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.     

Feeding and diel vertical migration cycles of Metridia gerlachei (Giesbrecht) in coastal waters of the Antarctic Peninsula

Diel vertical migration and feeding cycles of adult female Metridia gerlachei in the upper 290 m of a 335-m water column were measured during a total of 65 h in two periods of early summer (Dec 20–21 and Dec 25–26, 1991). Samples collected in eight depth strata by 35 MOCNESS tows (333-m mesh) were analyzed for abundance and mean individual gut pigment content. Most of the copepod population was concentrated in a 50-m depth interval at all times. Feeding began simultaneously with nocturnal ascent from a depth of 200–250 m at 18:00 h (local time), when the relative change in ambient light intensity was greatest. Ingestion rate increased exponentially (k_i = 0.988 h^–1) at double the gut evacuation rate (k_e = 0.488 h^–1) as the population moved upward at 22.3–26.5 m h^–1 through increasing concentrations of particulate chlorophyll-a. Although the bulk of the population did not move to depths shallower than 50 m, and began its downward migration at a rate of 20.8–31.7 mh^–1 in complete darkness, individual females continued to make brief excursions into chlorophyll-rich surface waters (4–8 g l^–1) during the first few hours of population descent. Ingestion rate diminished abruptly by one order of magnitude (k_i = 0.068 h^–1) at dawn ( 0330 h). Within four more hours, the population had reached its daytime depth and gut pigment content remained constant at a minimum value until the next migration cycle. No feeding appeared to take place at depth during the day. Ingestion by M. gerlachei females removed < 4% of daily primary production, with only 20% of this amount being removed from surface waters by active vertical transport.     

Feeding ecology of Antarctic fur seals at Cape Shirreff,South Shetlands,Antarctica

This study examined the diet of Antarctic fur seals, Arctocephalus gazella, from an active breeding colony at Cape Shirreff (62°28S, 60°48W), Livingston Island, South Shetland Archipelago, Antarctica. It analysed faecal samples from five consecutive years (1997–2001) and length distribution of krill taken by trawl nets in the vicinity of Livingston Island. Antarctic krill, Euphausia superba, was the most frequent prey item, followed by several myctophid species (Gymnoscopelus nicholsi, Electrona antarctica and Electrona carlsbergi), squid and penguin remains. From 1998 to 2001, a modal progression in krill size was evident, suggesting that A. gazella was depending on a strong krill cohort, at least over the study period. Analysis of size distribution and size selectivity of krill preyed upon by fur seals suggests a preference for larger krill (>34 mm), despite the broader size range of preys items available.     

Food and daily food consumption of southern minke whales in the Antarctic

Summary The stomach contents of 273 southern minke whales (Balaenoptera acutorostrata) taken in the region 55°S to the ice-edge and between 105°E and 115°E by a Japanese survey during 1987/88 were examined. The minke whales' dominant feeding ground coincided with a distinctive hydrographic front in the vicinity of the ice-edge at the mouth of Vincennes Bay. Krill (Euphausia superba) were the dominant food species comprising 100% and 94% by weight of stomach contents in the ice-edge and offshore zones, respectively. In the offshore zone, minke whales tend to feed on E. superba rather than Thysanoessa macrura, which is found more frequently than the former in net samples. Total food consumption by minke whales per day was estimated to be 1170 t (22.1 kg/km²) and 596 t (2.0 kg/km²) in the ice-edge and offshore zones, respectively. Feeding activity peaked in the early morning in the ice-edge zone, whereas it occurred irregularly throughout the day in the offshore zone. Two size modes of krill (25–28 mm in body length (1 year-old) and 41–48 mm (3–4 year-old)) dominated the diet. Although the former was numerically more abundant, the latter dominated the diet by weight, suggesting that larger krill are more important food for minke whales. Observed spatial pattern of krill populations in the study region suggests that movement of the subsurface cold water mass tended to carry krill offshore from the ice-edge.The earlier version of this paper was submitted to the 42nd Meeting of the Scientific Committee of International Whaling Commission as SC/42/SHM; 34.     

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.     

Variability of Euphausia superba populations near Elephant Island and the South Shetlands: 1981 vs. 1984

Summary March 1981 and 1984 Euphausia superba populations were compared using acoustics and net catches near Elephant Island, the South Shetlands, and in the Bransfield Strait. In 1981, krill tended to form large, thick swarms and in 1984, smaller, more dispersed, shallower patches. March body lengths of juvenile-adult krill were 22–59 mm in 1981 and 13–59 mm in 1984. Near Elephant I. in 1981, krill >45 mm were most numerous; in 1984 sizes <45 mm were dominant. In March 1984, the larger (>45 mm) body-size group was prereproductive and occurred from just west of Elephant I. westward into waters north of the South Shetlands; in 1981 the larger krill were postreproductive and more widely distributed in the Elephant I. area. Overall, netted postlarval krill, 1981 vs. 1984, averaged 73 vs. 48 individuals/m², or 54 vs. 16 g/m²; acoustic biomass estimates were 229 vs. 134–201 g/m². Larvae near Elephant I. averaged >2000/m² in 1981 vs. <1/m² in 1984—compatible with respective March reproductive states. Net-type comparisons revealed short-term (15 min to 6h) variability of a similar scale in both MOCNESS and bongo net catches, but bongo abundances averaged greater. Variation in maturity composition across 1981 swarms, patches, and random transects was like variation among the random 1984 tows; spatial distributions were more heterogeneous in 1984. The March 1984 krill of 20–44 mm (Year-2, mode 34 mm) relate to November 1983 krill of 9–30 mm (mode 21 mm), indicating growth averaging 12 mm during the season. Body-lengths and size-frequency modes of Year-2 and combined Years-3,3+ krill from comparable Feb-Mar data collected since 1968 suggest trends between times when (1) Year-2 krill average small and peak reproduction seems to be late in the season and/or weak (1979, 1982–1984), and (2) Year-2 krill are larger, and reproduction is possibly earlier and more successful (1976, 1980, 1981).     

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.     

Occurrence of cyclopoid copepods and faecal material in the Halley Bay region,Antarctica, during January–February 1991

Faecal material and cyclopoid copepods were collected during the expedition ANT IX/3, in the Halley Bay area (Weddell Sea, Antarctica), between January and February 1991. Faecal material comprised pellets produced by krill, copepods, ostracods and appendicularians. Cyclopoid copepods were represented by two genera, Oithona and Oncaea. In the Halley Bay area, higher concentrations of krill faecal material (420.9 mm³ m^–2) and chl.-a (39.3 mg m^–2) were found within the upper 200 m of the water column of the polynya than in ice-covered open-ocean areas (58.2 mm³m^–2 and 25.5 mg m^–2, respectively). At an ice-drift station, high concentrations of krill faecal strings under fest-ice were found. In addition, similarities between diatom assemblages in the pack-ice algae and krill faecal strings contents suggest an active utilization of ice-algae by krill populations. Sedimented material collected at 50 m depth by a sediment trap was dominated by krill faecal strings. Contents of small oval pellets (of probable cyclopoid copepod origin) resemble those of krill faecal pellets suggesting that coprophagy was involved. This suggestion is supported by: (1) The small quantity of food particles (other than krill faecal matter) available in the water column (< 0.3 g chl.-al^–1). (2) The negative in situ correlation between krill faecal strings and cyclopoid copepods. (3) The structure of cyclopoid copepod buccal appendages, which are more adapted for raptorial feeding.     

Feeding rate inhibition in crowded Daphnia pulex

Feeding rates of Daphnia pulex fed a range of levels of the alga Chlamydomonas reinhardi of 15 °C are strongly density-dependent. At lower densities, Daphnia (30 1^–1) fed at higher rates than crowded (270 1^–1) Daphnia which manifest a relatively depressed saturation feeding response. At 30 individuals/liter, Daphnia consumed 8.5 – 15.7 × 10⁴ cells d^–1h^–1 (on a volume basis, 12.1 – 22.2 × 10⁶ m³), at 270 L^–1 3.7 – 3.9 × 10⁴ (5.2 – 5.5 = 10⁶ m³ cells d^–1h^–1 when feeding on algae at 80 000 cells ml^–1 (11.3 × 10⁶ m³ ml^–1). The feeding rate data best fit an Ivlev feeding function. An autoallelopath might be causing the repression. Water preconditioned with crowded Daphnia completely repressed feeding in uncrowded Daphnia after six hours.     

Moulting interval and growth of juvenile Antarctic krill (Euphausia superba) fed different concentrations of the diatom Phaeodactylum tricornutum in the laboratory

Summary Juvenile Antarctic krill (Euphausia superba) were fed the diatom Phaeodactylum tricornutum at concentrations ranging from 0.0–5.0 mgCl^-1. Intermoult period (IP) decreased, but an increment of body length per moult (BL) of juvenile krill increased, up to a concentration of 1.0 mgCl^-1. No further effect of food concentrations on IP or BL was seen at concentration beyond 1.0 mgCl^-1. IP plateaued at 23.8 days and BL, 1.14 mm. The maximum daily growth rate (BL/IP) of juvenile krill was calculated to be 0.047 mm day^-1. BL and IP were correlated and the relationship is expressed as BL=-0.0066IP+2.47 (r=0.58, n=141, P<0.01). Growth conditions of krill in the Southern Ocean are discussed in terms of available food concentration in there.     

Interannual and between species comparison of the lipids, fatty acids and sterols of Antarctic krill from the US AMLR Elephant Island survey area

Antarctic euphausiids, Euphausia superba, E. tricantha, E. frigida and Thysanoessa macrura were collected near Elephant Island ¦ during 1997 and 1998. Total lipid was highest in E. superba small juveniles (16 mg g⁻¹ wet mass), ranging from 12 to 15 mg in other euphausiids. Polar lipid (56–81% of total lipid) and triacylglycerol (12–38%) were the major lipids with wax esters (6%) only present in E. tricantha. Cholesterol was the major sterol (80–100% of total sterols) with desmosterol second in abundance (1–18%). 1997 T. macrura and E. superba contained a more diverse sterol profile, including 24-nordehydrocholesterol (0.1–1.7%), trans-dehydrocholesterol (1.1–1.5%), brassicasterol (0.5–1.7%), 24-methylenecholesterol (0.1–0.4%) and two stanols (0.1–0.2%). Monounsaturated fatty acids included primarily 18:1(n−9)c (7–21%), 18:1(n−7)c (3–13%) and 16:1(n−7)c (2–7%). The main saturated fatty acids in krill were 16:0 (18–29%), 14:0 (2–15%) and 18:0 (1–13%). Highest eicosapentaenoic acid [EPA, 20:5(n−3)] and docosahexaenoic acid [DHA, 22:6(n−3)] occurred in E. superba (EPA, 15–21%; DHA, 9–14%), and were less abundant in other krill. E. superba is a good source of EPA and DHA for consideration of direct or indirect use as a food item for human consumption. Lower levels of 18:4(n−3) in E. tricantha, E. frigida and T. macrura (0.4–0.7% of total fatty acids) are more consistent with a carnivorous or omnivorous diet as compared with herbivorous E. superba (3.7–9.4%). The polyunsaturated fatty acid (PUFA) 18:5(n−3) and the very-long chain (VLC-PUFA), C₂₆ and C₂₈ PUFA, were not present in 1997 samples, but were detected at low levels in most 1998 euphausiids. Interannual differences in these biomarkers suggest greater importance of dinoflagellates or some other phytoplankton group in the Elephant Island area during 1998. The data have enabled between year comparisons of trophodynamic interactions of krill collected in the Elephant Island region, and will be of use to groups using signature lipid methodology.     

Experiments on the physiology of southern and northern krill,Euphausia superba and Meganyctiphanes norvegica,with emphasis on moult and growth – a review

Physiological data are needed for life history studies on krill, and as parameters for input into energy budgets and models. In conjunction with moult and growth data, these may also prove useful for assessing the fishable biomass of krill. Here, the development of physiological concepts in experimental krill research is briefly evaluated, with emphasis on the gaps to be filled. Krill growth is very flexible, as well as strongly temperature and nutrition dependent. The polar Antarctic krill Euphausia superba grows as fast as the boreal species Meganyctiphanes norvegica, at least during the first 2.5 years, and the species are comparable in terms of physiological plasticity. Accordingly, as krill appear to adjust quickly to specific laboratory conditions, short-term experiments are essential if field conditions are to be reflected as closely as possible. Furthermore, direct comparisons between laboratory experiments and swarming studies in the field are advantageous. For these, M. norvegica is particularly well-suited, as swarms can be followed over longer times and more easily than in E. superba. For example, processes of moult and reproduction were found to be highly coordinated in swarms and populations of Northern krill. For this species a conceptual model of reproduction was developed based on a combination of short-term laboratory observations coupled with field data on moult and ovary stages. In further physiological experiments krill should be studied as groups when swarming. Using proxies, that is applying physiological and/or biochemical methods side by side, is a promising way to enhance the reliability of life history data.     

Diel distribution of copepods across a channel of an overwash mangrove island

The distribution of copepod species and their nauplii was studied in a narrow, blind channel on an overwash mangrove island offshore of Belize. Copepodids were sampled with a pump at five stations across the channel during a diel cycle. Diel changes of copepodid stages II – VI were marked by horizontal dispersal of Dioithona oculata, the dominant species, from swarms in the prop roots along the shore during the day to the edge of the prop root habitat at night. Migration of copepodids back to the prop roots appeared to be controlled endogenously because change from a night to a daytime age structure began before first light. Mean copepodid stage at subsurface depths in the channel and prop root edge decreased from 4.2 (with 6.0 = all adults) to 2.9 at predawn to 1.1 during day. The oceanic Oithona nana and O. simplex, and the coastal zone O. fonsecae were evenly distributed with depth and distance from shore during day and night, with avoidance of prop root shoreline during day. These species were much less abundant than Dioithona oculata in the prop roots, but of comparable or greater abundance in the channel. Coastal zone Acartia spinata exhibited evidence of swarming. Nauplii, sampled with a 25m plankton net, were dominated by harpacticoid (50%) and cyclopoid (34%) nauplii, which generally were more abundant at 1m than at the surface and more abundant at night than the day. Lagrangian current measurements indicated velocities at ebb tide twice those of flood tide (1.9 vs. 0.8 cm s^–1) and a minimal residence time of 5 days, which could result in advection of D. oculata nauplii out of the Lair Channel before their recruitment into swarms as copepodid stage II. Previously reported maximum swimming speeds of swarming D. oculata copepodid stages (2.0 cm s^–1) and greater densities in prop roots and near the benthos may help copepodids avoid advection. The swarming behavior and diel horizontal migration (or dispersal) reported for D. oculata appears analogous to that of limnetic zooplankton, which may swarm among macrophytes along shorelines during the day to avoid visual predators and disperse or migrate away from the shoreline at night.     

Organic carbon in the upper layer and its sedimentation during the ice-retreat period in the Scotia-Weddell Sea, 1988

Summary The maximum of POC in the surface layer (200–400 g·l^–1) followed the retreating ice from end November to end December. In the upper 150 m DOC amounted to 10–20 times the POC content. Free floating sediment traps at 150 m showed a daily sedimentation of 0.6–11.6% of the standing stock of POC, 0.1–2.6% of the chlorophyll-a and 5–190% of the gross primary production. Maximum sedimentation occurred during grazing of a krill swarm, indicating the important role of krill swarms in the downward flux. Also at most other stations krill faecal strings formed a large part of the downward flux.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

Micronekton and macrozooplankton in the open waters near Antarctic ice edge zones (AMERIEZ 1983 and 1986)

Summary Micronekton and macrozooplankton assemblages (0–1000 m) were sampled from the open ocean in the vicinity of marginal ice zones in the southern Scotia and western Weddell Seas using midwater trawls. Small regional differences in species composition were found in the differing hydrographic settings with the Scotia Sea being slightly more diverse. Most species exhibited broad vertical ranges with no distinct pattern of vertical movement. Exceptions were mesopelagic fish and Salpa thompsoni which undertook diel vertical migrations. Biomass was high (2.4–3.1 g DW/m²), comparable to Pacific subarctic waters. Euphausia superba and Salpa tompsoni were the numerical and biomass dominants, representing over 50% of the total numbers and standing stocks. In terms of biomass, euphausiids were the most important group at shallow depths (0–200 m) but were surpassed by salps in the Scotia Sea and mesopelagic fish in the Weddell Sea when all depths down to 1000 m were considered. Pelagic fish biomass (3.3–4.4 g WW/m²) greatly exceeded published estimates for birds (0.025–0.070 g WW/m²), seals (0.068–0.089 g WW/m²) and whales (0.167 to 0.399 g WW/m²), making mesopelagic fish the most prevalent krill predators in the Antarctic oceanic system.     

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     

Colorimetry of the hepatopancreas in Antarctic krill,Euphausia superba

Summary Hepatopancreas color of freshly caught as well as starved Euphausia superba was examined by a chroma meter to express it quantitatively and compare it with plant pigment contents. There was a high positive correlation (r= 0.77) between plant pigment contents of hepatopancreas and the purity (vividness) of hepatopancreas color, and a negative correlation between the pigment contents and the luminosity (brightness) of the color. During the starvation experiment the purity decreased from 0.480 to 0.198 during the first 6 days in accordance with the decrease in plant pigment contents from 4.7 to 0.9 g krill^–1. An examination of hepatopancreas color by a chroma meter proved an easy and quantitative way of knowing the feeding condition of E. superba.     

Grazing by a dominant rotifer Conochilus unicornis Rousselet in a mountain lake: in situ measurements with synthetic microspheres

Grazing rates of zooplankton were analysed in the summer of 1999 in Yellow Belly Lake, an oligotrophic system in the Sawtooth Mountains of Idaho (U.S.A.). The colonial rotifer Conochilus unicornis was a dominant species in the epilimnion, with densities reaching 20 colonies l^–1 (ca. 400 ind. l^–1). Clearance rates were measured with an in situ Haney Grazing chamber and synthetic microspheres 5, 9 and 23m in diameter. At epilimnetic temperatures of around 14 °C, mean clearance rates for 5m particles ranged from 30 to 65 l ind.^–1 h ^–1. Clearance rates were 2–9 times higher on the 5m spheres than on the 9 m spheres, and C. unicornis almost never fed on the 23 m spheres. Grazing rates did not change over the diel cycle. Clearance rates declined more than 10-fold as temperatures declined from 14 °C in the epilimnion to 7 °C in the metalimnion. In the epilimnion, grazing by C. unicornis was more important than grazing by crustaceans in the community, at least on particles 9m. The results show the importance of grazing by rotifers in lakes, and the significance of spatial variations that influence grazing rates.     

Breeding success and chick provisioning in Wilson’s storm-petrels <Emphasis Type="Italic">Oceanites oceanicus</Emphasis> over seven years: frequent failures due to food shortage and entombment

We examined the natural variation in parameters of the breeding success of Wilsons storm-petrels on King George Island, South Shetland Islands, during seven breeding seasons, and determined the relative influence of krill abundance and climatic conditions. Feeding frequencies, feeding rates and meal sizes varied between years, and feeding rates were correlated with data of the Antarctic krill Euphausia superba abundance in the Elephant Island region. The overall breeding success of breeding pairs was low, ranging from 0 to 34%. Snow storms and wind conditions depressed breeding success even in years of good food availability, and snow storms were the most important determinant of the breeding success over the seven seasons. Snow storms during the breeding season occurred during periods of prevailing easterly winds. In addition, the distribution of wind directions was found to strongly influence the krill availability to storm-petrels.     

Synchronous moulting of krill,<Emphasis Type="Italic"> Euphausia superba</Emphasis>, in the Bransfield Strait (Antarctica)

A large mass of cast exoskeletons of the Antarctic krill Euphausia superba was fished in February 2001 at one station in the Bransfield Strait between 411 and 153 m. We assume that the mass of cast exoskeletons originated from a synchronous moult of thousands of individuals. Depending on the size of the krill moulted, the amount of exoskeletons corresponded to a krill abundance of 1,480–48,000 individuals per 1,000 m³.