Proximity of krill and salps in an Antarctic coastal ecosystem: evidence from penguin-mounted cameras

Antarctic krill (Euphausia superba) and salps (mainly Salpa thompsoni) are main components of Southern Ocean ecosystem, but little is known about their coastal distribution at a fine scale (<1 km). We deployed miniaturised cameras on breeding chinstrap (n = 9 birds) and gentoo penguins (n = 9 birds) in the Antarctic Peninsula region and obtained 2,333 krill images, 93 salp images and 609 sea floor images from 1,843 dives. 51.2 % of penguin dives that had salps present in the images occurred near the dives with krill images (within 5 min). The vertical distribution of salp images showed overlap with the upper depth zone of krill images. While 16.3 % of dives with krill images were associated in time with the sea floor, only 1.2 % of dives with salp images did. These results revealed close proximity between krill and salps within the penguin’s foraging range in an Antarctic coastal ecosystem. These results also imply that krill patches were common in both pelagic and benthic habitat, whereas salps were common mainly in pelagic habitat. If the effects of deployments are similar between the years or regions, inter-annual or regional comparison using the penguin-mounted camera will be valid for characterising prey environment in the penguin foraging area.     

Distribution of salps near the South Shetland Islands during austral summer, 1990–1991 with special reference to krill distribution

Distribution and biomass of salps and Antarctic krill (Euphausia superba) were investigated near the South Shetland Islands during austral summer 1990–1991. Salp biomass ranged between 0 and 556 mgC·m^–3 and was greatest at a station in the Bransfield Strait in late December 1990. Salp biomass was lower than that of E. superba. Two species of salps; Salpa thompsoni and Ihlea racovitzai were found, and the former was dominant numerically. Spatial distribution and generation composition of these two species was different. Spatial distributions of salps and E. superba did not overlap particularly so the January–February period. While E. superba was found mainly in the coastal area which showed high-chlorophyll a values, salps exhibited high biomass in the oceanic area with low chlorophyll a concentrations. Predation by salps on small krill and the competitive removal of food by them, are discussed as potential reasons for the relatively low abundance of E. superba at the stations where salps were present in great numbers.     

Lipids of Antarctic salps and their commensal hyperiid amphipods

Antarctic salps (Salpa thompsoni and Ihlea racovitzai) and their commensal hyperiid amphipods (Vibilia antarctica, Cyllopus lucasii and C. magellanicus) were collected near Elephant Island, in the South Shetland Islands, during 1997 and the salp-rich year 1998. The sterol composition of aggregate S. thompsoni and I. racovitzai (mostly 24-methyl-5,22E-dien-3β-ol, 24-nordehydrocholesterol, cholesterol and trans-dehydrocholesterol) was reflected in the sterol composition of the commensal amphipods and was consistent with a herbivorous planktonic diet. This was not the case for solitary S. thompsoni, with 24-methylenecholesterol as the major sterol. There was a greater abundance of aggregate salp stanols in 1997 (11.7% total sterols) than 1998 (5.2%) and these different stanol levels were reflected in the commensal amphipods. Eicosapentaenoic acid [20:5(n-3)] and docosahexaenoic acid [22:6(n-3)] were the major polyunsaturated fatty acids (PUFA) in all organisms. Octadecapentaenoic acid [18:5(n-3)] comprised 0.4–5.8% (of total fatty acids) in all 1998 salps and amphipods, but was absent in 1997 samples. This suggests a greater presence of dinoflagellates or other species rich in 18:5(n-3) in the “salp year” 1998. Very long chain PUFA (C₂₄, C₂₆, C₂₈) were also only detected in 1998 samples (up to 5.3%), reflecting commensalism and greater presence of dinoflagellates or species containing very long chain PUFA. Examination of the biomarker lipids has provided an indication of trophic interactions for these Antarctic salps and their commensal hyperiid amphipods. Accepted: 8 November 1999     

Discovery of ciliates reproducing in the gut of Antarctic krill

During a recent Antarctic research cruise (December 1994/February 1995), dissection of fresh Antarctic krill (Euphausia superba) on board ship revealed live motile ciliates in the gut of krill. Further observation of gut samples by scanning electron microscopy indicated that the ciliates were symbionts located within the gut. We inferred that the ciliates may have enabled the krill to digest a wider range of food items, and as a consequence, this had became an important strategy for Antarctic krill's survival in the Antarctic ecosystem. Received: 1 August 1996 / Accepted: 1 December 1996     

The diet of Antarctic fur sealsArctocephalus gazella during the breeding season at South Georgia

 The diet of lactating female Antarctic fur seals Arctocephalus gazella at South Georgia was investigated during the pup-rearing period (January/March) of 1991–1994. Antarctic krill Euphausia superba was the main prey item, occurring in 88% of all scats (n=497), whereas fish occurred in 47% and squid in 5% of all scats. There was considerable intra- and inter-annual variation in the characteristics of krill taken by fur seals. The distribution of krill sizes taken suggests that fur seals are not actively selecting particular sizes of krill and, therefore, that the krill in the diet reflects the krill available around South Georgia. The absence of group 3 krill (44–48 mm in length) in the South Georgia area, as indicated by their absence in the diet of seals, is suggested as a possible reason for low availability of krill and the subsequent reproductive failure among krill predators. The frequency of occurrence of fish was much higher than in previous studies; the pattern of fish consumption showed a consistent seasonal pattern in 3 of the 4 years studied. Of the total number of the myctophid Protomyctophum choriodon, the most numerous fish taxon, 98% were taken between early February and the middle of March. Champsocephalus gunnari and Lepidonotothen larseni agg., which both feed on krill, dominated the fish component of the diet outside this period and together constituted 94% of the total estimated biomass of fish consumed. The intra- and inter-annual variability in the diet of Antarctic fur seals emphasise the need for diet studies to be conducted during the entire pup-rearing periods of several years. Received: 15 March 1995/Accepted: 13 May 1995     

Comparative role of salps and other zooplankton in the cycling and transport of selected elements and natural radionuclides in Mediterranean waters

Salps, salp fecal pellets and other zooplankton species were analyzed for a suite of elements and natural radionuclides to assess their role in the biogeochemical cycling of nuclides in oceanic waters. The nuclide/Al ratios in organisms normalized to the same ratio in crustal rock indicated that Ca, Sr, Zn, Cu, U,²¹⁰Po, and²¹⁰Pb are enriched in the organisms. The concentrations of Fe, Al, Th isotopes and²¹⁰Pb in salps and fecal pellets were about an order of magnitude higher than those in salps, whereas Ca, Cu, Zn, Mn and Po were higher by factors of about 2–5. Fluxes via salp defecation were higher than those which have been measured in crustacean zooplankton species, a result primarily due to the high defecation rates characteristic of salps. High nuclide levels in salp fecal pellets coupled with high defecation rates and presumed high salp biomass in many areas underscore the importance of these indiscriminate filter feeders in packaging and transporting to depth particulate-associated nuclides in surface waters.     

Biology of Salpa thompsoni in waters adjacent to the Ross Sea, Southern Ocean, during austral summer 2008

Depth-stratified vertical sampling was carried out during the New Zealand International Polar Year cruise to the Ross Sea on board the RV Tangaroa in February–March 2008. The distribution (horizontal and vertical), density and population biology of Salpa thompsoni were investigated. Salps were found at two of the four major sampling locations, e.g. near the continental slope of the Ross Sea and in the vicinity of seamounts to the north of the Ross Sea. Both abundance and biomass of S. thompsoni were highest near the seamounts in the Antarctic Circumpolar Current reaching ~2,500 ind 1,000 m⁻³ and 8.2 g dry wt 1,000 m⁻³ in the water column sampled. The data showed that S. thompsoni populations were able to utilize horizontal and vertical discontinuities in water column structure, in particular the warm Circumpolar Deep Water (CDW), to persist in the high Antarctic. Although salps appeared to continue migrating to the surface colder layers to feed, both aggregate chain and young embryo release seem to be restricted to the CDW. This study for the first time has provided evidence that low Antarctic salp species may successfully reproduce in the hostile high Antarctic realm.     

Melatonin and its possible role in mediating seasonal metabolic changes of Antarctic krill, Euphausia superba

Melatonin, the chief secretory product of the vertebrate pineal gland is suspected to be a ubiquitous molecule principally involved in the transduction of photoperiodic information. Besides vertebrates, melatonin has been detected throughout phylogeny in numerous non-vertebrate taxa. In the present study, the occurrence of melatonin in Antarctic krill Euphausia superba and its possible role in mediating seasonal metabolic changes was evaluated. Melatonin was quantified by enzyme linked immunosorbent assay (ELISA) in high performance liquid chromatography (HPLC) purified extracts of eyestalks and hemolymph of krill sampled in the Lazarev Sea during the Antarctic winter and summer. In addition, oxygen uptake rates and the activities of the metabolic enzyme malate dehydrogenase (MDH) were recorded to assess the metabolic status of krill. Validation of melatonin measurements was carried out on the basis of three different extraction methods with parallel determination of melatonin by ELISA in crude extracts and in HPLC purified extracts, and after derivatization of melatonin under alkaline conditions in the presence of hydrogen peroxide. A significantly higher respiration rate and MDH activity was found in summer krill than in winter krill indicating that krill was in a state of reduced metabolic activity during winter. However, neither during winter nor during summer there were detectable melatonin concentrations in the visual system or hemolymph of krill. Based on these results, we question a mediating role of melatonin in the control of seasonal metabolic changes in Antarctic krill.     

Krill (Euphausia superba) density, proportional and absolute recruitment and biomass in the Elephant Island region (Antarctic Peninsula) during the period 1977 to 1997

The Elephant Island region (Antarctic Peninsula) was selected as a long-term monitoring site to describe the interannual variability of important krill stock parameters. The analysis reviewed and updated krill density and proportional recruitment indices. Krill absolute recruitment and biomass from net sampling surveys are introduced as additional indices from this time series. New survey results from the past two seasons indicate a very successful 1994/1995 year-class and slightly below average proportional recruitment of the 1995/1996 krill year-class. Absolute recruitment of the 1995/1996 year-class was high compared to preceding years, because total stock size was relatively high in 1996/1997. After a period of low krill density and biomass in the area for almost a decade, krill density and biomass have increased. Caution is expressed as to whether this observed increase represents a long-term recovery of the stock. Received: 15 October 1997 / Accepted: 5 January 1998     

Environmental influences on zooplankton and micronekton distribution in the Bransfield Strait and adjacent waters

The spatial distribution, biomass and size structure of key mesozooplankton species and micronekton in the Bransfield Strait (Antarctica) are described in relation to environmental variables during the austral summer 2001. Stratified (BIONESS) biological sampling at five depth-ranges and CTD casts were performed at 40 stations, including a cross-Strait transect. Six families, 11 genera and 16 species composed the total catch of larval and juvenile fish, which were more abundant in the upper 75 m. Trematomus scotti was the most abundant fish and large individuals dominated at depths >75 m. The fish distribution patterns were associated with the main water masses in the area and with bottom depth. Spatial segregation in density and biomass of krill (Euphausia superba) and salps (Salpa thompsoni) was observed at a relatively small scale, in relation to the main water masses. Also, size-related spatial segregation was found in krill. The present study suggests that the spatial distribution of krill, salps and early life stages of fish, are influenced by the main water masses in the Strait, and that species may adapt their reproductive strategies not only to seasonal production peaks, but to transport processes within water bodies that maximise their fitness through optimum temperature and/or trophic environments.     

Biometry and trophodynamics of Salpa thompsoni foxton (Tunicata: Thaliacea) near the Antarctic Peninsula in austral summer, 1983–1984

Summary Salpa thompsoni aggregates have a biometric profile similar to that of mid-latitude salps, being approximately 96% water and only about 1% organic matter. Of its ash-free dry weight about 13.6% is carbon and 3.1% is nitrogen. All weight parameters were related to body length by a power function of 2.3. S. thompsoni was a dominant member of the zooplankton community in waters near the Antarctic Peninsula in the austral summer of 1983–1984, with biomass in mid-March ranging from 49 to 671 mgCm^–2 and 9.6 to 146 mgNm^–2. Three subpopulations occurred in separate regions: (1) the Bransfield Strait, (2) north of the South Shetland Islands, and (3) west of Elephant Island; these were dominated by 20, 30 and 40 mm individuals, respectively. Biomass was greatest in the Bransfield Strait and least near the South Shetland Islands. Clearance rate measurements of S. thompsoni on naturally occurring particulate matter were significantly lower than those previously reported for mid-latitude salps, a result which we attribute primarily to the effect of low habitat temperature (ca. 1°C). Direct measurements suggest that fecal production by 21 mm blastozoids is equivalent to 10.2% body carbon d^–1 and 6.6% body nitrogen d^–1. Grazing by S. thompsoni may have removed a majority of the daily primary production in March, but <1% in January. The comparatively low biomass of krill larvae in 1983–1984 may be attributed partially to competitive removal of food by salps, but an equally important effect may have been direct predation.     

Penguins, fur seals, and fishing: prey requirements and potential competition in the South Shetland Islands, Antarctica

Antarctic and sub-Antarctic seabirds, marine mammals, and human fisheries concentrate their foraging efforts on a single species, Antarctic krill (Euphausiasuperba). Because these predators may have a significant effect on krill abundance, we estimated the energy and prey requirements of Adelie (Pygoscelisadeliae), chinstrap (Pygoscelisantarctica), and gentoo (Pygoscelispapua) penguins and female Antarctic fur seals (Arctocephalusgazella) breeding on the South Shetland Islands, Antarctica and compared these estimates with catch statistics from the Antarctic krill fishery. Published data on field metabolic rate, population size, diet, prey energy content, and metabolic efficiency were used to estimate prey requirements of these breeding, adult, land-based predators and their dependent offspring. Due to their large population size, chinstrap penguins were the most significant krill predators during the period examined, consuming an estimated 7.8 × 10⁸ kg krill, followed by Adelie penguins (3.1 × 10⁷ kg), gentoo penguins (1.2 × 10⁷ kg), and Antarctic fur seals (3.6 × 10⁶ kg). Total consumption of all land-based predators on the South Shetland Islands was estimated at 8.3 × 10⁸ kg krill. The commercial krill fishery harvest in the South Shetland Island region (1.0 × 10⁸ kg) was approximately 12% of this. Commercial harvest coincides seasonally and spatially with peak penguin and fur seal prey demands, and may affect prey availability to penguins and fur seals. This differs from the conclusions of Ichii et al. who asserted that the potential for competition between South Shetland predators and the commercial krill fishery is low. Received: 26 August 1997 / Accepted: 16 December 1997     

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

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

Spatial and temporal variability in reproductive timing of Antarctic krill (Euphausia superba Dana)

Spawning dates of Antarctic krill, Euphausia superba Dana, were calculated from larval stage compositions, and corrected using data on maturity stage composition of the adult krill. Both original and literature data obtained from the Antarctic Peninsula-Bellingshausen Sea area and around the Antarctic continent were used. A time series (1975/76–1986/87) for several subareas of the Antarctic Peninsula-Bellingshausen Sea area indicates considerable variation in the krill spawning start, maxima and completion. In particular years (1975/76, 1980/81), krill spawning in the western Atlantic sector began relatively early, was intensive, and completed early. Some years (1977/78, 1981/82) were characterised by long and non-synchronised krill spawning. Compiled data sets for the Atlantic sector (1980/81), the entire Antarctic (1983/84) and the east Indian-west Pacific Antarctic waters (1981–85) reveal some spatial patterns in krill reproductive timing. In relation to spawning timing variation, the habitats of the krill population fall into five categories: (1) areas with an early beginning (late Novemberearly December) and a variable, but normally long, duration (3–3.5 months) of krill spawning; this is generally the southern boundary of the Antarctic Circumpolar Current, (2) areas with an early beginning, but a short duration of krill spawning (Gerlache Strait), (3) areas with a highly variable (within 1–1.5 months) beginning and a relatively long duration (ca. 3 months) of krill spawning (Bransfield Strait, Palmer Archipelago), (4) areas with a late beginning (late December–January) and a long duration of krill spawning (Bellingshausen Sea, D'Urville Sea, and Balleny Islands area), and (5) areas with a delayed beginning, but a very short duration (ca. 1.5 months) of krill spawning (Ross Sea slope, probably the Coastal Current area off the Lasarev Sea shelf and in the south-eastern Weddell Sea. These patterns can be partly explained by peculiarities of the ice regime in particular areas and by routes of krill movement within water circulation systems.     

Beaching of the tunicate<Emphasis Type="Italic"> Salpa thompsoni</Emphasis> at high levels of suspended particulate matter in the Southern Ocean

A mass death event of the pelagic tunicate Salpa thompsoni, which occurred in April 2002 in Potter Cove near the Argentinean/German Antarctic station Jubany (62°14′S–58°40′W), King George Island, South Shetland Islands, is described. Salps appeared on the beach 2 days after very strong (>80 km.h⁻¹) winds were registered, which accumulated particulate material in the inner part of the cove and probably also re-suspended bottom sediments. The sharp increase in particulate-matter concentrations in Potter Cove caused clogging of salp mucous filtering nets, and probably a combination of clogging, winds and tides caused dying salps to be washed out onto the beach.     

Spatial distributions and population dynamics of two salp species, Ihlea racovitzai and Salpa thompsoni, in the waters north of Lützow-Holm Bay (East Antarctica) during austral summers of 2005 and 2006

Information on the sequences of generations and reproductive states of salps in the Southern Ocean is essential for an improved understanding of salp population growth, although changes in distribution patterns of two species of salps, Salpa thompsoni and Ihlea racovitzai, in the Southern Ocean have the potential to alter the Southern Ocean ecosystem. We used stratified, quantitative sampling from the surface to 2,000 m depth with an RMT 8 net in January of 2005 and 2006 to determine the distribution and population structure of salps in the north of Lützow-Holm Bay, East Antarctica. Ihlea racovitzai occurred in both 2005 and 2006, but S. thompsoni was found only in 2005. Ihlea racovitzai occurred abundantly along the ice edge where Antarctic Winter Water was well developed, whereas S. thompsoni was more abundant at northern stations affected by warm Modified Circumpolar Deep Water. Small solitary stages of I. racovitzai dominated in 2005, but they had declined significantly by 2006. The S. thompsoni population was composed of small immature aggregates and mature solitary stages, suggesting that the solitary stages were reproducing. We did not find mature aggregate and immature solitary stages in the present study and thus suggested that S. thompsoni was unable to complete its life cycle in the north of Lützow-Holm Bay because of failure of sexual reproduction in the aggregate stage. The S. thompsoni population was therefore probably transported to our study area by advection.     

Food and feeding ecology of emperor penguins in the eastern Weddell Sea

Summary The diet of the emperor penguin Aptenodytes forsteri in the eastern Weddell Sea, Antarctica was studied during October and November 1986 by stomach content analysis. Emperor penguins fed mainly on Antarctic krill Euphausia superba, Antarctic silverfish Pleuragramma antarcticum and squid Psychroteuthis glacialis. Benthic prey was not found. The prey composition suggests two different feeding strategies, shallow dives exploring the rugged underside of sea ice where krill is taken, and deep dives when mesopelagic fish and squid are consumed. Chicks were fed on average every 1.44 days.     

A comparison of Antarctic petrel (Thalassoica antarctica) diets with net samples of Antarctic krill (Euphausia superba) taken from the Prydz Bay region

Samples of the stomach contents of Antarctic petrels (Thalassoica antarctica) were obtained on board ship in the Prydz Bay region of Antarctica from birds which had spontaneously regurgitated. The weight of food and the species composition of the stomach contents were measured. Antarctic krill,Euphausia superba, the sole prey item taken, were compared to krill obtained by nets in the same region as part of a large-scale krill survey. Krill from petrel stomach samples were larger in mean size than those sampled by nets. This may be attributed to selection of the larger sized krill by the petrels, it may be caused by the nets sampling different populations of krill or it may be due to net avoidance by the larger krill.     

The biochemical composition, energy content, and chemical antifeedant defenses of the common Antarctic Peninsular sea stars Granaster nutrix and Neosmilaster georgianus

The sea stars Granaster nutrix and Neosmilaster georgianus are conspicuous members of benthic communities along the Antarctic Peninsula. An analysis of the proximate composition of somatic body components of nonreproductive adults indicates the nutrient storage organs (pyloric caeca) are rich in both protein (60.7 and 60.6% mean dry wt, respectively) and lipid (25.4 and 29.8% mean dry wt, respectively). Body-wall tissues, while containing small inconspicuous skeletal ossicles, are also comprised of significant levels of organic matter (33.5 and 55.7% mean dry wt, respectively), attributable primarily to protein. Both the pyloric caeca and body-wall tissues are relatively rich in energy (mean energy levels=24.8 and 26.5 kJ g⁻¹ dry wt; 8.4 and 14.1 kJ g⁻¹ dry wt, respectively). Despite the availability of these nutrients and energy neither sea star is preyed upon by the sympatric omnivorous sea star Odontaster validus, a common predator of other Antarctic sea stars. Laboratory feeding bioassays indicate that O. validus rejects live intact individuals and body-wall tissues of both sea star species while readily consuming dried krill. Alginate food pellets containing a krill powder and tissue level concentrations of organic methanol extracts of body-wall tissues were also rejected by O. validus. Moreover, the copious mucus released from the body wall of N. georgianus was deterrent in O. validus food pellet bioassays. Thus, both sea stars evidently possess defensive secondary metabolites that defend against predation and are likely to play a role in mediating materials and energy transfer in the Antarctic benthos.     

Aspects of the foraging behaviour of the Antarctic Tern Sterna vittata gaini at Harmony Point, South Shetland Islands

During January and February of 2002 and 2003, we studied the diet of the Antarctic Tern Sterna vittata gaini at two colonies in Nelson Island, South Shetland Islands, by identifying the prey fed to chicks by breeders. The fish Notothenia coriiceps was the main prey in both seasons, followed by the myctophid Electrona antarctica, Antarctic krill Euphausia superba and gammarid amphipods. The contribution of fish to the diet increased as chicks grew older. Fish and amphipods were brought to chicks during the day, whereas adults brought Antarctic krill at sunrise and sunset. Both the duration of the feeding trips and the number of trips per foraging bout varied according to the type of prey caught. Preliminary information suggests that, among other causes, the foraging strategy is strongly influenced by the predation pressure of skuas on chicks. Results are compared with the only two previous study on the diet of the Antarctic Tern at the South Shetland Islands.