Comparative lipid dynamics of euphausiids from the Antarctic and Northeast Pacific Oceans

To better understand the feeding and reproductive ecology of euphausiids (krill) in different ocean environments, lipid classes and individual lipid components of four different species of euphausiids from Northeast Pacific (temperate species) and Southern Ocean (Antarctic species) were analyzed in animals from multiple life stages and seasons. The dominant krill species in the Northeast Pacific Euphausia pacifica and Thysanoessa spinifera, were compared to the two major Antarctic species, Euphausia superba and E. crystallorophias. Analysis comprised total lipid and lipid classes together with individual fatty acid and sterol composition in adults, juveniles, and larvae. Antarctic krill had much higher lipid content than their temperate relatives (10–50 and 5–20% of dry mass for Antarctic and temperate species, respectively) with significant seasonal variations observed. Phospholipids were the dominant lipid class in both temperate krill species, while neutral storage lipids (wax esters and triacylglycerols for E. crystallorophias and E. superba, respectively) were the major lipid class in Antarctic krill and accounted for up to 40% of the total lipid content. Important fatty acids, specifically 16:0, 18:1ω9, 20:5ω3, and 22:6ω3, were detected in all four krill species, with minor differences between species and seasons. Detailed lipid profiles suggest that krill alter their lipid composition with life stage and season. In particular, larval Antarctic krill appear to utilize alternate food resources (i.e., sea-ice associated organisms) during austral winter in contrast to juveniles and adults (i.e., seston and copepods). Lipid dynamics in krill among krill in both systems appear closely linked to their life cycle and environmental conditions including food availability, and can provide a more complete comparative ecology of euphausiids in these environmentally distinct systems.     

Growth of Antarctic krill Euphausia superba at South Georgia

 Antarctic krill Euphausia superba has a central role in the ecosystem of the Southern Ocean and knowledge of its growth rate is central to determining the factors influencing population dynamics. The length of Antarctic krill in the diet of Antarctic fur seals Arctocephalus gazella at South Georgia revealed a consistent increase in size between ca. 42 and ca. 54 mm over the period October–March, indicating growth rates much higher than predicted by existing models. Geographical variation in growth rate may result in 2-year-old krill at South Georgia attaining the same size as 3-year-old krill in the Antarctic Peninsula region. The effect of geographical variation in growth rate on the population structure of krill has important implications for comparing the fate of individual cohorts over large scales and in the interpretation of krill life-cycles. Received: 20 May 2000 / Accepted: 11 August 2000     

Lipid metabolism of the Antarctic krill Euphausia superba and its ecological implications

 Various developmental stages (early larvae to adults) of Euphausia superba have been collected in different seasons in the Weddell Sea, the Lazarev Sea and off the Antarctic Peninsula to investigate the role of lipids and fatty acids in the life cycle of the Antarctic krill. The total-lipid data for E. superba exhibited seasonal variations, with low lipid levels in late winter/early spring and the highest levels in autumn. Seasonal changes were most pronounced in the immature and adult specimens, increasing from about 10% lipid of dry mass to more than 40%. The fatty-acid compositions of the younger stages were dominated by 20:5(n-3), 22:6(n-3) and 16:0. These are typical phospholipid fatty acids, which are major biomembrane constituents. The phospholipid composition was similar in the older stages. With increasing storage of triacylglycerols in the lipid-rich immature and adult stages, the fatty acids 14:0, 16:0 and 18:1(n-9) prevailed, comprising about 70% of total triacylglycerol fatty acids. The trophic-marker fatty acids 16:1(n-7) and 18:4(n-3), indicating phytoplankton ingestion, were less abundant. They reflected, however, the dependence of the larvae on phytoplankton as well as the seasonal changes in algal composition. The generally close linear relationships between fatty acids and lipid suggest that the fatty-acid compositions of the collected specimens were largely independent of the respective developmental stage, season and region. The linear fit indicates that triacylglycerol accumulation started at a level of about 5% of total lipid. Considering the various overwintering scenarios under discussion, the life cycle and reproductive strategies of krill are discussed in the context of the lipid metabolism and fatty-acid composition of E. superba. Lipid production is effective enough to accumulate large energy reserves for the dark season, but E. superba does not exhibit the sophisticated biosynthetic pathways known from other Antarctic euphausiids and copepods. Although important, lipid utilisation appears to be just one of several strategies of E. superba to thrive under the extreme Antarctic conditions, and this pronounced versatility may explain the success of this species in the Southern Ocean. Received: 16 June 2000 / Accepted: 18 December 2000     

Reproduction in the externally brooding sea anemone Epiactis georgiana in the Antarctic Peninsula and the Weddell Sea

External parental care is uncommon among actiniarians but common in Epiactis species. Here, several aspects of reproduction are analyzed for of one of them, Epiactis georgiana. Samples were collected in December, January, February, March, and April in the Antarctic Peninsula and the eastern Weddell Sea, during 1998, 2000, 2002, and 2003. Most sexually mature individuals of E. georgiana are male or female, but some are hermaphrodites. This is the first report of hermaphroditism in E. georgiana, which is the third species of the genus with this sexual pattern. The results suggest that oogenesis starts in December and that at least two generations of oocytes overlap; a third generation is often brooded externally. Putative fertilization is likely internal, and larvae and/or embryos are externally brooded on the distal part of the adult column until an advanced developmental stage. Apparently E. georgiana reproduces seasonally, probably releasing the embryos/larvae in the last months of the austral spring (December). Inter-individual variability was observed in gametogenesis. In addition, specimens from the Antarctic Peninsula were larger than those from the Weddell Sea. This study represents the first step in understanding the reproductive mode of E. georgiana.     

Growth of larval krill,<Emphasis Type="Italic"> Euphausia superba</Emphasis>, in fall and winter west of the Antarctic Peninsula

A quantitative characterization of growth-related parameters for larval and juvenile Antarctic krill, Euphausia superba (Dana), is essential to understanding the early life history of this key species in the Southern Ocean ecosystem. To this end, instantaneous growth rate experiments were conducted with larval and juvenile krill to determine growth increments and molting frequency in situ. All experiments were carried out during fall and winter months (April through September) on nine separate cruises west of the Antarctic Peninsula between 1987 and 1999. A consistent seasonal pattern across years was observed: growth rates decreased during fall (April/May), were minimal in early winter (June), and increased to maximum rates by late winter (September). Habitat-specific differences (water column vs under-ice) in growth rates of larvae collected on the same cruise were not observed in early winter (June 1993; within-year comparison). However, in a between-year comparison, larvae from the under-ice habitat (June/July 1987) had significantly higher growth increments than larvae from the open-water habitat (July 1989). The difference between these two comparisons may be a function of the degree of contrast in food availability in the water column and the sea ice at different times in the winter. Daylength at the time of collection explained 74% of the variation in larval and juvenile growth rates. This correlation may be an indirect effect of the influence of diel cycles on krill behavior and/or primary production in both the water column and ice.Communicated by J.P. Grassle, New Brunswick     

Dynamic intra-seasonal habitat use by Antarctic fur seals suggests migratory hotspots near the Antarctic Peninsula

Antarctic fur seals (Arctocephalus gazella) are major secondary consumers in the Southern Ocean, placing them in potential competition with commercial fisheries and requiring research to understand their seasonal habitat use. Using the data obtained during 14 shipboard surveys sampled on a fixed grid (150 K km²) during mid- to late summer, I quantified the spatial distribution and intra-seasonal variability of fur seal sightings relative to distance to land and hydrographic boundaries. I test the hypothesis that fur seals display an increase in their at-sea abundance during mid- to late summer near the Antarctic Peninsula as they prepare to take up wintering grounds. I also test whether abundances of their potential prey, krill and myctophids, exhibit intra-seasonal variability. During midsummer, high-abundance areas are located near major breeding colonies; however, during late summer, there is an order-of-magnitude increase in fur seal abundance, coinciding with an increase in the number of high-abundance areas located in Bransfield Strait. Coincidently, abundance of Euphausia superba decreased and the myctophid Electrona antarctica increased between mid- and late-summer surveys. High-abundance areas of fur seals are not associated with the southern Antarctic Circumpolar Current front but are concentrated within 100 km from land, potentially indicating the location of haul out and important coastal habitat use areas. The dynamic increase in the number and location of high-abundance areas during late summer represents a considerable amount of mammalian predators entering the Antarctic Peninsula marine ecosystem. This information is important for understanding the seasonal impact of fur seals on regional marine food webs and their potential interaction with the autumn–winter krill fishery.     

Krill evolution and the Antarctic ocean currents: evidence of vicariant speciation as inferred by molecular data

The phylogenetic relationships of the Antarctic krill Euphausia superba, the key species in the Antarctic food web, and other Antarctic and sub-Antarctic cuphausiids have been investigated using the 16S ribosomal mitochondrial gene. The phylogenetic reconstructions indicated that the Antarctic species form a monophyletic clade separated by the non-Antarctic species. The results revealed a large genetic divergence between the Antarctic (E. superba and E. crystallorophias) and sub-Antarctic species (E. vallentini). The time of separation between these species, estimated from the molecular data, is around 20 million years ago, which is comparable with the geological time of the formation of a circum-Antarctic water circulation and the Antarctic Polar Frontal Zone. The euphausiid molecular phylogeny therefore represents evidence for vicariant speciation.     

Larval distribution and reproductive ecology of Thysanoessa macrura (Crustacea: Euphausiacea) in the Scotia Sea

In the Scotia Sea, the larvae of Thysanoessa macrura G.O. Sars, 1885 were scarce in spring (September 27 to October 6, 1971) and mainly present at the Calyptopis I stage, while in summer (December 17, 1974 to January 3, 1975) they were much more abundant and much further developed (up to Furcilia VI). The spawning season commences as early as September and, as indicated by the presence of numerous nauplii and metanauplii in summer, is rather long. T. macrura begins to spawn much earlier in West-Wind-Drift waters than in waters of the Weddell Sea. It starts to spawn simultaneously with Euphausia frigida and E. triacantha, but significantly earlier than E. superba. These timings of the breeding process are related to differential biogeographical characteristics of these species. Plankton collections made at depths of 500 or 1000 m have, for T. macrura, revealed a developmental ascent analogous to that of E. superba. Ontogenetical deep-water migrations are, consequently, characteristic not only for E. superba, but also for other Antarctic euphausiids (in particular E. frigida, but possibly not for inshore E. crystallorophias) and, as correctly postulated by several other investigators, for all oceanic euphausiid species.     

Linking predator and prey behaviour: contrasts between Antarctic fur seals and macaroni penguins at South Georgia

Antarctic fur seals Arctocephalus gazella and macaroni penguins Eudyptes chrysolophus are the two main land-based krill Euphausia superba consumers in the northern Scotia Sea. Using a combination of concurrent at-sea (predator observations, net hauls and multi-frequency acoustics), and land-based (animal tracking and diet analysis) techniques, we examined variability in the foraging ecology of these sympatric top predators during the austral summer and autumn of 2004. Krill availability derived from acoustic surveys was low during summer, increasing in autumn. During the breeding season, krill occurred in 80% of fur seal diet samples, with fish remains in 37% of samples. Penguin diets contained the highest proportion of fish in over 20 years of routine monitoring (46% by mass; particularly the myctophid Electrona antarctica), with krill (33%) and amphipods (Themisto gaudichaudii; 21%) also occurring. When constrained by the need to return and feed their offspring both predator species foraged to the northwest of South Georgia, consistent with an area of high macrozooplankton biomass, but fur seals were apparently more successful at exploiting krill. When unconstrained by chick-rearing (during March) penguins foraged close to the Shag Rocks shelf-break, probably exploiting the high daytime biomass of fish in this area. Penguins and seals are able to respond differently to periods of reduced krill abundance (in terms of variability in diet and foraging behaviour), without detriment to the breeding success of either species. This highlights the importance of myctophid fish as an alternative trophic pathway for land-based predators in the Scotia Sea ecosystem.     

Modelling the effects of UV radiation on the survival of Antarctic krill (Euphausia superba Dana) in the face of limited data

Antarctic krill, Euphausia superba, is a keystone species of the Antarctic ecosystem. A fishery for krill may compete with land-based predators (penguins and seals), particularly during the breeding season. The Commission for the Conservation of Antarctic Marine Living Resources (CCAMLR) is moving towards management in small scale units. The management models specify predation and fishing mortality as space and time dependent but do not yet include non-predation natural mortality. Krill are known to be highly susceptible to ultraviolet radiation (UV) but there are limited empirical data. We develop a model for krill mortality caused by UV and parameterize and assess it by comparison with experimental data. The analysis allows us to identify key parameters that should be measured in future experiments and also leads to suggestions about modification of experimental procedure. We illustrate the method for krill found in the Livingston Island area and show that (a) it is possible to estimate the component of natural mortality due to UV-induced damage and (b) that cohorts born in 1979, 1984, or 1997 have different survival in the first 5 years of life, associated with differential UV exposure. In particular, those born in 1997 may have experienced as much as 10% lower survival than those born in 1979. The method developed here allows a potentially important source of krill mortality to be incorporated into the management models and suggests key experiments and field work in the future.     

Chlordane residues in krill,fish and Weddell seal from the Antarctic

Chlordane compounds (CHLs) were quantitated in krill (Euphausia superba), benthic fish (Trematomus bernacchii) and Weddell seal (Leptonychotes weddelli) collected around the Japanese Antarctic Research Station (Syowa Station: 69°00'S, 39°35'E). The concentrations of ZCHL (cis‐, trans‐chlordane+ cis‐, fraws‐nonachlor + oxych‐lordane) in krill was much lower than that of benthic fish. The ratio of SCHL to sum of SCHL, SDDT and PCBs decreased with the trophic levels. These results indicate that CHLs appear to be easily degraded in higher organisms.     

Antarctic shallow-water mega-epibenthos: shaped by circumpolar dispersion or local conditions?

The mega-epibenthos of two different geographic areas, the Antarctic Peninsula and the high Antarctic (eastern Weddell Sea), were investigated using underwater video. The distribution of the marine fauna at shallow depths between 55 and 160 m in these two areas was investigated to determine whether there are any zoogeographic differences at the community level. A total of 237 taxa represented by 85,538 individuals was identified. Multivariate analyses revealed significant faunal differences between northern Marguerite Bay (western Antarctic Peninsula) and the stations from the Weddell Sea, Atka Bay and Four-Seasons Bank. Echinoderms, especially ophiuroids, dominated Marguerite Bay, bryozoans and ascidians were abundant at Atka Bay, and hydroids and gorgonians were well represented at Four-Seasons Bank. These clear differences can mainly be explained by the influence of local environmental conditions that are probably the primary feature responsible in shaping the Antarctic shallow-water epifauna and not an intensive exchange with larger depths or a limited dispersion due to scarce and isolated shallow areas. In addition, modes of reproduction and characteristics of the early life history (e.g. brooding, viviparity or budding) of key taxa may also shape patterns of species distribution in shallow benthic Antarctic communities.Communicated by O. Kinne, Oldendorf/Luhe     

On the biology of the Antarctic krillEuphausia superba

Results are described of observations on the distribution and biology of the Antarctic krillEuphausia superba Dana in the Scotia Sea and off South Georgia, conducted on board the R.V. Academic Knipovich from february 13 to April 2, 1968. Commercial concentrations of krill were observed between the South Orkneys and the South Sandwich Islands. Trial commercial fishing resulted in 136 t/10 days, with an average catch of 2 t/h and a maximum catch of 7 t/h. Catches decreased in the morning, from 4 a.m. to 12 a.m. Northwest of South Georgia concentrations of krill were also observed. Krill tend to concentrate in areas with circular water movement, but the problem of prompt location of commercial concentrations has not yet been solved. According to size composition curves, krill become mature only at 3 years of age and breed in the fourth year of their life. Only a few individuals survive to the age of 4 years.     

Stable isotopes identify an ontogenetic niche expansion in Antarctic krill (Euphausia superba) from the South Shetland Islands, Antarctica

Antarctic krill (Euphausia superba) occupy a key position in the Southern Ocean linking primary production to secondary consumers. While krill is a dominant grazer of phytoplankton, it also consumes heterotrophic prey and the relative importance of these two resources may differ with ontogeny. We used stable isotope analyses to evaluate body size-dependent trophic and habitat shifts in krill during the austral summer around the South Shetland Islands, Antarctica. We found evidence for an asymmetric, ontogenetic niche expansion with adults of both sexes having higher and more variable δ¹⁵N values but consistent δ¹³C values in comparison with juveniles. This result suggests that while phytoplankton likely remains an important life-long resource, krill in our study area expand their dietary niche to include higher trophic food sources as body size increases. The broader dietary niches observed in adults may help buffer them from recent climate-driven shifts in phytoplankton communities that negatively affect larval or juvenile krill that rely predominately on autotrophic resources.     

Layers of Antarctic krill, Euphausia superba : are they just long krill swarms?

Antarctic krill, Euphausia superba Dana, a major component in the southern ocean food web, typically occur in aggregations that range from small, discrete swarms and schools through to layers and superswarms that extend horizontally for several kilometres. A large Longhurst–Hardy plankton recorder has been used to obtain high-resolution serial samples from within two layers (up to 4 km in length) that were found near Elephant Island, north of the Antarctic Peninsula. Krill length, sex and maturity stage, net and acoustic estimates of number density are shown to vary significantly within these large layers. The variation occurring within a single layer is comparable with that occurring in a set of 38 swarms sampled contemporaneously with the layers. Thus, unlike a krill swarm, a whole krill layer may account for a substantial amount of the variation in the local krill population, although individual parts of the layer cannot be considered in this way. The layers play an important role in the ecology of the krill within the area. Firstly, these large layers may contain a significant proportion of the biomass within an area. Secondly, the structure of the layers gives some insight into the ways in which krill swarm formation and dispersal may be occurring. Received: 28 March 1997 / Accepted: 3 December 1997     

Are penguins and seals in competition for Antarctic krill at South Georgia?

The Antarctic fur seal (Arctocephalus gazella) and macaroni penguin (Eudyptes chrysolophus) are sympatric top predators that occur in the Southern Ocean around South Georgia where they are, respectively, the main mammal and bird consumers of Antarctic krill (Euphausia superba). In recent years the population of fur seals has increased, whereas that of macaroni penguins has declined. Both species feed on krill of similar size ranges, dive to similar depths and are restricted in their foraging range at least while provisioning their offspring. In this study we test the hypothesis that the increased fur seal population at South Georgia may have resulted in greater competition for the prey of macaroni penguins, leading to the decline in their population. We used: (1) satellite-tracking data to investigate the spatial separation of the Bird Island populations of these two species whilst at sea during the breeding seasons of 1999 and 2000 and (2) diet data to assess potential changes in their trophic niches between 1989 and 2000. Foraging ranges of the two species showed considerable overlap in both years, but the concentrations of foraging activity were significantly segregated spatially. The size of krill taken by both species was very similar, but over the last 12 years the prevalence of krill in their diets has diverged, with nowadays less krill in the diet of macaroni penguins than in that of Antarctic fur seals. Despite a significant degree of segregation in spatial resource use by the study populations, it is likely that the South Georgia populations of Antarctic fur seal and macaroni penguin exploit the same krill population during their breeding season. For explaining the opposing population trends of the two species, the relative contributions of independent differential response to interannual variation in krill availability and of interspecies competition cannot be resolved with available evidence. The likely competitive advantage of Antarctic fur seals will be enhanced as their population continues to increase, particularly in years of krill scarcity.     

Spatial assessment of fin whale hotspots and their association with krill within an important Antarctic feeding and fishing ground

Fin whale (Balaenoptera physalus quoyi) habitat use and its relationship to environmental conditions are generally unknown in the Southern Ocean, presenting challenges for predicting their seasonal occurrence and potential effects of fishing pressure and climate change on this endangered species. Using biological data collected during 14 shipboard surveys off the northern Antarctic Peninsula and oceanographic data from satellite remote sensing, we mapped the distribution of fin whale hotspots, Antarctic krill abundance (biomass from acoustics, concentrations from nets) and ocean conditions during mid- and late-summer to investigate the environmental determinants of whale hotspots. Generalized additive models (GAM) were used to test the hypothesis that intra-seasonal changes in fin whale hotspot distribution relate to sea surface temperature (SST), krill abundance and eddy kinetic energy (EKE). More whale hotspots (sightings and individuals) are observed during late- than mid-summer surveys. During mid-summer, hotspots occurred near Elephant Island while in late-summer they were distributed throughout the slope region in proximity to the mean location of the southern Antarctic Circumpolar Current Front. The spatial mean of EKE did not differ between mid- and late-summer surveys, but the spatial mean of SST was significantly warmer during late-summer. The GAM for mid-summer indicates that fin whale hotspots were positively related to SST, EKE and acoustically determined krill biomass. The GAM for late-summer indicates the hotspots were negatively related to net-based krill abundance and positively related to acoustic krill biomass and EKE. This study is important because environmental determinants of fin whale hotspots may be used as reference points for implementing future conservation plans for their recovering populations.     

Validation and quantification of extractable age pigments for determining the age of Antarctic krill (<Emphasis Type="Italic">Euphausia superba</Emphasis>)

Antarctic krill, Euphausia superba, hatched from eggs and maintained for four years, were sampled periodically for age-pigment analysis. Extractable pigments from the eye and eyestalk ganglia were quantified using fluorescence intensity and standardised against protein. Three peak fluorescence intensities were detected at wavelengths of excitation 280 nm, emission 625 nm (pigment 1); excitation 355 nm, emission 510 nm (pigment 2); and excitation 463 nm, emission 620 nm (pigment 3). There was a positive correlation between the quantity of pigments 1 and 3 and the age of Antarctic krill. A model was developed to predict age from pigment 3 and to compare it with other age proxies (carapace length and eyeball diameter). The quantity of pigment 3 was the best predictor of age. The pigment method can discriminate between similar sized krill aged 12 and 36 months. Age pigments provide an improved tool for age estimation in Antarctic krill, particularly if used in conjunction with other demographic information.     

Effect of temperature on the molting,growth and maturation of the antarctic krill Euphausia superba (Crustacea: Euphausiacea) under laboratory conditions

The effect of temperature on molting, growth, and maturation rates was studied on laboratory-maintained Euphausia superba. The length of intermolt periods (IMP's) was inversely proportional to temperature (20.10 d, SD=1.60, at 0.12°C; 16.87 d, SD=1.68, at 0.97°C; and 12.48 d, SD=0.90, at 4.48°C), and directly proportional to krill size at 0.12°C and 0.97°C. For individually maintained krill the maximum growth rate at 4.48°C (0.068 mm d^-1) was nearly twice that at 0.68°C (0.037 mm d^-1). There was no observable temperature effect on maturation rates. The maturation changes of juveniles at all temperatures indicated that more than two years are probably required to reach maturity. Mature males and females regressed to immature forms, suggesting that E. superba may reproduce in successive years. These results and previously reported field and laboratory data for E. superba and other euphausiid species suggest a 4+ year life span for this species.This work was supported by NSF grant DPP 76-23437     

Contribution of heterotrophic material to diet and energy budget of Antarctic krill, Euphausia superba

A novel approach was used to estimate the heterotrophic carbon component in the diet of the Antarctic krill Euphausia superba. Over 200 specimens from seven samples collected in the Lazarev Sea (January 1993 and 1995), at the Antarctic Polar Front (January 1993), and around South Georgia (February/March 1994) were dissected, and the total carbon content of their stomachs was estimated with a CHN-analyser. Gut-pigment contents were also measured by the gut-fluorescence technique in specimens collected at the same time, and the equivalent amount of their gut carbon was then subtracted from the total organic carbon content of guts from the same samples. The remaining carbon was assumed to originate entirely from heterotrophic food sources. This heterotrophic component accounted for a substantial proportion of the total food consumed by Antarctic krill, ranging from 17.4 to 98.9% of the mass of the gut contents (mean = 78.8% ± 21.2 SD). The results make an important contribution to the elucidation of the energy budget of krill and its daily carbon ration. With a few exceptions, previous estimates were largely calculated from a solely autotrophic carbon source, and were unable to account for the metabolic requirements of E. superba. Krill plays an important role in Antarctic food webs, as it often constitutes ≃50% of the total biomass of the zooplankton, and produces fast-sinking, dense faecal pellets which are important in the vertical transport of organic carbon from the euphotic layer to the deep ocean. High consumption rates of smaller heterotrophic organisms by krill suggest that this large microphage may be more important than previously believed in re-packaging micro- and mesozooplankton into a longer-lasting and more easily sequestered carbon pool. Received: 26 October 1998 / Accepted: 14 October 1999