Functional biodiversity of lipids in Antarctic zooplankton: Calanoides acutus, Calanus propinquus, Thysanoessa macrura and Euphausia crystallorophias

Zooplankton samples were collected in January 1993 off Dronning Maud Land along a transect from open waters to the marginal ice zone close to the Antarctic ice shelf. Thysanoessa macrura was caught in open waters while Calanoides acutus and Calanus propinquus were mainly sampled between ice floes in the marginal ice zone. The “ice-krill”Euphausia crystallorophias was found over the shelf directly associated with ice floes. T. macrura had a lipid content up to 36% of its dry weight with the dominant lipid class, wax ester, accounting for 45–50% of the total lipid. The predominance of 18:1 fatty alcohols is the striking characteristic of the wax esters. Small specimens of E. crystallorophias had lipid levels up to 26% of their dry weight with, unexpectedly, triacylglycerols being the dominant lipid (up to 41% of total lipid). The small levels of wax esters in these animals (3–6% of total lipid) had phytol as a major constituent. Large specimens of E. crystallorophias had up to 34% of their dry weight as lipid, with wax esters (47% of total lipid) dominated by 16:0 and 14:0 fatty alcohols as the major lipid. Calanus propinquus had lipid levels of up to 34% of their dry weight, with triacylglycerols (up to 63% of total lipid) being the dominant lipid. High levels of 22:1 (n-9) fatty acid were present in the triacylglycerols. Calanoides acutus had lipid levels up to 35% of the dry weight with wax esters accounting for up to 83% of total lipid. High levels of (n-3) polyunsaturated fatty acids were recorded with 20:5(n-3), 22:6(n-3) and 18:4(n-3) being the dominant moieties. On the basis of their lipid compositions we deduce that: (1) Calanoides acutus is the strictest herbivore among the four species studied, heavily utilizing the typical spring bloom; (2) T. macrura is essentially omnivorous, probably utilizing the less defined bloom situations found in oceanic waters; (3) E. crystallorophias is an omnivore well adapted to utilize both a bloom situation and to feed on ice algae and micro-zooplankton associated with the ice; (4) Calanus propinquus seems to be the most opportunistic feeder of the four species studied, probably grazing heavily on phytoplankton during a bloom and, during the rest of the year, feeding on whatever material is available, including particulates, flagellates and other ice-associated algae. We conclude that the different biochemical pathways generating large oil reserves of different compositions, enabling species to utilize different ecological niches, are major determinants of biodiversity in polar zooplankton. Accepted: 22 June 1998     

Physiological characteristics of the antarctic copepod Calanoides acutus during late summer in the Weddell Sea

Ingestion, respiration and excretion rates as well as lipid and protein body content of the dominant Antarctic copepod Calanoides acutus (CIV to adult females) were studied during the period covering the end of phytoplankton bloom (February) to the beginning of transition to overwintering (March-April). Daily rations measured with gut fluorescence varied from 2.2 to 2.7% in surface C. acutus. Weight-specific respiration and excretion rates in deep C. acutus decreased by a factor of 11 and 3.5–3.8 compared to their surface counterparts. High lipid (up to 455 µg ind^–1) and protein (198 µg ind^–l) content was observed in surface C. acutus CV in February; a month later the animals with similar lipid and protein content were found in the depth (500–1000 m layer). Their lipid reserves were enough to overwinter and probably to ascend, molt and reproduce. At the same time some of the deep CVs had much lower protein and lipid content and could survive only for 4–5 months. Our own and literature data led to the conclusion that females of C. acutus reach adulthood at the age of more than one year while development of males could be completed in one year.     

Gut evacuation rates of Antarctic copepods during austral spring

The results of gut evacuation experiments performed on Antarctic copepods during the austral spring are presented and discussed. Four species of large copepods commonly occurring in the Indian sector of the Antarctic Ocean were studied: Calanus propinquus, Calanoides acutus, Rhincalanus gigas and Pleuromamma robusta. For each species two experiments were carried out, one in daytime and one in night-time, except for Calanoides acutus, which was only studied at night. P. robusta showed pigment gut retention in all experiments. The results showed that all species studied had a longer gut passage time than that previously recorded and that gut evacuation rate appears to decrease during daytime. Accepted: 1 October 1998     

Egg hatching times of Antarctic copepods

Egg hatching times were determined at a range of temperatures for four species of commonly occurring Antarctic copepods. At a given temperature the eggs of Rhincalanus gigas took longest to hatch, up to 9 days at 0°C, followed by those of Calanoides acutus, Calanus propinquus and Calanus simillimus. A Bělehrádeks temperature function with the parameter b fixed at −2.05 accounted for >95% of the variance for each species. There was an approximate doubling in hatching times between 5°C and 0°C for R. gigas and for the other species the increase in embryonic duration was 40–50% at the lower temperature. Received: 27 March 1997 / Accepted: 17 August 1997     

The occurrence of Rhincalanus gigas,Calanoides acutus,and Calanus propinquus (Copepoda: Calanoida) in late May in the area of the Antarctic Peninsula

Summary The Antarctic copepod species Rhincalanus gigas, Calanoides acutus and Calanus propinquus were studied in the area of the Antarctic Peninsula in May 1986. Research was focussed on vertical distribution and stage composition of the populations. Rhincalanus gigas occurred in greatest densities in the upper layers of the water column, and copepodite stages CI and CII and nauplii dominated the population. Gut content analyses suggest that R. gigas was actively feeding. Copepodite stage CV dominated the Calanoides acutus population. At two deep basin stations (water depth>1000 m) the C. acutus population occurred below 500 m, whereas at shallower stations the majority was found above 300 m. Most specimens had empty guts. Calanus propinquus occurred in low densities, mainly in the upper water layers, and copepodite stage CV dominated. Most individuals of stage V had food in their guts. Our results suggest that C. acutus had ceased feeding and was overwintering in a resting stage (diapause), while C. propinquus and R. gigas were still active, the latter species having finished a late autumn spawning.     

Winter distribution and overwintering strategies of the Antarctic copepod species Calanoides acutus,Rhincalanus gigas and Calanus propinquus (Crustacea,Calanoida) in the Weddell Sea

During the Winter Weddell Gyre Study in September–October 1989, the horizontal and vertical distribution, stage composition and feeding condition of the three antarctic copepod species Calanoides acutus, Rhincalanus gigas and Calanus propinquus were studied. The data indicate that C. acutus and R. gigas have the bases of their distributional ranges (sensu Makarov et al. 1982) in the Antarctic Circumpolar Current (ACC) and in the Warm Deep Water (WDW) entering the Weddell Gyre (WG). C. propinquus lived mainly in the cold WG south of the ACC. C. acutus overwintered mainly in the WG as stage IV copepodites (C). The species mainly inhabited the layers below the T_max stratum and down to 2000 m, but C V and females occurred slightly higher than C III and IV. Males prevailed over females and were confined to a rather narrow layer between 500 and 1000 m. Feeding experiments suggested all deep-living stages to be resting. However, if this species spawns in late autumn the younger C I–II can stay in the Winter Water (WW). R. gigas inhabited mainly the T_max stratum. In the eastern part of the WG, R. gigas breed in the WDW in autumn and hibernate as C I–III and C V–VI in the first and second winter, respectively. In the ACC zone, however, its life cycle is different and winter breeding of overwintered adults occurs. Most of the C. propinquus population overwintered in the WG as C III–V, inhabiting the WW. In the upper water layers in the interior of the WG, C III dominated with upto 18,000 individuals 1,000 m³. Shallow living C. propinquus were in the active, feeding state. Persistence of active feeding zooplankton populations in the WW of the WG can be an important factor influencing processes of phytoplankton development and the particle flux.     

Qualitative models of the life cycles of Calanoides acutus,Calanus propinquus,and Rhincalanus gigas

Summary The life cycle of Calanoides acutus, Calanus propinquus, and Rhincalanus gigas was analyzed in samples collected by five expeditions from 1963 to 1984 in the Scotia and Weddell Seas. The result of this analysis, and of published data suggest that C. acutus and C. propinquus have a one year life cycle. Males of C. acutus are present in the population for a short period of time during late winter and fertilization occurs in deep waters. C. propinquus mates in the surface and males are present in the population throughout the summer. Part of its population may remain in the surface waters in winter. R. gigas has a more flexible, one or two year life cycle. Late summer spawning may occur in this species.This is contribution No. 95 of the Alfred-Wegener-Institut für Polar- und Meeresforschung     

Testing hypotheses on life-cycle models for Antarctic calanoid copepods, using qualitative, winter, zooplankton samples

We analysed qualitative winter zooplankton samples, collected from the ocean surface, to test portions of proposed life-cycle models for Calanus propinquus. Nine zooplankton hauls were taken in the Bransfield Strait area during June 1996. Results show that C. propinquus is present in the ocean surface in winter. However, its presence seems to be related to ice coverage below 30%. Received: 31 October 1997 / Accepted: 23 February 1998     

Trophic ecology of Calanoides acutus in Gerlache Strait and Bellingshausen Sea waters (Antarctica, December 2002)

We measured ingestion rates of Calanoides acutus on different microbial components of the Gerlache Strait (GE) and Bellingshausen Sea (BE) waters during December 2002. At the time of the study the abundance of both zooplankton (42–133 ind m⁻³) and phytoplankton (0.76–1.5 µg chlorophyll a l⁻¹) were low, indicating that the spring phytoplankton bloom was still not fully developed. C. acutus showed high clearance rates along the study (up to 432 ml ind⁻¹ day⁻¹), selecting for large motile organisms such as ciliates and the dinoflagellate Gyrodinium spp., although their feeding impact was always <0.1% of the standing stock of any of their prey. The total daily rations were low (∼2% body carbon per day), mostly the result of phytoplankton consumption (except for station GE3 in which heterotrophic flagellates contributed to 73% of the diet), and barely enough to cover metabolic demands. Based on the relationship between oxygen (carbon) consumption and ammonia excretion (considered as indicative of the metabolic substrate) it seems that standard metabolic demands were supplied, apart from the diet, by the use of their own non-structural proteins, whereas the remaining reserve-lipids were used to produce eggs.     

Unexpected weight gain in resident Antarctic Terns Sterna vittata during the austral winter

Antarctic Terns Sterna vittata (Fig. 1) may display two very different migratory behaviours. According to Cooper (1976), the terns breeding on Tristan da Cunha in the South Atlantic fly thousands of kilometers to the coasts of Africa where they moult. Antarctic Terns that inhabit the more southern and colder Antarctic Peninsula region moult on their breeding grounds. An early account of these seemingly resident peninsular terns (Holdgate 1963) leaves little doubt that at least part of the tern population wintered in the vicinity of Arthur Harbor (64°46'S 64°03'W), Anvers Island, west of the Antarctic Peninsula. Watson (1975) stated that adult Antarctic Terns are generally sedentary around many insular breeding stations, moving only to the nearest open water in winter. During the years 1975 through 1978, 19 tern specimens of different sexes and ages were collected at Arthur Harbor in the non-breeding season near U.S. Palmer Station. An additional eight terns were collected at sea a short distance from Anvers Island during the non-breeding season in 1985 (Pietz & Strong, in press). I found that the adult birds taken at the height of winter weighed significantly more than the 150–180 g of a normal breeding Antarctic Tern. This unexpected discovery prompted me to examine the weights of an additional 34 specimens that had been collected at or near Anvers Island during several breeding seasons. By comparing the weights of adult terns by sex, age, and collection date (Fig. 2, Table 1) I found that both males and females weighed significantly more in the winter non-breeding season (April-September) than in the summer breeding season (October-March) (t₂₄ = 6·57, P < 0·001, and t₁₆ = 5·71, P < 0·001, respectively). No significant differences were detected between male and female weights in summer (t₂₃ = 0·76, P > 0·20) or winter (t₁₇ = 1·16, P > 0·20). In short, it appears that body-weights of adult terns rise rather dramatically following breeding, attain a peak in mid-winter, and then fall at the approach of the next breeding season (Fig. 2). I suggest that this increased body-weight is an adaptation to the austral winter rather than simply a recovery from weight loss due to energetic costs of breeding and moulting.     

Unexpected weight gain in resident Antarctic Terns Sterna vittata during the austral winter

Antarctic Terns Sterna oittata (Fig. 1) may display two very different migratory behaviours. According to Cooper (1976), the terns breeding on Tristan da Cunha in the South Atlantic fly thousands of kilometers to the coasts of Africa where they moult. Antarctic Terns that inhabit the more southern and colder Antarctic Peninsula region moult on their breeding grounds. An early account of these seemingly resident peninsular terns (Holdgate 1963) leaves little doubt that at least part of the tern population wintered in the vicinity of Arthur Harbor (64°46′S 64°03′W), Anvers Island, west of the Antarctic Peninsula. Watson (1975) stated that adult Antarctic Terns are generally sedentary around many insular breeding stations, moving only to the nearest open water in winter. During the year 1975 through 1978, 19 tern specimens of different sexes and ages were collected at Arthur Harbor in the non-breeding season near U.S. Palmer Station. An additional eight terns were collected at a sea short distance from Anvers Island during the non-breeding season in 1985 (Pietz & Strong, in press). I found that the adult birds taken at the height of winter weighed significantly more than the 150-180g of a normal breeding Antarctic Tern. This unexpected discovery prompted me to examine the weights of an additional 34 specimens that had been collected at or near Anvers Island during several breeding seasons. By comparing the weights of adult terns by sex, age, and collection date (Fig. 2, Table 1) I found that both males and females weighed significantly more in the winter non-breeding season (April-September) than in the summer breeding season (October-March) (t₂₄= 6.57, P < 0.001, and t₁₆= 5.71, P < 0.001, respectively). No significant differences were detected between male and female weights in summer(t₂₃=0.76, P >0.20) or winter (t₁₇=1.16, P0.20). In short, it appears that body-weights of adult terns rise rather dramatically following breeding, attain a peak in mid-winter, and then fall at the approach of the next breeding season (Fig. 2). I suggest that this increased body-weight is an adaptation to the austral winter rather than simply a recovery from weight loss due to energetic costs of breeding and moulting.     

Aspects of the study of the life cycles of Antarctic copepods

Different approaches to the study of life cycle strategies of Antarctic copepods are described in an attempt to shed new light on our present knowledge. To date, most studies were carried out on abundance, horizontal and vertical distribution and stage composition during different seasons and in various regions. Hence, the seasonal pictures had to be compiled from different years and sampling regions. The physiological method includes measurements on e.g. egg production, feeding, respiration and excretion rates, C:N and O:N ratios, lipid and protein contents. However, both physiological and biochemical data are still rare. Results of field observations are given in this paper for investigations conducted within the last 15 years in the eastern Weddell Sea, while data of physiological parameters are based on a broader geographical region. In the eastern Weddell Sea, eight copepod species account for about 95% of copepod abundance and for more than 80% of copepod biomass. Within the calanoids, the small species Microcalanus pygmaeus dominates by numbers with 66%, while the large species Calanoides acutus and Calanus propinquus comprise together 52% of the biomass. Species abundance is lowest in winter and highest in summer/autumn, however, seasonal changes in the abundance of M. pygmaeus are small and this species occurs in similar quantities throughout the year. All copepod species show a distinct seasonal vertical distribution pattern and they occur in upper water layers in summer, in contrast to the other seasons. However, the depth layers of maximum concentration differ between species. The ontogenetic vertical migration is most pronounced in C. acutus and relatively weak in C. propinquus. The age structure also shows seasonal differences with the youngest population observed in summer for C. acutus, C. propinquus, Ctenocalanus citer or autumn for Metridia gerlachei, whereas the M. pygmaeus population is oldest during summer. The youngest copepodite stage and the males are not always present in C. acutus and C. propinquus. In contrast, all developmental stages and both sexes occur throughout the year in M. gerlachei, M. pygmaeus and C. citer. Gonad maturation in the dominant calanoid species proceeds well before the onset of phytoplankton production in the eastern Weddell Sea. However, the highest portion of females with ripe gonads and hence highest egg production rates coincide with the productive period in spring and summer. In autumn, ovaries of the three larger species C. acutus, C. propinquus and M. gerlacheiare all spent. In contrast, the percentage of ripe females of the two smaller species, C. citer and M. pygmaeus, stays high in autumn. Egg production rates are highly variable within one region and species. Many copepods accumulate large depots of lipid, mainly wax esters. In contrast, five species (C. propinquus, C. simillimus, Euchirella rostromagna, Stephos longipes and Paralabidocera antarctica) almost exclusively synthesise triacylglycerols and not wax esters. The lipid content exhibits distinct seasonal patterns, and is highest in autumn. A seasonal difference is also obvious in metabolic activities with lowest rates during the dark season. The adaptation to the pronounced seasonality in the Southern Ocean differs greatly between copepod species, and most Antarctic copepods stay active during the dark season. Calanoides acutus seems to be the only true diapause species. Calculations of summer developmental rates and winter mortality rates of the large species C. acutus and C. propinquus suggest that both species have a 1-year life cycle with few females overwintering and probably spawning a second time. In contrast, a 2-year life cycle is more likely in R. gigas. However, life cycle durations of all species studied are still uncertain and regional differences are very probable.     

Observations of blue whales feeding in Antarctic waters

There are no published accounts of blue whales (Balaenoptera musculus) feeding in Antarctic waters. This note describes the behaviour of two groups of blue whales feeding in Antarctic pelagic waters. Whales were observed during the 18th IWC/IDCR southern hemisphere minke whale assessment cruise. Feeding behaviour in both cases resembled those described previously for both northern hemisphere blue whales and fin whales (B. physalus). These observations suggest that a programme of comparative behavioural observations could be developed to test the “feeding competition” hypothesis, which suggests that recovery of populations of blue whales will be impeded by feeding competition with sympatric minke whales. Accepted: 29 April 1999     

Reproductive response of the copepods Calanoides carinatus and Calanus agulhensis to varying periods of starvation in the southern Benguela upwelling region

Egg production by the calanoid copepods Calanoides carinatusand Calanus agulhensis fed excess Thalassiosira weissflogiiwas monitored in the laboratory following starvation periodsof 1, 3, 5, 7 and 9 days. Following short (1–3 day) periodsof starvation, egg production by C.agulhensis returned to thesatiated rate (51.1 eggs {female} day^–1) more rapidly(after 0.9–2.4 days of excess food) than that of Ca. carinatus(after 2.8–3.1 days). However, following longer (5–9day) periods of starvation, Ca. carinatus regained satiatedlevels of egg production (55.8 eggs {female}^–1 day^–1)more rapidly (after 3.1–4.0 days of excess food) thanC. agulhensis (after 3.8–5.2 days of feeding following5–7 days of starvation). Moreover, many C. agulhensisfemales did not regain normal rates of egg production after9 days of starvation. For both species, the time required foregg production to recover was proportional to the starvationperiod, although only up to 7 days for C. agulhensis, and wasthe same following 4.25 days of starvation. Previously fed Ca.carinatus terminated egg production more rapidly than C. agulhensiswhen starved. The ability of Ca. carinatus to tolerate, andrecover rapidly from, prolonged periods of starvation, combinedwith a comparatively fast development time and high rate ofegg production, provides this species with a strong competitiveadvantage over C. agulhensis in the highly pulsed food environmentof the southern Benguela upwelling region.     

Antarctic Ornithological Observations made during Bellingshausen's Voyage of Circumnavigation in 1819–1821.

I n the year 1819 His Imperial Majesty Alexander Pavlovich, Emperor of Russia, despatched an expedition to explore high southern latitudes, with the special object of proceeding as far as possible towards the South Pole. This expedition was led by Captain Thaddeus von Bellingshausen, who had under his command the corvette 'Vostok' and the sloop 'Mirnyi'.     

Lipid content and composition of three species of Antarctic fish in relation to buoyancy

Summary The lipip content and composition of various tissues from three species of nototheniid fish from McMurdo Sound, Antarctic have been examined in relation to their habitat and buoyancy. The pelagic midwater Dissostichus mawsoni is neutrally buoyant. It is rich in lipid which is located subcutaneously, as adipose tissue associated intimately with white muscle, and as lipid droplets within the cells of various tissues. White muscle, red muscle and liver are particularly lipid-rich, although the liver is not positively buoyant. The amount of lipid stored in the white muscle increases towards the centre of buoyancy of the fish. These deposits are documented at the anatomical, histological and ultrastructural levels. Tissues of Pagothenia borchgrevinki contain less lipid than D. mawsoni, but liver, red muscle and white muscle are still very rich in lipid. This species is cryopelagic, that is it spends most of the time in the water column just beneath the surface ice layer. It is not neutrally buoyant, but has a low weight in seawater. The tissues of the benthic Trematomus bernacchii contain only normal levels of lipid. The lipid class compositions of all three species are dominated by triacylglycerol, particularly when lipid contents are high. Serum lipids are an exception in containing high levels of the transport lipid sterol ester. The reason why Antarctic fish use triacylglycerols for buoyancy rather than was esters (as used by many myctophids) or squalene (as used by some sharks) is unclear.     

Measurements of buoyancy for some Antarctic notothenioid fishes from the South Shetland Islands

Buoyancy was measured for 258 specimens representing 13 species of adult and sub-adult nototheniids, bathydraconids, and channichthyids from the South Shetland Islands. Measurements were expressed as percentage buoyancy (%B)=W_water/W_air᎒². There were no neutrally buoyant species and mean values for %B were 3.07-6.11%, with channichthyids at the low end and benthic nototheniids and bathydraconids at the high end. All species showed an ontogenetic decrease in %B with increasing body weight. With the exception of Champsocephalus gunnari, there was no sexual dimorphism in %B within this sample. With some exceptions, values for %B were consistent with life-history information. Sub-adult Dissostichus mawsoni were not neutrally buoyant, as are large adults. Notothenia rossii had a significantly lower %B than closely related N. coriiceps. Benthic Gobionotothen gibberifrons had a lower %B than semipelagic Lepidonotothen larseni. Although they exhibit some diversification in life history, the four channichthyids in the sample were similar in %B. Neutral buoyancy is rare in notothenioids and may be confined to a single nototheniid clade.