Trophic importance of the chaetognaths Eukrohnia hamata and Sagitta gazellae in the pelagic system of the Prince Edward Islands (Southern Ocean)

Trophodynamics and predation impact of the 2 dominant chaetognaths Eukrohnia hamata and Sagitta gazellae were investigated at 19 stations in the vicinity of the Prince Edward Islands and at a 24-h station occupied at the sub-Antarctic Front in late summer (April/May) 1996. During the entire investigation, the zooplankton assemblages were numerically dominated by copepods with densities ranging from 21 to 170 ind. m⁻³. Amongst the copepods, Clausocalanus brevipes, Metridia gerlachei and M. lucens dominated accounting for >90% of the total. Generally, chaetognaths were identified as the second most important group composing at times up to 30% (mean = 14.7%) of total zooplankton abundance. Of the two chaetognath species, E.␣hamata was generally numerically dominant. Gut content analysis showed that both chaetognath species are opportunistic predators generally feeding on the most abundant prey, copepods. No feeding patterns were evident during the 24-h station, suggesting that both species feed continuously. The feeding rates of E. hamata ranged from 0 to 0.50 prey ind. day⁻¹ and between 0 and 0.90 prey ind. day⁻¹ for S. gazellae. The maximum total predation impact of E. hamata was equivalent to 5.2% of the copepod standing stock or up to 103% of copepod production per day. For S. gazellae the predation impact was lower, reaching a level of 3.2% of the copepod standing stock or 63% of the daily copepod production. Chaetognaths can, therefore, be regarded as an important pelagic predator of the Prince Edward Islands subsystem. Received: 27 March 1997 / Accepted: 11 September 1997     

Distribution and feeding of chaetognaths in the epipelagic zone of the Lazarev Sea (Antarctica) during austral summer

This study seeks to determine the effects of local hydrography on the distribution, abundance and feeding of chaetognaths in the Lazarev Sea, an area strongly controlled by physical processes which has been held responsible for initiating the Weddell Polynya. Zooplankton samples were taken at 39 stations on four transects located between 6°W and 3°E and from 60°S to 70°S between surface and 350 m. The dominant species, Eukrohnia hamata, accounted for 86.5% of all chaetognaths, followed by Sagitta gazellae (8.1%) and Sagitta marri (5.4%). These three species showed distinct vertical and horizontal distribution patterns. While E. hamata and S. marri had maximum abundances below 250 m depth, S. gazellae showed a narrow distribution band in the upper 150 m depth. The distribution pattern was strongly modified at the Greenwich meridian with an upward transport of a high abundance of deep dwelling organisms (S. marri and E. hamata) and a displacement of S. gazellae to the surface, likely coupled with the rise of the warm, saline halo around the Maud Rise. Small copepods were the main prey of all three chaetognath species. Feeding rates (FR) varied among species and depth. Sagitta marri showed the highest FR with 0.38 prey d⁻¹, followed by S. gazellae and E. hamata (0.22 and 0.07 prey d⁻¹). Feeding rates were usually highest in the 25–80-m stratum. Size distribution and maturity of E. hamata revealed a dominance of small and immature organisms along all depths and stations, suggesting that this area might be acting as an important source of recently spawned organisms to the surface.     

Feeding of Sagitta enflata and vertical distribution of chaetognaths in relation to low oxygen concentrations

Zooplankton samples were collected in Mejillones Bay, northernChile (23°00'15'S, 70°26'43'W ). Sampling was conductedat 4 h intervals, for 24 h during three seasons, austral spring(October 2000), summer ( January 2001) and winter (August 2001)at three different strata (0–25, 25–50 and 50–100m). Five species of chaetognaths were collected. Sagitta enflatawas the most abundant species, representing up to 65% of allchaetognaths in total numbers, followed by Sagitta bierii, makingup 34% of the total abundance of chaetognaths. S. enflata wasdistributed mainly above the Oxygen Minimum Zone, while S. bieriiremained below this zone. Feeding rates were relatively constantwithin the upper layer (0–25 m depth), for each samplingdate, averaging 1.2 prey S. enflata day^–1, and decreasingwith depth. Gut content analyses demonstrated that predationwas principally focused on small copepods (<1500 µm),with greatest feeding activity occurring at night. The dailypredation impact on the total standing stock of small copepodsvaried seasonally between 6% in spring and 0.4% in winter. Thispercentage may represent a negligible impact on the entire copepodcommunity, but it is relevant at the species or genus level,since S. enflata removed more than 20% of the standing stockof Centropages brachiatus and Corycaeus sp. Thus, during someperiods of the year, chaetognaths may strongly influence theabundance and size distribution of copepods in coastal upwellingecosystems.     

Chaetognaths of the United States South Atlantic Bight: Distribution, abundance and potential interactions with newly spawned larval fish

Chaetognaths are one of the most numerous organisms in the zooplankton community off the coast of North Carolina. During two years of offshore sampling in the late winter to early spring, sixteen chaetognath species were identified, four of which had not previously been reported in the waters of the United States South Atlantic Bight. Offshore samples were dominated by Sagitta enflata Grassi, 1881, one of the larger species, which contributed > 61% of total chaetognath abundance while dominant coastal species were S. tenuis Conant, 1896 and S. hispida Conant, 1895. Abundances, body sizes and spatial distributions were determined for the most abundant chaetognath species along with the overall abundance of three common co-occurring larval fish species (spot, Atlantic croaker and Atlantic menhaden). In addition, laboratory feeding experiments were conducted using S. tenuis and S. hispida to estimate the potential impact of chaetognath predation on representative North Carolina larval fish which spawn offshore and subsequently migrate into local estuaries. Feeding rates (no. prey items day^− 1) varied with prey type and the condition (starved/fed) of the chaetognath. Weight specific daily rations (SDR) were found to vary inversely with chaetognath size, decreasing exponentially with an increase in chaetognath length. The observed abundance and distribution data indicate that wintertime chaetognath populations in offshore waters of Onslow Bay, North Carolina have the potential to interact with recently spawned larval fish and may simultaneously act as competitors, predators, or prey. Furthermore, calculations using published values of chaetognath abundance, zooplankton standing crops, and our SDR estimates indicate that chaetognaths in a representative North Carolina estuary would require a minimum of 5.96 cal m^− 3 day^− 1 to sustain their biomass. Allowing for the reported spatial and temporal variability in zooplankton abundance in these systems, chaetognaths should consume 4.4%-20.9% of the estimated total zooplankton production day^− 1. This further emphasizes the role of chaetognaths not only as predators, but also as competitors with larval fish for zooplankton food stocks in southeastern United Sates estuaries.     

Meso- and bathypelagic distribution and abundance of chaetognaths in the Atlantic sector of the Southern Ocean

We conducted multinet sampling during winter and summer in the Southern Ocean (Atlantic sector) to investigate the effect of water mass, season and water depth on abundance and species composition of meso- and bathypelagic chaetognaths. Eukrohnia hamata (mean 115 ind. 1,000 m⁻³) and Sagitta marri (mean 51 ind. 1,000 m⁻³) were dominant, complemented by E. bathypelagica (mean 19 ind. 1,000 m⁻³) and E. bathyantarctica (mean 19 ind. 1,000 m⁻³) below 1,000 m. A further six species were identified, among them the rare bathypelagic species Heterokrohnia fragilis and the subtropical Eukrohnia macroneura that is new to the Antarctic. Water depth and season were the principal determinants of abundance and species composition patterns, indicating vertical seasonal migration and vertical segregation of species. The life cycles of E. hamata and S. marri were studied additionally. Their maturity stages were vertically segregated and prolonged reproductive periods are suggested for both species.     

The feeding rates ofPleurobrachia (ctenophora) andSagitta (chaetognatha), with notes on the potential seasonal role of planktonic predators in the dynamics of north sea zooplankton communities

The zooplankton off the north-east coast of England has been the subject of a number of studies focusing on its productivity. It has also been shown to be representative of the zooplankton of much of the western North Sea. The community contains a number of predatory species, three of which are widely described as ‘voracious’, the ctenophorePleurobrachia pileus, the chaetognathSagitta elegans and the hyperiid amphipodThemisto compressa (≡ Parathemisto gaudichaudi). This study investigates the role of these planktonic predators in this community, with special reference to the seasonal changes in predation pressure. The functional response ofPleurobrachia pileus feeding onAcartia was determined from laboratory experiments. It was found to be linear at prey densities typical of UK coastal waters, although the linear relationship appeared to break down at low and high prey densities. Feeding rate data forSagitta elegans were obtained from gut content analysis and published laboratory derived estimates of digestion time. Of the 1,789 individuals examined 198 (11.1%) had food remains in the gut. A linear relationship betweenSagitta length and prey size was established and the daily feeding rate ofSagitta elegans was estimated to be 0.4 prey items d⁻¹ ind⁻¹. For comparative purposes, the proportion of the copepod standing stock removed bySagitta elegans, Pleurobrachia pileus andThemisto gaudichaudi was estimated for each month of the year. From this model it was shown thatThemisto applied the most predation pressure, andSagitta elegans applied the least predation pressure of the three planktonic predators considered. The impact ofPleurobrachia will be to a large extent offset due to its peak of seasonal abundance coinciding with the zooplankton peak in the summer.     

Predator-limited population growth of the copepod Pseudocalanus sp.

The impact of predators on population growth of Pseudocalanussp. was investigated in Dabob Bay, Washington. Mortality ofPseudocalanus sp. was determined from stage-specific survivorship,from seasonal changes in mortality rates of adult males andfemales and from incidence of injuries to adult copepods. Theprincipal predators of adult Pseudocalanus were identified asthe predatory copepod Euchaeta elongata, the omnivorous euphausiidEuphausia pacifica and the chaetognath Sagirta elegans. Predatorattack rates - and prey mortality rates - are highly density-dependentand thus sensitive to prey dispersion in the water column, particularlyto layering in the vertical plane. Predation rates by the threeprincipal predators exceeded 100% of the recruitment rate toadult Pseudocalanus sp. beginning in early summer, thus restrictingpopulation growth. Planktivorous fish predation (by adult three-spinestickleback, Gasterosteus aculeatus, and juvenile chum salmon,Oncorhynchus keta) on Pseudocalanus sp. adults was estimatedto be two orders of magnitude lower than consumption rates bypredatory zooplankton, at a deep water station in July. Analysisof seasonal changes in prey ingested by Sagitta elegans revealedthat Pseudocalanus sp. was the major prey item of S. elegansin April (61.0% of prey) and in June (67.0% of prey), thereafterdeclining seasonally in importance. Predation by S. elegansvaried seasonally with changes in chaetognath stage structure,vertical distribution and diapause, not size structure alone.Although chaetognath recruitment and population growth appearto be directly coupled to the abundance of Pseudocalanus sp.,predation by S. elegans has little reciprocal impact on Pseudocalanussp. population growth; hence asymmetries may occur in the interaction of planktonic prey and predators.     

Diel vertical migration and feeding of chaetognaths in coastal waters of the eastern Mediterranean

This study investigates the diel vertical distribution and the diet of the most important chaetognath species found in the 0–50 m surface layer of a coastal area in the eastern Mediterranean during a 24-hour period in July 2004. Among the recorded chaetognaths, Sagitta enflata was the most abundant species (41.6%), followed by S. minima (32.5%) and S. serratodentata (20.8%). Those three species exhibited a “twilight migration” pattern, with only small differences among them. Vertical separation was found between S. enflata and S. minima. Both species preyed mainly on cladocerans, although copepods were the most abundant group in the zooplankton assemblage. The chaetognath species followed partially the diel vertical migration of their prey. S. enflata showed high feeding intensity at different times in both day and night, while S. minima fed more intensively at midday (12:00) and at dusk (20:00), and S. serratodentata in the morning (08:00). It seems that in order to coexist in an area of low productivity the chaetognath species follow the basic ecological rules of space, time and food-type separation, in order to reduce the inter- and intra-specific competition. The high preference of S. minima and especially of S. enflata for the cladocerans made them probably the most important predators of cladocerans during summer.     

The life history and vertical distribution of the chaetognath, Sagitta elegans, in Dabob Bay, Washington

Seasonal changes in the population structure and patterns ofvertical distribution and migration of the dominant chaetognath,Sagitta elegans, in a fjord off Puget Sound were determinedwith samples collected at approximately semi-monthly intervalsfrom November 1972 to November 1973. S. elegans bred continuouslyfrom March through August, with major recruitment in June andJuly. Seasonal changes in the vertical distribution of the populationand in the proportion of the population that vertically migratedreflected the stage of maturity and size of the individuals.Small chaetognaths were non-migratory and distributed in thetop 100 m. Stage II individuals were dispersed throughout thewater column; some did migration was evident. Breeding stageswere restricted to a layer between SO and 100 m during the dayand migrated to the surface layer at night, enhancing the probabilityof successful reproduction.     

Mesoscale spatial distribution of chaetognaths along hydrographic gradients in the South Scotia Sea (Antarctica)

Mesoscale distribution of chaetognaths in the South Scotia Sea was studied in January 1994. Three transects (ice margin to Elephant Island, ice margin to the Weddell-Scotia Confluence, and the Drake Passage) and a station grid in the East Bransfield Strait were laid out, seeking the greatest possible heterogeneity in hydrographic conditions. Six species were collected, with the most abundant species, Eukrohnia hamata, accounting for 83% of the total number of individuals, followed by Sagitta gazellae (9.7%). High spatial variability was observed both along the transects and over the mesoscale station grid. The station grid allowed two assemblages to be distinguished on the basis of species abundance and demographic population structure. Both assemblages were distributed through areas with distinct hydrographic characteristics. Use of a smaller sampling scale and a more detailed definition of development stages than in earlier studies revealed a higher level of spatial heterogeneity than previously suggested for chaetognaths in the Southern Ocean. Accepted: 11 April 1999     

Notes on feeding of Chaetognatha in Guanabara Bay, Brazil

A preliminary analysis, by means of the gut content, of thediet of Sagitta friderici and Sagitta enflata occurring in GuanabaraBay was carried out. The samples were collected at differenttimes over a 4day period in September 1995, during verticaltows with a conical net of 200 µm mesh size at a fixedstation (30 m depth). In total, 1000 individuals were examined.Copepoda were the numerically dominant group of prey of S.fridericiand S.enflata (67.2 and 74.6%, respectively); both species behaveas opportunistic carnivores, feeding mainly on the more abundantcopepod species throughout the water column. Adult individualsof S.fnderici and S.enflata (stages III and IV) seem to havefood requirements different from juveniles: adults fed on otheritems (chaetognaths, crus tacean larvae, hydromedusae and luciferidae),and the food containing ratio (FCR) was higher in indi vidualsof stage Ill. In general, only one prey per gut was registered.Chaetognaths may have chosen their prey in relation to theirsize, since the size of the prey was proportional to the sizeof their predators. Feeding intensity (NPC) was higher in individualscollected during the night periods.     

Planktivory in Pagothenia borchgrevinki (Pisces: Nototheniidae) in McMurdo sound,Antarctica

Summary Specimens of the Antarctic fish P. borchgrevinki were collected from just beneath the sea-ice in McMurdo Sound over the period 8 November to 6 December, 1985. The stomach contents of 200 fish were analysed and compared with the co-occurrence of the prey species in the plankton. Numerically, the dominant prey was a thecosomatid pteropod Limacina helicina (83%), followed by a hyperiid amphipod Hyperiella dilatata (5%). Other prey species in decreasing order of occurrence were Euphausia crystallorophias, Euchaeta antarctica, other copepods, a decapod crustacean larva, chaetognaths, the amphipods Orchomene plebs and Epimiriella macronyx, and unidentified juvenile fish. The order of frequency of occurrence of these prey in the guts of all the fish was H. dilatata (in 73% of the fish), L. helicina (71%), calanoid copepods (55%), chaetognaths (51%), E. crystallorophias (42%), decapod crustacean larva (32%), O. plebs (27%), juvenile fish (20%), and E. macronyx (14%). In volumetric terms, the dominant diet contributors were O. plebs (38%), L. helicina (17%) and chaetognaths (15%). All prey except O. plebs and E. macronyx were taken in concurrent plankton samples. Although planktivory of Pagothenia borchgrevinki is confirmed, it is not certain whether feeding is confined to the immediate sub-ice environment.     

Composition, spatial and temporal variations of Chaetognatha in Guanabara Bay, Brazil

This paper presents the specific composition and distributionof adult chaetognaths in Guanabara Bay, Brazil. A comparisonhas been made between chaetognath and copepod distribution patterns.Three areas were distinguished within Guanabara Bay: an externalarea, influenced by oceanic waters; an internal area, influencedby fluvial waters; and a transitional area between the two.Monthly samples were collected by horizontal surface hauls usinga conical net with 200µm mesh. Three species were identified:Sagitta friderici, S.enflata and S.hispida. Sagitta fridericiwas the dominant species and occurred throughout the year atthe three areas. Sagitta enflata and S.hispida were limitedto the more saline waters. Generally, greater densities of chaetognathswere observed following copepod peakden sities. Temperatureand salinity associated with copepod abundance were the mostimportant factors affecting the distribution pattern and theabundance of chaetognath species in Guanabara Bay.     

Vertical distribution and abundance of juvenile chaetognaths in the Weddell Sea (Antarctica)

The composition, abundance and vertical distribution of chaetognaths were analysed along a transect in the Weddell Sea during late spring. Three species were identified: Eukrohnia hamata (90.8%), Sagitta marri (6.4%) and S. gazellae (2.8%). Only juvenile stages were collected in the samples, a result related both to the type of sampling gear employed (mesh size: 100 μm) and the species' life-cycles. The vertical distributions showed that the juvenile stages of these species tended to aggregate at considerable depth (1000–500 m). It is postulated that this pattern may be related to the life-cycles of these species in association with seasonal Antarctic conditions, similar to the pattern postulated for krill and other polar crustaceans. Accepted: 10 July 2000     

Distribution, abundance, behavior and metabolism of Periphylla periphylla, a mesopelagic coronate medusa in a Norwegian fjord

The distribution, behavior and metabolism of the mesopelagic jellyfish, Periphylla periphylla (Péron & Lesueur), were investigated in Lurefjorden, Norway. Field studies, conducted in 1998–1999 with plankton nets and a remotely operated vehicle, indicated that 80-90% of the dense (up to 2.5 m^–3) population migrated 200–400 m vertically each day throughout the year. In situ observations with red light revealed that swimming rates and feeding activity varied with age and time of day. Detection of turbulence and contact with surfaces caused this medusa to conceal one or all of its tentacles in the stomach or to shed nematocyst-laden tissue from the tentacles. Stomachs of medusae collected with nets were often full of prey entangled with the sloughed tissue. Stomachs of medusae captured individually with ROV samplers were empty or contained only a few prey in their stomachs (typically, 1–4 copepods Calanus spp. or chaetognaths Eukrohnia hamata Möbius per medusa). Low rates (0.4–5.6 l O₂ mg C^–1 h^–1) of oxygen consumption of P. periphylla suggested that this species was sustained by relatively few (1–34) prey d^–1.     

Interannual variability in a predator-prey interaction: climate, chaetognaths and copepods in the southeastern Bering Sea

The interaction of the chaetognath Sagitta elegans with thecopepod community of the southeast Bering Sea middle shelf wasexamined in relation to environmental conditions during 1995–1999.Predation impact was estimated for 2 years, 1995 and 1997, usinggut content analysis, experimentally derived digestion time(DT) and abundances of chaetognaths and prey. Pseudocalanusconcentrations correlated with water temperature and Calanusmarshallae with sea ice extent. Sagitta elegans were less abundantbut individuals were larger in 1995, when C. marshallae predominated,compared to 1997, when Pseudocalanus and Acartia were the primaryprey. Predation by S. elegans removed <1% standing stockday^–1 of Pseudocalanus or C. marshallae in 1995 and 1.7to 2.3% of Pseudocalanus in 1997. The percent of the copepodcommunity biomass required by chaetognaths was estimated tobe <1% in 1995 compared with 8–12% in 1997. Calanusmarshallae may be more vulnerable than Pseudocalanus to cumulativepredation effects because of its reproductive strategy. Theeffect of chaetognath predation on the copepod community dependson which copepod species is predominant and its susceptibilityto cumulative predation effects, as well as on daily predationimpact, both of which varied between years with different climaticconditions.     

Prey selection and diel feeding of the freshwater jellyfish, Craspedacusta sowerbyi

1. The diet of the invasive freshwater jellyfish, Craspedacusta sowerbyi (Lankester), was assessed by analysing its stomach contents. 2. The medusae ingested various zooplankton and benthic prey in the 0.1–3.0 mm size range. The selectivity indices for prey showed that larger zooplankton (0.4–1.4 mm) and active prey such as copepods were preferred; small potential prey, and the loricate rotifer Keratella cochlearis in particular, were almost never consumed. Measurements suggest that spacing between the tentacle branches could determine prey selection. Another possibility is that some prey are too small or slow to activate nematocysts after collision with the predator. Prey > 1.4 mm can probably escape or are otherwise too large to be handled. 3. The medusae of Craspedacusta medusae stayed in deep water during daytime and migrated upwards at night. The greatest number of prey in freshwater jellyfish stomachs was detected at night. 4. Calculation of consumption rates, based on mean stomach contents and digestion times of 4–5 h, showed that, on average, one medusa takes 190 zooplankton prey day^??1 at a mean prey density of about 100 L^??1. At the observed medusae density of 1 m^??3, the cropping rate was about 0.2 zooplankton prey L^??1 day^??1.     

Seasonal variation and life history of the pelagic chaetognatha, Sagitta elegans Verrill, in Toyama Bay, southern Japan Sea

The seasonal variation and life history of the pelagic chaetognatha,Sagitta elegans Verrill, in Toyama Bay, southern Japan Sea,were investigated using a time series of 0–500 m verticalhauls with a Norpac net from 1 February 1990 to 30 January 1991.Nine species of one genus occurred in Toyama Bay throughoutthe year, including the cold-water species S.elegans. Therewas no remarkable variation in abundance throughout the year,although many individuals were collected in August. Juvenilesoccurred mostly in spring and summer, from late March to August.Adults (Stage 3) occurred in all seasons, except summer. Thebody length of adults ranged from 26 to 30 mm. There were twoprincipal spawning periods. One was in March-May and the otherwas in August. Life spans of both cohorts were 10–12 months.Copepods were major prey of S.elegans inhabiting Toyama Bay.The annual mean food-containing ratio (FRC) was 6.1%, but values>10% occurred in February, April, May, July and September.     

Feeding ecology of the lantern fish Benthosema glaciale in a subarctic region

Summary The myctophid, Benthosema glaciale, was found to occur in the region of Davis Strait, but not in northern Baffin Bay. The factors that limited its northern range were believed to be the cold temperature and the long periods of day light in Baffin Bay. During the day B. glaciale was found at a depth of between 300 and 900 m with the population concentrating at about 500 m. At night about 46% of the population migrated to the upper 65 m to feed on copepods, primarily Calanus finmarchicus stage V and C. hyperboreus stage IV for which they showed a high degree of selectivity. The weight of stomach contents of fish with 100% full stomachs was equal to about 3% of their body weight. Feeding occurred mainly at night in the upper layers above the coldest layer of water with a temperature of about 0.5 °C. The biomass of zooplankton and myctophids m^-2 were similar to those for these groups on the Nova Scotia slope during June. The fish fed on the same size range of prey in Davis Strait as on the Nova Scotia slope even though the zooplankton species composition was different in the two regions.     

Polar night ecology of a pelagic predator,the chaetognath <Emphasis Type="Italic">Parasagitta elegans</Emphasis>

The annual routines and seasonal ecology of herbivorous zooplankton species are relatively well known due to their tight coupling with their pulsed food source, the primary production. For higher trophic levels of plankton, these seasonal interactions are less well understood. Here, we study the mid-winter feeding of chaetognaths in high-Arctic fjord ecosystems. Chaetognaths are planktivorous predators which comprise high biomass in high-latitude seas. We investigated the common species Parasagitta elegans around the Svalbard archipelago (78–81°N) during the winters of 2012 and 2013. Our samples consisted of individuals (body lengths 9–55 mm) from three fjords, which were examined for gut contents (n = 903), stable isotopes, fatty acid composition, and maturity status (n = 352). About a quarter of the individuals contained gut contents, mainly lipid droplets and chitinous debris, whilst only 4 % contained identifiable prey, chiefly the copepods Calanus spp. and Metridia longa. The δ¹⁵N content of P. elegans, and its average trophic level of 2.9, confirmed its carnivorous position and its fatty acid profile [in particular its high levels of 20:1(n-9) and 22:1(n-11)] confirmed carnivory on Calanus. Observations of undeveloped gonads in many of the larger P. elegans, and the absence of small individuals <10 mm, suggested that reproduction had not started this early in the year. Its average feeding rate across fjords and years was 0.12 prey ind.^?1 day^?1, which is low compared to estimates of spring and summer feeding in high-latitude environments. Our findings suggest reduced feeding activity during winter and that predation by P. elegans had little impact on the mortality of copepods.