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.     

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.     

Spatial assessment of fin whale hotspots and their association with krill within an important Antarctic feeding and fishing ground

Fin whale (Balaenoptera physalus quoyi) habitat use and its relationship to environmental conditions are generally unknown in the Southern Ocean, presenting challenges for predicting their seasonal occurrence and potential effects of fishing pressure and climate change on this endangered species. Using biological data collected during 14 shipboard surveys off the northern Antarctic Peninsula and oceanographic data from satellite remote sensing, we mapped the distribution of fin whale hotspots, Antarctic krill abundance (biomass from acoustics, concentrations from nets) and ocean conditions during mid- and late-summer to investigate the environmental determinants of whale hotspots. Generalized additive models (GAM) were used to test the hypothesis that intra-seasonal changes in fin whale hotspot distribution relate to sea surface temperature (SST), krill abundance and eddy kinetic energy (EKE). More whale hotspots (sightings and individuals) are observed during late- than mid-summer surveys. During mid-summer, hotspots occurred near Elephant Island while in late-summer they were distributed throughout the slope region in proximity to the mean location of the southern Antarctic Circumpolar Current Front. The spatial mean of EKE did not differ between mid- and late-summer surveys, but the spatial mean of SST was significantly warmer during late-summer. The GAM for mid-summer indicates that fin whale hotspots were positively related to SST, EKE and acoustically determined krill biomass. The GAM for late-summer indicates the hotspots were negatively related to net-based krill abundance and positively related to acoustic krill biomass and EKE. This study is important because environmental determinants of fin whale hotspots may be used as reference points for implementing future conservation plans for their recovering populations.     

Differences between swarms of Antarctic krill and some implications for sampling krill populations

Antarctic krill, Euphausia superba Dana, were sampled from 28 discrete swarms in a small area south west of Elphant Island, South Shetlands, over a period of 14 d (24 February-9 March 1985). Four biological characteristics of the krill (length, sex, moult, gut fullness) were examined in a study of variation between swarms. Analysis of these characteristics indicated extensive differences between swarms and no single characteristic, or combination of characteristics, emerged as consistently accounting for the observed heterogeneity. There was no relationship between the degree of heterogeneity and the physical or temporal proximity of swarms. The variability between even close swarms strongly suggests that swarms are the basic unit of organization of krill populations. This conclusion has important implications for sampling programmes aimed at estimating population parameters. Thus, in the population studied here, we calculate that it was necessary to sample 23 swarms to estimate mean length and 10 swarms to establish a reliable estimate for the proportion of females in the population.     

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     

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.     

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.     

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.     

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     

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.     

Feeding ecology and niche segregation in oceanic top predators off eastern Australia

We examined the feeding ecology and niche segregation of the ten most abundant fish species caught by longline operations off eastern Australia between 1992 and 2006. Diets of 3,562 individuals were examined. Hook timer data were collected from a further 328 fish to examine feeding behaviour in relation to depth and time of day. Prey biomass was significantly related to predator species, predator length and year and latitude of capture. Although the fish examined fed on a mix of fish, squid and crustacea, fish dominated the diet of all species except small albacore (Thunnus alalunga) which fed mainly on crustacea and large swordfish (Xiphias gladius) and albacore which fed mainly on squid. Cannibalism was observed in lancetfish (Alepisaurus spp.). Multidimensional scaling identified three species groups based on their diet composition. One group consisted of yellowfin tuna (T. albacares), striped marlin (Tetrapturus audax) and dolphinfish (Coryphaena hippurus); a second group consisted of bigeye tuna (T. obesus), swordfish and albacore; and a third consisted of southern bluefin tuna (T. maccoyii) and blue shark (Prionace glauca). Of note was the separation of mako shark (Isurus oxyrhynchus) and lancetfish from all other predators. Prey length generally increased with increasing predator length although even large predators fed on a wide range of prey lengths including very small prey. Overall, differences in prey type and size, feeding times and depths were noted across the range of species examined to the extent that predators with overlapping prey, either in type or size, fed at different times of the diel period or at different depths. Taken together these data provide evidence for feeding niche segregation across the range of oceanic top predators examined.     

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.     

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     

The Scotia Sea krill fishery and its possible impacts on dependent predators: modeling localized depletion of prey

The nature and impact of fishing on predators that share a fished resource is an important consideration in ecosystem-based fisheries management. Krill (Euphausia superba) is a keystone species in the Antarctic, serving as a fundamental forage source for predators and simultaneously being subject to fishing. We developed a spatial multispecies operating model (SMOM) of krill-predator fishery dynamics to help advise on allocation of the total krill catch among 15 small-scale management units (SSMUs) in the Scotia Sea, with a goal to reduce the potential impact of fishing on krill predators. The operating model describes the underlying population dynamics and is used in simulations to compare different management options for adjusting fishing activities (e.g., a different spatial distribution of catches). The numerous uncertainties regarding the choice of parameter values pose a major impediment to constructing reliable ecosystem models. The pragmatic solution proposed here involves the use of operating models that are composed of alternative combinations of parameters that essentially try to bound the uncertainty in, for example, the choice of survival rate estimates as well as the functional relationships between predators and prey. Despite the large uncertainties, it is possible to discriminate the ecosystem impacts of different spatial fishing allocations. The spatial structure of the model is fundamental to addressing concerns of localized depletion of prey in the vicinity of land-based predator breeding colonies. Results of the model have been considered in recent management deliberations for spatial allocations of krill catches in the Scotia Sea and their associated impacts on dependent predator species.     

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.     

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.     

Stable isotopes identify an ontogenetic niche expansion in Antarctic krill (Euphausia superba) from the South Shetland Islands, Antarctica

Antarctic krill (Euphausia superba) occupy a key position in the Southern Ocean linking primary production to secondary consumers. While krill is a dominant grazer of phytoplankton, it also consumes heterotrophic prey and the relative importance of these two resources may differ with ontogeny. We used stable isotope analyses to evaluate body size-dependent trophic and habitat shifts in krill during the austral summer around the South Shetland Islands, Antarctica. We found evidence for an asymmetric, ontogenetic niche expansion with adults of both sexes having higher and more variable δ¹⁵N values but consistent δ¹³C values in comparison with juveniles. This result suggests that while phytoplankton likely remains an important life-long resource, krill in our study area expand their dietary niche to include higher trophic food sources as body size increases. The broader dietary niches observed in adults may help buffer them from recent climate-driven shifts in phytoplankton communities that negatively affect larval or juvenile krill that rely predominately on autotrophic resources.     

Kill before being killed: an experimental approach supports the predator-removal hypothesis as a determinant of intraguild predation in top predators

Intraguild predation (IGP) has been explained in terms of competitor-removal, food-stress and predator-removal hypotheses. Only the first two hypotheses have been fairly well studied. To test the predator-removal hypothesis as a force determining IGP in avian predators, we performed a field experiment to simulate the presence of an IG predator (eagle owl Bubo bubo dummy) in the surrounding of the nests of four potential IG prey (black kite Milvus migrans, red kite Milvus milvus, booted eagle Aquila pennata and common buzzard Buteo buteo). To discard the possibility that an aggressive reaction towards the eagle owl was not related to the presence of the IG predator, we also presented a stuffed tawny owl Strix aluco, which is a potential competitor but cannot be considered an IG predator of the studied diurnal raptors considered in the experiment. While almost always ignoring the tawny owl, raptors chiefly showed an interspecific aggressive behaviour towards their IG predator. Our results seem to support the predator-removal hypothesis, as the IG prey may take advantage of the diurnal inactivity of the IG predator to remove it from their territory. However, the recorded behaviour may be also considered as a special variety of mobbing (i.e. a prey’s counter-strategy against its predator), where the mobber is sufficiently powerful to escalate predator harassment into deliberate killing attempts. In their turn, eagle owls can respond with an IG predatory behaviour aimed at removing IG prey species which are highly aggressive mobbers.     

Change in the gross biochemistry and mineral content accompanying the moult cycle in the Antarctic krill Euphausia superba

Concentrations of water, ash, protein, chitin, lipid, calcium, magnesium, sodium, potassium, strontium and copper were measured in individuals from a laboratory population of Antarctic krill, Euphausia superba Dana, over the course of a moult cycle. Significant changes in all variables were encountered. Total ash, lipid, calcium, magnesium and strontium all increased in concentration following moulting. Water, protein and copper concentrations all decreased following ecdysis and increased again towards the end of the moult cycle. The major ions sodium and potassium fluctuated around mean levels. Cast moults of E. superba were shown to be a drain on the ionic load of the krill, and the losses inherent in exuviation could account for much of the variation observed during the moult cycle.