Recruitment variability in North Atlantic cod and match-mismatch dynamics

Background

Fisheries exploitation, habitat destruction, and climate are important drivers of variability in recruitment success. Understanding variability in recruitment can reveal mechanisms behind widespread decline in the abundance of key species in marine and terrestrial ecosystems. For fish populations, the match-mismatch theory hypothesizes that successful recruitment is a function of the timing and duration of larval fish abundance and prey availability. However, the underlying mechanisms of match-mismatch dynamics and the factors driving spatial differences between high and low recruitment remain poorly understood.

Methodology/Principal Findings

We used empirical observations of larval fish abundance, a mechanistic individual-based model, and a reanalysis of ocean temperature data from 1960 to 2002 to estimate the survival of larval cod (Gadus morhua). From the model, we quantified how survival rates changed during the warmest and coldest years at four important cod spawning sites in the North Atlantic. The modeled difference in survival probability was not large for any given month between cold or warm years. However, the cumulative effect of higher growth rates and survival through the entire spawning season in warm years was substantial with 308%, 385%, 154%, and 175% increases in survival for Georges Bank, Iceland, North Sea, and Lofoten cod stocks, respectively. We also found that the importance of match-mismatch dynamics generally increased with latitude.

Conclusions/Significance

Our analyses indicate that a key factor for enhancing survival is the duration of the overlap between larval and prey abundance and not the actual timing of the peak abundance. During warm years, the duration of the overlap between larval fish and their prey is prolonged due to an early onset of the spring bloom. This prolonged season enhances cumulative growth and survival, leading to a greater number of large individuals with enhanced potential for survival to recruitment.     

Reproduction,grazing, and development of the large subarctic calanoid <Emphasis Type="Italic">Eucalanus bungii</Emphasis>: is the spring diatom bloom the key to controlling their recruitment?

The response of large calanoid, Eucalanus bungii, to environmental fluctuation, particularly in relation to the spring diatom bloom in the Oyashio region, western subarctic Pacific Ocean, was examined by investigating egg production, grazing, development and starvation tolerance. Mean in situ egg production rate increased with ambient chlorophyll-a concentration, ranging from 0 to 47 eggs female⁻¹ d⁻¹, while no diurnal synchronous spawning behavior was observed. Under the spring bloom condition, E. bungii showed prey preference for less mobile and larger-sized prey (≥30 μm ESD) and bloom-forming diatom Thalassiosira spp. accounted for >80% of ingested carbon. In the laboratory, E. bungii was successfully reared from newly hatched nauplii to adult with the diatom, Thalassiosira nordenskioldi, as a food resource. Nauplii newly hatched from eggs reached the adult stage in ca. 150 days (5°C) with a sigmoidal developmental pattern and no sexual difference in development pattern. Starvation experiments indicated that the starved copepodids (C1–C4) became more vulnerable to high temperature with the progression of developmental stage, suggesting that the post-bloom condition with low food availability and increased temperature is harsh for their copepodids. The results of this study in conjunction with previous findings suggest that E. bungii is well adapted to utilize large-sized phytoplankton, such as a bloom-forming diatoms and, therefore, their recruitment processes, including egg production, development and mortality would be strongly affected by the duration and intensity of the spring bloom.     

The influence of turbidity on growth and survival of fish larvae: a numerical analysis

Growth and survival through the early larval phase probably limit the production potential of many commercially important fish stocks. Attempts to increase the production of these stocks by restocking of juveniles have generally failed. Here, we analyse how enhanced concentrations of phytoplankton and zooplankton affect the survival of fish larvae during their early life stages. The analysis is developed for larvae feeding on copepod eggs and nauplii, with fish and invertebrates as major predators. A model of feeding and growth of fish larvae is applied to assess the benefit of enhanced phytoplankton and zooplankton abundance. The analysis shows that the shading effect of higher phytoplankton concentration may reduce predation rates on fish larvae substantially. This `top-down' effect may be more important for the cohort survivorship than the `bottom-up' mechanism in situations when larval food is sufficiently abundant. However, while increased algal biomass will improve recruitment at high zooplankton concentration, it may also reduce recruitment at low zooplankton concentrations and shallow mixing depths. Both the larvae and their vertebrate predators are dependent on light to detect their prey, and the longer reactive distance of the predators make them more susceptible than the larvae to reduced light levels and increased turbidity. We discuss the implications of reduced predation and increased zooplankton abundance on recruitment and production of fish larvae, and point at environmental conditions where changing algal biomass is likely to affect recruitment success.     

The effect of sedimentation and microbial nitrogen regeneration in a plankton community: a simulation investigation

A computer simulation model was developed to investigate nitrogenfluxes associated with microbial interactions in plankton communities.A short time scale was used, appropriate to the build-up anddecline of phytoplankton blooms in temperate shelf waters aftera mixing or upwelling event. The model depicts a continuum ofevents, many of which have been observed in coastal, upwellingand oceanic systems, including two phytoplankton peaks correspondingto ‘new production’ and ‘regenerated production’.It predicts that nitrogen loss through sedimentation of phytoplanktonand faeces may result in a smaller bloom with a delayed onsetand prolonged duration. Microbial regeneration of nitrogen wasfound to be important in sustaining the middle stages of a phytoplanktonbloom, whereas micro- and meso-zooplankton regeneration occurredtowards the end of the bloom.     

Larval release in a sub-euphotic zone invertebrate triggered by sinking phytoplankton particles

This paper reports field results showing that the larval releaseof the snow crab, Chionoecetes opilio, in a subarctic fjordis phased with the sedimentation peak of phytoplankton particlesoriginating from the surface spring bloom, but with a lag of2–3 weeks. Further laboratory experiments with egg-bearingfemales showed a significant induction of larval release whenthey were exposed to senescent cultures of different phytoplanktonspecies, as opposed to controls and females exposed to freshcultures. These results are interpreted and discussed in thecontext of possible strategies evolved by the snow crab to enhanceits reproductive success. The field and laboratory evidencepresented here represents the first instance indicating thatphyto-detritus can act as a chemical cue for initiating andsynchronizing reproductive activities in sub-euphotic invertebrates.Selection for control of larval release by downward flux ofalgal particles may be especially important for invertebratepopulations in high-latitude areas, where the development ofplankton communities shows a high level of seasonality.     

Climate change impacts on mismatches between phytoplankton blooms and fish spawning phenology

Substantial interannual variability in marine fish recruitment (i.e., the number of young fish entering a fishery each year) has been hypothesized to be related to whether the timing of fish spawning matches that of seasonal plankton blooms. Environmental processes that control the phenology of blooms, such as stratification, may differ from those that influence fish spawning, such as temperature‐linked reproductive maturation. These different controlling mechanisms could cause the timing of these events to diverge under climate change with negative consequences for fisheries. We use an earth system model to examine the impact of a high‐emissions, climate‐warming scenario (RCP8.5) on the future spawning time of two classes of temperate, epipelagic fishes: “geographic spawners” whose spawning grounds are defined by fixed geographic features (e.g., rivers, estuaries, reefs) and “environmental spawners” whose spawning grounds move responding to variations in environmental properties, such as temperature. By the century's end, our results indicate that projections of increased stratification cause spring and summer phytoplankton blooms to start 16 days earlier on average (±0.05 days SE) at latitudes >40°N. The temperature‐linked phenology of geographic spawners changes at a rate twice as fast as phytoplankton, causing these fishes to spawn before the bloom starts across >85% of this region. “Extreme events,” defined here as seasonal mismatches >30 days that could lead to fish recruitment failure, increase 10‐fold for geographic spawners in many areas under the RCP8.5 scenario. Mismatches between environmental spawners and phytoplankton were smaller and less widespread, although sizable mismatches still emerged in some regions. This indicates that range shifts undertaken by environmental spawners may increase the resiliency of fishes to climate change impacts associated with phenological mismatches, potentially buffering against declines in larval fish survival, recruitment, and fisheries. Our model results are supported by empirical evidence from ecosystems with multidecadal observations of both fish and phytoplankton phenology.     

Plankton distribution across a slope current-induced front in the southern Bay of Biscay

Relatively warm (12.50–12.75°C) and high-salinity[<35.640 practical salinity units (PSU)] water flowing eastwardwas detected at the shelf-break during a cruise carried Outin the southern Bay of Biscay in Spring 1987. The slope currentinduced the formation of a convergent front separating well-mixedoceanic waters from haline-stratified coastal waters. Very highconcentrations of dissolved oxygen (295 µmol kg^–1)and chlorophyll a(>4.5 mg m^–3) were found at the outeredge of the frontal boundary. Small autotrophic flagellatesdominated the phytoplankton community. Primary production peakedat the boundary region. Estimated phytoplankton growth ratesindicated that active growth was taking place, with lower turnovertimes integrated over the water column at the frontal station(2.5–5 days) than at coastal (1.5–2.8 days) or oceanic(1.5–3.5 days) stations. The lowest doubling times (1–2days) were calculated for surface frontal populations. Accumulationof zooplankton was also observed associated with the convergentphysical structure, although this relationship was less markedthan for phytoplankton. Copepods, mainly Paracalanus parvus,Acartia clausi and Oithona helgolandica, formed the bulk ofthe mesozooplankton biomass. Compatibility between the sizeof phytoplankton cells and copepod size spectra indicate highfood availability for these animals, particularly in the vicinityof the front. The distribution of fish eggs and fish larvaewas also coupled with the slope current-induced front. Sardinelarvae were more abundant at the coastal side of the front,whereas larval stages of blue whiting reached the highest densitiesat off-shelf stations. Larvae of lamellibranch molluscs andbryozoa were restricted to nearshore waters, as the frontalboundary prevented larval dispersion to the open ocean. Theresults presented in this paper suggest that the Iberian slopecurrent and its associated shelf-break frontal structure werecrucial in controlling phytoplankton primary production, activityof grazers, distribution of larvae of fishes and benthic invertebrates,and ultimately in determining the structure of the pelagic foodweb in the southern Bay of Biscay during the seasonal periodof vertical mixing.     

Modeling microzooplankton and macrozooplankton dynamics within a coastal upwelling system

A simple nutrient–phytoplankton–zooplankton (NPZ)pelagic ecosystem model coupled to a two-dimensional primitiveequation circulation model with explicit mixed-layer physicsis configured in a coastal setting to study the biological responseto idealized wind-driven upwelling conditions. Conventionalecosystem model parameterization, which assumes macrozooplanktonas the target grazers, leads to upwelling-induced phytoplanktonblooms that exhaust available nutrient supply and whose zonalscale increases with wind duration. Offshore zooplankton maximaresult from upwelled water with greater total nitrogen concentrationsthan initial ambient surface water. Substantial vertical mixingin the surface boundary layer sets the vertical scale of theproductivity. Phytoplankton sinking contributes to a nearshoreaccumulation of total nitrogen, and enhances the magnitude andduration of the phytoplankton bloom. The system responds differentlywhen the zooplankton are parameterized to represent microzooplankton.The phytoplankton and zooplankton maxima have more limited zonalextent, are more independent of the duration of wind forcing,and near-surface nutrient levels remain high over most of thedomain. When winds are relaxed, the diminished offshore transportreveals the underlying biological oscillations in the microzooplankton-parameterizedecosystem, and reduced vertical mixing decouples surface fromsubsurface dynamics. In contrast, the macrozooplankton systemrelaxes to a steady state supporting limited phytoplankton andlarge zooplankton levels in the upwelling region.     

Climate-induced variability in Calanus marshallae populations

Calanus marshallae is the dominant mesozooplankton copepod speciesover the south-eastern Bering Sea middle shelf. Climate-inducedchanges in the magnitude and timing of production by C. marshallaemay affect the living marine resources of the Bering Sea shelfecosystem. We examined springtime abundance, gonadal maturityand stage distributions of C. marshallae copepodites duringfive consecutive years (1995–1999) that spanned the rangeof variability observed over the past 34 years in terms of watertemperature and ice cover. We compared our results with previouswork conducted during cool (1980) and warm (1981) years [ Smith,S. L. and Vidal, J. (1986) Cont. Shelf Res., 5, 215–239].The spring phytoplankton bloom began relatively early in associationwith ice (1995, 1997, 1999), but began late when ice was absentor retreated early (1996, 1998). Egg production began well beforethe bloom and continued over a long duration. Copepodites, however,were recruited during a relatively short period, coincidentwith the spring phytoplankton bloom. The relationship betweenbrood stock and spring-generation copepodite abundances wasweak. Copepodite concentrations during May were greatest inyears of most southerly ice extent. Copepodite populations werehighly variable among years, reflecting interannual variabilityin the atmosphere–ice–ocean system.     

Phytoplankton blooms: a 'loophole' in microzooplankton grazing impact?

Phytoplankton size and relations between phytoplankton and microzooplankton(ciliates and heterotrophic dinoflagellates) biomass are analysedin 12 globally distributed areas. In view of the results, ahypothesis is posed where blooming species are those able toescape control by microzooplankton through a combination ofpredation avoidance mechanisms (e.g. larger size, colonies,spines, and toxic compounds) at the beginning of the bloom.Factors that help to enhance subsequent bloom development includepositive feedback from the poor nutritional status of the phototrophicprey which adversely affects predation, inter-microzooplanktongrazing and top–down grazing by mesozooplankton on microzooplankton.Blooming conditions are interpreted as physical or chemicalperturbations disrupting the predator–prey controls thatnormally operate at the level of the microbial loop, opening‘loopholes’ into which some phytoplankton speciespopulations can explode.     

Meteorological factors affecting the bloom of Anabaenopsis raciborskii Wolosz. (Cyanophyta:Hormogonales) in the shallow Lake Balaton, Hungary

There was a heavy bloom of a nitrogen-fixing blue-green algaAnabaenopsis raciborskii Wolosz. in Lake Balaton in September-October1982. The authors studied the dynamics of the phyto- and bacterio-plankton of the lake by daily and weekly samplings. Some meteorologicalfactors were also considered. It could be concluded that thecauses of the bloom were accumulating nutrients being enrichedjust before the bloom-period suddenly washed into the lake byheavy rains, followed by windless warm days (water temperatures26–29°C). An expressed correlation between bacterio-and phytoplankton indicates their close trophic relationship.     

Springtime microprotozoan abundance and biomass in the southeastern Bering Sea and Shelik of Strait, Alaska

We surveyed springtime biomass and abundance of the >20 µmmicroprotozoa in surface waters of the SE Bering Sea and Shelikof Strait, Alaska. This study was part of the Fisheries OceanographyCoordinated Investigations (FOCI) program examining processeswhich affect the recruitment variability of walleye pollock(Theragra chalcogramma). Microprotozoa are a potential preyresource for larval pollock which has not been previously examined.In both areas, the >20 µm microprotozoa were predominantlydinoflagellates and ciliates. At the time of sampling (May 1990in Shelikof Strait and April 1992 in the SE Bering Sea), thespring diatom bloom was under way in Shelik of Strait, but notin the SE Bering Sea. Heterotrophic dinoflagellates dominatedthe microprotozoan assemblage in Shelik of Strait, but not inthe SE Bering Sea. In the SE Bering Sea. total microprotozoanabundances ranged from 300 to 6233 organisms 1^–1 and biomassfrom 0.58 to 9.73 µg C 1^–1. In Shelik of Strait,abundance and biomass were higher, ranging from 850 to 14 960organisms 1^–1 and from 1.29 to 70.73 µg C 1^–1,respectively. These biomass levels are comparable to those reportedfrom other coastal and oceanic regions. Microprotozoan biomasslevels were sufficient to support the estimated metabolic needsof first-feeding larval walleye pollock. It remains to be shownwhether larval pollock use this resource.     

Phytoplankton studies in the fjords of West Spitzbergen: physical environment and production in spring and summer

The phytoplankton in the fjords of West Spitzbergen was investigatedfrom 1978 to 1985. The spring bloom lasted longer than at theNorwegian coast: from mid March to early June. There is no delayin the onset of the spring bloom in Spitzerbergen fjords relativeto fjords of northern Norway. This is probably related to therapid daylength increase at high latitudes. The phytoplanktonspecies composition during spring was comparable to that alongthe coast of northern Norway. Annual primary production in theinvestigated area was calculated to be 150 g C m^–2 year^–1.     

Experimental test of the effect of maternal hormones on larval quality of a coral reef fish

Maternal hormones can play an important role in the development of fish larvae. Levels of the stress hormone, cortisol, in females are elevated by social interactions and transferred directly to the yolk of eggs, where they may influence developmental rates. In some vertebrates, prenatal exposure to high levels of testosterone determine early growth rates, social status and reproductive success. The present study examined whether post-fertilization exposure of eggs of the tropical damselfish, Pomacentrus amboinensis (Pomacentridae), to natural levels of cortisol or testosterone directly affects larval morphology at hatching. Maternal and egg levels of cortisol and testosterone varied widely among clutches of eggs from local populations around Lizard Island on the Great Barrier Reef. The morphology of larvae produced by these local fish populations also varied widely and differed significantly among sites (e.g., standard length: 2.6–3.4 mm; yolk sac area: 0.01–0.13 × 10⁻² mm²). Laboratory experiments showed that elevated cortisol levels in the egg reduced larval length at hatching, while slight elevations in testosterone increased yolk sac size. The influence of testosterone, and to a smaller extent cortisol, on larval morphology differed among egg clutches. These differences were partly explained by differences in initial egg hormone levels. Morphological changes induced by experimental hormonal regimes encompassed the entire range of variability in body attributes found in field populations. It is unclear whether cortisol influences growth alone or development rate or both. Testosterone appears to influence yolk utilization rates, and has no significant effect on growth, in contrast to its role in later developmental stages. Maternally derived cortisol and testosterone are important in regulating growth, development, and nutritive reserves of the embryo and larvae of this fish species. Factors that influence the maternal levels of cortisol and testosterone may have a major impact on larval mortality schedules and, therefore, on which breeding individuals contribute to the next generation. Received: 19 August 1998 / Accepted: 16 November 1998     

Fish-induced changes in zooplankton grazing on phytoplankton and bacterioplankton: a long-term study in shallow hypertrophic Lake Sobygaard

The impact of fish-mediated changes on the structure and grazingof zooplankton on phytoplankton and bacterioplankton was studiedin Lake Søbygaard during the period 1984–92 bymeans of in vitro grazing experiments (¹⁴C-labelled phytoplankton,³H-labelled bacterioplankton) and model predictions. Measuredzooplankton clearance rates ranged from 0–25 ml l^–1h^–1 on phytoplankton to 0–33 ml l^–1 h^–1on bacterioplankton.The highest rates were found during thesummer when Daphnia spp. were dominant. As the phytoplanktonbiomass was substantially greater than that of bacterioplanktonthroughout the study period, ingestion of phytoplankton was26-fold greater than that of bacterioplankton. Multiple regressionanalysis of the experimental data revealed that Daphnia spp.,Bosmina longirostris and Cyclops vicinus, which were the dominantzooplankton, all contributed significantly to the variationin ingestion of phytoplankton, while only Daphnia spp. contributedsignificantly to that of bacterioplankton. Using estimated meanvalues for clearance and ingestion rates for different zooplankters,we calculated zooplankton grazing on phytoplankton and bacterioplanktonon the basis of monitoring data of lake plankton obtained duringa 9 year study period. Summer mean grazing ranged from 2 to4% of phytoplankton production and 2% of bacterioplankton productionto maxima of 53 and 88%, respectively. The grazing percentagedecreased with increasing density of planktivorous fish caughtin August each year using gill nets and shore-line electrofishing.The changes along a gradient of planktivorous fish abundanceseemed highest for bacterioplankton. Accordingly, the percentagecontribution of bacterioplankton to the total ingestion of thetwo carbon sources decreased from a summer mean value of 8%in Daphnia-dominated communities at lower fish density to 0.7–1.1%at high fish density, when cyclopoid copepods or Bosmina androtifers dominated. Likewise, the percentage of phytoplanktonproduction channelled through the bacteria varied, it beinghighest (5–8%) at high fish densities. It is argued thatthe negative impact of zooplankton grazing on bacterioplanktonin shallow lakes is highest at intermediate phosphorus levels,under which conditions Daphnia dominate the zooplankton community.     

Recruitment patterns in Antarctic Peninsula shelf sediments: evidence of decoupling from seasonal phytodetritus pulses

Summer phytoplankton blooms on the West Antarctic Peninsula (WAP) shelf result in episodic deposition of labile food material for benthic detritivores. This summer deposition is thought to enhance benthic recruitment of macro- and megafauna. To explore seasonality in benthic recruitment, juvenile invertebrates (>100 μm) were collected in a seasonal time series at three stations on the WAP continental shelf. 4,098 juveniles were collected (average densities 2,000–7,000 m⁻²), with polychaetes dominant (2,581 individuals). The majority of polychaetes showed evidence of enhanced recruitment prior to the summer bloom, although patterns varied across stations. Additional taxa showed recruitment peaks in summer, but again, patterns varied among stations. Based on observed patterns, polychaete taxa are classified as “seasonal”, “marginally seasonal”, and “non-seasonal” recruiters, with the latter two patterns predominating. The year-round presence of small juveniles in most taxa suggests that recruitment occurs continuously, with periodic enhancement. Year-round recruitment is consistent with the presence of a persistent “food bank” of labile organic material in WAP shelf sediments, allowing recruitment to be largely decoupled from seasonal bloom dynamics.     

Seasonal and fine-scale spatial variations in egg production and triacylglycerol content of the copepod Acartia tonsa in a river-dominated estuary and its coastal plume

We measured egg production rates of the estuarine calanoid copepodAcartia tonsa in Mobile Bay, an estuary in the northern Gulfof Mexico. Two stations were sampled approximately monthly,one at the mouth of the bay and the other just beyond the mouthin the salinity front between bay and coastal waters. Over thewhole year, temperature was the most important environmentalvariable controlling egg production. Rates increased with temperatureup to 30°C and 140 eggs female^–1 day^–1. We foundno evidence of food limitation. There was no correlation betweenegg production and phytoplankton abundance, nor increased eggproduction in response to supplements of phytoplankton addedto natural food, suggesting that non-phytoplankton food wasimportant in the diet. At the highest egg production rates,the amount of the storage lipid triacylglycerol (TAG) in adultfemales was greatly diminished, to <50 ng female^–1.This suggests that lipids in the diet can be very tightly coupledto egg production. Both egg production and TAG content of femalesshowed significant variability on spatial scales of 3–15km, especially in relation to the salinity front separat ingwater outwelling from the bay from open coastal water. For organismsthat are using copepods and their eggs as food, this variabilitywould result in a heterogeneous food environment, both in termsof the abundance of food and its nutritional content.     

The role of solid waste materials as habitats for macroinvertebrates in a lowland dam reservoir

Eight hypereutrophic phytoplankton dominated ponds from the Brussels Capital Region (Belgium) were biomanipulated (emptied with fish removal) to restore their ecological quality and reduce the risk of cyanobacterial bloom formation. Continuous monitoring of the ponds before and after the biomanipulation allowed the effects of the management intervention on different compartments of pond ecosystems (phytoplankton, zooplankton, submerged vegetation and nutrients) to be assessed. Fish removal resulted in a drastic reduction in phytoplankton biomass and a shift to the clear-water state in seven out of eight biomanipulated ponds. The reduction in phytoplankton biomass was associated with a marked increase in density and size of large cladocerans in six ponds and a restoration of submerged macrophytes in five ponds. The phytoplankton biomass in the ponds with extensive stands of submerged macrophytes was less affected by planktivorous fish recolonisation of some of the ponds later in the summer. The two non-vegetated ponds as well as one pond with sparse submerged vegetation showed a marked increase in phytoplankton biomass associated with the appearance of fish. Phytoplankton biomass increase coincided with the decrease in large Cladocera density and size. One pond lacking submerged macrophytes could maintain very low phytoplankton biomass owing to large Cladocera grazing alone. The results of this study confirmed the importance of large zooplankton grazing and revegetation with submerged macrophytes for the maintenance of the clear-water state and restoration success in hypereutrophic ponds. They also showed that large Cladocera size is more important than their number for efficient phytoplankton control and when cladocerans are large enough, they can considerably restrain phytoplankton growth, including bloom-forming cyanobacteria, even when submerged vegetation is not restored. The positive result of fish removal in seven out of eight biomanipulated ponds clearly indicated that such management intervention can be used, at least, for the short-term restoration of ecological water quality and prevention of noxious cyanobacterial bloom formation. The negative result of biomanipulation in one pond seems to be related to the pollution by sewage water. Guest editors: B. Oertli, R. Cereghino, A. Hull & R. Miracle Pond Conservation: From Science to Practice. 3rd Conference of the European Pond Conservation Network, Valencia, Spain, 14–16 May 2008     

Nitrate uptake by size-fractionated phytoplankton on the Scotian Shelf (Northwest Atlantic): spatial and temporal variability

New (nitrate) phytoplankton production was estimated monthlyduring 1 year (March 1991–March 1992) at three stationson the Scotian Shelf, Northwest Atlantic. Samples were sizefractionated to assess the uptake of nitrate by small (<5µm) and large (>5 µm) phytoplankton. The biomassof small phytoplankton remained relatively constant over theyear, whereas that of the large size fraction was high in earlyspring and low during the remainder of the year. Monthly variationsin nitrate uptake were similar for the two size fractions, suggestingthat both small and large phytoplankton used nitrate when available.It follows that, outside the spring bloom, new production waslargely due to the small fraction. Our results do not supportthe notion that new production is associated with large phytoplanktonand regenerated production with small phytoplankton.     

Life strategy and diet of Calanus glacialis during the winter�Cspring transition in Amundsen Gulf, south-eastern Beaufort Sea

The copepod Calanus glacialis plays a key role in the lipid-based energy flux in Arctic shelf seas. By utilizing both ice algae and phytoplankton, this species is able to extend its growth season considerably in these seasonally ice-covered seas. This study investigated the impacts of the variability in timing and extent of the ice algal bloom on the reproduction and population success of C. glacialis. The vertical distribution, reproduction, amount of storage lipids, stable isotopes, fatty acid and fatty alcohol composition of C. glacialis were assessed during the Circumpolar Flaw Lead System Study. Data were collected in the Amundsen Gulf, south-eastern Beaufort Sea, from January to July 2008 with the core-sampling from March to April. The reduction in sea ice thickness and coverage observed in the Amundsen Gulf in 2007 and 2008 affected the life strategy and reproduction of C. glacialis. Developmental stages CIII and CIV dominated the overwintering population, which resulted in the presence of very few CV and females during spring 2008. Spawning began at the peak of the ice algal bloom that preceded the precocious May ice break-up. Although the main recruitment may have occurred later in the season, low abundance of females combined with a potential mismatch between egg production/development to the first feeding stage and phytoplankton bloom resulted in low recruitment of C. glacialis in the early summer of 2008.