Sea ice and the onshore–offshore gradient in pre-winter zooplankton assemblages in southeastern Beaufort Sea

Zooplankton communities were studied in southeastern Beaufort Sea (Arctic Ocean) in September–October 2002. Cluster analysis and non-metric multidimensional scaling revealed three distinct mesozooplankton assemblages. A neritic assemblage occurred on the Mackenzie Shelf and in Franklin Bay, while distinct off-shelf assemblages prevailed in the Cape Bathurst Polynya and on the Beaufort Slope respectively. Over 95% of the mesozooplankton was comprised of eight copepod taxa. Pseudocalanus spp. contributed predominantly to the discrimination of the three assemblages and was the only significant indicator of the Shelf assemblage. Oithona similis, Oncaea borealis, Metridia longa and Calanus hyperboreus were indicators of the Polynya assemblage. Cyclopina sp. and Microcalanus pygmaeus were indicative of the overall off-shelf community (Polynya and Slope assemblages). The importance of omnivores and carnivores increased from the shelf to the polynya and the slope. Station depth and duration of reduced ice conditions during summer (< 50% ice concentration) underpinned the distribution of the assemblages (r² = 0.71 and 0.45 respectively). The abundance of Pseudocalanus spp. was independent of depth and increased with the duration of reduced ice conditions (r_s = 0.438). The abundance of Cyclopina sp., M. pygmaeus and other indicators of the offshore assemblages followed the opposite trend (r_s = − 0.467 and − 0.5 respectively). Under continued climate warming, a reduction of the ice cover will affect the biogeography of mesozooplankton on and around the Mackenzie Shelf, to the potential advantage of Pseudocalanus spp. and other calanoid herbivores.     

Suspended sediment and erosion dynamics in Kugmallit Bay and Beaufort Sea during ice-free conditions

The Mackenzie River is the largest river on the North American side of the Arctic and its huge freshwater and sediment load impacts the Canadian Beaufort Shelf. Huge quantities of sediment and associated organic carbon are transported in the Mackenzie plume into the interior of the Arctic Ocean mainly during the freshet (May to September). Changing climate scenarios portend increased coastal erosion and resuspension that lead to altered river-shelf-slope particle budgets. We measured sedimentation rates, suspended particulate matter (SPM), particle size and settling rates during ice-free conditions in Kugmallit Bay (3–5 m depth). Additionally, measurements of erosion rate, critical shear stress, particle size distribution and resuspension threshold of bottom sediments were examined at four regionally contrasting sites (33–523 m depth) on the Canadian Beaufort Shelf using a new method for assessing sediment erosion. Wind induced resuspension was evidenced by a strong relationship between SPM and wind speed in Kugmallit Bay. Deployment of sediment traps showed decreasing sedimentation rates at sites along an inshore–offshore transect ranging from 5400 to 3700 g m^− 2 day^− 1. Particle settling rates and size distributions measured using a Perspex settling chamber showed strong relationships between equivalent spherical diameter (ESD) and particle settling rates (r² = 0.91). Mean settling rates were 0.72 cm s^− 1 with corresponding ESD values of 0.9 mm. Undisturbed sediment cores were exposed to shear stress in an attempt to compare differences in sediment stability across the shelf during September to October 2003. Shear was generated by vertically oscillating a perforated disc at controlled frequencies corresponding to calibrated shear velocity using a piston grid erosion device. Critical (Type I) erosion thresholds (u) varied between 1.1 and 1.3 cm s^− 1 with no obvious differences in location. Sediments at the deepest site Amundsen Gulf displayed the highest erosion rates (22–54 g m^− 2 min^− 1) with resuspended particle sizes ranging from 100 to 930 µm for all sites. There was no indication of biotic influence on sediment stability, although our cores did not display a fluff layer of unconsolidated sediment. Concurrent studies in the delta and shelf region suggest the importance of a nepheloid layer which transports suspended particles to the slope. Continuous cycles of resuspension, deposition, and horizontal advection may intensify with reduction of sea ice in this region. Our measurements coupled with studies of circulation and cross-shelf exchange allow parameterization and modeling of particle dynamics and carbon fluxes under various climate change scenarios.     

Primary productivity and export fluxes on the Canadian shelf of the Beaufort Sea: A modelling study

We present a coupled sea ice–ocean-biological (including ice algae) model in the Arctic Ocean. The 1D model was developed and implemented on the Canadian Beaufort Sea shelf to examine the importance of different physical processes in controlling the timing and magnitude of primary production and biogenic particle export over an annual cycle (1987). Our results show that the snow and sea ice cover melt and/or break-up controls the timing of the phytoplankton bloom but primary producers (ice algae and phytoplankton) on the outer shelf are essentially nutrient limited. The total annual primary production (22.7 to 27.7 g-C m^? 2) is thus controlled by nutrient “pre-conditioning” in the previous fall and winter and by the depth of wind mixing that is controlled in part by the supply of fresh water at the end of spring (ice melt or runoff). The spring bloom represents about 40% of the total annual primary production and occurs in a period of the year when sampling is often lacking. Time interpolation of observed values to obtain total annual primary production, as done in many studies, was shown to lead to an underestimation of the actual production. Our simulated ratios of export to primary production vary between 0.42 and 0.44.     

Physical and biological correlates of virus dynamics in the southern Beaufort Sea and Amundsen Gulf

As part of the Canadian Arctic Shelf Exchange Study (CASES), we investigated the spatial and seasonal distributions of viruses in relation to biotic (bacteria, chlorophyll-a (chl a)) and abiotic variables (temperature, salinity and depth). Sampling occurred in the southern Beaufort Sea Shelf in the region of the Amundsen Gulf and Mackenzie Shelf, between November 2003 and August 2004. Bacterial and viral abundances estimated by epifluorescence microscopy (EFM) and flow cytometry (FC) were highly correlated (r² = 0.89 and r² = 0.87, respectively), although estimates by EFM were slightly higher (FC = 1.08 × EFM + 0.12 and FC = 1.07 × EFM + 0.43, respectively). Viral abundances ranged from 0.13 × 10⁶ to 23 × 10⁶ ml^− 1, and in surface waters were ~ 2-fold higher during the spring bloom in May and June and ~ 1.5-fold higher during July and August, relative to winter abundances. These increases were coincident with a ~ 6-fold increase in chl a during spring and a ~ 4-fold increase in bacteria during summer. Surface viral abundances near the Mackenzie River were ~ 2-fold higher than in the Mackenzie Shelf and Amundsen Gulf regions during the peak summer discharge, concomitant with a ~ 5.5-fold increase in chl a (up to 2.4 μg l^− 1) and a ~ 2-fold increase in bacterial abundance (up to 22 × 10⁵ ml^− 1). Using FC, two subgroups of viruses and heterotrophic bacteria were defined. A low SYBR-green fluorescence virus subgroup (V2) representing ~ 71% of the total viral abundance, was linked to the abundance of high nucleic acid fluorescence (HNA) bacteria (a proxy for bacterial activity), which represented 42 to 72% of the bacteria in surface layers. A high SYBR-green fluorescence viral subgroup (V1) was more related to high chl a concentrations that occurred in surface waters during spring and at stations near the Mackenzie River plume during the summer discharge. These results suggest that V1 infect phytoplankton, while most V2 are bacteriophages. On the Beaufort Sea shelf, viral abundance displayed seasonal and spatial variations in conjunction with chl a concentration, bacterial abundance and composition, temperature, salinity and depth. The highly dynamic nature of viral abundance and its correlation with increases in chl a concentration and bacterial abundance implies that viruses are important agents of microbial mortality in Arctic shelf waters.     

Linking mercury exposure to habitat and feeding behaviour in Beaufort Sea beluga whales

Mercury (Hg) levels in the Beaufort Sea beluga population have been increasing since the 1990's. Ultimately, it is the Hg content of prey that determines beluga Hg levels. However, the Beaufort Sea beluga diet is not understood, and little is known about the diet Hg sources in their summer habitat. During the summer, they segregate into social groups based on habitat use leading to the hypothesis that they may feed in different food webs explaining Hg dietary sources. Methyl mercury (MeHg) and total mercury (THg) levels were measured in the estuarine-shelf, Amundsen Gulf and epibenthic food webs in the western Canadian Arctic collected during the Canadian Arctic Shelf Exchange Study (CASES) to assess their dietary Hg contribution. To our knowledge, this is the first study to report MeHg levels in estuarine fish and epibenthic invertebrates from the Arctic Ocean. Although the Mackenzie River is a large source of Hg, the estuarine-shelf prey items had the lowest MeHg levels, ranging from 0.1 to 0.27 μg/g dry weight (dw) in arctic cisco (Coregonus autumnalis) and saffron cod (Eleginus gracilis) respectively. Highest MeHg levels occurred in fourhorn sculpin (Myoxocephalus quadricornis) (0.5 μg/g dw) from the epibenthic food web. Beluga hypothesized to feed in the epibenthic and Amundsen Gulf food webs had the highest Hg levels matching with high Hg levels in associated food webs, and estuarine-shelf belugas had the lowest Hg levels (2.6 μg/g dw), corresponding with the low food web Hg levels, supporting the variation in dietary Hg uptake. The trophic level transfer of Hg was similar among the food webs, highlighting the importance of Hg sources at the bottom of the food web as well as food web length. We propose that future biomagnification studies incorporate predator behaviour with food web structure to assist in the evaluation of dietary Hg sources.     

Picophytoplankton and nanophytoplankton abundance and distribution in the southeastern Beaufort Sea (Mackenzie Shelf and Amundsen Gulf) during Fall 2002

The distribution of picophytoplankton (0.2–2 µm) and nanophytoplankton (2–20 µm) in the Beaufort Sea–Mackenzie Shelf and Amundsen Gulf regions during autumn, 2002 is examined relative to their ambient water mass properties (salinity, temperature and nutrients: nitrate + nitrite, phosphate, and silicate) and to the ratio of variable to maximum fluorescence, Fv/Fm. Total phytoplankton and cell abundances (< 20 µm) were mainly correlated with salinity. Significant differences in picophytoplankton cell numbers were found among waters near the mouth of the Mackenzie River, ice melt waters and the underlying halocline water masses of Pacific origin. Picophytoplankton was the most abundant phytoplankton fraction during the autumnal season, probably reflecting low nitrate concentrations (surface waters average ~ 0.65 µM). The ratio Fv/Fm averaged 0.44, indicating that cells were still physiologically active, even though their concentrations were low (max Chl a = 0.9 mg m^− 3). No significant differences in Fv/Fm were evident in the different water masses, indicating that rate limiting conditions for photosynthesis and growth were uniform across the whole system, which was in a pre-winter stage, and was probably already experiencing light limitation as a result of shortening day lengths.     

Bacterial production and microbial food web structure in a large arctic river and the coastal Arctic Ocean

Globally significant quantities of organic carbon are stored in northern permafrost soils, but little is known about how this carbon is processed by microbial communities once it enters rivers and is transported to the coastal Arctic Ocean. As part of the Arctic River-Delta Experiment (ARDEX), we measured environmental and microbiological variables along a 300 km transect in the Mackenzie River and coastal Beaufort Sea, in July–August 2004. Surface bacterial concentrations averaged 6.7 × 10⁵ cells mL^− 1 with no significant differences between sampling zones. Picocyanobacteria were abundant in the river, and mostly observed as cell colonies. Their concentrations in the surface waters decreased across the salinity gradient, dropping from 51,000 (river) to 30 (sea) cells mL^− 1. There were accompanying shifts in protist community structure, from diatoms, cryptophytes, heterotrophic protists and chrysophytes in the river, to dinoflagellates, prymnesiophytes, chrysophytes, prasinophytes, diatoms and heterotrophic protists in the Beaufort Sea.Size-fractionated bacterial production, as measured by ³H–leucine uptake, varied from 76 to 416 ng C L^− 1 h^− 1. The contribution of particle-attached bacteria (> 3 µm fraction) to total bacterial production decreased from > 90% at the Mackenzie River stations to < 20% at an offshore marine site, and the relative importance of this particle-based fraction was inversely correlated with salinity and positively correlated with particulate organic carbon concentrations. Glucose enrichment experiments indicated that bacterial metabolism was carbon limited in the Mackenzie River but not in the coastal ocean. Prior exposure of water samples to full sunlight increased the biolability of dissolved organic carbon (DOC) in the Mackenzie River but decreased it in the Beaufort Sea.Estimated depth-integrated bacterial respiration rates in the Mackenzie River were higher than depth-integrated primary production rates, while at the marine stations bacterial respiration rates were near or below the integrated primary production rates. Consistent with these results, P_CO2 measurements showed surface water supersaturation in the river (mean of 146% of air equilibrium values) and subsaturation or near-saturation in the coastal sea. These results show a well-developed microbial food web in the Mackenzie River system that will likely convert tundra carbon to atmospheric CO₂ at increasing rates as the arctic climate continues to warm.     

Protist entrapment in newly formed sea ice in the Coastal Arctic Ocean

Protist abundance and taxonomic composition were determined in four development stages of newly formed sea ice (new ice, nilas, young ice and thin first-year ice) and in the underlying surface waters of the Canadian Beaufort Sea from 30 September to 19 November 2003. Pico- and nanoalgae were counted by flow cytometry whereas photosynthetic and heterotrophic protists ≥ 4 µm were identified and counted by inverted microscopy. Protists were always present in sea ice and surface water samples throughout the study period. The most abundant protists in sea ice and surface waters were cells < 4 µm. They were less abundant in sea ice (418–3051 × 10³ cells L^− 1) than in surface waters (1393–5373 × 10³ cells L^− 1). In contrast, larger protists (≥ 4 µm) were more abundant in sea ice (59–821 × 10³ cells L^− 1) than in surface waters (22–256 × 10³ cells L^− 1). These results suggest a selective incorporation of larger cells into sea ice. The ≥ 4 µm protist assemblage was composed of a total number of 73 taxa, including 12 centric diatom species, 7 pennate diatoms, 11 dinoflagellates and 16 flagellates. The taxonomic composition in the early stage of ice formation (i.e., new ice) was very similar to that observed in surface waters and was composed of a mixed population of nanoflagellates (Prasinophyceae and Prymnesiophyceae), diatoms (mainly Chaetoceros species) and dinoflagellates. In older stages of sea ice (i.e., young ice and thin first-year ice), the taxonomic composition became markedly different from that of the surface waters. These older ice samples contained relatively fewer Prasinophyceae and more unidentified nanoflagellates than the younger ice. Diatom resting spores and dinoflagellate cysts were generally more abundant in sea ice than in surface waters. However, further studies are needed to determine the importance of this winter survival strategy in Arctic sea ice. This study clearly shows the selective incorporation of large cells (≥ 4 µm) in newly formed sea ice and the change in the taxonomic composition of protists between sea ice and surface waters as the fall season progresses.     

On the distribution of dissolved lead in the Loire estuary and the North Biscay continental shelf, France

The dissolved lead was studied in the whole salinity gradient of the system composed of the Loire estuary and the North Biscay continental shelf. About 130 samples were collected in winter 2001 and spring 2002 during Nutrigas and Gasprod campaigns (Programme PNEC-Golfe de Gascogne, RV Thalassa) and metal measurements were conducted on board by Potentiometric Stripping Analysis. In the Loire estuary, levels of dissolved lead ranged from 0.15 to 0.24 nM and from 0.04 to 0.26 nM in winter and spring, respectively. Compared to the concentrations reported in 1987 and 1990 (0.4–1.7 nM; Boutier, B., Chiffoleau, J.F., Auger, D., Truquet, I., 1993. Influence of the Loire river on dissolved lead and cadmium concentrations in coastal waters of Brittany. Estuar. Coast. Shelf S., 36:133–143, Estuarine, Coastal and Shelf Science 36, 133–143) our study indicated much lower values. The fall in concentration in the estuary could be attributed to the stopping of activity of Octel, a big manufacturer of tetra alkyl lead. Discharge in dissolved metal to the continental shelf by the Loire river was assessed as 7.5 and 1.9 kg day^− 1 for winter and spring, respectively. On the continental shelf, levels of dissolved lead varied within 0.06 and 0.27 nM in winter (0.15 ± 0.06 nM, sd = 1.96, n = 49), whereas concentrations measured in spring were in the range 0.06–0.17 nM (0.09 ± 0.03 nM, sd = 1.96, n = 60). This difference in metal concentration was related to the amounts of rainfall that have fallen over the continental shelf: estimations of inputs by this way (74 and 32 kg day^− 1 in winter and spring, respectively) appeared to be significantly higher than inputs from the Loire river (7.5 and 1.9 kg day^− 1 in winter and spring, respectively). The distributions of dissolved metal in the surface waters highlighted the role of suspended particular matter (SPM) for a rapid “trapping” of lead near the mouth of the estuary. The vertical distributions showed, in the stratified area, a biological transfer of lead between winter and spring from surface waters to the halocline.     

Distribution of dinoflagellate cysts in surface sediments of the Mackenzie Shelf and Amundsen Gulf, Beaufort Sea (Canada)

In order to document long-term climate cycles and predict future climate trends for the Arctic, we need to look at the geological records to establish the link between historical and pre-historical sea-surface parameters. Dinoflagellate cysts (dinocysts) are used as proxy indicators of sea-surface parameters (temperature, salinity, sea-ice cover, primary productivity) jointly with transfer functions and a modern dinocyst reference database, to reconstruct the evolution of sea-surface conditions at decadal and millennial timescales. Here we present the surface distribution of recent dinocyst assemblages from 34 surface sediment samples collected on the Mackenzie Slope/Amundsen Gulf during the 2004 CASES (Canadian Arctic Shelf Exchange Study) cruise. Dinocyst concentrations in surface sediments are relatively high outside the Mackenzie plume area and increase gradually eastward toward Amundsen Gulf. The cysts of autotrophic dinoflagellates are dominant throughout the study area, while the maximum abundance of heterotrophic taxa is found within the Mackenzie plume. Hierarchical clustering analyses allowed defining two dinocyst assemblages. Assemblage I is located on the Mackenzie Slope and southern Amundsen Gulf, while Assemblage II is located within the Cape Bathurst Polynya area in northern Amundsen Gulf. Both assemblages are dominated by Operculodinium centrocarpum, but are distinguished on the basis of the relative abundance of Islandinium minutum, a taxon generally associated with sea ice. I. minutum is found in lower abundance in the Cape Bathurst Polynya.     

Mesozooplankton species distribution in the NW and N Iberian shelf during spring 2004: Relationship with frontal structures

We have analysed the mesozooplankton community structure in the southern Bay of Biscay shelf and its relationship with the hydrographic conditions during spring 2004. According to thermohaline characteristics, we observed two frontal zones of distinct origin along the shelf (around 7° and 3°W), that allowed us to differentiate three different hydrographic domains. The westernmost part of the shelf (WC), defined by the presence of relatively warm and salty water related to the presence of the Iberian Poleward Current (IPC), the easternmost region (EC), characterized by colder and fresher water and subject to the influence of freshwater inputs from the Adour river in the French coast, and a region in the Central Cantabrian Sea (CC), where thermohaline characteristics were intermediate between these two extremes. The mixing layer depth (MLD) regime in these areas was also different: the WC region was characterized by a mixed water column, whereas in the EC region the river discharges produces stratification of the upper meters of the water column (< 10 m); in the CC region, we found a distinct vertical mixing regime that separated coastal (stratification) from shelf (mixed water column) stations, giving rise to a notorious across-shelf front. We found a good match between the aforesaid hydrographic regions and the distribution of mesozooplankton species composition and community assemblages: the Mantel correlation between physical variables and mesozooplankton distribution was highly significant (n = 63, r = 0.70, α < 0.001). In the WC region, the community was dominated by Paracalanus parvus, Oithona helgolandica, Acartia clausi and Clausocalanus pergens, while in the EC region the most dominant species were Noctiluca scintillans, Oncaea media and Temora longicornis. The CC region showed similar composition of copepods than the WC region, but larvaceans (Oikopleura spp. and Fritillaria spp.) were more abundant in the CC region than in the WC region. Within each zone, the relative abundances of the dominant species differed between coastal and shelf locations.     

Ecophysiological growth characteristics and modeling of the onset of the spring bloom in the Baltic Sea

The onset of spring bloom in temperate areas is a transition period where the low productive, winter phytoplankton community is transformed into a high productive spring community. Downwelling irradiance, mixing depth and the ability of the phytoplankton community to utilize the light, are key parameters determining the timing of the onset of the spring bloom. Knowing these parameters would thus provide tools for modeling the spring bloom and enhance our knowledge of ecophysiological processes during this period.Our main objective with this study was to provide data for the growth characteristics of some key species forming the spring bloom in the Gulf of Finland, and to apply those results in a simple dynamic model for the onset of the spring bloom, in order to test if the timing of the spring bloom predicted by the models corresponds to field observations. We investigated the photosynthetic characteristics of three diatoms and two dinoflagellates (Chaetoceros wighamii, Melosira arctica, Thalassiosira baltica, Scrippsiella hangoei and Woloszynskia halophila), at low temperatures (4–5 °C). All of these species are common during spring bloom in the Baltic Sea.Cultures of these species were acclimated to different irradiance regimes prior to measurements of photosynthesis, respiration, pigment concentration and light absorption. We did not find a positive relationship between respiration and growth rate, and we hypothesize that this relationship, which is well established at higher temperatures, is negligible or absent at low temperatures (< 10 °C). Photosynthetic maximum (P_m), and maximum light utilization coefficient (α) was lowest and respiration (R) highest in the dinoflagellates.We made a model of the onset of the spring bloom in the western part of Gulf of Finland, using the obtained data together with monitoring data of mixing depth and water transparency from this area. Model results were compared to field observations of chlorophyll-a (Chl-a) concentration. There was a good agreement between the model predictions and the observed onset of the spring bloom for the diatoms. S. hangoei, however, was not able to reach positive production in the model, and W. halophila had the similar growth characteristics as S. hangoei. Consequently, these species must have other competition strategies enabling them to exist and grow during spring bloom.     

Seasonal productivity dynamics in the pelagic central Benguela System inferred from the flux of carbonate and silicate organisms

Flux of bulk components, carbonate- and silicate-bearing skeleton organisms, and the δ¹⁵N-isotopic signal were investigated on a 1-year time-series sediment trap deployed at the pelagic NU mooring site (Namibia Upwelling, ca. 29°S, 13°E) in the central Benguela System. The flux of bulk components mostly shows bimodal seasonality with major peaks in austral summer and winter, and moderate to low export in austral fall and spring. The calcium carbonate fraction dominates the export of particulates throughout the year, followed by lithogenic and biogenic opal. Planktonic foraminifera and coccolithophorids are major components of the carbonate fraction, while diatoms clearly dominate the biogenic opal fraction. Bulk δ¹⁵N isotopic composition of particulate matter is positively correlated with the total mass flux during summer and fall, while negatively correlated during winter and spring. Seasonal changes in the intensity of the main oceanographic processes affecting the NU site are inferred from variations in bulk component flux, and in the flux and diversity patterns of individual species or group of species. Influence from the Namaqua (Hondeklip) upwelling cell through offshore migration of chlorophyll filaments is stronger in summer, while the winter flux maximum seems to reflect mainly in situ production, with less influence from the coastal and shelf upwelling areas. On a yearly basis, dominant microorganisms correspond well with the flora and fauna of tropical/subtropical waters, with minor contribution of near-shore organisms. The simultaneous occurrence of species with different ecological affinities mirrors the fact that the mooring site was located in a transitional region with large hydrographic variability over short-time intervals.     

CO2 partial pressure in Northeast Atlantic and adjacent shelf waters: Processes and seasonal variability

CO₂ partial pressure in surface water was measured in the Northeast Atlantic and in the Hebride Shelf/North Sea area during a cruise with R.V. Poseidon in June 1991. A mean p_CO2 of 303 μatm was found in the Atlantic between 50°N and 60°N. For an atmospheric CO₂ content of 357.5 ppm(v) this corresponds to a partial pressure difference of −55 μatm. This supports the view that the subarctic Atlantic is a significant sink within the CO₂ cycle between the ocean and the atmosphere. A comparison of our measurements with other data reveals that the p_CO2 distribution changes significantly during May/June. This explained by seasonal warming, CO₂ exchange with the atmosphere and biomass production. The contribution by each of these processes to the seasonal variations is calculated. It was found that during a plankton bloom the production of biomass is the dominating factor and may lower seawater p_CO2 by almost 100 μatm. The shelf areas are charactrized by strong p_CO2 gradients which are explained by water exchange with the Atlantic, temperature effects and biomass production.     

Plankton distribution and the impact of copepod grazing on primary production in Fram Strait, Greenland Sea

Two hydrobiological transects across the East Greenland Shelf and the open waters of Fram Strait in summer were chosen to illustrate the distribution and production of phyto- and zooplankton in relation to water masses and ice cover. The parameters used were temperature and salinity, inorganic nutrients, chlorophyll a, primary production, phytoplankton species composition, abundance of the dominant herbivorous copepods Calanus finmarchicus, C. glacialis, C. hyperboreus, Metridia longa and egg production of C. finmarchicus and C. glacialis. Grazing impact of copepodites and adults of these four species was modelled for each station by using egg production rates as an index of growth. Seasonal development of plankton communities was closely associated with the extent of the ice cover, hydrographic conditions and the water masses typical of the different hydrographic domains. Four regions were identified from their biological activities and physical environment: The Northeast Water polynya on the East Greenland Shelf, with a springbloom of diatoms and active reproduction of herbivorous copepods. The pack ice region, dominated by small flagellates and negligible grazing activities. The marginal ice zone, with high variability and strong gradients of autotroph production related to eddies and ice tongues, an active microbial loop and low egg production. The open water, with high station-to-station variability of most of the parameters, probably related to hydrographic mesoscale activities. Here, Phaeocystis pouchetii was a prominent species in the phytoplankton communities. Its presence may at least partly be responsible for the generally low egg production in the open waters. Grazing impact on primary production was always small, due to low zooplankton biomass in the polynya and due to low ingestion in the remaining regions.     

Nutrient and organic matter patterns across the Mackenzie River, estuary and shelf during the seasonal recession of sea-ice

Suspended material, nutrients and organic matter in Mackenzie River water were tracked along a 300 km transect from Inuvik (Northwest Territories, Canada), across the estuarine salinity gradient in Kugmallit Bay, to offshore marine stations on the adjacent Mackenzie Shelf. All particulates measured (SPM, POC, PN, PP) declined by 87–95% across the salinity gradient and levels were generally below conservative mixing. Organic carbon content of suspended material decreased from 3.1% in the river to 1.7% in shelf surface waters while particulate C:N concurrently decreased from 17.1 to 8.6. Nitrate and silicate concentrations declined by more than 90% across the salinity gradient, with nitrate concentrations often below the conservative mixing line. Phosphate concentrations increased from 0.03 μmol/L in the river to 0.27 μmol/L over shelf waters, thereby changing the inorganic nutrient regime downstream from P to N limitation. Dissolved organic carbon decreased conservatively offshore while dissolved organic N and P persisted at high levels in the Mackenzie plume relative to river water, increasing 2.7 and 25.3 times respectively. A deep chlorophyll-a maximum was observed at two offshore stations and showed increases in most nutrients, particulates and organic matter relative to the rest of the water column. During river passage through the Mackenzie estuary, particulate matter, dissolved organic carbon and inorganic nutrients showed sedimentation, dilution and biological uptake patterns common to other arctic and non-arctic estuaries. Alternatively, inorganic content of particles increased offshore and dissolved organic N and P increased substantially over the shelf, reaching concentrations among the highest reported for the Arctic Ocean. These observations are consistent with the presence of a remnant ice-constrained (‘stamukhi’) lake from the freshet period and a slow flushing river plume constrained by sea-ice in close proximity to shore. Nutrient limitation in surface shelf waters during the ARDEX cruise contributed to the striking deep chlorophyll-a maximum at 21 m where phytoplankton communities congregated at the margin of nutrient-rich deep ocean waters.     

Observations on colony formation by the cosmopolitan phytoplankton genus Phaeocystis

Few marine phytoplankton have heteromorphic life cycles and also often dominate the ecosystems in which they occur. The class Prymnesiophyceae contains a notable exception: the genus Phaeocystis includes three species that form gelatinous colonies but also occur within their ranges as solitary cells. Phaeocystis antarctica and P. pouchetii are exclusively high latitude taxa, and are notable for regionally tremendous blooms of the colony stage. P. globosa occurs circumglobally, yet its colony blooms primarily are confined to colder waters within its range. Three additional species are warm water forms that have been reported only as solitary cells or loose aggregations that bear little resemblance to the organized colonies of the other taxa. Interpretation of existing data indicates that resource availability (light, temperature and nutrients) by itself is not sufficient to explain this distinction between cold-water colony-forming taxa and warm water solitary cell taxa, nor why colony development in P. globosa is essentially a spatially restricted phenomenon within a much broader geographic range. Colony development by P. globosa in situ has been observed at temperatures ≥20 °C, but only rarely and generally under conditions of seasonally or anthropogenically elevated nutrient supply. Data presented here demonstrate colony development at 20–22 °C in natural plankton communities from oligotrophic waters that were pre-screened through 63 μm mesh (i.e. lacking mesozooplankton and large microzooplankton), but not in unscreened communities containing microzooplankton and >63 μm zooplankton. Reduction of colony proliferation at higher temperatures by mesozooplankton grazing remains as an intriguing possibility that is consistent with available evidence to help explain differences in latitudinal extent of in situ colony development. These data are interpreted within a theoretical framework regarding the potential advantages and disadvantages of the two life cycle stages.     

Microbial food web responses to light and nutrients beneath the coastal Arctic Ocean sea ice during the winter–spring transition

We measured the abundance and biomass of phototrophic and heterotrophic microbes in the upper mixed layer of the water column in ice-covered Franklin Bay, Beaufort Sea, Canada, from December 2003 to May 2004, and evaluated the influence of light and nutrients on these communities by way of a shipboard enrichment experiment. Bacterial cell concentrations showed no consistent trends throughout the sampling period, averaging (± SD) 2.4 (0.9) × 10⁸ cells L^− 1; integrated bacterial biomass for the upper mixed layer ranged from 1.33 mg C m^− 3 to 3.60 mg C m^− 3. Small cells numerically dominated the heterotrophic protist community in both winter and spring, but in terms of biomass, protists with a diameter > 10 µm generally dominated the standing stocks. Heterotrophic protist biomass integrated over the upper mixed layer ranged from 1.23 mg C m^− 3 to 6.56 mg C m^− 3. Phytoplankton biomass was low and variable, but persisted during the winter period. The standing stock of pigment-containing protists ranged from a minimum value of 0.38 mg C m^− 3 in winter to a maximal value of 6.09 mg C m^− 3 in spring and the most abundant taxa were Micromonas-like cells. These picoprasinophytes began to increase under the ice in February and their population size was positively correlated with surface irradiance. Despite the continuing presence of sea ice, phytoplankton biomass rose by more than an order of magnitude in the upper mixed layer by May. The shipboard experiment in April showed that this phototrophic increase in the community was not responsive to pulsed nutrient enrichment, with all treatments showing a strong growth response to improved irradiance conditions. Molecular (DGGE) and microscopic analyses indicated that most components of the eukaryotic community responded positively to the light treatment. These results show the persistence of a phototrophic inoculum throughout winter darkness, and the strong seasonal response by arctic microbial food webs to sub-ice irradiance in early spring.     

Hydrographic structure and zooplankton abundance and diversity off Paita, northern Peru (1994 to 2004) — ENSO effects, trends and changes

The objective of the present study was to verify possible spatial, seasonal, and inter-annual changes in the zooplankton off Paita (northern Peru), an upwelling area located closely to the limits of cold Humboldt Current and warm Equatorial Surface Waters. Zooplankton was sampled at subsurface on 53 occasions from August 1994 to December 2004 at four stations located 2 to 30 km offshore with a WP-2 net (300 µm). Extremely high surface water temperatures combined with low salinities were observed during the 1997/98 El Niño up to 29.0 °C) and in April 2002 (up to 25.0 °C). Temperatures more than 2 °C above monthly average were also observed in October 1994, in April 2000, and in November 2004. Significant trends were observed for oxygen concentration (increase) and several horizontal and vertical gradients. Among the copepods (72% of all individuals), the most abundant species were Paracalanus parvus (28%), Acartia tonsa (26%), and Calanus sp. (10%). The strong 1997–98 El Niño (EN) event led to drastic changes in species composition that were reversed during the 1998–99 La Niña (LN) event. Community parameters such as total abundance, diversity, equitability and species richness displayed marked variations associated with the 1997–98 EN and long-term trends. Long-term trends were significant for several vertical and horizontal temperature and oxygen gradients, indicating an increase in upwelling intensity at the shelf during the study period. 10-year-trends were also significant for total zooplankton abundance (increase) and community evenness (J, decline). Our data confirmed the importance of the weak EN in 2002/03 for the study region. Within the trend of increasing zooplankton abundance, a sharp step or shift was observed from 1999 to 2000. When using sequential t-tests to detect shifts in (x + 1) transformed abundance data, a significant rupture was found between the last sampling in 1999 and the first sampling in 2000. Also, a substantial decrease in diel variability occurred after 1999, probably due to changes in vertical migration patterns. The considerable increase in zooplankton abundance over the study period, the ENSO effect, and the 1999–2000 transition are discussed with regard to synchronicity with other zooplankton time series. The present study contributes with the first evidence from an important area located in the Humboldt Current for synchronous trends and changes that were previously observed elsewhere in the Pacific. Our results demonstrated the importance of long-term zooplankton monitoring studies in upwelling areas, and confirms the idea of dramatic changes in pelagic ecosystem structure occurring in the East Pacific.