Genetic variation and population structure of krill (Euphausia superba) in the Atlantic sector of Antarctic waters and off the Antarctic peninsula

Summary Samples of Antarctic krill Euphausia superba from 16 locations in the Weddell Sea, Scotia Sea and on the west coast of the Antarctic Peninsula were analysed for protein variation using enzyme electrophoresis techniques. Analysis of allele distributions indicate that samples from all locations are part of a single breeding population of krill. Significant departures from random mating expectations were observed in the distribution of electrophoretic phenotypes of the enzymes esterase and glucose phosphate isomerase. Possible explorations of these results are advanced, but they do not materially affect conclusions concerning population structure. The spatial and temporal distribution of electrophoretically detected genetic variation in Atlantic and Pacific sector krill stocks is discussed by comparing the results of this study with those of previous authors working with E. superba.     

Genetic homogeneity of krill (Euphausia superba Dana) in the Southern Ocean

Summary Development of a comprehensive picture of the genetic population structure of the Antarctic krill (Euphausia superba) has been hampered by a lack of genetic data from two major areas of the species' distribution, the Bellingshausen Sea and the Ross Sea. Evidence from earlier studies of a discrete Bellingshausen Sea population was based on anomalous allele frequencies in two sample sets that were collected near the west coast of the Antarctic Peninsula rather than in the Bellingshausen Sea proper. In this paper we describe the first biochemical genetic data obtained on krill from the central Bellingshausen Sea and from the Ross Sea. Analyses of eight polymorphic loci in samples from these two areas have failed to provide any evidence of population structuring within the Pacific sector of the Southern Ocean, and have indicated that Pacific sector krill cannot be genetically discriminated from Atlantic sector krill or Indian Ocean sector krill. These findings further support the hypothesis of a single circumpolar breeding population of Antarctic krill.     

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.     

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     

Krill maintenance and experimentation at the australian antarctic division

Live Antarctic krill, Euphausia superba, have been maintained for experimental purposes at the Australian Antarctic Division since 1981. This population has been replenished on an annual basis with animals taken from the wild. Techniques used to capture and maintain live krill are discussed here, with particular reference given to the development of systems for their maintenance. Details are also provided for specific experimental systems that have been used to conduct research into the behaviour and physiology of krill both at-sea and in shore-based laboratories.     

Antarctic krill population genomics: apparent panmixia,but genome complexity and large population size muddy the water

Antarctic krill (Euphausia superba; hereafter krill) are an incredibly abundant pelagic crustacean which has a wide, but patchy, distribution in the Southern Ocean. Several studies have examined the potential for population genetic structuring in krill, but DNA‐based analyses have focused on a limited number of markers and have covered only part of their circum‐Antarctic range. We used mitochondrial DNA and restriction site‐associated DNA sequencing (RAD‐seq) to investigate genetic differences between krill from five sites, including two from East Antarctica. Our mtDNA results show no discernible genetic structuring between sites separated by thousands of kilometres, which is consistent with previous studies. Using standard RAD‐seq methodology, we obtained over a billion sequences from >140 krill, and thousands of variable nucleotides were identified at hundreds of loci. However, downstream analysis found that markers with sufficient coverage were primarily from multicopy genomic regions. Careful examination of these data highlights the complexity of the RAD‐seq approach in organisms with very large genomes. To characterize the multicopy markers, we recorded sequence counts from variable nucleotide sites rather than the derived genotypes; we also examined a small number of manually curated genotypes. Although these analyses effectively fingerprinted individuals, and uncovered a minor laboratory batch effect, no population structuring was observed. Overall, our results are consistent with panmixia of krill throughout their distribution. This result may indicate ongoing gene flow. However, krill's enormous population size creates substantial panmictic inertia, so genetic differentiation may not occur on an ecologically relevant timescale even if demographically separate populations exist.     

A surplus no more? Variation in krill availability impacts reproductive rates of Antarctic baleen whales

The krill surplus hypothesis of unlimited prey resources available for Antarctic predators due to commercial whaling in the 20th century has remained largely untested since the 1970s. Rapid warming of the Western Antarctic Peninsula (WAP) over the past 50 years has resulted in decreased seasonal ice cover and a reduction of krill. The latter is being exacerbated by a commercial krill fishery in the region. Despite this, humpback whale populations have increased but may be at a threshold for growth based on these human-induced changes. Understanding how climate-mediated variation in prey availability influences humpback whale population dynamics is critical for focused management and conservation actions. Using an 8-year dataset (2013–2020), we show that inter-annual humpback whale pregnancy rates, as determined from skin-blubber biopsy samples (n = 616), are positively correlated with krill availability and fluctuations in ice cover in the previous year. Pregnancy rates showed significant inter-annual variability, between 29% and 86%. Our results indicate that krill availability is in fact limiting and affecting reproductive rates, in contrast to the krill surplus hypothesis. This suggests that this population of humpback whales may be at a threshold for population growth due to prey limitations. As a result, continued warming and increased fishing along the WAP, which continue to reduce krill stocks, will likely impact this humpback whale population and other krill predators in the region. Humpback whales are sentinel species of ecosystem health, and changes in pregnancy rates can provide quantifiable signals of the impact of environmental change at the population level. Our findings must be considered paramount in developing new and more restrictive conservation and management plans for the Antarctic marine ecosystem and minimizing the negative impacts of human activities in the region.     

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.     

Spatial and temporal variability in reproductive timing of Antarctic krill (Euphausia superba Dana)

Spawning dates of Antarctic krill, Euphausia superba Dana, were calculated from larval stage compositions, and corrected using data on maturity stage composition of the adult krill. Both original and literature data obtained from the Antarctic Peninsula-Bellingshausen Sea area and around the Antarctic continent were used. A time series (1975/76–1986/87) for several subareas of the Antarctic Peninsula-Bellingshausen Sea area indicates considerable variation in the krill spawning start, maxima and completion. In particular years (1975/76, 1980/81), krill spawning in the western Atlantic sector began relatively early, was intensive, and completed early. Some years (1977/78, 1981/82) were characterised by long and non-synchronised krill spawning. Compiled data sets for the Atlantic sector (1980/81), the entire Antarctic (1983/84) and the east Indian-west Pacific Antarctic waters (1981–85) reveal some spatial patterns in krill reproductive timing. In relation to spawning timing variation, the habitats of the krill population fall into five categories: (1) areas with an early beginning (late Novemberearly December) and a variable, but normally long, duration (3–3.5 months) of krill spawning; this is generally the southern boundary of the Antarctic Circumpolar Current, (2) areas with an early beginning, but a short duration of krill spawning (Gerlache Strait), (3) areas with a highly variable (within 1–1.5 months) beginning and a relatively long duration (ca. 3 months) of krill spawning (Bransfield Strait, Palmer Archipelago), (4) areas with a late beginning (late December–January) and a long duration of krill spawning (Bellingshausen Sea, D'Urville Sea, and Balleny Islands area), and (5) areas with a delayed beginning, but a very short duration (ca. 1.5 months) of krill spawning (Ross Sea slope, probably the Coastal Current area off the Lasarev Sea shelf and in the south-eastern Weddell Sea. These patterns can be partly explained by peculiarities of the ice regime in particular areas and by routes of krill movement within water circulation systems.     

Modelling individual growth of the Antarctic krill Euphausia superba Dana

Summary Growth of the Antarctic krill, Euphausia superba, is not easily determined from net catches nor from laboratory experiments. Therefore, in support of these methods, a phenomenological model was constructed which in its present state describes the growth of a single krill specimen under periodically limiting food conditions with summer seasons of variable lengths. Published data of krill body length vs. age and of the annual cycle of primary production of algae in the Drake Passage were used to formulate equations and to calculate growth curves. At 1,000 days after hatching, the model predicts a body length of 63 mm, growth being delayed by 380 days compared with constant, optimal feeding conditions. Final length, weight and time delay are related to the amount of food supplied and compared with published population growth curves.     

Adult antarctic krill feeding at abyssal depths

Antarctic krill (Euphausia superba) is a large euphausiid, widely distributed within the Southern Ocean [1], and a key species in the Antarctic food web [2]. The Discovery Investigations in the early 20(th) century, coupled with subsequent work with both nets and echosounders, indicated that the bulk of the population of postlarval krill is typically confined to the top 150 m of the water column [1, 3, 4]. Here, we report for the first time the existence of significant numbers of Antarctic krill feeding actively at abyssal depths in the Southern Ocean. Biological observations from the deep-water remotely operated vehicle Isis in the austral summer of 2006/07 have revealed the presence of adult krill (Euphausia superba Dana), including gravid females, at unprecedented depths in Marguerite Bay, western Antarctic Peninsula. Adult krill were found close to the seabed at all depths but were absent from fjords close inshore. At all locations where krill were detected they were seen to be actively feeding, and at many locations there were exuviae (cast molts). These observations revise significantly our understanding of the depth distribution and ecology of Antarctic krill, a central organism in the Southern Ocean ecosystem.     

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     

Relative changes in krill abundance inferred from Antarctic fur seal

Antarctic krill Euphausia superba is a predominant species in the Southern Ocean, it is very sensitive to climate change, and it supports large stocks of fishes, seabirds, seals and whales in Antarctic marine ecosystems. Modern krill stocks have been estimated directly by net hauls and acoustic surveys; the historical krill density especially the long-term one in the Southern Ocean, however, is unknown. Here we inferred the relative krill population changes along the West Antarctic Peninsula (WAP) over the 20th century from the trophic level change of Antarctic fur seal Arctocephalus gazella using stable carbon (δ(13)C) and nitrogen (δ(15)N) isotopes of archival seal hairs. Since Antarctic fur seals feed preferentially on krill, the variation of δ(15)N in seal hair indicates a change in the proportion of krill in the seal's diets and thus the krill availability in local seawater. For the past century, enriching fur seal δ(15)N values indicated decreasing krill availability. This is agreement with direct observation for the past ～30 years and suggests that the recently documented decline in krill populations began in the early parts of the 20th century. This novel method makes it possible to infer past krill population changes from ancient tissues of krill predators.     

Length–weight relationships of five fish species associated with krill fishery in the Atlantic sector of the Southern Ocean

Length–weight relationships (LWRs) of five fish species commonly associated with Antarctic krill fishery, were determined. Samples were collected with Chinese krill trawler (codend mesh size 2 cm) in the Atlantic sector of the Southern Ocean from January to August 2016. Data on LWRs and the relationships between weight and standard length as well as weight and total length of those species were updated for the database of FishBase. Those data on fish species derived from Antarctic krill fishery will be very helpful in understanding the interaction between krill fishery and the associated fish community.     

Environmental response of upper trophic-level predators reveals a system change in an Antarctic marine ecosystem

Long-term changes in the physical environment in the Antarctic Peninsula region have significant potential for affecting populations of Antarctic krill (Euphausia superba), a keystone food web species. In order to investigate this, we analysed data on krill-eating predators at South Georgia from 1980 to 2000. Indices of population size and reproductive performance showed declines in all species and an increase in the frequency of years of low reproductive output. Changes in the population structure of krill and its relationship with reproductive performance suggested that the biomass of krill within the largest size class was sufficient to support predator demand in the 1980s but not in the 1990s. We suggest that the effects of underlying changes in the system on the krill population structure have been amplified by predator-induced mortality, resulting in breeding predators now regularly operating close to the limit of krill availability. Understanding how krill demography is affected by changes in physical environmental factors and by predator consumption and how, in turn, this influences predator performance and survival, is one of the keys to predicting future change in Antarctic marine ecosystems.     

Discovery of ciliates reproducing in the gut of Antarctic krill

During a recent Antarctic research cruise (December 1994/February 1995), dissection of fresh Antarctic krill (Euphausia superba) on board ship revealed live motile ciliates in the gut of krill. Further observation of gut samples by scanning electron microscopy indicated that the ciliates were symbionts located within the gut. We inferred that the ciliates may have enabled the krill to digest a wider range of food items, and as a consequence, this had became an important strategy for Antarctic krill's survival in the Antarctic ecosystem. Received: 1 August 1996 / Accepted: 1 December 1996     

Assessing population-level variation in the mitochondrial genome of <Emphasis Type="Italic">Euphausia superba</Emphasis> using 454 next-generation sequencing

The Antarctic krill (Euphausia superba Dana 1852) is widely distributed throughout the Southern Ocean, where it provides a key link between primary producers and upper trophic levels and supports a major commercial fishery. Despite its ecological and commercial importance, genetic population structure of the Antarctic krill remains poorly described. In an attempt to illuminate genetic markers for future population and phylogenetic analysis, five E. superba mitogenomes, from samples collected west of the Antarctic Peninsula, were sequenced using new 454 next-generation sequencing techniques. The sequences, of lengths between 13,310 and 13,326 base pairs, were then analyzed in the context of two previously-published near-complete sequences. Sequences revealed relatively well-conserved partial mitochondrial genomes which included complete sequences for 11 of 13 protein-coding genes, 16 of 23 tRNAs, and the large ribosomal subunit. Partial sequences were also recovered for cox1 and the small ribosomal subunit. Sequence analysis suggested that the cox2, nad5, and nad6 genes would be the best candidates for future population genetics analyses, due to their high number of variable sites. Future work to reveal the noncoding control region remains.     

Diet of the Antarctic starry skate Amblyraja georgiana (Rajidae, Chondrichthyes) at South Georgia (Southern Ocean)

Stomach contents were identified from 206 Antarctic starry skate (Amblyraja georgiana) that were collected during three groundfish surveys (September 2007, April 2008 and January 2009) at South Georgia, Southern Ocean. The diet of A. georgiana varied with skate size and between years. Preferred prey included fish (particularly for larger individuals) and Antarctic krill, Euphausia superba, as well as amphipods, polychaetes and other benthic fauna. The skate A. georgiana appears to be an opportunistic predator, and the clear presence of Antarctic krill in this demersal predator’s diet may indicate a benthic habit of this euphausiid species, which has hitherto mainly been considered as occupying a purely pelagic niche.     

A comparison of Antarctic petrel (Thalassoica antarctica) diets with net samples of Antarctic krill (Euphausia superba) taken from the Prydz Bay region

Samples of the stomach contents of Antarctic petrels (Thalassoica antarctica) were obtained on board ship in the Prydz Bay region of Antarctica from birds which had spontaneously regurgitated. The weight of food and the species composition of the stomach contents were measured. Antarctic krill,Euphausia superba, the sole prey item taken, were compared to krill obtained by nets in the same region as part of a large-scale krill survey. Krill from petrel stomach samples were larger in mean size than those sampled by nets. This may be attributed to selection of the larger sized krill by the petrels, it may be caused by the nets sampling different populations of krill or it may be due to net avoidance by the larger krill.     

A Southern Ocean archipelago enhances feeding opportunities for a krill predator

Productivity in the oceans is heightened around oceanographic and bathymetric features such as fronts and islands. This can have a flow-on effect, providing increased food availability for higher trophic level species. Using data from a 5-day combined visual and acoustic survey, we examined the hypothesis that higher Antarctic krill (Euphausia superba) density provides a lucrative resource for humpback whales (Megaptera novaeangliae) at a remote Antarctic feeding area, the Balleny Islands (67^oS, 164°E). We assessed whale presence at the feeding area in relation to prey (krill), productivity and environmental variables using density surface modeling. We found stark differences between krill swarms near the islands and those in adjacent open water. Swarms were twice as dense and three times more numerous near the Balleny Islands compared to an open water pelagic environment, suggesting that the islands offered a profitable feeding opportunity. At the feeding area, whales were found in deeper and more productive waters with medium krill densities. These relationships, along with the high krill availability around the islands, may be the result of the Island Mass Effect.