Ciliates as a food source for marine planktonic copepods

Copepods of the genusEurytemora, isolated from the Patuxent River, a tributary of Chesapeake Bay, were fed suspensions of the ciliateUronema isolated from the Rhode River, a subestuary of Chesapeake Bay. Grazing by copepods was determined by the decrease in numbers of ciliates, which were monitored by both direct counting and particle size analysis. Results from both methods of analysis showed significant reduction in the numbers ofUronema in the suspension whenEurytemora was present. Survival of copepods with ciliates added as food source was significantly longer than without ciliates. Analysis of field samples collected in the fall showed that ciliates comprised approximately 20% of the total plankton biomass at selected sampling sites. The results of the laboratory and field studies indicate that copepods can feed on ciliates and suggest that, in nature, ciliates may comprise an important source of food for copepods.     

Intrashoot distributions of leaf dwelling harpacticoid copepods on the seagrass Zostera marina L.: implications for sampling design

The distribution of leaf dwelling harpacticoid copepods within seagrass (Zostera marina L.) shoots was investigated on four dates in 1986 and 1987. Copepods were found to be non-uniformly distributed on shoots, with higher abundances observed on older leaves. Patterns of abundance within shoots could not be explained by the surface area of individual leaves except on the sampling date with the highest copepod densities. It is suggested that harpacticoid copepod distributions on seagrass shoots are primarily determined by the pattern of epiphytic biomass. However, at high population densities, habitable surface area may be the limiting factor. Increased habitat complexity at high epiphyte loads does not seem to be the cause of the copepod distributions observed in this study. An accurate method for estimating seagrass copepod abundance per unit sediment area using intrashoot distributions is described and compared to existing methodology.     

Light primes the escape response of the calanoid copepod, Calanus finmarchicus

The timing and magnitude of an escape reaction is often the determining factor governing a copepod's success at avoiding predation. Copepods initiate rapid and directed escapes in response to fluid signals created by predators; however little is known about how copepods modulate their behavior in response to additional sensory input. This study investigates the effect of light level on the escape behavior of Calanus finmarchicus. A siphon flow was used to generate a consistent fluid signal and the behavioral threshold and magnitude of the escape response was quantified in the dark and in the light. The results show that C. finmarchicus initiated their escape reaction further from the siphon and traveled with greater speed in the light than in the dark. However, no difference was found in the escape distance. These results suggest that copepods use information derived from multiple sensory inputs to modulate the sensitivity and strength of the escape in response to an increase risk of predation. Population and IBM models that predict optimal vertical distributions of copepods in response to visual predators need to consider changes in the copepod's behavioral thresholds when predicting predation risk within the water column.     

Copepod foraging and predation risk within the surface layer during night-time feeding forays

Vertical distribution data seem to indicate that certain speciesof diel vertical migrating copepods avoid the surface high chlorophyll(Chl) region within coastal and estuarine environments, evenduring the night. Copepods may avoid this layer to reduce predationmortality, avoid advective loss or to avoid consuming too muchtoxic algae. We hypothesize that copepods make several intermittentfeeding ‘forays’ into shallow surface layers duringthe night, returning to intermediate depths between forays.Using an individual-based model (IBM) of Calanus pacificus,we examined the implications of this behavior on feeding successand mortality risk, and tested whether a practical field-samplingscheme would be able to detect foray-like behavior. In somecases, mortality of the foray-foraging copepods was up to 50%less than that of randomly behaving controls, for a given amountof food ingested. The trapping scheme devised should be ableto detect the occurrence of foray behavior (FB) in the fieldand should show differences in the gut contents of copepodsentering and leaving the uppermost food-rich layer. The presenceor absence of foray-like behavior significantly altered therelative concentration of copepods within various surface strataand thus could influence the temporal availability of copepodsas prey for the larvae and juveniles of several important managedfish species. This paper is one of six on the subject of the role of zooplanktonpredator–prey interactions in structuring plankton communities.     

Effects of food nitrogen content and concentration on the forms of nitrogen excreted by the calanoid copepod, Acartia tonsa

Nitrogen excreted as ammonium, urea, and dissolved primary amines (DPA), and nitrogen ingested by the planktonic calanoid copepod, Acartia tonsa, were measured while fed 4 foods with different N/C ratios in high (500 μg C l^− 1) and low (50 μg C l^− 1) concentrations. Adult copepods were fed the ciliate, Uronema marinum (N/C = 0.26), the diatom, Thalassiosira weissflogii, in log-phase growth (N/C = 0.20), and in senescent-phase growth (N/C = 0.12), and detritus derived from the saltmarsh grass, Spartina alterniflora, (N/C = 0.04). Total nitrogen excreted ranged from 0.06 to 0.18 μg N copepod^− 1 d^− 1 whereas nitrogen ingested exhibited considerably more variation (0.01 to 0.39 μg N copepod ^− 1d ^− 1). Ammonium was the dominant form of nitrogen excreted and was influenced by both food concentration and N/C ratio. Copepods fed foods with N/C ratios resembling their own body composition (log-phase diatoms and ciliates) excreted more ammonium when fed higher concentrations of food. In contrast, copepods fed foods with lower N/C ratios than their own body composition excreted more ammonium when fed lower concentrations of food, suggesting that they were catabolizing body protein for survival. Excretion of urea varied with food N/C ratio, with more urea excreted when the copepods were fed higher N/C foods. The excretion of DPA did not vary with either food concentration or food N/C ratio. Homeostasis serves to conserve the N/C ratio of copepods. Thus nitrogen excretion by healthy copepods should be expected to increase with ingestion only when copepods have high quantities of nitrogen-rich foods relative to the body composition of the copepods.     

Zooplankton community changes in Lake Kinneret (Israel) during 1969–1985

Long-term records (1969–1985) of zooplankton density in Lake Kinneret indicated significant reduction of biomass (Copepods, Cladocera, Rotifera) and production (Copepods, Cladocera). Nauplius and adult copepod densities decreased but those of copepodites did not change. Mesocyclops was suppressed more than the smaller Thermocyclops and males of both genera became more abundant relative to the larger females. Ratios of small/large Cladocera densities became higher. Numbers of total cladocerans were stable, and therefore reduction of Cladocera grazing capacity is assumed. The abundant Keratella spp. were reduced. It is likely that intensification of fish visual-attack-predation pressure shifted the size-class structure towards smaller adult copepods and cladocerans. Reduction of Keratella spp. and copepod nauplii was probably affected by increasing pressure of fish filter-feeders. Data on fish food consumption, feeding behaviour and fisheries management suggested their direct impact on long-term changes of zooplankton in Lake Kinneret.     

Feeding response of a benthic copepod to ciliate prey type,prey concentration and habitat complexity

1. Protozoans are important consumers within microbial food webs and, in turn, they represent potential prey for small metazoans. However, feeding interactions within these food webs are rarely characterised and this is especially true for freshwater sediments. 2. We aimed to quantify the feeding links between a freshwater meiofaunal copepod and ciliates in two laboratory experiments. The first experiment addressed the response of Eucyclops serrulatus towards ciliate density and type (two ciliate species of the same genus differing in terms of body size). A second experiment assessed the effect of habitat structure on feeding rates by introducing different structural complexity into the feeding arena. In contrast to the first experiment, which was run only for one time period, this experiment also tested three different total feeding times (4, 7 and 9 h). 3. Eucyclops serrulatus exhibited high ingestion rates, with 3–69 ciliates copepod^?1 h^?1 consumed depending on food concentration, food type and habitat complexity. Copepods exhibited a preference for the smaller ciliate when total ciliate concentration was low, but selected both ciliates equally when food concentrations were medium or high. However, at very high food concentration, Eucyclops preferred the larger ciliate (which was 1/3 of its own body size), suggesting that the longer handling times of the larger prey are rewarding when the large prey is present in high numbers. In terms of total numbers consumed, copepods fed on more small ciliates, but in terms of carbon units both ciliates were selected equally when total prey concentration was low or medium. However, copepods derived more carbon from the larger prey at high and very high prey concentrations (up to 0.7 μgC out of a maximum of 1.1 μgC copepod^?1 h^?1). Habitat complexity influenced the feeding of copepods when it was observed over time. 4. The copepod–ciliate link is well known from the pelagic zone of both marine and freshwater habitats. We have shown its potential importance within the benthos, where it can be influenced by food identity, food quantity and possibly by habitat complexity.     

Food selection by calanoid copepods in response to between-lake variation in food abundance

1. Food selection experiments were conducted by acclimating calanoid copepods (Eudiaptomus spp.) in suspensions of natural seston and then adding pairs of dual-labelled (¹⁴C/³²P) algae. Each feeding trial measured selectivity between a small, high-quality reference alga, Chlamydomonas reinhardii, and a test alga that differed in size and/or food quality. The influence of food concentration on food selection was tested by using seston from two lakes with contrasting food abundance and by including treatments with filtered lake water ('starved’) and seston diluted with filtered water or enriched with cultured algae. 2. Copepods that had been starved or acclimated to natural seston with low food abundance preferred the larger of two labelled algae, regardless of the nutritional quality of the algae. In agreement with the predictions of an optimal diet model, however, copepods that were acclimated to high food conditions discriminated against low-quality foods, including digestion-resistant algae and dead algae. 3. Selectivity coefficients showed excellent agreement with a previous study involving the same taxa of copepods and labelled algae but in which the copepods had been acclimated to pairs of cultured algae rather than natural seston. Thus, these comparisons emphasize the importance of food availability in modulating copepod selectivity for foods that differ in nutritional quality and suggest that such behaviour occurs in nature.     

The role of photoreception in the swarming behavior of the copepod Dioithona oculata

The copepod Dioithona oculata forms dense swarms near mangrove prop roots that are centered around shafts of light penetrating the mangrove canopy. Swarms can be created in the laboratory within light shafts created with a fiber optic light pipe. Laboratory observations of swarming behavior were recorded using video cameras, and the swimming behavior of the copepods and density of the swarms were quantified using video‐computer motion and image analysis techniques. Swarm formation results from a combination of phototactic and klino‐kinetic behavior. Dark adapted copepods initially exhibit a photophobic response to a light shaft, but become positively phototactic within 3–5 min after exposure to the light. Copepod aggregation rates under the light fit a saturation model, suggesting that copepods are attracted independently to the swarm marker. Copepods reverse their swimming direction when they encounter light intensity gradients near the edge of a light shaft, which aids in maintaining the swarm. Swarm formation can occur in the laboratory at light intensities as slow as 0.1 μM photons m^‐2 s^‐1, which is similar to light intensities at dawn when they are first observed to form in nature. Swarm formation appears to have an endogenous rhythm, as copepods will not form swarms at night under a light shaft.     

Induction and costs of tail spine elongation in Daphnia hyalina×galeata: reduction of susceptibility to copepod predation

1. Seasonal field data showed a positive correlation between the tail spine length of Daphnia hyalina × galeata and the density of the copepod Acanthocyclops robustus . Laboratory experiments were designed to assess the mechanisms underlying the induction of tail spine elongation and to test whether this morphology reduced predation on Daphnia juveniles.
2. Both the elongated tail spine morph and control spine morph produced progeny with elongated tail spine when exposed to copepods. Instar increments, calculated from individual body length, showed that individual growth decreased in the presence of copepods. This decrease in individual growth was less pronounced in spined morphs relative to control morphs.
3. Copepods exhibited significantly higher feeding rates on control morph juveniles compared to elongated tail spine morph juveniles.
4. All Daphnia matured at instar 5 even though the control morph neonates took 24–30 more hours to mature in the presence of copepods than in the absence of copepods. Enlarged body and spine lengths of progeny coincided with reduced progeny number during the first six reproductive instars. The disadvantage of the reduced offspring number produced per female was balanced by enhanced survivorship of progeny subjected to copepod predation.     

Spatial distribution and trophic ecology of dominant copepods associated with turbidity maximum along the salinity gradient in a highly embayed estuarine system in Ariake Sea, Japan

The present study aimed to investigate into the feeding ecology of the dominant copepods along a salinity gradient in Chikugo estuary. Copepod composition was studied from samples collected from stations positioned along the salinity gradient of the estuary. Copepod gut pigment concentrations were measured by fluorescence technique and hydrographical parameters such as temperature, salinity, transparency, suspended particulate matter (SPM); pigments such as chlorophyll-a (Chl-a), phaeopigment; and particulate nutrients such as particulate organic carbon (POC) and particulate organic nitrogen (PON) were measured. Two distinct zones in terms of nutrient and pigment concentrations as well as copepod distribution and feeding were identified along the estuary. We identified a zone of turbidity maximum (TM) in the low saline upper estuary which was characterized by having higher SPM, higher POC and PON but lower POC:PON ratios, higher pigment concentrations but lower Chl-a/SPM ratios and higher copepod dry biomass. Sinocalanus sinensis was the single dominant copepod in low saline upper estuary where significantly higher concentrations of nutrients and pigments were recorded and a multispecies copepod assemblage dominated by common coastal copepods such as Acartia omorii, Oithona davisae and Paracalanus parvus was observed in the lower estuary where nutrient and pigment concentrations were lower. Copepods in the estuary are predominantly herbivorous, feeding primarily on pigment bearing plants. However, completely contrasting trophic environments were found in the upper and the lower estuary. It was speculated from the Chl-a and phaeopigment values that copepods in the upper estuary receive energy from a detritus-based food web while in the lower estuary an algal-based food web supports copepod growth. Overall, the upper estuary was identified to provide a better trophic environment for copepod and is associated with higher SPM concentrations and elevated turbidity. The study demonstrates the role of estuarine turbidity maximum (ETM) in habitat trophic richness for copepod feeding. The study points out the role of detritus-based food web as energy source for the endemic copepod S. sinensis in the upper estuary, which supports as nursery for many fish species.     

Copepods act as a switch between alternative trophic cascades in marine pelagic food webs

A recent meta‐analysis indicates that trophic cascades (indirect effects of predators on plants via herbivores) are weak in marine plankton in striking contrast to freshwater plankton ( Shurin et al. 2002 , Ecol. Lett., 5, 785–791). Here we show that in a marine plankton community consisting of jellyfish, calanoid copepods and algae, jellyfish predation consistently reduced copepods but produced two distinct, opposite responses of algal biomass. Calanoid copepods act as a switch between alternative trophic cascades along food chains of different length and with counteracting effects on algal biomass. Copepods reduced large algae but simultaneously promoted small algae by feeding on ciliates. The net effect of jellyfish on total algal biomass was positive when large algae were initially abundant in the phytoplankton, negative when small algae were dominant, but zero when experiments were analysed in combination. In contrast to marine systems, major pathways of energy flow in Daphnia‐dominated freshwater systems are of similar chain length. Thus, differences in the length of alternative, parallel food chains may explain the apparent discrepancy in trophic cascade strength between freshwater and marine planktonic systems.     

Fatty acid profiles of algae mark the development and composition of harpacticoid copepods

1. The value of algal fatty acids (FA) as diet biomarkers for benthic harpacticoid copepods was investigated. A high proportion of 18:1ω9 and 18:2ω6 FA was observed in the lipid reserve fraction of copepods fed with cyanobacteria. In contrast, a high proportion of 16:1ω7 and ω3 FA (including eicosapentaenoic) was present in the lipid reserve fraction of copepods grown on diatoms. 2. Copepods that were grown on cyanobacteria showed reduced survival and took 26% more time to develop from the first copepodid stage to adult than copepods that were grown on diatoms. Copepods feeding on the cyanobacteria showed reduced FA content when compared with animals fed with diatoms. This reduction in FA content was more pronounced in the apolar lipid fraction (mainly reserve lipids) than in the polar (mainly structural) lipid fraction. 3. The FA profiles of algae were used to calculate a function discriminating between diatoms and cyanobacteria. This function was applied to the FA profiles in the reserve lipid fraction of copepods and correctly classified copepod diet. 16:1ω7, 18:2ω6 and 20:5ω3 were the most important FA in the discriminant function. The suitability of this chemometric method to infer copepod diet was further tested by using algal class FA data from literature to derive the discriminant functions. The correct classification of the diet when the functions were applied to FA composition of the copepod reserve lipids suggests that this method may be employed in trophic web studies. 18:3ω3, 18:1ω9 and 16:1ω7 were the most important FA in the functions discriminating diatoms, cyanobacteria and green algae. The identification and quantification of the whole suit of 16:1ω7, 18:1ω9, 18:2ω6, 18:3ω3 and 20:5ω3 in trophic web studies is therefore of paramount importance to infer diet origin of aquatic herbivores. 4. The FA profile of copepod polar lipids did not reflect that of the diet. The presence of long chain polyunsaturated FAs in the polar lipid fraction of copepods feeding on the cyanobacterium suggests that C18 FAs from the diet may be elongated and desaturated by the copepod. The ability to elongate and desaturated FAs may reduce the importance of some FAs as diet biomarkers while it may turn the copepods into valuable trophic intermediaries in transferring organic matter from microorganisms to higher trophic levels.     

Observing copepods through a genomic lens

Background

Copepods outnumber every other multicellular animal group. They are critical components of the world's freshwater and marine ecosystems, sensitive indicators of local and global climate change, key ecosystem service providers, parasites and predators of economically important aquatic animals and potential vectors of waterborne disease. Copepods sustain the world fisheries that nourish and support human populations. Although genomic tools have transformed many areas of biological and biomedical research, their power to elucidate aspects of the biology, behavior and ecology of copepods has only recently begun to be exploited.

Discussion

The extraordinary biological and ecological diversity of the subclass Copepoda provides both unique advantages for addressing key problems in aquatic systems and formidable challenges for developing a focused genomics strategy. This article provides an overview of genomic studies of copepods and discusses strategies for using genomics tools to address key questions at levels extending from individuals to ecosystems. Genomics can, for instance, help to decipher patterns of genome evolution such as those that occur during transitions from free living to symbiotic and parasitic lifestyles and can assist in the identification of genetic mechanisms and accompanying physiological changes associated with adaptation to new or physiologically challenging environments. The adaptive significance of the diversity in genome size and unique mechanisms of genome reorganization during development could similarly be explored. Genome-wide and EST studies of parasitic copepods of salmon and large EST studies of selected free-living copepods have demonstrated the potential utility of modern genomics approaches for the study of copepods and have generated resources such as EST libraries, shotgun genome sequences, BAC libraries, genome maps and inbred lines that will be invaluable in assisting further efforts to provide genomics tools for copepods.

Summary

Genomics research on copepods is needed to extend our exploration and characterization of their fundamental biological traits, so that we can better understand how copepods function and interact in diverse environments. Availability of large scale genomics resources will also open doors to a wide range of systems biology type studies that view the organism as the fundamental system in which to address key questions in ecology and evolution.     

Accumulation of mercury in estuarine food chains

To understand the accumulation of inorganic mercury and methylmercury at the base of the estuarine food chain, phytoplankton (Thalassiosira weissflogii) uptake and mercury speciation experiments were conducted. Complexation of methylmercury as methylmercury-bisulfide decreased the phytoplankton uptake rate while the uptake rate of the methylmercury-cysteine and -thiourea complexes increased with increasing complexation by these ligands. Furthermore, our results indicated that while different ligands influenced inorganic mercury/methylmercury uptake by phytoplankton cells, the ligand complex had no major influence on either where the mercury was sequestered within the phytoplankton cell nor the assimilation efficiency of the mercury by copepods. The assimilation efficiency of inorganic mercury/methylmercury by copepods and amphipods feeding on algal cells was compared and both organisms assimilated methylmercury much more efficiently; the relative assimilation efficiency of methylmercury to inorganic mercury was 2.0 for copepods and 2.8 for amphipods. The relative assimilation is somewhat concentration dependent as experiments showed that as exposure concentration increased, a greater percentage of methylmercury was found in the cytoplasm of phytoplankton cells, resulting in a higher concentration in the copepods feeding on these cells. Additionally, food quality influenced assimilation by invertebrates. During decay of a T. weissflogii culture, which served as food for the invertebrates, copepods were increasingly less able to assimilate the methylmercury from the food, while even at advanced stages of decay, amphipods were able to assimilate mercury from their food to a high degree. Finally, fish feeding on copepods assimilated methylmercury more efficiently than inorganic mercury owing to the larger fraction of methylmercury found in the soft tissues of the copepods.     

Feeding rates of planktonic copepods from a tropical sea

The feeding rates of the marine planktonic copepods, Eucalanus subcrassus Giesbrecht, Tortanus gracilis (Brady), Calanopia elliptica (Dana) (both male and female), Temora turbinata (Dana), and Paracalanus aculeatus Giesbrecht (only female) from tropical inshore waters have been studied. Newly hatched Artemia nauplii (for Eucalanus, Tortanus, and Calanopia), Dunaliella (for Temora), and Skeletonema (for Paracalanus) were used as food.Feeding rates were measured for a single individual through successive incubations once or twice a day until death to determine changes in feeding rate after collection. Copepods survived from a few days to three weeks. In general, feeding rates varied from day to day, but were less variable than the differences between day and night rates. In some cases, feeding rate consistently decreased up to the death of the copepod. Daily ration, estimated in terms of percentage body weight, was in the range of 28–329 ‰ Using the results together with the those of other workers, gives the relation between daily ration (Y, % body weight) and body weight of copepods (X; μg dry weight) at 20 °C as, logY = 2.531?0.377 logX.Copepods given Artemia nauplii as food killed more nauplii than were eaten. This phenomenon, tentatively called ‘over-hunting’, is possibly an important feeding behaviour for carnivorous copepods.     

Differential Impacts of Copepods and Cladocerans on Lake Seston,and Resulting Effects on Zooplankton Growth

In an enclosure study in Schöhsee, a small mesotrophic lake in Northern Germany, the impact of copepods and daphniids on the seston community was studied. In general, these two guilds differ in their feeding behaviour. Copepods actively select their food, with a preference for larger particles, whereas most cladocerans are unselective filter-feeders. In this study we investigate how the impact of the two different grazers affects zooplankton growth. We combine results obtained in the laboratory with results measured in situ in the enclosures. Copepods and cladocerans were cultured on seston from enclosures that were inhabited by density gradients of copepods or daphniids. We observed that Daphnia grew faster on seston that was pre-handled by copepods than on seston that was pre-handled by daphniids, and that somatic growth decreased with increasing densities of daphniids in the enclosures. In contrast, we observed no differences in development rates for copepods grown on the different media. The population growth rates of Daphnia in the Daphnia treatments were determined in the enclosures. Growth differences in both somatic- and population growth of Daphnia were correlated to food quality aspects of the seston. In the laboratory we found that Daphnia growth was correlated with several fatty acids. The strongest regression was with the concentration of 20:43 (r ²= 0.37). This particular fatty acid also showed the highest correlation with growth after normalisation of the fatty acids to the carbon content of the enclosures (r ²= 0.33). On the other hand, in the enclosure the population growth correlated most to the particulate nitrogen content (r ²= 0.78) and only to the N:C ratio, when normalised to carbon (r ²= 0.51).     

Does Acartia clausi (Copepoda: Calanoida) use an area-restricted search foraging strategy to find food?

The swimming behavior of Acartia clausi within small aquaria containing phytoplankton was videotaped, along with unfed controls. From these videos, aspects such as swimming speed, distance traveled, and 2D (horizontal, X, and vertical, Z) headings were then measured from over 75000 X Z positional data in order to quantify their swimming functional response. There was generally no significant difference in swimming behavior between different food levels, although there was a large difference between feeding and unfed controls. Because feeding bouts moved the copepods a short distance at a slow speed, net vertical displacement decreased as feeding-bout frequency increased. Non-feeding individuals showed longer periods of sinking, interspersed with longer-distance, high-speed jumps, which displaced them farther distances than filter-feeding copepods over the same time interval. For both fed and unfed control copepods, a 1D unbiased random walk was adequate to describe their displacement over these very short time periods (10 s). This behavior is consistent with an area-restricted search foraging strategy. The net vertical displacements predicted by combining the experimental data with a 1D random walk model suggest that A. clausi would be retained within even small 10 cm phytoplankton patches long enough to fill their gut. However, copepods outside of phytoplankton patches must rely on other means to find patches of food greater than 50 cm apart. Potentially, knowledge of a copepod's swimming functional response could enable predictions about the typical spatial and temporal patchiness of its food in situ.     

The food and feeding habits of the silver pomfret, Pampus argenteus (Euphrasen), in Kuwait waters

The food habits of the silver pomfret, Pampus argenteus (Euphrasen 1788), in Kuwait waters were investigated by examining the stomach contents of 738 specimens collected from May 1996 to April 1997. Their diet consisted of a broad spectrum of food types, but Crustacea were dominant, with copepods and their eggs constituting 39 % other non-copepod Crustacea constituted 16 %. The next major food group was Bacillariophyta (21 %), followed by Mollusca (11 %), fish scales (10 %) and, finally, fish eggs and larvae (3 %). In summer the species fed on a wider variety of food items than in winter. Copepods, other non-copepod crustaceans, and molluscs tended to occur in the stomachs in higher frequencies with an increase in P. argenteus size (up to 18.5–20.4 cm), while the bacillariophytes tended to increase in stomachs at fish sizes between 22.5–24.5 cm. Analysis of monthly variations in stomach fullness indicated that feeding intensity fluctuated throughout the year, with a low during August and September, corresponding to the spawning period.     

Experimental evidence of the grazing impact of <Emphasis Type="Italic">Boeckella poppei</Emphasis> on phytoplankton in a maritime Antarctic lake

The zooplankton biomass of Lake Boeckella (Hope Bay, Antarctic Peninsula) is strongly dominated by the calanoid copepod Boeckella poppei Mrázek. This work analyses the grazing impact of this copepod on the two dominant fractions of phytoplankton, pico- and nanoplankton, and on the bacterioplankton. By means of in-situ experiments using microcosms, the following hypotheses were tested: (a) the early stages of the copepod mainly graze on phytoplankton; (b) the pre-adult and adult stages graze on phytoplankton and benthic algae. Copepods were separated into two groups of maturity: early stages, and pre-adult and adult stages. The following treatments were performed: (1) only nano- and picoplankton, (2) nano- and picoplankton+periphyton, and (3) only periphyton, for each one of the two copepod maturity groups, and (4) control (without copepods). The variation in nano-phytoplankton density was analysed after 2 and 4 days. The results determined a significant grazing on the nano-phytoplankton fraction in all microcosms containing copepods of both maturity groups (P<0.01). The effect on the nano-phytoplankton was greater when the copepods did not have another source of food (P<0.01). No significant differences between the maturity groups were observed (P>0.05). We also conclude that the copepods use the periphyton as an alternative source of food, which was corroborated by the analysis of gut content. In general, no significant differences among treatments were recorded for the pico-sized fraction (pico-phytoplankton and bacterioplankton), which would suggest that no direct grazing on this fraction exists.