Abundance of plankton in the upper 200 m layer in tropical oligotrophic regions of the Pacific and Indian Oceans

Quantitative distribution of plankton (mostly mesoplankton) is studied in the upper 200 m layer of oligotrophic waters in tropical anticyclonic gyres of the Pacific and Indian Oceans. Some general features of its trophic and taxonomic structures and vertical distribution are described.

Marine diatoms in deep sea sediments

Long-term evolution is thought to take opportunities that arise as a consequence of mass extinction (as argued, for example, by Gould, 2002) and the following biotic recovery, but there is absolutely no evidence for this being the case. However, our study shows that eutrophication by oceanic mixing also played a part in the enhancement of several evolutionary events amongst marine organisms, and these results could indicate that the rates of oceanic biodiversification may be slowed if upwelling becomes weakened by future global warming. This paper defines three distinct evolutionary events of resting spores of the marine diatom genus Chaetoceros, to reconstruct past upwelling through the analysis of several DSDP, ODP and land-based successions from the North, South and equatorial Pacific as well as the Atlantic Ocean during the past 40 million years. The Atlantic Chaetoceros Explosion (ACE) event occurred across the E/O boundary in the North Atlantic, and is characterized by resting spore diversification that occurred as a consequence of the onset of upwelling following changes in thermohaline circulation through global cooling in the early Oligocene. Pacific Chaetoceros Explosion events-1 and -2 (PACE-1 and PACE-2) are characterized by relatively higher occurrences of iron input following the Himalayan uplift and aridification at 8.5 Ma and ca. 2.5 Ma in the North Pacific region. These events not only enhanced the diversification and increased abundance of primary producers, including that of Chaetoceros, other diatoms and seaweeds, but also stimulated the evolution of zooplankton and larger predators, such as copepods and marine mammals, which ate these phytoplankton and plants. Current thinking suggests new evolutionary niches open up after a mass extinction, but our study finds that eutrophication can also stimulate evolutionary diversification. Moreover, in the opposite fashion, our results show that as thermohaline circulation abates, global warming progresses and the ocean surface becomes warmer, many marine organisms will be affected by the environmental degradation.

(Table 1) Biomass (as energy) of different elements of the planktonic community in the Equatorial Pacific in the 0-150 m layer

An analysis was made of composition and content of nutrients, salts, particulate and dissolved organic matter, and various plankton groups in a series of samples collected by a 140-liter sampling bottle to depth up to 150 m at 4 equatorial stations between 97° and 154°W. Large and small phytoplankton, bacteria (aggregated and dispersed), heterotrophic flagellates, infusorians, radiolarians, foraminifers, fine filter-feeders, small and large, mostly herbivorous copepods, cyclopoids, predatory calanoids, and other predators were investigated separately. Trophic relations between these elements are established from personal and published data, and rate of their metabolism and some other physiological parameters are determined. Such functional characteristics as extent of satisfaction of food requirements of organisms belonging to various trophic groups, intensity of trophic relations, balance between production and consumption by individual elements of the community, ecological efficiency, and net and specific production of the groups distinguished, of individual trophic levels, of total zooplankton, and of the community as a whole are calculated. Variations of these characteristics along the equator with decreasing upwelling intensity are examined and their possible causes and mechanisms are discussed.

Zoobenthos and oceanographical parameters in the Kemskaya Guba (Bay) - estuary of the Kem' River entering the Onega Bay, White Sea

Data on distribution of zoobenthos in the Kemskaya Guba (or Kemskaya Bay - the estuary of the Kem' River entering the Onega Bay of the White Sea), which is strongly influenced by river runoff, are presented. The number of species at sampling stations varied from 4 to 65. Density of communities and zoobenthos biomass varied from 342±68 to 4293±96 #/m**2 and from 0.418±0.081 to 1975.22±494.36 g/m**2, respectively. Shannon index values varied between 1.19 to 4.7 bit/ind. At the upper part of the estuary, detritivores dominated, while in the central part and at outlets sestonophages prevailed. Changes in quantitative parameters of the zoobenthos along gradient of water salinity were traced, and relations of these parameters with seven other environmental factors were revealed. It was found that species composition, biodiversity, and trophic structure of the zoobenthos significantly correlated with some of parameters mentioned above. Multiple regression analysis was used to assess combined effect of factors, and it revealed which of them played a determining role in Kemskaya Guba: for species composition - depth, water color, and total concentration of suspended matter; for number of species - contents of <0.01 mm grain size (pelite) fraction and organic carbon in bottom sediments. Biomass depended on water salinity, water chromaticity, and organic carbon contents in bottom sediments and suspended matter. Values of the Shannon index of diversity are determined by water color, and contents of organic carbon and pelite fraction in bottom sediments. Calculations of ecological stress values revealed two zones with unstable state of the zoobenthos.

Carbon/Nitrogen ratio of plant tissues and biomass of belowground consumers after 11 years of experimental treatment, Toolik lake

Theory and observation indicate that changes in the rate of primary production can alter the balance between the bottom-up influences of plants and resources and the top-down regulation of herbivores and predators on ecosystem structure and function. The Exploitation Ecosystem Hypothesis (EEH) posited that as aboveground net primary productivity (ANPP) increases, the additional biomass should support higher trophic levels. We developed an extension of EEH to include the impacts of increases in ANPP on belowground consumers in a similar manner as aboveground, but indirectly through changes in the allocation of photosynthate to roots. We tested our predictions for plants aboveground and for phytophagous nematodes and their predators belowground in two common arctic tundra plant communities subjected to 11 years of increased soil nutrient availability and/or exclusion of mammalian herbivores. The less productive dry heath (DH) community met the predictions of EEH aboveground, with the greatest ANPP and plant biomass in the fertilized plots protected from herbivory. A palatable grass increased in fertilized plots while dwarf evergreen shrubs and lichens declined. Belowground, phytophagous nematodes also responded as predicted, achieving greater biomass in the higher ANPP plots, whereas predator biomass tended to be lower in those same plots (although not significantly). In the higher productivity moist acidic tussock (MAT) community, aboveground responses were quite different. Herbivores stimulated ANPP and biomass in both ambient and enriched soil nutrient plots; maximum ANPP occurred in fertilized plots exposed to herbivory. Fertilized plots became dominated by dwarf birch (a deciduous shrub) and cloudberry (a perennial forb); under ambient conditions these two species coexist with sedges, evergreen dwarf shrubs, and Sphagnum mosses. Phytophagous nematodes did not respond significantly to changes in ANPP, although predator biomass was greatest in control plots. The contrasting results of these two arctic tundra plant communities suggest that the predictions of EEH may hold for very low ANPP communities, but that other factors, including competition and shifts in vegetation composition toward less palatable species, may confound predicted responses to changes in productivity in higher ANPP communities such as the MAT studied here.

Biomass of nanoprotozooplankton in the Atlantic sector of the Southern Ocean

The dynamic of early spring nanoprotozoa was investigated in three characteristic water masses of the Southern Ocean: the Marginal Ice Zone, the intermediate waters of the Antarctic Circumpolar Current and the Polar Frontal Zone. Biomass and feeding activities of nanoprotozoa were measured, as well as the biomass of their potential prey-bacteria and phototrophic flagellates-on the 6°W meridian in the Southern Ocean along three repetitive transects between 47 and 60° South from October to November 1992. On average, nanoprotozooplankton biomass accounted for 77% of the combined biomass of bacteria and phototrophic flagellates, and was dominated by dinoflagellates and flagellates smaller than 5 µm. As a general trend, low protozoan biomass of 2 mg C/m**3 was typical of the ice covered area, while significantly higher biomasses culminating at 15 mg C/m**3 were recorded at the Polar Front. Biomasses of bacteria and total phytoplankton were distributed accordingly, with larger values at the Polar Front. Phototrophic flagellates did not show any geographical trend. No seasonal trend could be identified in the Marginal Ice Zone and in the intermediate waters of the Antarctic Circumpolar Current. On the other hand, at the Polar Front region a three-fold increase was observed within a 2-month period for nanoprotozooplankton biomass. Such a biomass increase was also detected for bacterioplankton and total phytoplankton biomass. Half-saturation constants and maximum specific ingestion of nanoprotozoan taxons feeding on bacteria and phototrophic flagellates were determined using the technique of fluorescent labelled bacteria (FLB) and algae (FLA) over a large range of prey concentrations. Maximum ingestion rates ranged between 0.002 and 0.015/h for bactivorous nanoprotozoa and heterotrophic flagellates larger than 5 µm feeding on phototrophic flagellates. The markedly high maximum ingestion rates of 0.4/h characterising nanophytoplankton ingestion by dinoflagellates evidenced the strong ability of dinoflagellates for feeding on nanophytoplankton. Daily ingestion rates were calculated from nanoprotozoan grazing parameters and carbon biomass of prey and predators. This indicated that nanoprotozoa ingestion of daily bacterioplankton and phytoplankton production in early spring ranged from 32 to 40%.

Barnacle recurit density, biomass, and reproductive output measured in rocky intertidal habitats in Glasgow Head, on the Atlantic coast of Nova Scotia, Canada

Through a field experiment, we show that a predator has negative nonconsumptive effects (NCEs) on different life-history stages of the same prey species. Shortly before the recruitment season of the barnacle Semibalanus balanoides (May-June), we established experimental cages in rocky intertidal habitats in Nova Scotia, Canada. The cages were used to manipulate the presence and absence of dogwhelks, Nucella lapillus, the main predators of barnacles. At the centre of each cage, we installed a tile where barnacle pelagic larvae could settle and the resulting recruits grow. Mesh prevented caged dogwhelks from accessing the tiles, but allowed waterborne dogwhelk cues to reach the tiles. Results in May indicated that barnacle larvae settled preferentially on tiles from cages without dogwhelks. In November, at the end of the dogwhelk activity period and once the barnacle recruits had grown to adult size, barnacle body mass was lower in the presence of dogwhelks. This limitation may have resulted from a lower barnacle feeding activity with nearby dogwhelks, as found by a previous study. The observed larval and adult responses in barnacles are consistent with attempts to decrease predation risk. November data also indicated that dogwhelk cues limited barnacle reproductive output, a possible consequence of the limited growth of barnacles. Overall, this study suggests that a predator species might influence trait evolution in a prey species through NCEs on different life-history stages.

Biometrics, fatty acid composition and stable isotopic ratios of five seabird species from Kongsfjorden, Svalbard

Marine birds are important predators in the marine ecosystem, and dietary studies can give useful information about their feeding ecology, food webs and oceanographic variability. The aim of this study was to increase our understanding of the diet and trophic level of the seabirds breeding in Kongsfjorden, Svalbard. We have used fatty acids and stable isotopes, both of which integrate diet information over space and time, to determine trophic relationships in marine food webs. Fatty acid compositions of muscle from Little auk (Alle alle), Brünnich's guillemot (Uria lomvia), Black-legged kittiwake (Rissa tridactyla), Northern fulmar (Fulmarus glacialis) and Glaucous gull (Larus hyperboreus) were determined and compared with their prey species. Canonical analysis (CA) showed that fatty acid composition differed among the five seabird species. Little auk, Black-legged kittiwake and Northern fulmar had high levels of the Calanus markers 20:1n9 and 22:1, indicating that these seabirds are a part of the Calanus food chain. Brünnich's guillemot differed from the other species with much lower levels of 20:1n9 and 22:1. Brünnich's guillemot is a pursuit diver feeding on fish and amphipods deeper in the water column, below 30 m. Glaucous gull also differed from the other seabird species, with a larger variation in the fatty acid composition indicating a more diverse diet. Trophic level analysis placed Little auk at the lowest trophic level, Brünnich's guillemot and Black-legged kittiwake at intermediate levels and Glaucous gull and Northern fulmar at the highest trophic level.

Chlorophyll a concentration, and proportion of time in the water column obtained by instrumented bowhead whales

Sixty hours of direct measurements of fluorescence were collected from six bowhead whales (Balaena mysticetus) instrumented with fluorometers in Greenland in April 2005 and 2006. The data were used to (1) characterize the three-dimensional spatial pattern of chlorophyll-a (Chl-a) in the water column, (2) to examine the relationships between whale foraging areas and productive zones, and (3) to examine the correlation between whale-derived in situ values of Chl-a and those from concurrent satellite images using the NASA MODIS (Moderate Resolution Imaging Spectroradiometer) EOS-AQUA satellite (MOD21, SeaWifs analogue OC3M and SST MOD37). Bowhead whales traversed 1600 km**2, providing information on diving, Chl-a structure and temperature profiles to depths below 200 m. Feeding dives frequently passed through surface waters ( >50 m) and targeted depths close to the bottom, and whales did not always target patches of high concentrations of Chl-a in the upper 50 m. Five satellite images were available within the periods whales carried fluorometers. Whales traversed 91 pixels collecting on average 761 s (SD 826) of Chl-a samples per pixel (0-136 m). The depth of the Chl-a maximum ranged widely, from 1 to 66 m. Estimates of Chl-a made from the water-leaving radiance measurements using the OC3M algorithm were highly skewed with most samples estimated as <1 mg/m**3 Chl-a, while data collected from whales had a broad distribution with Chl-a reaching >9 mg/m**3. The correlation between the satellite-derived and whale-derived Chl-a maxima was poor, a linear fit explained only 10% of the variance.

Dive depth and dive duration data of female southern elephant seals from Marion Island between 2004 and 2008 with links to datasets

The at-sea behaviour of marine top predators provides valuable insights into the distribution of prey species and strategies used by predators to exploit patchily distributed resources. We describe the water column usage and dive strategies of female southern elephant seals from Marion Island tracked between 2004 and 2008. Dives representing increases in forage effort were identified using a method that combines dive type analyses and the calculation of relative amounts of time that animals spend in the bottom phases of dives. Results from this analysis indicate that female elephant seals from Marion Island tend to display lower levels of forage effort closer to the island and display intensive opportunistic forage bouts that occur at a minimum distance of approximately 215 km from the island. Females from Marion Island dived deeper and for longer periods of time, compared to females from other populations. Most animals displayed positive diel vertical migration, evidently foraging pelagically on vertically migrating prey. A few animals displayed periods of reverse (negative) diel vertical migration, however, diving to deeper depths at night, compared to daytime. This behaviour is difficult to explain and prey species targeted during such periods unknown. Our results illustrate plasticity in foraging behaviour of southern elephant seals, as well as inter-population differences in forage strategies.

(Table 1) Movement parameters of nine adult female snowy owls (Bubo scandiacus) tracked during the winter period 07/08 and 08/09 around Hudson Strait, Canada

Top predators of the arctic tundra are facing a long period of very low prey availability during winter and subsidies from other ecosystems such as the marine environment may help to support their populations. Satellite tracking of snowy owls, a top predator of the tundra, revealed that most adult females breeding in the Canadian Arctic overwinter at high latitudes in the eastern Arctic and spend several weeks (up to 101 d) on the sea-ice between December and April. Analysis of high-resolution satellite images of sea-ice indicated that owls were primarily gathering around open water patches in the ice, which are commonly used by wintering seabirds, a potential prey. Such extensive use of sea-ice by a tundra predator considered a small mammal specialist was unexpected, and suggests that marine resources subsidize snowy owl populations in winter. As sea-ice regimes in winter are expected to change over the next decades due to climate warming, this may affect the wintering strategy of this top predator and ultimately the functioning of the tundra ecosystem.

(Table 2) Mercury and selenium concentration in the brain stem of wild polar bears (Ursus maritimus), east Greenland

Polar bears (Ursus maritimus) are exposed to high concentrations of mercury because they are apex predators in the Arctic ecosystem. Although mercury is a potent neurotoxic heavy metal, it is not known whether current exposures are of neurotoxicological concern to polar bears. We tested the hypotheses that polar bears accumulate levels of mercury in their brains that exceed the estimated lowest observable adverse effect level (20 µg/g dry wt) for mammalian wildlife and that such exposures are associated with subtle neurological damage, as determined by measuring neurochemical biomarkers previously shown to be disrupted by mercury in other high-trophic wildlife. Brain stem (medulla oblongata) tissues from 82 polar bears subsistence hunted in East Greenland were studied. Despite surprisingly low levels of mercury in the brain stem region (total mercury = 0.36 ± 0.12 µg/g dry wt), a significant negative correlation was measured between N-methyl-D-aspartate (NMDA) receptor levels and both total mercury (r = -0.34, p < 0.01) and methylmercury (r = -0.89, p < 0.05). No relationships were observed among mercury, selenium, and several other neurochemical biomarkers (dopamine-2, gamma-aminobutyric acid type A, muscarinic cholinergic, and nicotinic cholinergic receptors; cholinesterase and monoamine oxidase enzymes). These data show that East Greenland polar bears do not accumulate high levels of mercury in their brain stems. However, decreased levels of NMDA receptors could be one of the most sensitive indicators of mercury's subclinical and early effects.

(Table 1) Foraging trip characteristics of male Adélie penguins (Pygoscelis adeliae) during their first incubation trip at Pointe Géologie

Knowledge of habitat use by top marine predators in response to environmental conditions is crucial in the current context of global changes occurring in the Southern Ocean. We examined the at-sea locations of male Adelie penguins (Pygoscelis adeliae) breeding at Dumont d'Urville during their first, long incubation trip. Compared with the chick-rearing period, penguins performed longer trips, going to oceanic waters as far as 320 km from the colony. We observed 3 strategies: (1) five individuals covered large distances to the north, targeting open-ocean areas and following the currents of two persistent eddies; (2) five individuals foraged to the north-west, close to the Antarctic shelf slope at the limit of the pack ice; and (3) three individuals covered much shorter distances (northwards or eastwards). The foraging range also seemed to be limited by the body condition of the penguins before their departure to sea.