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This study couples observations of krill (Thysanoessa inermis, Thysanoessa longicaudata, Thysanoessa rashii) from Tucker trawl nets and cameras, trying to test hypothesis that in the glaciated fjords of Svalbard most of the euphausiids biomass is located in near-bottom habitat and explains why in this region there are a substantial part of the krill population near the sea floor. Photographic material from the summers of 2013–2017 shows large numbers of near-bottom euphausiids (39% of the total krill biomass in Hornsund and 41% in Kongsfjorden), which reached a maximum density of 751 ± 224 indiv. m−3 in Kongsfjorden, 731 ± 198 indiv. m−3 in Hornsund, and 426 ± 124 indiv. m−3 in Adventfjorden. Regional distribution of near-bottom aggregations of krill seem to be associated with close proximity to the glacier front rather than with depth. The highest densities were located in the glacial bays. Where and why these aggregations occur is probably complicated and dependent on many environmental factors acting together. However, the dominant factors seem to be sedimentation and estuarine circulation. No krill aggregations were found during the winter cruise. The dominating species was T. inermis which made up 90% of the community. Other krill species—T. rashii and T. longicaudata, made up 6% and 4%, respectively. In the summer, aggregations of other macrozooplankton were also observed: amphipods of the genus Themisto and chaetognaths of the genera Eukrohnia and Parasagitta. Euphausiid densities in the water column (from Tucker trawl hauls) were an order of magnitude lower (0.33 indiv. m−3 for Kongsfjorden and 0.61 indiv. m−3 for Hornsund) than those of the near-bottom aggregations observed on cameras system. At most stations, the krill exhibited a behaviour, known as “nose diving” in the sediment, which is likely related to feeding. Observation of this phenomenon may indicate that krill (mostly T. inermis), found near the bottom of Spitsbergen fjords, is looking for food there. Near-bottom aggregations of zooplankton, mainly krill, are common in glacial bays and can be important in the function of the fjord ecosystem. Our research proves that the zooplankton biomass can be highly underestimated if only Tucker trawl sampling is done, due to neglecting the near-bottom layer in this type of method.

This study sought to estimate the effect of an anthropogenic and climate-driven change in prey availability on the degree of individual and population specialization of a large marine predator, the fin whale (Balaenoptera physalus). We examined skin biopsies from 99 fin whales sampled in the St. Lawrence Estuary (Canada) over a nine year period (1998–2006) during which environmental change was documented. We analyzed stable isotope ratios in skin and fatty acid signatures in blubber samples of whales, as well as in seven potential prey species, and diet was quantitatively assessed using Bayesian isotopic models. An abrupt change in fin whale dietary niche coincided with a decrease in biomass of their predominant prey, Arctic krill (Thysanoessa spp.). This dietary niche widening toward generalist diets occurred in nearly 60% of sampled individuals. The fin whale population, typically composed of specialists of either krill or lipid-rich pelagic fishes, shifted toward one composed either of krill specialists or true generalists feeding on various zooplankton and fish prey. This change likely reduced intraspecific competition. In the context of the current “Atlantification” of northern water masses, our findings emphasize the importance of considering individual-specific foraging tactics and not only population or group average responses when assessing population resilience or when implementing conservation measures.

In polar ecology, zooplankton diets and survival rates vary according to the seasonality of solar radiation and oceanographic conditions. Each zooplankton species has evolved feeding strategies to survive in the diet-limited conditions of the “polar night.” Many zooplankton studies have reported seasonal adaptations in feeding activity during polar night based on their trophic niches. Nitrogen isotope analysis of amino acids has provided improved accuracy in estimates of trophic position (TP) in various marine species. In this study, field work was conducted in Kongsfjorden before (October 2017) and after polar night (April 2018). As representative zooplankton, an amphipod (Themisto abyssorum), euphausids (Meganycitiphanes norvegica and Thysanoessa sp.), a chaetognath (Parasagitta elegans), and copepods (Calanus spp. and Oithona similis) were collected. trophic position values of each taxon were estimated using the nitrogen isotope ratio of glutamic acid (δ15NGlu) and phenylalanine (δ15NPhe). Results showed that TP values of P. elegans were relatively constant, averaging 3.2 in both seasons, likely due to continuous feeding activity during polar night. Trophic position values were also constant for Calanus spp., ranging 2.5–3.0 in both seasons, due to their ability to utilize stored high-energy wax. In contrast, average TP values for O. similis, an omnivorous zooplankton, were 2.9 in October and 2.3 the following April. Trophic position values for O. similis before polar night can be attributed to the relatively high availability of algae during longer periods of daylight. We found that TP variation in zooplankton before and after polar night differed according to feeding activities in diet-restricted circumstances.

Source at http://dx.doi.org/10.1002/lno.10519 Zooplankton vertical migration enhances the efficiency of the ocean biological pump by translocating carbon (C) and nitrogen (N) below the mixed layer through respiration and excretion at depth. We measured C and N active transport due to diel vertical migration (DVM) in a Svalbard fjord at 79°N. Multifrequency analysis of backscatter data from an Acoustic Zooplankton Fish Profiler moored from January to September 2014, combined with plankton net data, showed that Thysanoessa spp. euphausiids made up > 90% of the diel migrant biomass. Classical synchronous DVM occurred before and after the phytoplankton bloom, leading to a mismatch with intensive primary production during the midnight sun. Zooplankton DVM resulted in C respiration of 0.9 g m−2 and ammonium excretion of 0.18 g N m−2 below 82 m depth between February and April, and 0.2 g C m−2 and 0.04 g N m−2 from 11 August to 9 September, representing > 25% and > 33% of sinking flux of particulate organic carbon and nitrogen, respectively. Such contribution of DVM active transport to the biological pump in this high-Arctic location is consistent with previous measurements in several equatorial to subarctic oceanic systems of the World Ocean. Climate warming is expected to result in tighter coupling between DVM and bloom periods, stronger stratification of the Barents Sea, and northward advection of boreal euphausiids. This may increase the role of DVM in the functioning of the biological pump on the Atlantic side of the Arctic Ocean, particularly where euphausiids are or will be prevalent in the zooplankton community.

Mesopelagic sound scattering layers (SSL) are ubiquitous in all oceans. Pelagic organisms within the SSL play important roles as prey for higher trophic levels and in climate regulation through the biological carbon pump. Yet, the biomass and species composition of SSL in the Arctic Ocean remain poorly documented, particularly in winter. A multifrequency echosounder detected a SSL north of Svalbard, from 79.8 to 81.4°N, in January 2016, August 2016, and January 2017. Midwater trawl sampling confirmed that the SSL comprised zooplankton and pelagic fish of boreal and Arctic origins. Arctic cod dominated the fish assemblage in August and juvenile beaked redfish in January. The macrozooplankton community mainly comprised the medusa Cyanea capillata, the amphipod Themisto libellula, and the euphausiids Meganyctiphanes norvegica in August and Thysanoessa inermis in January. The SSL was located in the Atlantic Water mass, between 200–700 m in August and between 50–500 m in January. In January, the SSL was shallower and weaker above the deeper basin, where less Atlantic Water penetrated. The energy content available in the form of lipids within the SSL was significantly higher in summer than winter. The biomass within the SSL was >12-fold higher in summer, and the diversity of fish was slightly higher than in winter (12 vs. 9 species). We suggest that these differences are mainly related to life history and ontogenetic changes resulting in a descent toward the seafloor, outside the mesopelagic layer, in winter. In addition, some fish species of boreal origin, such as the spotted barracudina, did not seem to survive the polar night when advected from the Atlantic into the Arctic. Others, mainly juvenile beaked redfish, were abundant in both summer and winter, implying that the species can survive the polar night and possibly extend its range into the high Arctic. Fatty-acid trophic markers revealed that Arctic cod mainly fed on calanoid copepods while juvenile beaked redfish targeted krill (Thysanoessa spp.). The relatively high biomass of Arctic cod in August and of redfish in January thus suggests a shift within the SSL, from a Calanus-based food web in summer to a krill-based food web during winter.

We synthesize recent research on variation in annual production of copepods (Calanus spp.), euphausiids (Thysanoessa spp.), and juvenile walleye pollock (Gadus chalcogrammus) in the southeastern Bering Sea. We reach five conclusions: 1) the timing of the spring bloom is more important than the amount of annual primary production for the transfer of primary to secondary production (i.e., timing matters); 2) summer and fall, not just spring, matter: organisms must maximize energy intake devoted to somatic growth and storage of lipids and minimize energy expenditures during each season; 3) stored lipids are important for the overwinter survival of both zooplankton and age-0 walleye pollock; 4) variation in ice extent and timing of ice retreat affect the spatial distributions of phytoplankton, zooplankton, and age-0 walleye pollock; when these spatial distributions match in late-ice-retreat years, the annual production of copepods, euphausiids, and juvenile walleye pollock often increases (i.e., location matters); 5) if years with late ice retreat, which favor copepod, euphausiid, and juvenile walleye pollock production, occur in succession, top–down control increases. These conclusions help to explain annual variation in production of copepods, euphausiids and juvenile walleye pollock. Copepods and euphausiids often are more abundant in cold years with late ice retreat than in warm years with early ice retreat due to bloom timing and the availability of ice algae during years with late ice retreat. As a consequence, age-0 walleye pollock consume lipid-enriched prey in cold years, better preparing them for their first winter and their overwinter survival is greater. In addition, there is a spatial match of primary production, zooplankton, and age-0 walleye pollock in cold years and a mismatch in warm years.

Euphausiids are an important component of the eastern Bering Sea marine ecosystem. We synthesized information on the ecological roles of two species, Thysanoessa raschii, which predominates over the Middle and Inner Shelf Domains, and Thysanoessa inermis, which predominates over the Outer Shelf Domain. Although estimates of euphausiid biomass across the shelf are not well constrained, we estimated that, between April and July, 2004–2010, euphausiid biomass was 3.08–5.25 g C m−2 on the outer shelf and 1.95–3.92 g C m−2 on the middle shelf. Modeled estimates of euphausiid production, for spring and summer combined, varied between 0.043 g C m−2 d−1 and 0.051 g C m−2 d−1, depending on location, with a mean of 0.048 g C m−2 d−1. Recently reported field measurements of annual primary production over the southeastern Bering Sea in 2008–2009 vary between 0.06 and 6.65 g C m−2 d−1, with a mean of 1.262 g C m−2 d−1±2.049 g C m−2 d−1 in spring and summer combined, a level sufficient to support euphausiids, at least on an annualized basis. Walleye pollock (Gadus chalcogrammus, hereafter pollock) is the single most important consumer of euphausiids over the eastern Bering Sea shelf. We estimated that pollock consumed between 0.0042 and 0.019.7 g C m−2 d−1 of euphausiids, depending on year, with a mean of 0.011 g C m−2 d−1 in summer averaged over 1999–2009. This consumption is equivalent to between 17% and 29% of summer euphausiid production, depending on location. Over the period for which data were available (2004–2012), we observed a strong negative relationship between euphausiid biomass as determined in acoustic surveys and pollock biomass as estimated in the eastern Bering Sea pollock stock assessment (r2=0.82). During this time period, sea-surface temperature was the second strongest predictor of euphausiid biomass, (r2=0.63). However, for the period 2004–2010, bottom temperature (r2=0.94) was the strongest predictor, followed by pollock biomass from the pollock stock assessment (r2=0.82), and sea-surface temperature (r2=0.81). Mean pollock density in the acoustic surveys was not a powerful predictor of euphausiid biomass during either period. In spatially explicit multiple regression analyses for the periods 2004–2012 and 2004–2010 those formulations that included sea-surface and bottom temperatures as well as survey estimates of pollock had the greatest explanatory value. However, when either or both temperature terms were dropped, the explanatory value of the models dropped considerably. When pollock biomass was dropped from the models, there was little change in explanatory value compared to the full model. Euphausiid production and pollock consumption data coupled with a negative relationship between euphausiid biomass and stock assessment estimates of pollock biomass indicate a top-down predation effect. However, strong negative relationships between euphausiid biomass and water temperatures indicate the influence of a bottom-up mechanism. The apparent differences in these results may relate to the different spatial and temporal scales used to assess the pollock biomass used in the analyses. Alternatively, euphausiid biomass may be strongly controlled during a restricted portion of the year, such as spring, if critical food needs are not met in some years. We lack the data necessary to resolve these alternative hypotheses.

The polar night in the Arctic is characterized by up to six months of darkness, low temperatures and limited food availability. Biological data on species composition and abundance during this period are scarce due to the logistical challenges posed when sampling these regions. Here, we characterize the plankton community composition during the polar night using water samplers and zooplankton net samples (50, 64, 200, 1500 μm), supplemented by acoustics (ADCPs, 300 kHz), to address a previously unresolved question–which species of zooplankton perform diel vertical migration during the polar night? The protist community (smallest plankton fraction) was mainly represented by ciliates (Strombidiida). In the larger zooplankton fractions (50, 64, 200 μm) the species composition was represented primarily by copepod nauplii and small copepods (e.g., Microcalanus spp., Pseudocalanus spp. and Oithona similis). In the largest zooplankton fraction (>1500 μm), the euphausiid, Thysanoessa inermis, was the most abundant species followed by the chaetognath Parasagitta elegans. Classical DVM was not observed throughout the darkest parts of the polar night (November–mid-January), although, subtle vertical migration patterns were detected in the acoustic data. With the occurrence of a more distinct day–night cycle (i.e., end of January), acoustical DVM signals were observed, paralleled by a classical DVM pattern in February in the largest fractions of zooplankton net samples. We suggest that Thysanoessa spp. are main responsible for the acoustical migration patterns throughout the polar night, although, chaetognaths and copepods may be co-responsible.

We studied seasonal and regional change in vertical distribution and DVM patterns of four euphausiid species (Euphausia pacifica, Thysanoessa inspinata, Thysanoessa longipes, and Tessarabrachion oculatum) from two years of surveys using MOCNESS above 1500 m depth across a transect in 3 regions of the northwestern (NW) Pacific, off east of Japan; Oyashio, Kuroshio, and Oyashio–Kuroshio Mixed Water Regions (MWR). The four euphausiid species exhibited a regional change in vertical distribution, i.e., slightly deeper in the MWR and much deeper in the Kuroshio region than in the Oyashio region. They found in higher and wider temperature ranges in the MWR than in the Oyashio region, which demonstrated that the four species were able to adapt to different temperatures in different regions. In the MWR and Oyashio regions, E. pacifica is a surface migrant (differences between day and night mean median depths, D-N, were ca. 300 m) and T. oculatum is a moderate subsurface migrant that performs short DVM in the upper mesopelagic zone (D-N ca. 100 m). The other two morphologically similar Thysanoessa species (T. inspinata and T. longipes) segregated vertically between E. pacifica and T. oculatum at night in the Oyashio region, suggesting vertical habitat partitioning with the former two species but not with themselves. However, a seasonal pattern was observed in the vertical distribution and DVM of T. longipes in the Oyashio region. It behaves as a surface migrant in May, whereas most of individuals were found in the mesopelagic layer in September. In contrast, T. inspinata did not exhibit a clear DVM throughout the year (i.e., a moderate subsurface migrant). This seasonal difference might be a strategy to minimize competition between related species. Among the four species, only E. pacifica was found in higher temperatures at night than during the daytime, and the highest temperatures at the median depth varied among species (from 7.5 °C to 13.7 °C) although the lowest temperature did not vary greatly (from 1.0 °C to 1.8 °C), which indicates high temperatures act as a limiting factor as opposed to low temperatures. Furthermore, the integrated chlorophyll a values exhibited significant negative correlation with median depths of only E. pacifica at night. These results indicate a strategy which makes E. pacifica the dominant species in the area, that is, it has a trade-off of long migrations and a warmer environment that accelerates metabolism, in return for obtaining a food-rich environment.

Euphausiid (krill) and amphipod dynamics were studied during 2006–2011 by use of plankton nets in Kongsfjorden (79°N) and adjacent waters, also including limited sampling in Isfjorden (78°N) and Rijpfjorden (80°N). The objectives of the study were to assess how variations in physical characteristics across fjord systems affect the distribution and abundance of euphausiids and amphipods and the potential for these macrozooplankton species to reproduce in these waters. The abundances of euphausiids and amphipods were higher in Kongsfjorden than in Rijpfjorden and Isfjorden, and the highest abundances were observed at the innermost stations of Kongsfjorden, where Thysanoessa inermis and Themisto libellula dominated. The Atlantic species Thysanoessa longicaudata, Meganyctiphanes norvegica and Themisto abyssorum dominated at the outside Kongsfjorden. Inter-annual and seasonal variability in abundances of euphausiids and amphipods were evident. The presence of ripe euphausiids outside Kongsfjorden indicates that they may reproduce in these areas. Mature individuals of T. abyssorum were recorded mainly outside Kongsfjorden, whereas no mature or ripe T. libellula were present in both the inner and outer parts of this fjord. If the warming trend persists, as seen during the last decade, this would favour the Atlantic/boreal euphausiid species, while Arctic species, such as the amphipod T. libellula, may decline. Euphausiids and amphipods are major food of capelin (Mallotus villosus) and polar cod (Boreogadus saida), respectively, in this region, and changes in prey abundance will likely have an impact on the feeding dynamics of these important fish species.

The high-Arctic Kongsfjord is influenced mainly by cold Arctic water but also by warmer Atlantic water masses. In recent years, the proportion of the Atlantic inflow from the south has increased. Concurrently, one temperate-boreal (Meganyctiphanes norvegica) and one subtropical-temperate (Nematoscelis megalops) krill species are now regularly found in the Kongsfjord, in addition to the previously prevailing arcto-boreal species Thysanoessa inermis and T. raschii. In view of the recent changes in these species’ biogeographic distributions, we compared their physiological tolerances. Using non-invasive optical oxygen sensors, respiration measurements served to characterize metabolic responses to temperature variations. Thysanoessa spp. appear more cold-stenotherm than the other 2 krill species: the upper level of respiratory capacity is reached at 12°C and they are less tolerant to decreasing oxygen concentrations. This finding is consistent with their arcto-boreal distribution. In contrast, M. norvegica and N. megalops showed a higher tolerance to temperature changes, a robust nutritional condition and sexual maturity. Such physiological plasticity may explain the recent northward expansion of their geographic range.

The distribution, abundance, biomass, and community structure of macrozooplankton were investigated in relation to oceanographic conditions in the Barents Sea during summer 2013, using an IKS-80 net (0.5 m2 mouth opening, 550 μm mesh). Overall, we found 20 taxa (11 in Murmansk Coastal Water-MCW and in Atlantic Water-AW, 13 in Arctic Water-ArW, and 19 in Barents Sea Water-BSW). Total macrozooplankton abundance varied between 36 and 1116 ind. m−2 (0.2–4.6 ind. m−3) with a mean value of 238 ± 34 ind. m−2 (1.0 ± 0.1 ind. m−3) for the entire area. Although the catch was fairly diverse, a small number of species accounted for the majority of the total abundance and biomass. The most abundant taxa were Parasagitta elegans (MCW, AW, BSW, ArW), Thysanoessa spp. larvae (MCW, AW), Mertensia ovum (BSW), Aeginopsis laurentii (BSW), Dimophyes arctica (ArW), and Themisto libellula (ArW). Total macrozooplankton biomass ranged from 17 mg dry mass (DM) m−2 (0.1 mgDM m−3) to 6710 mgDM m−2 (23.1 mgDM m−3), averaging 1058 ± 240 mgDM m−2 (4.3 ± 0.9 mgDM m−3) across the study area, with Parasagitta elegans dominant by biomass. Cluster analysis revealed three communities (southern, central and northern) which demonstrated clear variations with latitude. Canonical correspondence analysis showed that location (latitude) and environmental conditions (mean temperature, salinity, mean phytoplankton abundance and biomass) explained 36% of the spatial variability in macrozooplankton communities. In BSW, there was a strong positive correlation between total macrozooplankton biomass and mesozooplankton stock suggesting an important role for food availability in controlling macrozooplankton variability. Predator–prey interactions also might play an important role in influencing the spatial distribution of macrozooplankton. The results may be useful as basis for future investigations of macrozooplankton in order to detect possible influence of changing environment on plankton communities of the Barents Sea during modern warming trend in the Arctic.

Myctophid fish play an important role in the Southern Ocean pelagic food web. The lanternfish Krefftichthys anderssoni is one of the most common myctophids in the region, but its ecology is poorly known. This study examines spatial and temporal patterns in the species distribution of density, life cycle, population structure and diet using samples collected by mid-water trawl nets deployed in different seasons across the Scotia Sea. Virtually absent from the sea-ice zone, the species was most abundant in the northern Scotia Sea around the Georgia Basin at depths below 400 m that are associated with the Circumpolar Deep Water. The species migrated during night from waters deeper than 700 m to waters above the 400 m following their main prey species: the copepods Rhincalanus gigas and Calanoides acutus and euphausiids of the Thysanoessa genus. Larvae length distribution and post-larvae length-frequency analyses suggested a life cycle of ~ 3 years with spawning and recruitment strongly connected with APF and the South Georgia shelf. Our results show that species spatial distribution, population structure and diet changed both seasonally and ontogenetically. This study is the most comprehensive examination of the ecology of K. anderssoni in the Southern Ocean to date and contributes to resolving how pelagic food webs and ecosystems operate in the region.

We analysed summer diet and fledging mass of macaroni penguins Eudyptes chrysolophus breeding at Bird Island, South Georgia, during the crèche period (January and February) between 1989 and 2010. Crustaceans were the main prey accounting, for over 90% of the diet by mass. Antarctic krill Euphausia superba was the main prey, in 17 out of 22 years. Amphipods Themisto gaudichaudii were the main prey in 1994 and 2009, fish in 2004, and the euphausiids Thysanoessa spp. and Euphausia frigida in 2000. There was no clearly dominant prey group in 1999. Prey diversity and the frequency occurrence of T. gaudichaudii both increased with a decreasing proportion of E. superba in the diet. There was strong evidence that macaroni penguins have a sigmoidal functional response, indicating that this kind of response should be accounted for when devising ecosystem-based management reference points for seabirds. The energy and mass of all euphausiids combined (rather than E. superba in particular) in the diet were the most reliable predictors of chick fledging mass; the correlation between model-predicted and observed values was 0.84. The gross energy content of individual meals was often above average in years when the diets contained fewer euphausiids, but fledging mass was always below average in these years. Although macaroni penguins are able to feed on a variety of prey types, chick growth was always severely impacted by a shortage of euphausiids due to higher energy or time costs associated with feeding on alternative prey types. Given their reliance on euphausiids, macaroni penguins would be particularly vulnerable to potential climate-driven declines in krill stocks.

Since the Discovery Investigations of the 1920s and 1930s, seawater temperatures have increased in the Atlantic sector by ∼1 °C; greater than the global mean rise. The aims of this paper were first to rescue the Discovery macrozooplankton data, second to provide quantitative “baseline” distribution maps, relating these to indices of temperature and food. Our third aim was to use the relationships we derived between abundance and temperature to project the potential affect of a 1 °C warming on the Discovery era distribution patterns. Based on the 1 m ringnet data retrieved from 615 stations (Nov–March), four taxa comprised >90% of the Antarctic macrozooplankton abundance: Rhincalanus gigas, Thysanoessa spp., Euphausia superba, and Chaetognaths. Most of the taxa, especially the more abundant ones, were warm water species penetrating into Antarctica and thus total macrozooplankton abundance decreased about 100-fold from 50°S to 70°S. While temperature correlated best with distribution at this large scale, food availability (proxied by a present-day satellite-based Chlorophyll a climatology) had a secondary effect, with the major euphausiids Euphausia superba and Thysanoessa spp. concentrated in high chl a areas. A modelled uniform 1 °C temperature rise produced a poleward shift for all taxa, but the Antarctic continent blocked this re-adjustment for the high latitude species, constricting their predicted range. More widespread polar/sub-polar species were predicted to increase their penetration into Antarctica by 4–12° in latitude, whereas the poleward shift in potential range of sub-Antarctic taxa were limited by the steep temperature gradient across the Antarctic Polar Front (APF). However, within the Scotia Sea the relatively warm temperatures of the northern Antarctic Zone, abundant food due to iron fertilisation and intense eddy activity provide a “gateway” for northern species to penetrate south of the APF. Our model predictions, based on measured distributional ranges and observed temperature increases, provide a yardstick with which to compare modern day data compilations and assess the potential effects of future temperature increases.

Fecal pellets of zooplankton can be important carriers of organic matter from surface to deep waters of the oceans, The summer cruise ANTARES 4 gave us the opportunity to collect zooplankton from the frontal subantarctic zone (station 7 or s-ant) and adjacent subtropical waters (station 8 or s-trop). Triglyceride fatty acid profiles were used to assess the trophic position of two copepods (Metridia lucens and Pleuromamma spp.), three euphausiids (Euphausia spinifera, Nematoscelis megalops, and Thysanoessa sp) and a decapod Acanthephyra sp. The euphausiid E. spinifera was studied at the two sites and its reaction to contrasting trophic environments and hydrology was highlighted by variations in its lipid composition. Most species showed various degree of omnivory: E. spinifera from subantarctic showed the highest degree of herbivory on phytoplankton (high proportion of 16:4(n-1), 18:5(n-3), 16:1(n-7)), while the same species at the subtropical station, as well as Thysanoessa sp and M lucens from the subantarctic station displayed more omnivorous feeding characteristics, partly linked to the abundance of microzooplankton cells. Acanthephirasp, N. megalops and Pleuromammasp from station 8 (s-trop) appear to be essentially carnivorous (high 18:1 n-9/18:1n-7 ratio, high levels of 20: 1 (n-9) or 22: 1 (n-11)). The trophic position influenced the composition of feces, with higher PUFA percentages in fecal pellets from carnivorous species. Hence, the surface faunal assemblage and the season controlled not only the quantity but also the quality of the food supply as PUFA exported towards the benthic domain. (c) 2007 Elsevier B.V. All rights reserved.

Myctophids are the most abundant mesopelagic fishes in the Southern Ocean, although their trophic role within the predominantly krill-based food web in regions south of the Antarctic Polar Front (APF) is poorly resolved. This study therefore examined the diets of 10 species of myctophid fishes: Electrona antarctica, E. carlsbergi, Gymnoscopelus braueri, G. fraseri, G. nicholsi, Krefftichthys anderssoni, Protomyctophum bolini, P. tenisoni, P. choriodon and Nannobrachium achirus, in the Scotia Sea, together with their predatory impact on the underlying zooplankton community. Myctophids and their prey were sampled in different seasons by scientific nets deployed across the Scotia Sea from the sea-ice zone to the APF. Based on the percentage index of relative importance, myctophids had high overlap in their diets, although the data indicate dietary specialisation in some species. There was also a distinct switch in diet, from copepods to euphausiids and amphipods, with increasing myctophid size. Myctophid predation impacted daily copepod production by between 0.01 and 5%, with Calanus simillimus being most impacted. Total annual consumption of copepods was around 1.5 million t (Mt) per year. All myctophids preyed upon the euphausiid Thysanoessa spp., consuming ~12% of its daily productivity and around 4 Mt per year. However, only larger myctophid species preyed upon Antarctic krill Euphausia superba, consuming 2% of its daily productivity, which could amount to as much as 17 Mt per year. Themisto gaudichaudii was also an important dietary component, with 4% of its daily productivity being consumed, amounting to around 2 Mt per year. This study demonstrates that myctophids link secondary productivity to higher predators both through krill-dependent and krill-independent trophic pathways.

Knowledge of crustacean moulting is derived mainly from benthic decapods, which often show profound changes in physiology and behaviour through the moult cycle. In contrast, euphausiids are suggested to be little impaired by moulting, enabling a swarming pelagic life. The aim of this study was to quantify moult cycle-related changes in the feeding activity of 2 euphausiids, Meganyctiphanes norvegica and Thysanoessa spp. Late furcilia larvae and early postlarvae were kept individually over 6 to 7 wk and fed with either a high or low concentration of Artemia salina nauplii or particulate fish food. The intermoult period, similar to9 d for M norvegica and similar to8 d for Thysanoessa spp., increased with body weight, but did not differ with food source. Moulting was partially synchronised, with up to 50 % of the individuals moulting within 48 h of each other. Daily feeding rates on A, salina decreased on the day before moulting, but increased during the next few days with highest values on Days 1 to 3 after moulting. The deviation from the mean feeding rate over the whole moult was more pronounced at the higher food concentration, reaching up to 40 %. Likewise, the defecation volume was reduced on the moulting day and the following day to similar to50 % of the mean, but increased to 180 % of the mean on Day 3 after moulting. Thus, the moult cycle induces significant changes in feeding rates of larval euphausiids with a similar succession of events and intensity as observed in decapods. Feeding rates, extrapolated from spot measurements on a few individuals, are unlikely to represent average values over the whole moult cycle, especially when populations moult synchronously. We propose a protocol to increase the precision of field estimates on feeding rates.

The use of stable isotopes to study food webs has increased rapidly, but there are still some uncertainties in their application. We examined the d 15 N and d 13 C values of Antarctic euphausiids and copepods from the Polar Front, Lazarev Sea, and Marguerite Bay against their foodweb baseline of particulate organic matter (POM). Interpretations of trophic level were helped by comparison with other approaches and by calibration experiments with Euphausia superba fed known diets. Results for well-known mesozooplankters (e.g., Calanoides acutusand Metridia gerlachei) were internally consistent and corresponded to those derived from independent methods. This gave confidence in the isotope approach for copepods and probably larval euphausiids. Among the dominant yet poorly known species, it suggested mainly herbivory for Rhincalanus gigas but omnivory for Calanus simillimus and furcilia larvae of Thysanoessa spp. and Euphausia frigida. The d 15 N values of adult copepods were up to 3‰ higher than those of early copepodites, pointing to ontogenetic shifts in diet. In the Lazarev Sea in autumn, the isotopic signals of E. superba larvae suggested pelagic, mainly herbivorous, feeding rather than feeding within the ice. In contrast to the mesozooplankton, some anomalous results for postlarval krill species indicated problems with this method for micronekton. The experiments showed that postlarval E. superba did not equilibriate with a new diet within 30 d. We suggest that the slower turnover of these larger species, partly in combination with their ability to migrate, has confounded trophic effects with those of a temporally/spatially changing food-web baseline. Interpretations of food sources of micronekton could be helped by analyzing their molts or fecal pellets, which responded faster to a new diet.

Four krill species with overlapping functional biology coexist in Greenland waters. Here, we used stable isotopes to investigate and discuss their trophic role and mode of coexistence. Bulk carbon (δ13C) and nitrogen (δ15N) stable isotope analyses of Thysanoessa longicaudata, T. inermis, T. raschii and Meganyctiphanes norvegica sampled in June 2010 in Godthåbsfjord, SW Greenland, revealed new insight into the species’ trophic roles and positions. There was a general positive correlation between body length and trophic position. The largest species, M. norvegica, had the highest trophic position (TP [mean ± SE] = 2.8 ± 0.2) indicating carnivory, while T. inermis (TP = 2.4 ± 0.3) had a more omnivorous diet. In turn, T. longicaudata and T. raschii (TP = 2.2 ± 0.2) were herbivorous. Along the fjord, plankton composition affected trophic position. T. longicaudata was more omnivorous offshore than inshore, where it had the same trophic position as the baseline primary consumer Calanus spp. Similarly, T. raschii and T. inermis had higher trophic positions in the mouth of the fjord compared with the inner fjord. Regardless of spatial variations in potential food and the overlap in diet, typical of opportunistic species, body size appears as the key factor determining the role and position of krill in the food web., Project BOFYGO (Danish Centre for Marine Research), EUROBASIN (7FP EU ENV.2010.2.2.1-1 grabt agreement 264933), Greenland Climate Centre (Project 6505), 2,4830

As part of a joint project on the fate of phytoplankton in Balsfjorden in Northern Nor- way, we investigated the trophic fate and sedimentation potential of Phaeocystis pouchetii by tracing the transition of biomarker patterns from a phytoplankton bloom to sediment traps and during a gut- passage experiment. The phytoplankton biomass during the spring bloom 1996 was dominated by colonial P. pouchetii (ca 85%) and 4 members of the diatom family Thalassiosiraceae (ca 10%). Par- ticulate organic carbon in sediment traps largely consisted of fecal material from the Arctic krill Thysanoessa sp. Sterol and fatty acid biomarker patterns in the phytoplankton bloom could be repro- duced by combining the individual biomarker patterns of the isolated phytoplankters P. pouchetii and Thalassiosira decipiens in a ratio of ca 75:25. In a laboratory experiment, Arctic krill (Thysanoessa raschii) fed with similar efficiency on P. pouchetii colonies and the Thalassiosiraceae. During gut passage, the abundance of Thalassiosiraceae biomarkers in fecal strings increased relative to P. pouchetii biomarkers, while biomarkers from krill became dominant. This transition of biomarker patterns due to gut passage in T. raschii closely resembled the biomarker transition from the surface bloom to material in sediment traps at 40 to 170 m depth, which was mainly composed of krill fecal strings. We conclude that krill grazed efficiently on P. pouchetii colonies in Balsfjorden and caused sedimentation of P. pouchetii-derived organic matter below the euphotic zone via fecal strings. Hence, both transfer to higher trophic levels and sedimentation of Phaeocystis spp.-derived organic matter can be more effective than is commonly believed.

Chromosome number and morphology were studied in gonadal tissue from eight euphausiid species, using an air-drying technique and Giemsa staining. Among Mediterranean species, haploid chromosome numbers were: n = 19 in Euphausia hemigibba, Euphausia brevis, and Nematoscelis megalops, n = 18 in Euphausia krohni, and n = 11 in Nyctiphanes couchi. Among Antarctic species, haploid chromosome numbers were: n = 17 in Euphausia superba, n = 20 in Thysanoessa macrura, and n = 13 in Thysanoessa vicina. Chromosome morphology was further studied for three species after chromosome measurements: Euphausia hemigibba karyotype showed 17 metacentric and 2 submetacentric chromosome pairs, Euphausia krohni 17 metacentric and 1 submetacentric chromosome pairs, and Nematoscelis megalops 17 metacentric and 2 submetacentric chromosome pairs. Three species sharing the same chromosome number of n = 19, i.e., Euphausia hemigibba, Nematoscelis megalops, and Meganyctiphanes norvegica were distinguished by a different slope of the decrease in chromosome size and by the presence/absence of metacentric and submetacentric chromosome pairs differing in number and position. Chromosome morphology of other species was derived from the observation of meiotic metaphases II. A majority of metacentric chromosomes was also displayed. Cytotaxonomic differences in haploid chromosome number allow the discrimination of morphologically neighboring sympatric species (e.g., Meganyctiphanes norvegica and Nyctiphanes couchi; Thysanoessa vicina and T. macrura). Euphausiids share common chromosomal features which clearly separate them from other eucarids, i.e., a narrow range of haploid chromosome number, a large chromosome size, and a majority of metacentric chromosomes.

International audience; Despite the large biomass of macaroni penguins Eudyptes chrysolophus in the Southern Ocean, their feeding ecology is poorly known at some important breeding localities. We investigated the diving behaviour and diet of female macaroni penguins feeding small chicks on Marion Island (468520S, 37850E), South Africa, one of the species' most northerly breeding sites, supporting 4% of their global population. We then compared our results with similar studies from other localities. In December 2008, we collected information on 12 foraging trips from 6 individuals using time-depth recorders, as well as diet from 42 individuals. Median trip duration was 22.8 h (5.6-80.8 h). Penguins performed 42.8 ± 15.9 dives per hour at sea, with dive depths averaging 24.6 ± 8.6 m and lasting 40.8 ± 12.1 s, although 74.3% of dives were \10 m. Euphasids dominated their diet (86% by mass), mainly Thysanoessa vicina. A second peak in dive depths at 55-80 m might reflect the 12% of fish in their diet. The substantial proportion of shallow night dives (30% of total dives) suggests some foraging occurs at night. Differences in diving patterns of individual macaroni penguins in this study confirmed the behavioural flexibility of these birds reported from other breeding localities. However, most other studies assumed that dives\3-5 m were commuting dives whereas our study suggests that at least some prey are caught during shallow dives. We highlight how different analytical methods can change the outcome of studies. Despite macaroni penguins' apparent flexibility in foraging behaviour during the breeding season, their numbers are decreasing globally. Further investigations of their foraging behaviour are needed to assess potential competition with other predators and krill fisheries.

Samples obtained during austral summer (February) were utilized to study the morphological differences of Thysanoessa macrura and the total lipid content, in relation to sex and geographic location. This species shows an evolution in morphometry, its carapace length increases slightly with sexual maturation, and its second thoracic leg becomes bigger and stronger, as well as its dactylus and especially the setae in the dactylus, which are stronger in a bigger animal allowing them to feed on other lipid-rich zooplankton. All these characteristics seem to support the hypothesis that animals change their diet during this period, allowing them to grow faster during summer, and accumulate lipids as a reserve material. This hypothesis is supported by a steeper slope in the length-weight regression during summer and an even steeper slope of the regression between carapace length and total lipids, especially adults. Great differences in lipid content were found between subadults and adults during this season, being higher in adults and, especially females. Lipids were higher in the northern stations in frontal area. A significant difference in lipid content was found among the stations sampled. Also a significant difference in lipid content was found between subadults, males and females. The species has a longer reproductive season than previously thought, which could depend on local conditions. Its development is accelerated during the summer bloom. Life history strategies of subadults and adults are apparently quite different, according to our lipid results and morphometrics data. Lipids reserves in subadults are not accumulated during summer, thus. T. macrura must accumulate massive amounts of lipids during autumn or late summer as a survival strategy of the species, as proposed for other species. (C) 2009 Elsevier Ltd. All rights reserved.

Krill from the Barents Sea were studied on six cruises from 1985 to 1989. Thysanoessa inermis and T. longicaudata were the dominant species, while T. raschii and Meganyctiphanes norvegica were rarer in the studied areas. The two dominant species T. inermis and T. longicaudata are mainly found in the Atlantic. Water and they do not to a large extent penetrate into Arctic water masses in the northern Barents Sea. M. norvegica is a more strict boreal species that does not occur as extensively in the Barents Sea as do the Thysanoessa species. The mean population abundance ranged from 1 to 61 individuals m?2 for T. inermis and from 2 to 52 ind. M?2 for T. longicaudata. The mean dry weight biomass of these two species ranged from 14 to 616 and from 19 to 105 mg?2. Length frequency distributions indicate a life span of just over two years for T. inermis and T. longicaudata. Growth took place from about April to autumn with no apparent growth during winter. Maturation and spawning seem to occur after two years for T. inermis and one year for T. longicaudata. Main spawning occurred from May to June coinciding with the spring phytoplankton bloom. Captive spawners of T. inermis (total length 17-22 mm) shed 30-110 eggs per female in a single batch.

The distribution and abundance of the euphausiids Thysanoessa raschi and Thysanoessa inermis in the shelf waters of the southeastern Bering Sea were investigated during spring and summer of 1980 and 1981. Experiments were conducted during the study to describe the reproduction, growth and development of these species. T. inermis was the dominant euphausiid species observed over the outer shelf region; it began spawning in early April while T. raschi dominated the euphausiids over the middle shelf and began spawning in mid or late May. The seasonal progression in spawning followed the seasonal development of temperature; however, spawning did not begin earlier in 1981 which was a warmer year than 1980. Average egg production of T. raschi ranged from 3.4% to 3.8% dry body weight of the female per day during the first three days after capture. Secondary production estimates for T. raschi females ranged from 4.2% to 5.2% dry body weight per day in 1980 and 5.9% to 6.0% dry body weight per day in 1981. A sharp decline in the abundance of adolescent and adult euphausiids over the middle shelf during the spring bloom period when food appeared to be abundant suggests that predation by diving birds, pollack. Tanner crabs, whales and seals effectively controls the euphausiid population.

The diet of Royal Eudyptes schlegeli and Rockhopper E. chrysocome Penguins, studied during the 1981/82 breeding season on Macquarie Island, is described. Quantitative analysis was performed on 29 stomach samples from Royal Penguins at Bauer Bay (west coast) and 21 samples from Sandy Bay (east coast). Samples were also obtained from 19 Rockhopper Penguins at Garden Cove (east coast). Principal dietary components were Euphausiids: Euphausia vallentini and Thysanoessa gregarra; Amphipods: Primno macropa and Parathemisto gaudichaudi; fish and squid, which were in general unidentifiable. Results of dietary analysis are presented as percent frequency of occurrence, and as percent frequency composition by weight. Considerable differences in dietary composition between the two coasts were observed with fewer euphausiids (45% cf 95%) and more amphipods (86% cf 38%) being taken on the west coast than on the east. Length of time that the parent was away from the chick and the fact that, with only minor exceptions, all dietary species were pelagic indicate that these penguins may feed at a considerable distance from the island.

Rates of predation by the invertebrates Aurelia aurita, Thysanoessa raschi and Euchaeta norvegica on larval stages of cod (Gadus morhua L.), flunder (Platichthys flesus L.), plaice (Pleuronectes platessa L.), herring (Clupea harengus L.), and turbot (Scophthalmus maximus L.) were determined. Experiments were conducted in late winter and early spring 1982 with predators collected in Loch Etive, Scotland and prey obtained from several locations in Great Britain. Early stages of the smallest species, cod, flounder and turbot, tended to be most vulnerable to all three predators, while the early stages of the larger species, plaice and herring, and older stages of all species, were less vulnerable. For all stages and species of larvae, predation rates by the three predators were most closely related to larval length and escape swimming speed. Larval length itself was closely correlated to indices of larval escape ability. Low predation rates on large larvae by E. norvegica could be due to handling difficulties, whereas for A. aurita and T. raschi these low rates were due to escape abilities of the larger larvae. Prey movement is an important stimulus eliciting predation in E. norvegica but not in A. aurita or T. raschi.

The diet of Royal Penguins Eudyptes schlegeli at Macquarie Island was studied in the 1984/85 and 1985/86 breeding seasons. The major food was euphausiid crustaceans (51% by weight), comprising Euphausia valentini (328 by weight) and Thysanoessa gregatia (10% by weight), together with the myctophld fish Krefftichthys anderssoni (24%). Other myctophid species such as Electrona carlsbergi, nototheniids and cephalopods were taken only in small amounts (25% by weight of the total diet). Most of the euphausiids eaten were adults, while the fish tended to be juveniles. Composition of the diet varied throughout the breeding season; E valentini was prevalent during incubation and moulting but the diet became more varied during chick feeding. It also varied between the east and west coasts of the island, penguins on the latter eating fewer E valentini and more E. gregaria than those on the east coast.

In the past, two euphausiid species prevailed in the high Arctic Kongsfjorden, the arcto-boreal Thysanoessa inermis (Kroeyer, 1846) and Thysanoessa raschii (Sars. 1964). Both were considered expatriates from the Barents Sea or Norwegian Sea and non-reproductive due to low temperatures. The macro-zooplankton of the fjord has been studied as a component in an ecosystem context since 2006, including baseline investigation of distribution and functional performance of key species. In recent years, three additional krill species were regularly detected in the fjord and are the focus of an intensive long-term study. Of these species, Thysanoessa longicaudata (Kroeyer, 1846) and Meganyctiphanes norvegica (Sars, 1857) are typical for the boreal Atlantic whereas Nematoscelis megalops (Sars, 1883) has a broad distribution in temperate to subtropical provinces. Their occurrence in the Kongsfjorden clearly indicates increasing Atlantic influence. During the 2011 campaign, T. raschii was observed spawning in the field for the first time and showed development up to the naupliar stage in the laboratory. Should more evidence of reproduction be encountered in any of the five krill species in the Kongsfjorden in the future, it will be taken as an indication of a changing environment concerning temperature and food web composition.