Genetic variation and population structure of krill (Euphausia superba dana) from the Prydz Bay region of Antarctic waters

Summary Samples of Antarctic krill E. superba from six locations near Prydz Bay were analysed electrophoretically to detect genetically-based protein variation. Analyses of allele distributions at four polymorphic loci indicate no evidence of significant heterogeneity, a result consistent with the hypothesis that all samples were derived from a single breeding population of krill. The results of this study agree closely with genetic data from other studies on krill in the Atlantic sector of the Southern Ocean, suggesting that krill stocks over at least 6000 km of Antarctic waters are derived from a single interbreeding population.     

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.     

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.     

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     

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     

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.     

Related antipodes: a comparative study on digestive endopeptidases from Northern krill and Antarctic krill (Euphausiacea)

The Antarctic krill, Euphausia superba, and the Northern krill, Meganyctiphanes norvegica, are closely related species but occupy significantly different trophic and climatic environments. E. superba holds a key position as a phytoplankton grazer in the Southern Ocean. The omnivorous M. norvegica is an important member of plankton communities in the Northeast Atlantic. Both species expressed high proteolytic activities which were dominated by serine proteinases. In the stomachs of Antarctic krill, activities of total proteinase, trypsin, and chymotrypsin were significantly higher than in Northern krill. In the midgut glands, however, total proteinase and trypsin activities were similar in both species, but chymotrypsin activity was significantly higher in Antarctic krill. Moreover, Antarctic krill expressed four trypsin isoforms while only one isoform appeared in Northern krill. Chymotrypsin was present in either species as one single isoform. Antarctic krill adapted to the low and patchy distribution of food by elevated enzyme activities and the expression of trypsin isoforms with slightly different catalytic properties. Presumably, these enzymes facilitate in concerted action the efficient utilization of proteins from phytoplankton, the major food. Northern krill, in contrast, seems not to be equipped to face food limitation. It expresses a “simple” or “basic” set of digestive enzymes for utilizing abundant and easily digestible prey.     

The diet of Antarctic fur sealsArctocephalus gazella during the breeding season at South Georgia

 The diet of lactating female Antarctic fur seals Arctocephalus gazella at South Georgia was investigated during the pup-rearing period (January/March) of 1991–1994. Antarctic krill Euphausia superba was the main prey item, occurring in 88% of all scats (n=497), whereas fish occurred in 47% and squid in 5% of all scats. There was considerable intra- and inter-annual variation in the characteristics of krill taken by fur seals. The distribution of krill sizes taken suggests that fur seals are not actively selecting particular sizes of krill and, therefore, that the krill in the diet reflects the krill available around South Georgia. The absence of group 3 krill (44–48 mm in length) in the South Georgia area, as indicated by their absence in the diet of seals, is suggested as a possible reason for low availability of krill and the subsequent reproductive failure among krill predators. The frequency of occurrence of fish was much higher than in previous studies; the pattern of fish consumption showed a consistent seasonal pattern in 3 of the 4 years studied. Of the total number of the myctophid Protomyctophum choriodon, the most numerous fish taxon, 98% were taken between early February and the middle of March. Champsocephalus gunnari and Lepidonotothen larseni agg., which both feed on krill, dominated the fish component of the diet outside this period and together constituted 94% of the total estimated biomass of fish consumed. The intra- and inter-annual variability in the diet of Antarctic fur seals emphasise the need for diet studies to be conducted during the entire pup-rearing periods of several years. Received: 15 March 1995/Accepted: 13 May 1995     

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.     

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.     

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.     

Electrophoretic investigation of genetic variation in two krill species Euphausia superba and E. crystallorophias (Euphausiidae)

Summary Specimens of Euphausia superba and of E. crystallorophias from different locations were analyzed electrophoretically for protein variation. The present study extends previous genetic studies on E. superba by analyzing samples from the East Wind Drift and repeat samples from the Bransfield Strait, Elephant Island and the Weddell Sea. E. crystallorophias was collected in the Weddell Sea and around the Antarctic Peninsula in order to provide information on the breeding structure of the species in this region. For all loki taking all sampling sites together for both species except GPI in E. crystallorophias no significant deviation of phenotype distributions from random mating expectations was observed. Furthermore, the allele distributions in all polymorphic loci for both species were found to be homogeneous. Estimates of genetic distance between samples within each species are low (0.0001 to 0.0003 in E. superba and 0.0001 to 0.0002 in E. crystallorophias), and are consistent with results expected for samples from a single interbreeding population. Estimate of genetic distance between these two species was 0.9729. These results suggest that for each species specimens from all locations investigated in the Bransfield Strait and Weddell Sea belong to a single genetically homogeneous population. A possible mechanism for maintaining such homogeneity and the implications for fishery management are discussed.     

Preliminary study on chromosomes of Antarctic krill,Euphausia superba Dana

Summary Meiotic chromosomes of antarctic krill, Euphausia superba Dana, collected at Admiralty Bay, are described and the method used to obtain this genetic material is reported. Forty seven dividing meiotic nuclei at diakinesis and metaphase I from seminal glands of four males were examined, all with n= 17 bivalents. The results were compared with those obtained by other workers using krill specimens from the opposite side of the Antarctic continent and apparent differences in chromosome numbers were observed. Possible reasons for these differences and the potential for using cytogenetic techniques to study populations of krill are discussed.     

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.     

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.     

Eugregarine infection within the digestive tract of larval Antarctic krill, Euphausia superba

The infection rate of eugregarine parasites, Cephaloidophora pacifica, within the digestive tract of larval Antarctic krill, Euphausia superba, was examined using samples collected from the Indian sector of the Southern Ocean. Immature and mature eugregarine gamont stages were found at all larval stages older than Calyptopis I stage. Eugregarine infection in 14.0% (N = 108) of the first feeding stage (Calyptopis I) suggested that krill larvae are at risk from being infected during physiological transition from non-feeding to feeding stages. Eugregarine prevalence and intensity of infection at the three calyptopis stages increased with stage/krill length. Statistical analysis showed that the intensity of C. pacifica infection also increased with host density. Thus, krill density is probably a key determinant of the intensity of infection. We found gamont stage eugregarines in the host hind-gut, blocking the passage of food. Eugregarine infections in larval krill may have a negative impact on digestion and absorption in the host digestive tract.     

Natural age-dependent mortality rates of Antarctic krill Euphausia superba Dana in the Indian sector of the Southern Ocean

Data on the size and age composition of Antarctic krill (Euphausia superba Dana) were collected in the Cooperation and Cosmonaut Seas (Indian sector of the Southern Ocean) from 1985 to 1990. The estimation of the age-dependent annual extinction rate of krill [=1-exp(-M)] was obtained using the Zikov and Slepokurov (1982) approach and results were fitted by a parabolic equation. The coefficients of instantaneous natural mortality (M) of E. superba derived with this approach range from 0.52 during the maturation period, to 1.1–2.41 during the first and last years of life.     

Synopsis of “The biology and ecology of the Antarctic krill,Euphausia superba Dana”

Summary The review summarises current knowledge on the ecology, biology and life history of the Antarctic krill (Euphausia superba Dana). This paper presents a synopsis of the contents and major conclusions of that review. Emphasis is placed on recent developments, particularly on the results of various national and international research programmes loosely co-ordinated under the auspices of BIOMASS (Biological Investigations of Marine Antarctic Systems and Stocks). Attention is focused on topics which are either directly or indirectly applicable to the effective management of krill exploitation within the provisions set out by the Convention for the Conservation of Antarctic Marine Living Resources (CCAMLR).