Genetic population structure of Euphausia superba DANA in the Atlantic Sector of the Southern Ocean as demonstrated by different electrophoretic techniques

Summary Electrophoretic analyses of allele distributions at nine polymorphic gene loci were conducted in samples of Euphausia superba from the Bransfield Strait, off the South Orkney Islands, and from the south-eastern part of the Weddell Sea. The aim was to determine whether reported phenotypic differences between krill stocks from these areas could be linked to genotypic differences. Despite minor deviations of genotype distributions from Hardy-Weinberg expectations, the data gave no evidence of genetic heterogeneity over the sampled area, and the hypothesis of a single genetically homogeneous krill population could not be rejected. Genetic data for four selected loci were verified by using two different electrophoretic methods. Results from these two techniques yielded no discrepancies in interpretation of the data.     

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.     

Larvae of euphausiids off Queen Maud Land

Summary In a March 1981 study of the Lazare Sea, a sector of the Southern Ocean along Queen Maud Land, Euphausia superba and Thysanoessa macrura larvae were abundant in the north and northwest part of the region near the frontal zone between the Weddell Warm Countercurrent (WWC) and Weddell Gyre, 5°W–15°E. Toward the south, E. superba and E. crystallorophias larvae were numerous in the frontal zone between the WWC and the Antarctic Coastal Current. Only T. macrura larvae occurred in the central part of the sea. The age composition of larvae was most advanced in T. macrura, probably due to the drift of larvae from the northwest. Because the ice cover diminishes later in the flow from the Weddell Gyre (northernmost area of the region) than in the south, E. superba larvae were younger in that oceanic subregion than near the shore. T. macrura larvae were sparse near the coast and an onshore-offshore age difference was not pronounced. Both frontal zones correspond to reproductive areas of E. superba and T. macrura. E. crystallorophias spawns only in the southern area.     

Comparisons of morphology and neritic distributions of <Emphasis Type="Italic">Euphausia crystallorophias</Emphasis> and <Emphasis Type="Italic">Euphausia superba</Emphasis> furcilia during autumn and winter west of the Antarctic Peninsula

Euphausia crystallorophias and E. superba larvae often overlap in distribution in Antarctic coastal regions. Here, we describe the morphology and ecology of E. crystallorophias furcilia stages F3–F6, with emphasis on characteristics that distinguish them from E. superba, based on samples collected west of the Antarctic Peninsula during autumn and winter 2001 and 2002. During autumn most E. crystallorophias occurred as F4s (53%) and F5s (35%), while E. superba occurred in all furcilia stages (F1–F6). During winter, F6 was the dominant stage (>67%) for both species. On average, body lengths of E. crystallorophias larval stages were significantly greater than those of E. superba. During autumn, densities of the two species were similar (range: 0.003–11.8 m^–3) at many on-shelf stations, with lower densities during winter. Where both species occurred, >58% of E. crystallorophias furcilia were collected between 50 and 100 m depth, while 82% of E. superba were shallower (25–50 m). Younger stages of E. crystallorophias occurred more frequently (54% of F3s) in water >100 m than older stages (11% of F6s). Thus, many larval E. crystallorophias were vertically segregated from E. superba, thereby reducing grazing competition between the young of these morphologically similar species.     

Effect of the abundance of three predominating copepod species on adequate sample volume and sample size in Bransfield Strait (Antarctic Peninsula) and waters north of the Weddell Sea

Summary The effect on adequate sample size and sample volume of the abundances of three predominant copepod species, Metridia gerlachei, Calanus propinquus and Calanoides acutus, were studied in Bransfield Strait (Antarctic Peninsula) in the austral summer of 1988–1989 and waters north of the Weddell Sea in 1989–1990. Copepod abundances were higher in the area north of the Weddell Sea, with the exception of Metridia gerlachei, which was evenly distributed over both areas. Local (intra-station) patchiness was not found, indicating random distribution over small areas. In the assessment of inter-station variability in Bransfield Strait, with standard error of the mean set arbitrarily at 20% of the average abundance and a sampling volume of 150 m³, the theoretical minimum sample size (number of sampling stations) ranged from 6 to 17 for juvenile copepods and from 11 to 25 for adults. The minimum number of stations in the area north of the Weddell Sea reached from 5 to 7, and from 7 to 10 respectively.     

A comparison of Antarctic euphausiids sampled by net and from geothermally heated waters: insights into sampling bias

The Antarctic euphausiids Euphausia superba and E. crystallorophias were sampled using an 8 m² rectangular midwater trawl inside the caldera of Deception Island. Specimens of the same species were collected on the same day from the strandline and sublittoral fringe within the caldera at a bay where these euphausiids had been incapacitated by high temperature geothermally heated waters. Differences between length frequency distributions of samples obtained by both methods were investigated once net-caught specimens had been subject to the simulated effects of geothermally heated waters. Differences between maturity stage/sex ratios for E. superba samples were also assessed. No significant differences between sizes of net-caught or beach-sampled E. crystallorophias were detected, but significant differences were found between both the sizes and sex/maturity stages for E. superba samples. Possible explanations for these differences are discussed. Accepted: 15 March 1999     

The role of notothenioid fish in the food web of the Ross Sea shelf waters: a review

The Ross Sea, a large, high-latitude (72–78°S) embayment of the Antarctic continental shelf, averages 500 m deep, with troughs to 1,200 m and the shelf break at 700 m. It is covered by pack ice for 9 months of the year. The fish fauna of about 80 species includes primarily 4 families and 53 species of the endemic perciform suborder Notothenioidei. This review focuses on the diet and role in the food web of notothenioids and top-level bird and mammal predators, and also includes new information on the diets of artedidraconids and bathydraconids. Although principally a benthic group, notothenioids have diversified to form an adaptive radiation that includes pelagic and semipelagic species. In the southern Ross Sea, notothenioids dominate the fish fauna at levels of abundance and biomass >90% and are, therefore, inordinately important in the food web. Antarctic krill (Euphausia superba) and mesopelagic fishes are virtually absent from the shelf waters of the Ross Sea. Of the four notothenioid families, nototheniids show the most ecological and dietary diversification, with pelagic, cryopelagic, epibenthic and benthic species. Neutrally buoyant Pleuragramma antarcticum constitutes >90% of both the abundance and biomass of the midwater fish fauna. Most benthic nototheniids are opportunistic and feed on seasonally or locally abundant zooplanktonic prey. Artedidraconids are benthic sit-and-wait predators. Larger bathydraconids are benthic predators on fish while smaller species feed mainly on benthic crustaceans. Channichthyids are less dependent on the bottom for food than other notothenioids. Some species combine benthic and pelagic life styles; others are predominantly pelagic and all consume euphausiids and/or fish. South polar skuas, Antarctic petrels, Adélie and emperor penguins, Weddell seals and minke and killer whales are the higher vertebrate components of the food web, and all prey on notothenioids to some extent. Based on the frequency of occurrence of prey items in the stomachs of fish, bird and mammal predators, P. antarcticum and ice krill E. crystallorophias are the key species in the food web of the Ross Sea. P. antarcticum is a component of the diet of at least 11 species of nototheniid, bathydraconid and channichthyid fish and, at frequencies of occurrence from 71 to 100%, is especially important for Dissostichus mawsoni, Gvozdarus svetovidovi and some channichthyids. At least 16 species of notothenioids serve as prey for bird and mammal predators, but P. antarcticum is the most important and is a major component of the diet of south polar skua, Adélie and emperor penguins and Weddell seals, at frequencies of occurrence from 26 to 100%. E. crystallorophias is consumed by some nototheniid and channichthyid fish and can be of importance in the diet of emperor and Adélie penguins, although in the latter case, this is dependent on location and time of year.Unlike the linear phytoplanktonE. superbaconsumers of the E. superba food chain hypothesized for much of the Southern Ocean, the food web of the Ross Sea shelf is non-linear, with complex prey-predator interactions. Notothenioid fish play a key role: as predators, they occupy most of the trophic niches available in the ecosystem, relying on benthic, zooplanktonic and nektonic organisms; as prey, they are important food resources for each other and for most top predators living and foraging on the shelf. They also constitute the major link between lower (invertebrates) and higher (birds and mammals) levels of the food web. This is especially true for P. antarcticum. Along with E. crystallorophias, its ecological role in the Ross Sea is equivalent to that of myctophids and E. superba elsewhere in the Southern Ocean.     

Zooplankton communities in the Southern Weddell Sea (Antartica)

Summary Standardized abundances of 40 frequently occurring macrozooplankton taxa collected by double-oblique bongo net hauls between the surface and 200 m depth were submitted to an agglomerative hierarchical cluster analysis, to characterize surface zooplankton communities in the southern and eastern Weddell Sea. The sensitivity of the analysis concerning reduction of the number of considered taxa (especially stages of species) was tested. Dominant taxa in the entire area over the whole period were copepods Metridia gerlachei and Calanoides acutus. Calanus propinquus, Appendicularia spp. and calyptopis-1 larvae of krill, Euphausia superba were abundant at 73°S/19°W. Euphausia crystallorophias and larval fish Pleuragramma antarcticum dominated at 77°S/40°W on the southern shelf. The cluster analysis revealed marked similarities between the southern shelf community and the January community further to the northeast. E. crystallorophias, and larval Pleuragramma antarcticum are thought to be transported in surface layers of the coastal current to the southwest. Oceanic species such as Thysanoessa sp. and E. superba are probably not transported in the same way. Possible mechanisms for the maintenance of the community structures are discussed. A simplified method of characterizing communities, based on analysis of euphausiids, is presented.     

Horizontal niche partitioning of humpback and fin whales around the West Antarctic Peninsula: evidence from a concurrent whale and krill survey

A dedicated aerial cetacean survey was conducted concurrently to a standardised net trawl survey for krill in order to investigate distribution patterns of large whales and different krill species and to investigate relationships of these. Distance sampling data were used to produce density surface models for humpback (Megaptera novaeangliae) and fin whales (Balaenoptera physalus) around the West Antarctic Peninsula (WAP). Abundance for both species was estimated over two strata in the Bransfield Strait and Drake Passage. Distinct distribution patterns suggest horizontal niche partitioning of the two whale species around the WAP, with fin whales aggregating at the shelf edge of the South Shetland Islands in the Drake Passage and humpback whales in the Bransfield Strait. Krill biomass estimated from the concurrent krill survey was used along with CTD data from the same expedition, bathymetric parameters and satellite data on chlorophyll-a and ice concentration to model krill distribution. Comparisons of the predicted distributions of both whale species with the predicted distributions of Euphausia superba, Euphausia crystallorophias and Thysanoessa macrura suggest a complex relationship rather than a straightforward correlation between krill and whales. However, results indicate that fin whales were feeding in an area dominated by T. macrura, while humpback whales were found in areas of higher E. superba biomass. Our results provide abundance estimates for humpback whales and, for the first time, fin whales in the WAP and contribute important information on feeding ecology and habitat use of these two species in the Southern Ocean.     

Distribution of salps near the South Shetland Islands during austral summer, 1990–1991 with special reference to krill distribution

Distribution and biomass of salps and Antarctic krill (Euphausia superba) were investigated near the South Shetland Islands during austral summer 1990–1991. Salp biomass ranged between 0 and 556 mgC·m^–3 and was greatest at a station in the Bransfield Strait in late December 1990. Salp biomass was lower than that of E. superba. Two species of salps; Salpa thompsoni and Ihlea racovitzai were found, and the former was dominant numerically. Spatial distribution and generation composition of these two species was different. Spatial distributions of salps and E. superba did not overlap particularly so the January–February period. While E. superba was found mainly in the coastal area which showed high-chlorophyll a values, salps exhibited high biomass in the oceanic area with low chlorophyll a concentrations. Predation by salps on small krill and the competitive removal of food by them, are discussed as potential reasons for the relatively low abundance of E. superba at the stations where salps were present in great numbers.     

Euphausia superba dominates in the diet of Adélie penguins feeding under fast sea-ice in the shelf areas of Enderby Land in summer

Adélie penguins Pygoscelis adeliae in Enderby Land, Antarctica feed mainly on Euphausia superba during the chick rearing season in shelf areas where fast sea-ice remains: indicating that E. superba is abundant under the fast sea-ice in these areas. The shelf areas in Enderby Land, therefore, are unique since the previous studies of Adélie penguin diet in Ross Sea, Adélie Land and Prydz Bay show that E. crystallorophias is the most abundant krill species in shelf areas in general.     

Stomach contents of the Antarctic Silverfish Pleuragramma antarcticum from the southern and eastern Weddell Sea (Antarctica)

Summary The stomach contents of 142 Pleuragramma antarcticum from the southernmost part of the Weddell Sea (Gould Bay: 77°19S) and from the eastern coast (72°–74°S) were analyzed. The fish were collected in February 1983 and stored frozen. Size range of the investigated specimens was from 6.4 cm SL (7.3 cm TL) to 21.9 cm SL (24.1 cm TL). The fish were caught by bottom trawl (Gould Bay) and Agassiz Trawl and pelagic RMT-m net in the upper 300 m layer at the eastern coast, respectively. Thirty-six stomachs were empty or contained sand only. The most important prey in terms of biomass were euphausiids, which occurred in 49 stomachs at a mean number of 2. Their dry weight was estimated to be 15–50 times that of the next important food items, which were copepods, gastropods and gammarids. Krill (Euphausia superba) constituted up to 14% of the euphausiids in the diet of fish caught in the eastern Weddell Sea. The rest was usually made up by Euphausia crystallorophias. This species was taken with preference even when the abundance of E. superba in accompanying plankton catches was 16 times higher. In the Gould Bay, gastropods were frequently eaten despite high numbers of copepods in the plankton, whereas in the eastern Weddell Sea, copepods were abundant both in plankton and in the stomach content. The almost complete absence of the early stages of fish in the diet of Pleuragramma antarcticum in the southern and eastern Weddell Sea is due to a pronounced vertical segregation of fish of different sizes. This distribution pattern is thought to be an adaptive mechanism to avoid intraspecific predation, as 90% of the ichthyoplankton in that area is Pleuragramma antarcticum.     

Light and productivity of Antarctic phytoplankton during austral summer in an ice edge region in the Weddell-Scotia Sea

The photosynthesis-irradiance relationships (P-I curves) ofnatural plankton samples were studied in the Weddell Sea ice-edgezone, between Elephant Island and South Orkney Islands, duringthe austral summer of 1988–89. Three water bodies weredistinguished in the region: Bellingshausen Sea waters modifiedafter flowing through Drake Passage and Bransfield Strait, WeddellSea waters and Weddell Sea waters modified by melting. The stationssituated in modified Bellingshausen waters showed a net phytoplanktoncomposition which was different from that of the other two waterbodies. Weddell Sea waters and Weddell Sea waters modified bymelting of sea ice had the same net phytoplankton composition.In the area of modified Weddell Sea waters, there was an accumulationof phytoplankton in the upper 40 m (>4 mg Chl m^–1).p^B, and     

Hydrography and biochemical indicators of microplankton biomass in the Bransfield Strait (Antarctica) during January 1994

 The relationships between hydrography and spatial distribution of several biochemical indicators of microplankton biomass (chlorophyll, protein and ATP) were studied in an area covering the eastern part of the Bransfield Strait and the northern part of the Weddell Sea, during Antarctic summer (January 1994). Four hydrographic zones were identified: (a) the northern part of the Bransfield Strait, covered by waters of Bellings- hausen Sea origin; (b) a Weddell Sea water mass that affected most of the study area; (c) the Weddell-Scotia Confluence waters, observed north of Elephant Island; and (d) waters influenced by ice melting, found towards the southeastern part of the sampled area. The highest values of biomass indicators (chlorophyll a, ATP and protein) were found in the zones affected by ice-melting processes and in waters from the Bellingshausen Sea. The lowest values of all biochemical parameters were found in the Weddell Sea and in the Weddell-Scotia Confluence waters. A high variability in the hydrographic structure and the distribution of biochemical indicators was observed. The degree of stabilization of the water column, the depth of the upper mixed layer and the grazing pressure of herbivorous zooplankton played a major role in the development, accumulation and spatial variability of microplankton biomass. Received: 15 August 1995/Accepted: 18 February 1996     

Horizontal and vertical distribution and demography of euphausiids in the Ross Sea and its adjacent waters in 2004/2005

The horizontal and vertical distribution and population structure of euphausiids in the Ross Sea and its adjacent waters were investigated during the summers of 2004/2005 using stratified towed samples. Nine species of euphausiids occurred in the survey area. Among them, Euphausia triacantha was dominant in biomass north of the southern boundary of the Antarctic circumpolar current (SB). Thysanoessa spp. was widely distributed north of the continental slope, while E. superba was distributed from the SB to the slope, where it showed the highest biomass. Juvenile E. superba was distributed offshore near the SB and remained at the surface, but gravid females were dominant in the slope and mainly occurred in the middle layers (400–600 m). Adult and juvenile E. crystallorophias were found at 200–300 m in the colder water of the continental shelf. In general, the peak biomass of euphausiids was found in the mid layers of the Ross Sea area. The life span and the number of spawns for major species are also discussed.     

Trophic plasticity of Antarctic echinoids under contrasted environmental conditions

Echinoids are common members of Antarctic zoobenthos, and different groups can show important trophic diversity. As part of the ANT-XXIX/3 cruise of RV Polarstern, trophic plasticity of sea urchins was studied in three neighbouring regions (Drake Passage, Bransfield Strait and Weddell Sea) featuring several depth-related habitats offering different trophic environments to benthic consumers. Three families with contrasting feeding habits (Cidaridae, Echinidae and Schizasteridae) were studied. Gut content examination and stable isotopes ratios of C and N suggest that each of the studied families showed a different response to variation in environmental and food conditions. Schizasteridae trophic plasticity was low, and these sea urchins were bulk sediment feeders relying on sediment-associated organic matter in all regions and/or depth-related habitats. Cidaridae consumed the most animal-derived material. Their diet varied according to the considered area, as sea urchins from Bransfield Strait relied mostly on living and/or dead animal material, while specimens from Weddell Sea fed on a mixture of dead animal material and other detritus. Echinidae also showed important trophic plasticity. They fed on various detrital items in Bransfield Strait, and selectivity of ingested material varied across depth-related habitats. In Weddell Sea, stable isotopes revealed that they mostly relied on highly ¹³C-enriched food items, presumably microbially reworked benthic detritus. The differences in adaptive strategies could lead to family-specific responses of Antarctic echinoids to environmental and food-related changes.     

The developmental rate ofEuphausia crystallorophias larvae in Ellis Fjord,Vestfold Hills,Antarctica

 Euphausia crystallorophias is the dominant zooplankton species in the neritic seas of Antarctica, where it occurs in similar abundances to those of Euphausia superba in more offshore areas. Despite its great abundance and probable ecological significance, few details are known of this species’ development, life history and ecology. This study found that E. crystallorophias spawned in Ellis Fjord from late November to early December and completed its larval development under the sea ice during the Antarctic winter. The mean time for E. crystallorophias eggs to develop to furcilia stage VI was 235.5 days, which is virtually identical to the developmental time already reported in the laboratory, but almost twice that of E. superba. This slow development rate is likely to be due either to the low water temperatures (<0^°C) in which E. crystallorophias lives, or to low levels of food being available over winter. Received: 30 August 1995/Accepted: 11 December 1995     

Larval fish distribution and abundance in the Antarctic Peninsula region and adjacent waters

The spatial distribution, species composition and abundance of ichthyoplankton in the Bransfield Strait and adjacent waters were studied during two cruises in the Antarctic spring 1991/92 and summer 1992/93 seasons. A multiple plankton net (Bioness) and a Bongo net were used to collect samples at 35 stations in 1991/92 and 75 stations in 1992/93. Early larval stages (14 species) and juveniles (13 species) representing the known Bransfield Strait ichthyofauna were present in the water column. The nototheniids predominated in the entire study area. The greatest species diversity was found in the uppermost 200 m of the water column in the Bransfield Strait. Notothenia gibberifrons and Nototheniops larseni dominated in spring, whilst in summer Pleuragramma antarcticum dominated in association with N. larseni. The dominant species in the Gerlache Strait were P. antarcticum and Notothenia kempi, while P. antarcticum and Trematomus scotti were predominant in the Bellingshausen Sea area.     

Geographic structure in the Southern Ocean circumpolar brittle star Ophionotus victoriae (Ophiuridae) revealed from mtDNA and single‐nucleotide polymorphism data

Marine systems have traditionally been thought of as “open” with few barriers to gene flow. In particular, many marine organisms in the Southern Ocean purportedly possess circumpolar distributions that have rarely been well verified. Here, we use the highly abundant and endemic Southern Ocean brittle star Ophionotus victoriae to examine genetic structure and determine whether barriers to gene flow have existed around the Antarctic continent. Ophionotus victoriae possesses feeding planktotrophic larvae with presumed high dispersal capability, but a previous study revealed genetic structure along the Antarctic Peninsula. To test the extent of genetic differentiation within O. victoriae, we sampled from the Ross Sea through the eastern Weddell Sea. Whereas two mitochondrial DNA markers (16S rDNA and COI) were employed to allow comparison to earlier work, a 2b‐RAD single‐nucleotide polymorphism (SNP) approach allowed sampling of loci across the genome. Mitochondrial data from 414 individuals suggested three major lineages, but 2b‐RAD data generated 1,999 biallelic loci that identified four geographically distinct groups from 89 samples. Given the greater resolution by SNP data, O. victoriae can be divided into geographically distinct populations likely representing multiple species. Specific historical scenarios that explain current population structure were examined with approximate Bayesian computation (ABC) analyses. Although the Bransfield Strait region shows high diversity possibly due to mixing, our results suggest that within the recent past, dispersal processes due to strong currents such as the Antarctic Circumpolar Current have not overcome genetic subdivision presumably due to historical isolation, questioning the idea of large open circumpolar populations in the Southern Ocean.     

Ophiuroid biodiversity patterns along the Antarctic Peninsula

Benthic ecological surveys using standardized methods are crucial for assessing changes associated with several threats in the Southern Ocean. The acquisition of data on assemblage structure over a variety of spatial scales is important to understand the variation of biodiversity patterns. During the ANT XXIX/3 (PS81) expedition of RV Polarstern, three different regions at the tip of the Antarctic Peninsula were sampled: the northwestern Weddell Sea, the Bransfield Strait, and the northern boundary of the South Shetland Archipelago in the Drake Passage. The aim of this study was to characterize the distribution and biodiversity patterns of ophiuroid assemblages in these regions and depths. We quantified different community parameters in terms of the number of species, abundance, and biomass. Additionally, we calculated various components of species diversity (alpha, beta, and gamma diversity) over the three regions. Based on the benthic surveys, we collected 3331 individuals that were identified to species level (17 species). Overall, species diversity, as measured based on rarefaction, species richness and evenness estimators, was higher in the Bransfield Strait compared to the Weddell Sea and Drake Passage. Two deep stations in the Weddell Sea showed high dominance only of Ophionotus victoriae. Significant differences in the patterns of alpha diversity were found among the regions but not between depth zones, whereas beta diversity showed no differences. Regarding the resemblance among the ophiuroid assemblages of each region, there was a significant gradient from east to west with a maximum distance between the stations in the Drake Passage and the Weddell Sea. This study provides a baseline for detecting potential effects related to climate change, and it furnishes a basis for the implementation of monitoring schemes of Antarctic assemblages.