Dynamics of coexisting Calanus finmarchicus, Calanus glacialis and Calanus hyperboreus populations in a high-Arctic fjord

We studied the population dynamics of Calanus finmarchicus, Calanus glacialis and Calanus hyperboreus in Billefjorden, Svalbard (78°40N). All three species reproduced in the fjord with different timing. The maximum abundance of Calanus spp. copepodite stages peaked on the 11th of July (29,000 ind m^–2). C. glacialis was the dominant species accounting for 60–80% of the total Calanus abundance. C. finmarchicus appear to thrive in the fjord despite the low temperatures (–1.86°C to 5°C) and accounted for 20–30% of the total population. C. hyperboreus contributed less to the total abundance (5–20%). A 1-year life cycle is suggested for C. finmarchicus and C. hyperboreus in the fjord, C. glacialis has a 1- to 2-year life cycle. Highest mortality rates were observed for copepodite stage CV in C. finmarchicus and C. glacialis (0.09 and 0.075 d^–1, respectively) and for females in C. hyperboreus (0.149 d^–1). Mortality of copepodite stages was substantially lower in C. glacialis than in the other species. This is particularly obvious in the early and numerous copepodite stages (CI + CII) during the period of recruitment to these stages. This suggests that differences in secondary production in Arctic pelagic ecosystems are controlled partly by population loss rates.     

Climate,copepods and seabirds in the boreal Northeast Atlantic – current state and future outlook

The boreal Northeast Atlantic is strongly affected by current climate change, and large shifts in abundance and distribution of many organisms have been observed, including the dominant copepod Calanus finmarchicus, which supports the grazing food web and thus many fish populations. At the same time, large‐scale declines have been observed in many piscivorous seabirds, which depend on abundant small pelagic fish. Here, we combine predictions from a niche model of C. finmarchicus with long‐term data on seabird breeding success to link trophic levels. The niche model shows that environmental suitability for C. finmarchicus has declined in southern areas with large breeding seabird populations (e.g. the North Sea), and predicts that this decline is likely to spread northwards during the 21st century to affect populations in Iceland and the Faroes. In a North Sea colony, breeding success of three common piscivorous seabird species [black‐legged kittiwake (Rissa tridactyla), common guillemot (Uria aalge) and Atlantic puffin (Fratercula arctica)] was strongly positively correlated with local environmental suitability for C. finmarchicus, whereas this was not the case at a more northerly colony in west Norway. Large seabird populations seem only to occur where C. finmarchicus is abundant, and northward distributional shifts of common boreal seabirds are therefore expected over the coming decades. Whether or not population size can be maintained depends on the dispersal ability and inclination of these colonial breeders, and on the carrying capacity of more northerly areas in a warmer climate.     

Vertical distribution and seasonal numerical abundance of the Calanidae in oceanic waters to the south-west of the British Isles

The vertical distribution and seasonal abundance of the copepodite and adult stages of Calanus finmarchicus, C. helgolandicus, C. tenuicornis, Neocalanus gracilis, Nannocalanus minor and Calanoides carinatus from a series of Longhurst Hardy Plankton Recorder hauls taken in the oceanic waters, off the continental shelf, to the south-west of the British Isles are described. The sampling area was selected because the geographical distributions of the major Calanidae copepods of the north-east Atlantic Ocean are shown to overlap in this region. It marks the southern boundary of the distribution of C. finmarchicus, the central area of C. helgolandicus and the approximate northern limit of distribution of C. tenuicornis, N. gracilis, N. minor and C. carinatus. These four southern species occasionally penetrate further north (60° N) in the open ocean but do not breed at these northern latitudes. In autumn and winter, when C. finmarchicus and C. helgolandicus were overwintering below 400 m primarily as Stage V copepodites, N. gracilis occurred in the upper 200 m of the water column in a breeding condition; all copepodite stages were present. This copepod reproduced throughout the year in this regions while C. tenuicornis was observed to breed primarily in spring and summer. The geographical and vertical distributions of the Calanidae are related to the observed seasonal temperatures of the North Atlantic and the breeding strategies of species are compared.     

Regional and seasonal variability of zooplankton collected using sediment traps in the southeastern Beaufort Sea, Canadian Arctic

Time-series sediment traps were deployed at six mooring sites in the southeastern Beaufort Sea from October 2003 to August 2004 during the cruise of the Canadian research vessel Amundsen within the framework of the Canadian Arctic Shelf Exchange Study (CASES). Trap-collected zooplankton (TCZ) at around 200 m water depth was dominated by copepods accounting for 74–93% of the total abundance throughout the year with increase in abundance at all sites during the fall. Seven distinct TCZ groups were identified through cluster analysis. Two marked seasonal shifts in TCZ composition from late fall to early winter and from spring to early summer were revealed at five sites at 200 m depth. The zooplankton was dominated by Oncaea spp., pteropods, and copepod nauplii in the late fall cluster and in the winter cluster, and by copepod nauplii in the summer cluster. A significant change in water temperature, salinity, and sea ice concentration was observed only with the spring–summer shift. The cluster analysis also revealed that TCZ composition at 200 m at a station located in the Cape Bathurst Polynya was markedly different from those at other sites through the study period by being characterized by the dominance of various copepodite stages of Metridia longa. This was probably due to a less prolonged period of sea ice cover, which provides favorable food conditions for the zooplankton community.     

Lipids and life strategies of Calanus finmarchicus, Calanus glacialis and Calanus hyperboreus in late autumn, Kongsfjorden, Svalbard

Stage IV and V copepodites were the dominant forms of Calanus finmarchicus, C. glacialis and C. hyperboreus in Kongsfjorden in late September 1997. Stage IV and V copepodites of C. glacialis and C. hyperboreus were rich in lipid, largely wax esters, and were well fitted to overwinter. Stage IV copepodites of C. finmarchicus were also rich in wax esters, but stage V copepodites of C. finmarchicus were less wax ester-rich. Large size increments between stage IV and V copepodites and between stage V copepodites and females were noted in C. finmarchicus. A very large increment between stage IV and V copepodites was noted for C. glacialis but the size difference between stage V copepodites and females was very small in this species. Particularly large increments were noted between stage IV and V copepodites of C. hyperboreus and also between stage V copepodites and females of this species. The very large, wax ester-rich C. hyperboreus is well adapted to survive the most extreme variations in the Arctic, in Arctic basin waters, whereas the smaller, wax ester-rich C. glacialis is adapted to survive less extreme Arctic variations, as in Arctic shelf waters. The smallest of the three, C. finmarchicus, is best adapted to survive the more predictable waters of the North Atlantic and the Barents Sea. Accepted: 3 January 2000     

Winter ecology of Arctic charr (<Emphasis Type="Italic">Salvelinus alpinus</Emphasis>) and brown trout (<Emphasis Type="Italic">Salmo trutta</Emphasis>) in a subarctic lake,Norway

We studied habitat choice, diet, food consumption and somatic growth of Arctic charr (Salvelinus alpinus) and brown trout (Salmo trutta) during the ice-covered winter period of a subarctic lake in northern Norway. Both Arctic charr and brown trout predominantly used the littoral zone during winter time. Despite very cold winter conditions (water temperature <1°C) and poor light conditions, both fish species fed continuously during the ice-covered period, although at a much lower rate than during the summer season. No somatic growth could be detected during the ice-covered winter period and the condition factor of both species significantly declined, suggesting that the winter feeding rates were similar to or below the maintenance requirements. Also, the species richness and diversity of ingested prey largely decreased from summer to winter for both fish species. The winter diet of Arctic charr <20 cm was dominated by benthic insect larvae, chironomids in particular, and Gammarus lacustris, but zooplankton was also important in December. G. lacustris was the dominant prey of charr >20 cm. The winter diet of brown trout <20 cm was dominated by insect larvae, whereas large-sized trout mainly was piscivorous, feeding on juvenile Arctic charr. Piscivorous feeding behaviour of trout was in contrast rarely seen during the summer months when their encounter with potential fish prey was rare as the small-sized charr mainly inhabited the profundal. The study demonstrated large differences in the ecology and interactions of Arctic charr and brown trout between the winter and summer seasons.     

Seasonal variations in biomass, abundance and composition of zooplankton in the subarctic waters north of Iceland

The seasonal variations in biomass, abundance and species composition of zooplankton in relation to hydrography and chlorophyll a were studied in the subarctic waters north of Iceland. The sampling was carried out at approximately monthly intervals from February 1993 to February 1994 at eight stations arranged along a transect extending from 66°16′N–18°50′W to 68°00′N–18°50′W. The mean temperature at 50 m depth showed a clear seasonal pattern, with lowest water temperatures in February (∼1.1°C) and the highest in July (∼5.4°C). The spring growth of the phytoplankton began in late March and culminated during mid-April (∼7.0 mg Chl a m⁻³). Both the biomass and the abundance of total zooplankton were low during the winter and peaked once during the summer in late May (∼4 g m⁻² and ∼38,000 individuals m⁻²). A total of 42 species and taxonomic groups were identified in the samples. Eight taxa contributed ∼90% of the total zooplankton number. Of these Calanus finmarchicus was by far the most abundant species (∼60% of the total zooplankton). Less important groups were ophiuroid larvae (∼9%), Pseudocalanus spp. (∼8%), Metridia longa (∼4%), C. hyperboreus (∼3%), Acartia longiremis (∼2%), chaetognaths (∼2%) and euphausiid larvae (∼2%). The dominant copepods showed two main patterns in seasonal abundance: C. finmarchicus, C. hyperboreus and C. glacialis had one annual peak in numbers in late May, while Pseudocalanus spp., M. longa and A. longiremis showed two maxima during the summer (July) and autumn (October/November). Ophiuroid larvae and chaetognaths (mainly Sagitta elegans) peaked during the middle of July, while the number of euphausiid eggs and larvae was greatest from May to July. The succession in population structure of C. finmarchicus indicated its main spawning to be in April and May, coincident with the phytoplankton spring bloom. A minor spawning was also observed sometime between August and October. However, the offspring from this second spawning contributed only insignificantly to the overwintering stock of C. finmarchicus. Received: 12 September 1997 / Accepted: 1 March 1998     

Lipid‐rich zooplankton subsidise the winter diet of benthivorous Arctic charr (Salvelinus alpinus) in a subarctic lake

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Interannual phenological variability in two North-East Atlantic populations of Calanus finmarchicus

Phenological variations of the marine copepod Calanus finmarchicus were studied in Svalbard and northern Iceland, where samples were collected in summer and spring, respectively, over two decades. Four phenological indices, developed for copepodite stage-structured data, were used: the proportion of CV to total abundance (CVT), the population development index (PDI), the average weighted stage (AWS), and the average age in days (AAD). The variation of these indices was compared within and between locations to evaluate their suitability for the analysis of phenological effects. For both populations, phenology was related to local temperature and spring bloom dynamics, influenced by Atlantic water inflow. Large-scale climate was related to phenological variation only in the Svalbard population. C. finmarchicus phenology advanced under warmer conditions in both locations. We conclude that vertical phenological indices, i.e. based on interannual changes in copepodite stage structure, are useful to investigate zooplankton phenology, especially when data series covering the whole life cycle are unavailable. We suggest that AWS and AAD can be applied irrespective of sampling time, while PDI and CVT should be applied for early and late sampling seasons, respectively. When multiple phenological indices are needed, AAD in combination with either CVT or PDI should be preferred.     

Consistency in the life history traits of four invasive pseudorasbora parva populations in Southern England

Life history plasticity can be a strong predictor of the establishment and invasion success of introduced fishes. Here, the life-history traits of four P. parva populations in adjacent ponds in Southern England were measured throughout 2013 to determine the timing and length of their reproductive season and the extent of trait plasticity. The relative abundance of the populations (as catch per unit effort) was similar, with low variability in their traits relating to reproductive effort and somatic growth. All the populations were male dominated. Both sexes matured at small body sizes, with fish as small as 30 mm being mature in both sexes, with the age at maturity for both sexes being age 1+ years. The peak spawning period, characterised by female fish investing heavily in reproduction when their gonado-somatic index (GSI) values were highest and declined thereafter, occurred in May and June, and after a mean of 212 ± 24 degree-days >12°C, and with water temperatures being 13.2–14.6°C. There were no further peaks in GSI, despite the species being considered to have prolonged spawning periods across spring and summer. These results also suggest that whilst the expression of life-history traits of invasive fishes often vary with density, they can be highly consistent between populations at similar abundances and locations.     

Timing and size of blooms of the ctenophore Mnemiopsis leidyi in relation to temperature in Narragansett Bay, RI

The ctenophore Mnemiopsis leidyi is at the northern extreme of its geographic range in Narragansett Bay, an estuary on the northeast coast of the United States. Blooms have typically been observed in late summer and fall according to records from 1950 to 1979. We document an expansion of the seasonal range of this important planktonic predator to include springtime blooms during the 1980s and 1990s. This shift to an earlier seasonal maximum is associated with increasing water temperature in Narragansett Bay. Temperatures in spring have risen, on average, 2 °C from 1950 to 1999 with warm years being associated with the positive phase of the North Atlantic Oscillation. During 1999, M. leidyi appeared earlier in spring and was more abundant than during any previous year for which records are available. Changes in the seasonal pattern and abundance of this predator are likely to have important effects on planktonic ecosystem dynamics of Narragansett Bay. These include reduction of zooplankton abundance in spring followed by increases in size and frequency of summer phytoplankton blooms. Earlier blooms of M. leidyi may also reduce survival of eggs and larvae of fish because, as in 1999, they coincide with the period of peak spawning.     

Ecological investigations on the zooplankton community of Balsfjorden,northern Norway: Generation cycles,and variations in body weight and body content of carbon and nitrogen related to overwintering and reproduction in the copepod Calanus finmarchicus (Gunnerus)

The generation cycles of Calanus finmarchicus (Gunnerus) are described together with the seasonal variations in length, wet wt, dry wt, carbon content, nitrogen content and $\text{C}\text{N}$ ratio in copepodite stage IV, V and stage VI males and females from Balsfjorden (69°21′N: 19°06′E), a subarctic fjord in northern Norway. C. finmarchicus overwinters in copepodite stage IV (≈ 20%) or V (≈80%) and produces one generation a year. Variations in body weight and body content of carbon and nitrogen in the different copepodite stages showed a pronounced seasonal pattern. For instance, the $\text{C}\text{N}$ ratio was lowest (4.9) in adult females during the spawning period. Copepodite stage IV and V had higher $\text{C}\text{N}$ values in summer and autumn (12 to 14) than in spring (8 to 10). Variations in length, weight and chemical composition revealed that the overwintering stock of C. finmarchicus went through two growth phases during this period. From September to January no significant changes in the measured variables were detected. During the second phase of the overwintering period, January to April, the different stages showed a profound decrease in weight and change in chemical composition. This seemed to be connected with the onset of sexual differentiation in stage V starting in January, subsequent moulting into adults and gonad maturation in these adults. These results are further discussed in relation to the different prevailing hypotheses concerning overwintering strategy in Calanus species.     

Breaking the ice: large-scale distribution of mesozooplankton after a decade of Arctic and transpolar cruises

Mesozooplankton collected during five summer expeditions to the Arctic Ocean between 1987 and 1991 was analysed for regional patterns in biomass and species distribution, distinguishing between an epipelagic (0–100 m) and a deeper (0–500 m) layer. A total of 58 stations was sampled mainly in the Nansen, Amundsen and Makarov Basins of the central Arctic Ocean and in areas of the Greenland Sea, West Spitsbergen Current and Barents Sea. Results from the different expeditions were combined to create a transect extending from the Fram Strait across the Eurasian Basin into the Makarov Basin. Mesozooplankton dry mass in the upper 500 m decreased from 8.4 g m⁻² in the West Spitsbergen Current to less than 2 g m⁻² in the high-Arctic deep-sea basins. In the central Arctic Ocean, biomass was concentrated in the upper 100 m and was dominated by the large copepods Calanus hyperboreus and C. glacialis. In contrast, the mesozooplankton in the West Spitsbergen Current was more evenly distributed throughout the upper 500 m, with C. finmarchicus as the prevailing species. The distribution of abundant mesopelagic species reflected the hydrographic regime: the calanoid copepod Gaetanus tenuispinus and the hyperiid amphipod Themisto abyssorum were most abundant in the Atlantic inflow, while Scaphocalanus magnus was a typical component of the high-Arctic fauna. The relatively high mesozooplankton biomass and the occurrence of boreal-Atlantic species in the central Arctic Ocean are indicators for the import of organic material from allochthonous sources, especially from the northern North Atlantic. Hence, in spite of its enclosure by land masses, the Arctic Ocean is characterized by an exchange of water masses and organisms with the North Atlantic, and advection processes strongly influence the distribution of plankton species in this high-latitude ecosystem. Received: 18 December 1997 / Accepted: 11 April 1998     

The effect of food limitation on gonad development and egg production of the planktonic copepod Calanus finmarchicus

Calanus finmarchicus, one of the dominant copepods species of the North Atlantic, often encounters low food concentration or quality during the reproductive period; however, our knowledge on the effect of these conditions on reproduction processes is scarce. The present study combines experiments with histological observations to describe the response of C. finmarchicus to limited food focussing on (1) oocyte maturation processes, (2) gonad morphology, (3) egg production rates as a function of spawning frequency and clutch size, and (4) the fuelling of egg production by measuring carbon and nitrogen content of the females. In the laboratory, C. finmarchicus females were exposed to 0, 10, 50, 150 or >300 μg C l⁻¹ for several days. To account for food quality and season, reproductive activity was compared in April and July 1999 between females feeding on diatoms or dinoflagellates. The effect of feeding history was studied in February with females fed and starving prior to the experiment. Feeding conditions had severe effects on oocyte maturation process. Hence, egg production varied significantly with food concentration and quality, season and feeding history due to variation in both clutch size and spawning frequency. Clutch size differed by a factor of 2-4 between food limited and well fed females, and is thus an important parameter for modelling egg production. Changes in clutch size were related to changes of the number of maturing oocytes in the females gonads indicating that the latter can be used to precise the prediction of egg production from preserved samples. The proportion of females carrying at least some mature oocytes was relatively high at low food availability. Apparently, these females used internal body reserves as the carbon and nitrogen content decreased significantly under these conditions. These results indicate that C. finmarchicus embarks on the strategy to enable reproduction in all or many females of a population at low rates when feeding conditions are unfavourable.     

Ecological investigations of the zooplankton community of Balsfjorden,northern Norway: Generation cycle,seasonal vertical distribution,and seasonal variations in body weight and carbon and nitrogen content of the copepod Metridia longa (Lubbock)

Metridia longa (Lubbock) has an annual life cycle in Balsfjorden (69°21'N:19°06'E) with spawning occurring from early to mid-May. Development through copepodite stages I-V takes place during summer. At the end of August copepodite stage V accounted for >80% of the population. Overwintering (October-March) was mainly performed by adults. At noon throughout the year copepodite stage V and adults were found within the bottom 30 m of the fjord. The majority of stage IV occurred within the bottom 80 m, while copepodite stages I-III were mainly concentrated within the surface 50 m. Body length, weight, carbon and nitrogen content, and $\text{C}\text{N}$ ratio were determined for copepodite stage V, adult males and females throughout the year. While body length was constant, weight and C and N content varied with season. Lowest $\text{C}\text{N}$ ratios (4–6) were found in copepodite stage V in July and in adults in late winter. Highest $\text{C}\text{N}$ ratios (16–20) were measured in adults from October through early winter. These results are discussed in relation to the life cycle of M. longa and the primary production cycle of the fjord. It is argued that M. longa overwinters in an active state compared with Calanus finmarchicus (Gunnerus) from the same locality.     

Spatial distribution and life-cycle timing of zooplankton in the marginal ice zone of the Barents Sea during the summer melt season in 1995

The marginal ice zone (MIZ) in the northern Barents Sea is ecologically important because it represents a highly productive area in Arctic water masses north of the Polar Front. During a multi-disciplinary cruise in 1995, ecological and oceanographic processes were investigated at four stations located in a north-south transect in the MIZ. This study was carried out in Arctic water masses north of the Polar Front where ice conditions varied from dense first-year pack ice to open water. Also, the phytoplankton development varied along the transect from a pre-bloom situation at the northern-most station to a post-bloom situation in the open water. This paper includes a study of the zooplankton community and population structure of some of the dominant copepod species. Numerically, the most important mesozooplankton components were the copepods Calanus glacialis, Pseudocalanus minutus and Oithona similis. Copepods of Atlantic origin, such as Calanus finmarchicus and Oithona atlantica, gave evidence of an advection of Atlantic water masses into the area. It is concluded that the occurrence of new cohorts of Arctic copepods coincides with the onset of the phytoplankton bloom in the MIZ, and, that therefore, the spawning relies on stored energy.      

Increasing picocyanobacteria success in shelf waters contributes to long‐term food web degradation

Continental margins are disproportionally important for global primary production, fisheries and CO₂ uptake. However, across the Northeast Atlantic shelves, there has been an ongoing summertime decline of key biota—large diatoms, dinoflagellates and copepods—that traditionally fuel higher tropic levels such as fish, sea birds and marine mammals. Here, we combine multiple time series with in situ process studies to link these declines to summer nutrient stress and increasing proportions of picophytoplankton that can comprise up to 90% of the combined pico‐ and nanophytoplankton biomass in coastal areas. Among the pico‐fraction, it is the cyanobacterium Synechococcus that flourishes when iron and nitrogen resupply to surface waters are diminished. Our field data show how traits beyond small size give Synechococcus a competitive edge over pico‐ and nanoeukaryotes. Key is their ability to grow at low irradiances near the nutricline, which is aided by their superior light‐harvesting system and high affinity to iron. However, minute size and lack of essential biomolecules (e.g. omega‐3 polyunsaturated fatty acids and sterols) render Synechococcus poor primary producers to sustain shelf sea food webs efficiently. The combination of earlier spring blooms and lower summer food quantity and quality creates an increasing period of suboptimal feeding conditions for zooplankton at a time of year when their metabolic demand is highest. We suggest that this nutrition‐related mismatch has contributed to the widespread, ~50% decline in summer copepod abundance we observe over the last 60 years. With Synechococcus clades being prominent from the tropics to the Arctic and their abundances increasing worldwide, our study informs projections of future food web dynamics in coastal and shelf areas where droughts and stratification lead to increasing nutrient starvation of surface waters.     

Spawning strategies in cypriniform fishes in a lowland river invaded by non-indigenous European barbel Barbus barbus

Spawning strategies of lowland river fishes include single spawning, where reproduction generally occurs in early spring to provide 0+ fish with an extended growth season through the summer, but with a high risk of stochastic mortality events occurring, such as early summer floods. This risk can be reduced by multiple or protracted spawning strategies, where 0+ fish are produced over an extended period, often into mid-summer, but with the trade-off being a shorter growth season. The spawning strategies of cypriniform fish were explored in the River Teme, a spate river in Western England, which has non-indigenous European barbel Barbus barbus present. Sampling 0+ fish in spring and summer and across three spawning periods, B. barbus, chub Squalius cephalus and minnow Phoxinus phoxinus always revealed multiple spawning events, with 0+ fish of < 20 mm present in samples collected from June to August. Fish below 20 mm in August remained relatively small by the end of their growth season (October). For dace Leuciscus leuciscus, only single spawning events were evident, but with 0+ dace always being relatively large. Therefore, multiple spawning appears to be a common strategy that provides resilience in 0+ fish against stochastic mortality events in lowland rivers.

     

Effects of Partially Anadromous Arctic Charr (Salvelinus alpinus) Populations on Ecology of Coastal Arctic Lakes

Little research has been conducted on effects of iteroparous anadromous fishes on Arctic lakes. We investigated trophic ecology, fish growth, and food web structure in six lakes located in Nunavut, Canada; three lakes contained anadromous Arctic charr (Salvelinus alpinus) whereas three lakes did not contain Arctic charr. All lakes contained forage fishes and lake trout (Salvelinus namaycush; top predator). Isotope ratios (δ¹³C, δ¹⁵N) of fishes and invertebrates did not differ between lakes with and without anadromous Arctic charr; if anadromous Arctic charr deliver marine-derived nutrients and/or organic matter to freshwater lakes, these inputs could not be detected with δ¹³C and/or δ¹⁵N. Lake trout carbon (C):nitrogen (N) and condition were significantly higher in lakes with Arctic charr (C:N = 3.42, K = 1.1) than in lakes without Arctic charr (C:N = 3.17, K = 0.99), however, and ninespine stickleback (Pungitius pungitius) condition was significantly lower in lakes with Arctic charr (K = 0.58) than in lakes without Arctic charr (K = 0.64). Isotope data indicated that pre-smolt and resident Arctic charr may be prey for lake trout and compete with ninespine stickleback. Linear distance metrics applied to isotope data showed that food webs were more compact and isotopically redundant in lakes where Arctic charr were present. Despite this, lake trout populations in lakes with Arctic charr occupied a larger isotope space and showed greater inter-individual isotope differences. Anadromous Arctic charr appear to affect ecology and feeding of sympatric freshwater species, but effects are more subtle than those seen for semelparous anadromous species.     

Seasonal variability in copepod ingestion and egg production on the Faroe shelf

Copepod community ingestion rates of Calanus finmarchicus, Temora longicornis, Acartia longiremis and Pseudocalanus spp., and egg production rates of C. finmarchicus and T. longicornis, were studied in relation to phytoplankton composition, abundance and biomass on the Faroe shelf during a one-year cycle. The phytoplankton community during winter was mainly composed of small flagellates and the copepods of Pseudocalanus spp. As the spring bloom progressed, diatoms increased in abundance and dominated the biomass throughout summer. C. finmarchicus increased in numbers in early spring, while T. longicornis and A. longiremis dominated the community during summer and autumn. While no response in ingestion rates was observed for A. longiremis and Pseudocalanus spp. with increasing diatom biomass, both ingestion rates and egg production of C. finmarchicus and T. longicornis, generally increased, showing a dependence upon diatoms for production. The daily ingestion for C. finmarchicus females was 7% and 22% of body biomass during the pre-bloom and bloom period, respectively, while for T. longicornis females it was 33% and 56% and for A. longiremis females 22% and 33%, respectively. C. finmarchicus accounted for more than 80% of the total copepod ingestion in May, but in mid- and late summer, T. longicornis and A. longiremis dominated, and represented 80–90% of the total copepod ingestion. The proportion of reproductively mature C. finmarchicus increased as the phytoplankton biomass increased. Most of the time there was good agreement between herbivorous ingestion rates and calculated carbon demand for the observed egg production. However, both species showed a peak in egg production prior to the phytoplankton spring-bloom. Handling editor: J. Padisak.