Moult in relation to some aspects of reproduction and growth in swarms of Antarctic krill,Euphausia superba

In a 14-d period 38 swarms of Antarctic krill,Euphausia superba Dana, were sampled in an area 55.5 x 55.5 km to the southwest of Elephant Island, at the north of the South Shetland Islands. Moult stage, maturity stage, and size of ca. 100 krill from each swarm were measured. Each of the characteristics varied greatly between swarms. Moulting krill were found in most swarms, but in one swarm all of the krill were just about to moult. Ways in which moulting may act as a possible sorting mechanism are discussed. Data for all the analysed krill (ca. 3000 specimens) were used to investigate the interdependence of moult rate, sexual maturation and growth. While all immature krill moulted at approximately the saine rate in the study, there were significant differences in the moulting rates of mature male and female krill. Gravid female krill continued to moult, although less frequently than mature males. As a consequence males had to attach spermatophores to females after each moult. It is likely that variation in moult rate of females and males was related to the energy expenditure required for ovary development in females and spermatophore production and searching behaviour in males. Spawning and moulting were only partly coupled in gravid females. Spawning appeared to take place predominantly during Moult Stage D₂. There was no evidence of intermoult growth by intersegmental dilation.     

Lipid metabolism of the Antarctic krill Euphausia superba and its ecological implications

 Various developmental stages (early larvae to adults) of Euphausia superba have been collected in different seasons in the Weddell Sea, the Lazarev Sea and off the Antarctic Peninsula to investigate the role of lipids and fatty acids in the life cycle of the Antarctic krill. The total-lipid data for E. superba exhibited seasonal variations, with low lipid levels in late winter/early spring and the highest levels in autumn. Seasonal changes were most pronounced in the immature and adult specimens, increasing from about 10% lipid of dry mass to more than 40%. The fatty-acid compositions of the younger stages were dominated by 20:5(n-3), 22:6(n-3) and 16:0. These are typical phospholipid fatty acids, which are major biomembrane constituents. The phospholipid composition was similar in the older stages. With increasing storage of triacylglycerols in the lipid-rich immature and adult stages, the fatty acids 14:0, 16:0 and 18:1(n-9) prevailed, comprising about 70% of total triacylglycerol fatty acids. The trophic-marker fatty acids 16:1(n-7) and 18:4(n-3), indicating phytoplankton ingestion, were less abundant. They reflected, however, the dependence of the larvae on phytoplankton as well as the seasonal changes in algal composition. The generally close linear relationships between fatty acids and lipid suggest that the fatty-acid compositions of the collected specimens were largely independent of the respective developmental stage, season and region. The linear fit indicates that triacylglycerol accumulation started at a level of about 5% of total lipid. Considering the various overwintering scenarios under discussion, the life cycle and reproductive strategies of krill are discussed in the context of the lipid metabolism and fatty-acid composition of E. superba. Lipid production is effective enough to accumulate large energy reserves for the dark season, but E. superba does not exhibit the sophisticated biosynthetic pathways known from other Antarctic euphausiids and copepods. Although important, lipid utilisation appears to be just one of several strategies of E. superba to thrive under the extreme Antarctic conditions, and this pronounced versatility may explain the success of this species in the Southern Ocean. Received: 16 June 2000 / Accepted: 18 December 2000     

Ecological implications of fecal pellets produced by the Antarctic krill Euphausia superba in the Antarctic Ocean

Organic analyses and electron microscopic observations on fecal pellets produced by the Antarctic krill Euphausia superba Dana showed that krill fed on choanoflagellates, the abundant heterotrophic flagellate in the Antarctic Ocean. Two new pathways of organic materials in the Antarctic ecosystem are proposed: (1) a new food chain including non-living particulate and dissolved organics, and bacteria-choanoflagellate-krill-vertebrate, which coexists with the traditional diatom-krill-vertebrate food chain; (2) non-phytoplanktonic organic materials in surface waters are transferred into choanoflagellates and are transported to deep water as fecal pellets which are still useful as nutrition for other organisms there.     

Sampling biological characteristics of krill: Effect of heterogeneous nature of swarms

An effect of the very patchy distribution of Antarctic krill (Euphausia superba Dana) in various forms of aggregation is that a single swarm may not provide an unbiased estimate of population parameters such as mean length of krill in the local area. Here, we analyse the number of samples required to estimate the characteristics of a local population as precisely as if there were no differences between krill swarms in terms of their biological composition. Krill were intensively sampled over different spatial and temporal scales around South Georgia in 1981 and 1982, and in the Bransfield Strait in 1985. These varied from replicate hauls at a single station over 24 h and repeat sampling in restricted areas over periods of 6 to 14 d to regional surveys around South Georgia and in the Bransfield Strait. Various biological characteristics were measured such as length, maturity, moult stage and feeding state. Depending upon the biological characteristic examined and the area covered by the sampling programme, the number of samples needed to obtain the same degree of precision as would be found in the absence of heterogeneity varied from 3 to > 80 samples. This has important implications for the design of net-sampling programme for monitoring krill populations.     

Growth of Antarctic krill Euphausia superba at South Georgia

 Antarctic krill Euphausia superba has a central role in the ecosystem of the Southern Ocean and knowledge of its growth rate is central to determining the factors influencing population dynamics. The length of Antarctic krill in the diet of Antarctic fur seals Arctocephalus gazella at South Georgia revealed a consistent increase in size between ca. 42 and ca. 54 mm over the period October–March, indicating growth rates much higher than predicted by existing models. Geographical variation in growth rate may result in 2-year-old krill at South Georgia attaining the same size as 3-year-old krill in the Antarctic Peninsula region. The effect of geographical variation in growth rate on the population structure of krill has important implications for comparing the fate of individual cohorts over large scales and in the interpretation of krill life-cycles. Received: 20 May 2000 / Accepted: 11 August 2000     

Omnivorous feeding behavior of the Antarctic krill Euphausia superba

Feeding experiments were conducted at Palmer Station from December 1985 to February 1986 to examine the potential role of copepod prey as an alternative food source for Euphausia superba. Copepod concentration, copepod size, phytoplankton concentration, the duration of krill starvation and the volume of experimental vessels were altered to determine effects on ingestion and clearance rates. Krill allowed to feed on phytoplankton and copepods in 50-litre tubs showed greatly increased feeding rates relative to animals feeding in the much smaller volumes of water traditionally used for krill-feeding studies. Clearance rates on copepods remained constant over the range of concentrations offered, but clearance rates on phytoplankton increased linearly with phytoplankton concentration. Feeding rates increased when larger copepods were offered and when krill were starved for two weeks prior to experiments. Clearance rates of krill feeding on copepods were higher than, but not correlated with, their clearance rates on phytoplankton in the same vessel. E. superba may have a distinct mechanism for capturing copepods, perhaps through mechanoreception. Although our observed clearance rate of 1055 ml krill^-1 h^-1 indicates that krill can feed very efficiently on copepod prey, such feeding would meet less than 10% of their minimum metabolic requirements at the typical copepod concentrations reported for Antarctic waters. However, substantial energy could be gained if krill fed on the patches of high copepod concentrations occasionally reported during the austral summer, or if krill and copepods were concentrated beneath the sea ice during the winter or spring months. Our results, indicating efficient feeding on zooplankton and higher clearance rates on phytoplankton than previously believed, represent a step towards balancing the energy budget of E. superba in Antarctic waters.     

Are penguins and seals in competition for Antarctic krill at South Georgia?

The Antarctic fur seal (Arctocephalus gazella) and macaroni penguin (Eudyptes chrysolophus) are sympatric top predators that occur in the Southern Ocean around South Georgia where they are, respectively, the main mammal and bird consumers of Antarctic krill (Euphausia superba). In recent years the population of fur seals has increased, whereas that of macaroni penguins has declined. Both species feed on krill of similar size ranges, dive to similar depths and are restricted in their foraging range at least while provisioning their offspring. In this study we test the hypothesis that the increased fur seal population at South Georgia may have resulted in greater competition for the prey of macaroni penguins, leading to the decline in their population. We used: (1) satellite-tracking data to investigate the spatial separation of the Bird Island populations of these two species whilst at sea during the breeding seasons of 1999 and 2000 and (2) diet data to assess potential changes in their trophic niches between 1989 and 2000. Foraging ranges of the two species showed considerable overlap in both years, but the concentrations of foraging activity were significantly segregated spatially. The size of krill taken by both species was very similar, but over the last 12 years the prevalence of krill in their diets has diverged, with nowadays less krill in the diet of macaroni penguins than in that of Antarctic fur seals. Despite a significant degree of segregation in spatial resource use by the study populations, it is likely that the South Georgia populations of Antarctic fur seal and macaroni penguin exploit the same krill population during their breeding season. For explaining the opposing population trends of the two species, the relative contributions of independent differential response to interannual variation in krill availability and of interspecies competition cannot be resolved with available evidence. The likely competitive advantage of Antarctic fur seals will be enhanced as their population continues to increase, particularly in years of krill scarcity.     

A refinement of the moult-staging technique for Antarctic krill (Euphasia superba)

The moult-staging technique for euphausiids developed by Buchholz was examined experimentally to obtain estimates of the relative duration of each of the constituent stages. The method entailed removing uropods from indiviually maintainedEuphausia superba Dana on the day of moulting and for each day post-moult, and examining them microscopically to determine the moult stage. The krill were kept alive following uropod removal until the next moult so that the stage observed in the excised uropod could be referenced to both ends of the moult cycle for each individual examined. The proportion of the moult cycle occupied by each stage at 0°C was thereby calculated.     

Metachronal swimming in Antarctic krill: gait kinematics and system design

Metachronal swimming, in which adjacent appendages stroke in sequence, is widespread among crustaceans inhabiting the transitional flow realm in which both viscosity and inertia effects are important. However, the design and operation of this propulsion system in response to various hydrodynamic, energetic, and behavioral needs have not been well investigated. We examine free-swimming Antarctic krill (Euphausia superba) as a model species and identify three distinct behavioral swimming gaits. The pleopod kinematics of these gaits, hovering, fast-forward swimming, and upside-down swimming, are quantified via image analysis of high-speed video. Pleopod stroke amplitude and frequency were found to vary significantly among these swimming modes. In order to increase swimming speed, krill were found first to increase stroke amplitude and secondarily to increase beat frequency. The kinematics of these distinct swimming modes provide insight as we consider multi-appendage metachronal swimming from a design standpoint. The ratio of the distance between adjacent appendage bases and appendage length is identified as a key parameter in metachrony, the value of which is constrained to a narrow range for a wide variety of species.     

Gut evacuation rates and pigment destruction in the Antarctic krill Euphausia superba

Krill grazing data collected during cruises in the region of the Antarctic Polar Front (S.A. Agulhas Voyage 70) and the South Georgia shelf (R.V. Africana Voyage 119) during the austral summer of 1993 were analyzed to estimate the variability of crucial parameters of the gut fluorescence technique in relation to food availability and krill feeding history. Gut evacuation rates (k) and passage or throughput times (1/k) varied in the ranges of 0.101 to 0.424 h^-1 and 2.3 to 9.9 h and were strongly correlated (p<0.001, r ²=0.98) to krill feeding activity (estimated as initial gut pigment content, G₀) but not to ambient chlorophyll a concentration. A significant differences was found when k values derived from incubations in filtered seawater and low charcoal particle concentrations (0.4 to 0.8 mgl^-1) were compared with values derived from krill fed high concentrations of charcoal (6 mgl^-1). The efficiency of gut pigment destruction was among the highest recorded for zooplankton organisms, 58.1 to 98.4%, and did not covary significantly (p>0.05) with ambient food concentration. However, the pigment lost per individual krill was strongly correlated with the total amount of pigment ingested (p<0.001, r ²=0.99). We suggest that both gut evacuation rates and pigment destruction efficiency may be realistically estimated only when krill is allowed to continue ingesting particles uninterruptedly. Charcoal particle concentration should be equivalent to the in situ wet weight of total seston per unit volume. An objective criterion for the standardization of the measurement and calculation of k values is also proposed.     

Chlordane residues in krill,fish and Weddell seal from the Antarctic

Chlordane compounds (CHLs) were quantitated in krill (Euphausia superba), benthic fish (Trematomus bernacchii) and Weddell seal (Leptonychotes weddelli) collected around the Japanese Antarctic Research Station (Syowa Station: 69°00'S, 39°35'E). The concentrations of ZCHL (cis‐, trans‐chlordane+ cis‐, fraws‐nonachlor + oxych‐lordane) in krill was much lower than that of benthic fish. The ratio of SCHL to sum of SCHL, SDDT and PCBs decreased with the trophic levels. These results indicate that CHLs appear to be easily degraded in higher organisms.     

Geospatial variability of krill and top predators within an Antarctic submarine canyon system

The spatial distribution patterns of krill, seabirds (penguin, petrel and albatross), fur seals and baleen whales were mapped in nearshore waters (<50 km from land) to investigate their habitat selection within two adjacent submarine canyons near Livingston Island, Antarctica. Three shipboard surveys were conducted (February 2005–2007), and an echosounder was used to measure the distribution and abundance of krill while simultaneously conducting visual surveys to map seabird and marine mammals. Using a multispecies approach, we test the hypothesis that spatial organization of krill and top predators co-vary according to fine-scale changes in bathymetry in the nearshore marine environment. GAMs are used to examine the effect of sea depth, slope and distance to isobaths on the spatial distribution and abundance of krill and predators. Spatial distribution patterns of krill and predators relate to fine-scale (1–10 km) changes in bathymetry and exhibit cross-shelf gradients in abundance. Krill were concentrated along the shelf-break and abundant within both submarine canyons. Predators exhibited different preferences for locations within the submarine canyon system that relates to their foraging behavior. Penguins concentrated closer to shore and within the head of the east submarine canyon immediately adjacent to a breeding colony. Whales were also concentrated over the head of the east canyon (overlapping with penguins), whereas albatrosses and fur seals were concentrated in the west canyon. Fur seals also showed preference for steep slopes and were concentrated along the shelf-break. Petrels exhibited peaks in abundance throughout both submarine canyons. Owing to their orientation, size and proximity to the coastline, submarine canyons provide important habitat heterogeneity for krill and a variety of predators. This study highlights the multispecies approach for studying spatial ecology of top predators and krill and has implications for marine spatial management of the Scotia Sea.     

Using a high-powered strobe light to increase the catch of Antarctic krill

Adult euphausiids are difficult to capture with the nets typically used by oceanographers due in part to avoidance of the net. A cruise of the Southern Ocean GLOBEC program (NBP0103) in April/May 2001 permitted an evaluation of the efficacy of a high-powered strobe light (150 W) mounted on a 1-m² multiple opening/closing net and environmental sampling system (MOCNESS) in reducing net avoidance. Three horizontal tows were made in Laubeuf Fjord, Marguerite Bay (western Antarctic Peninsula, 67.89°S; 68.30 W) on 28–29 May in an area that had a high abundance of adult krill (mostly Euphausia superba, Dana and a few E. crystallorophias, Holt and Tattersall). Each tow consisted of a series of paired down and up casts through a set depth interval (e.g. 50–90 m), with each successive net (335 µm mesh) sampling both a down and up cast. The strobe light was either on or off while each net was open, and when on, the light flashed at 4-s intervals. During a tow, four of the eight nets sampled with the strobe flashing and four sampled with the strobe off, in a random sequence. Total zooplankton displacement volume was significantly higher (P<0.05), on average by a factor of ~1.5, when the strobe light was on. The increased biovolume was due to the enhanced catch (factor of ~2) of adult krill (15–60 mm length). There was no enhanced catch of smaller krill (5–15 mm length: a mixture of E. superba, E. crystallorophias, and Thysanoessa macrura, GO Sars). In addition, the average length of the large krill fraction was not changed substantially with the strobe light on. These results suggest that krill avoidance of nets can be overcome by intense strobe lighting.Electronic Supplementary Material Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s00227-003-1228-z.Communicated by J.P. Grassle, New Brunswick     

Spatial methods for plot-based sampling of wildlife populations

Classical sampling methods can be used to estimate the mean of a finite or infinite population. Block kriging also estimates the mean, but of an infinite population in a continuous spatial domain. In this paper, I consider a finite population version of block kriging (FPBK) for plot-based sampling. The data are assumed to come from a spatial stochastic process. Minimizing mean-squared-prediction errors yields best linear unbiased predictions that are a finite population version of block kriging. FPBK has versions comparable to simple random sampling and stratified sampling, and includes the general linear model. This method has been tested for several years for moose surveys in Alaska, and an example is given where results are compared to stratified random sampling. In general, assuming a spatial model gives three main advantages over classical sampling: (1) FPBK is usually more precise than simple or stratified random sampling, (2) FPBK allows small area estimation, and (3) FPBK allows nonrandom sampling designs.     

Contribution of heterotrophic material to diet and energy budget of Antarctic krill, Euphausia superba

A novel approach was used to estimate the heterotrophic carbon component in the diet of the Antarctic krill Euphausia superba. Over 200 specimens from seven samples collected in the Lazarev Sea (January 1993 and 1995), at the Antarctic Polar Front (January 1993), and around South Georgia (February/March 1994) were dissected, and the total carbon content of their stomachs was estimated with a CHN-analyser. Gut-pigment contents were also measured by the gut-fluorescence technique in specimens collected at the same time, and the equivalent amount of their gut carbon was then subtracted from the total organic carbon content of guts from the same samples. The remaining carbon was assumed to originate entirely from heterotrophic food sources. This heterotrophic component accounted for a substantial proportion of the total food consumed by Antarctic krill, ranging from 17.4 to 98.9% of the mass of the gut contents (mean = 78.8% ± 21.2 SD). The results make an important contribution to the elucidation of the energy budget of krill and its daily carbon ration. With a few exceptions, previous estimates were largely calculated from a solely autotrophic carbon source, and were unable to account for the metabolic requirements of E. superba. Krill plays an important role in Antarctic food webs, as it often constitutes ≃50% of the total biomass of the zooplankton, and produces fast-sinking, dense faecal pellets which are important in the vertical transport of organic carbon from the euphotic layer to the deep ocean. High consumption rates of smaller heterotrophic organisms by krill suggest that this large microphage may be more important than previously believed in re-packaging micro- and mesozooplankton into a longer-lasting and more easily sequestered carbon pool. Received: 26 October 1998 / Accepted: 14 October 1999     

Restricted gene flow and evolutionary divergence between geographically separated populations of the Antarctic octopus Pareledone turqueti

Samples of the Antarctic octopus Pareledone turqueti were taken from three locations on the Scotia Ridge in the Southern Ocean. The genetic homogeneity of these populations was investigated using isozyme electrophoresis. Whilst panmixia appeared to be maintained around South Georgia (F _ST = 0) gene flow between this island and Shag Rocks, an island only 150 km away but separated by great depths, was extremely limited (F _ST = 0.74). These results are examined with respect to the discontinuous distribution of P. turqueti throughout Antarctica. An estimate of effective population size was also calculated (N _e = 3600). Received: 7 March 1997 / Accepted: 27 March 1997     

Submesoscale distribution of Antarctic krill and its avian and pinniped predators before and after a near gale

We conducted two ship-based surveys of the nearshore ecosystem north of Livingston Island, Antarctica during 2–10 February 2005. Between the two surveys, a low-pressure system (963 mbar) passed through the area providing the opportunity to measure ecosystem parameters before and after a near gale. A ship-based multiple-frequency acoustic-backscatter survey was used to assess the distribution and relative abundance of Antarctic krill (Euphausia superba). Net tows, hydrographic profiles, and meteorological data were collected to measure biological and physical processes that might affect the krill population. During the survey, the distribution and behavior of several krill predators [chinstrap penguins (Pygoscelis antarctica), cape petrels (Daption capense), and Antarctic fur seals (Arctocephalus gazella)] were measured from the vessel by visual observations. The survey encompassed an area of roughly 2,500 km², containing two submarine canyons with one to the west and one to the east of Cape Shirreff, which had different abundances of krill and predators. Several aspects of the nearshore ecosystem changed after the near gale including: hydrography of the upper 100 m of the water column, phytoplankton biomass, the abundance and distribution of krill, and the distribution of some krill predators. Differences in these parameters were also measured between the two canyons. These changes in the physical and biological environment during the survey period are quantified and show that the ecosystem exhibited significant changes over relatively short spatial (tens of kilometers) and time (tens of hours) scales.     

Laboratory and field estimates of the rate of faecal pellet production by Antarctic krill,Euphausia superba

The time course of faecal pellet production (egestion) was monitored in January 1985 for a population of Antarctic krill, Euphausia superba Dana, maintained in flowing seawater aquaria at Palmer Station, Antarctica. Following transfer to filtered seawater, krill produced faecal strings for roughly 40 min, after which time faecal egestion virtually ceased. Similar results were obtained for freshly-trawled krill at sea in February and March 1985. There were wide daily variations in total faecal egestion rate; mean rates varied from 0.54 to 1.66 mg dry wt h^-1 and individual rates from 0.25 to 2.35 mg h^-1 (all data corrected to a standard krill of 600 mg fresh weight). Despite these wide fluctuations in total faecal egestion, the loss of organic matter showed no significant daily variation, with a mean value of 0.13 mgh^-1. The relationship between faecal egestion rate and faecal organic content suggested that feeding rate was governed by food quality; when inorganic load was high, feeding rate increased to ensure sufficient energy intake. The data suggest that superfluous feeding does not occur in krill and that values of gut-clearance time calculated from time intervals greater than about 40 min will not be representative of previous feeding history. the rates of faecal egestion observed in this study indicate that the flux of faecal pellets from krill is substantial. They imply an energy intake in E. superba of 17 to 28% body weight per day, much higher than estimated previously for this species by summing known energy losses, but similar to estimates for other euphausiids.     

Design based estimation for ranked set sampling in finite populations

In this paper, we consider design-based estimation using ranked set sampling (RSS) in finite populations. We first derive the first and second-order inclusion probabilities for an RSS design and present two Horvitz–Thompson type estimators using these inclusion probabilities. We also develop an alternate Hansen–Hurwitz type estimator and investigate its properties. In particular, we show that this alternate estimator always outperforms the usual Hansen–Hurwitz type estimator in the simple random sampling with replacement design with comparable sample size. We also develop formulae for ratio estimator for all three developed estimators. The theoretical results are augmented by numerical and simulation studies as well as a case study using a well known data set. These show that RSS design can yield a substantial improvement in efficiency over the usual simple random sampling design in finite populations.