population. Results presented here, show that known aspects of krill morphology and behavior are actually adaptations to the ice habitat, suggesting that the dominance of krill in the Antarctic marine ecosystem is a result of its capacity to grow and reproduce in the water column in summer, and find both food and shelter in the ice cover during the rest of the year. This conclusion has far-reaching implications for our understanding of Southern Ocean biology and ecology.