Is diatom size selection by harpacticoid copepods related to grazer body size?

Copepods are known as important consumers of primary production and are eaten by larger animals. They therefore form a main link to higher trophic levels. While feeding pathways and specificity of planktonic copepods have been well studied, the selectivity of the benthic harpacticoid copepods is far less documented. A better knowledge of the functional ecology of harpacticoids as important grazers on primary producers may have consequences for the re-evaluation of basic energy flow in benthic ecosystems.We tested whether size selectivity for diatoms exists in harpacticoid copepods. We hypothesized that size selectivity of harpacticoid copepod species is strongly related to body size. Because of morphological constraints, we expected smaller copepods to prefer smaller diatoms while larger copepods should be able to consume both small and large diatoms. We tested this hypothesis in four harpacticoid copepod species of varied body size: Tigriopus brevicornis, Harpacticus obscurus, Amphiascus minutus and Paramphiascella fulvofasciata. As food source we used two ¹³C labelled strains of the benthic diatom Seminavis robusta with a four-fold difference in cell biovolume.Three out of four harpacticoid species showed size selectivity: H. obscurus and A. minutus preferred the larger Seminavis cells, while P. fulvofasciata selected the smaller Seminavis cells. Based on monoclonal treatments, there was no clear preference found for T. brevicornis although there was a small preference for large cells in the mixed treatments. Except for P. fulvofasciata, all species showed a lower uptake when offered the mixed diet (both small and large cells). Although most species showed a size selectivity, our results suggest that this selectivity was not related to their body size. However, the only species that ate significantly more of small diatoms was characterised by comparatively small mouthparts in relation to its body size.     

Food patch size, food concentration and grazing efficiency of the harpacticoid Paramphiascella fulvofasciata (Crustacea, Copepoda)

Harpacticoid copepods are known as important grazers on primary producers. The underlying factors for their food selectivity and grazing efficiency are however far from well known. For instance, their patchy distribution in the marine environment is well documented but how meiofaunal organisms cope with the spatial distribution/accessibility of the available food resources is less clear.In the present study a laboratory experiment was conducted to test the grazing efficiency of Paramphiascella fulvofasciata (Copepoda, Harpacticoida) on the epipelic diatom Seminavis robusta applied in recipients of different area and in various concentrations. Diatoms were enriched in the stable isotope ¹³C in order to trace food uptake and copepods were left to graze for 4 days.We found that the grazing efficiency of P. fulvofasciata was diatom concentration-dependent. A lower diatom uptake at lower diatom densities illustrated this clear functional response. On the contrary, there was no significant effect of the area per se where the copepods could graze upon. The lack of a significant effect of area is mainly due to the high variability in uptake that was recorded in some treatments. Although P. fulvofasciata is a very motile copepod, known as endobenthic and epibenthic species, it was able to concentrate on food uptake at the bottom of the experimental unit as there was no significant difference in uptake between treatments with different water heights in the units. In addition, it was found that a diatom concentration of about 140 000 cells/cm² favours egg production of P. fulvofasciata.     

Effect of food preservation on the grazing behavior and on the gut flora of the harpacticoid copepod Paramphiascella fulvofasciata

Harpacticoids owe their ‘reputation’ as primary consumers in aquatic food webs to their substantial grazing on diatoms, thus assuring an efficient energy flow to higher trophic levels. Due to the complex feeding behavior of harpacticoids, the nature and dynamics of diatom-harpacticoid trophic interactions remain poorly understood. In addition, there is a growing interest from aquaculture industry in mass-culturing harpacticoids with algal foods but the labor costs of maintaining algal stock cultures are high. This study focuses on the palatability of preserved diatoms for copepods and considers the possible role of bacterially mediated effects on diatom food uptake.The grazing of Paramphiascella fulvofasciata on a preserved freeze-dried diatom diet was tested and compared to the grazing on fresh cells. P. fulvofasciata assimilated the preserved diet, but assimilation of fresh cells was higher. When both cell types were mixed, no selective feeding was observed. Community fingerprinting of the bacteria associated with diatoms and fecal pellets suggests that the copepod gut flora was modified depending on the food source. Furthermore, the results suggest that the egestion of gut bacteria enriches the microenvironment and this can have an additional influence on the feeding behavior of the copepod.Experimental research using preserved foods must take into account that copepod grazing assimilations of fresh foods are likely to be significantly higher. Yet, the stated high assimilation of the mixed diet, encourages further exploration of the application of preserved ‘balanced’ foods for harpacticoid mass-culturing.     

Differential effects of nutrient-limited primary production on primary, secondary or tertiary consumers

Nutritional imbalances between predator and prey are the rule rather than the exception at the lower end of food webs. We investigated the role of different grazers in the propagation of nutritionally imbalanced primary production by using the same primary producers in a three-trophic-level food chain and a four-trophic-level food chain experimental setup. The three-trophic-level food chain consisted of a classic single-cell primary producer (Rhodomonas salina), a metazoan grazer (the copepod Acartia tonsa) and a top predator (the jellyfish Gonionemus vertens), while we added a protozoan grazer (Oxyrrhis marina) as primary consumer to the food chain to establish the four-trophic-level food chain. This setup allowed us to investigate how nutrient-limitation effects change from one trophic level to another, and to investigate the performance of two components of our experimental food chains in different trophic positions. Stoichiometry and fatty acid profiles of the algae showed significant differences between the nutrient-depleted [no N and no P addition (?P), respectively] and the nutrient-replete (f/2) treatments. The differences in stoichiometry could be traced when O. marina was the first consumer. Copepods feeding on these flagellates were not affected by the nutritional imbalance of their prey in their stoichiometry, their respiration rates nor in their developmental rates. In contrast, when copepods were the primary consumer, those reared on the ?P algae showed significantly higher respiration rates along with significantly lower developmental rates. In neither of our two experimental food chains did the signals from the base of the food chains travel up to jelly fish, our top predator.     

Trophic structure of common marine species in the Bohai Strait,North China Sea,based on carbon and nitrogen stable isotope ratios

Trophic relationships among common coastal species in the Bohai Strait, North China Sea, were investigated in this study using stable isotope ratios of carbon (¹³C/¹²C) and nitrogen (¹⁵N/¹⁴N) as tracers. During the research cruises of this study, 16 species of invertebrates and 18 species of fishes were collected, as were various food sources (phytoplankton, macroalgae, and sediment). The carbon and nitrogen isotope ratios of all collected samples were measured and used for trophic level analyses. The results showed that the grazers Oregonia gracilis, Notoacmea schrenckii, Chlorostoma rustica, and Chelon affinis, preferentially consumed Zostera marina (p < 0.01), whereas Anthocidaris crassispina preferred Ulva conglobata (p < 0.01). Two trophic models based on nitrogen isotope ratios were built to identify the trophic level of each species. The model that combined food sources was more appropriate than the model that used a single primary producer to identify the relative trophic positions of primary consumers. Scombermorus niphonius, an important fishery resource, was at the top trophic level. The seastar Asterias amurensis was at the highest level among the invertebrates and directly threatened production of shellfish. Based on the trophic level and food source relationships identified in this study, we gave some advice for ecological restoration, such as optimizing the structure of food source distribution, limiting seastar numbers, and improving the applicability of habitat for fishery species.     

Copepod Population-Specific Response to a Toxic Diatom Diet

Diatoms are key phytoplankton organisms and one of the main primary producers in aquatic ecosystems. However, many diatom species produce a series of secondary metabolites, collectively termed oxylipins, that disrupt development in the offspring of grazers, such as copepods, that feed on these unicellular algae. We hypothesized that different populations of copepods may deal differently with the same oxylipin-producing diatom diet. Here we provide comparative studies of expression level analyses of selected genes of interest for three Calanus helgolandicus populations (North Sea, Atlantic Ocean and Mediterranean Sea) exposed to the same strain of the oxylipin-producing diatom Skeletonema marinoi using as control algae the flagellate Rhodomonas baltica. Expression levels of detoxification enzymes and stress proteins (e.g. glutathione S-transferase, glutathione synthase, superoxide dismutase, catalase, aldehyde dehydrogenases and heat shock proteins) and proteins involved in apoptosis regulation and cell cycle progression were analyzed in copepods after both 24 and 48 hours of feeding on the diatom or on a control diet. Strong differences occurred among copepod populations, with the Mediterranean population of C. helgolandicus being more susceptible to the toxic diet compared to the others. This study opens new perspectives for understanding copepod population-specific responses to diatom toxins and may help in underpinning the cellular mechanisms underlying copepod toxicity during diatom blooms.     

Fish predation regimes modify benthic diatom community structures: Experimental evidence from an in situ mesocosm study

Diatoms are important primary producers in shallow water environments. Few studies have assessed the importance of biological interactions in structuring these communities. In the present study, benthic diatom community structure in relation to manipulated food webs was assessed using in situ mesocosms, whereby predator‐free environments and environments comprising two different fish species were assessed. Zooplankton abundance, settled algal biomass and the diatom community were monitored over a 12‐day period across each of the three trophic scenarios. Differences among treatments over time were observed in zooplankton abundances, particularly copepods. Similarly, the benthic diatom community structure changed significantly over time across the three trophic treatments. However, no differences in total algal biomass were found among treatments. This was likely the result of non‐diatom phytoplankton contributions. We propose that the benthic diatom community structure within the mesocosms was influenced by trophic cascades and potentially through direct consumption by the fish. The study highlights that not only are organisms at the base of the food web affected by predators at the top of the food web, but that predator identity is potentially an important consideration for predator–prey interaction outcomes with consequences for multiple trophic levels.     

Food web analysis of an eelgrass (Zostera marina L.) meadow and neighbouring sites in Mitsukuchi Bay (Seto Inland Sea,Japan) using carbon and nitrogen stable isotope ratios

The contribution of benthic microalgal production has been compared both within and outside a coastal eelgrass (Zostera marina L.) meadow. Carbon and nitrogen stable isotope ratios of suspended particulate organic matter (POM), epiphytic and epilithic organic matter (EOM), leaves of Z. marina (inside the meadow only) and two secondary consumer species (small crustaceans and fish) were measured inside and outside a meadow in Mitsukuchi Bay, Northwest Seto Inland Sea, Japan. Inside the meadow, primary producers (epiphyton) and consumers showed higher δ¹³C signatures than outside. Primary and secondary consumers inside the meadow were mainly dependent on epiphyton on the leaves of Z. marina, while consumer species outside the meadow were basically dependent on epilithon.     

Temperature-induced changes in fatty acid dynamics of the intertidal grazer Platychelipus littoralis (Crustacea,Copepoda, Harpacticoida): Insights from a short-term feeding experiment

Dietary lipids, and in particular the essential fatty acids (EFA), EPA (20:5ω3) and DHA (22:6ω3), guarantee the well-being of animals and are recognized for their potential bottom-up control on animal populations. They are introduced in marine ecosystems through primary producers and when grazed upon, they are consumed, incorporated or modified by first-level consumers. As the availability of EFA in the ecosystem is affected by ambient temperature, the predicted rise in ocean temperature might alter the availability of these EFA at the basis of marine food webs. Despite the FA bioconversion capacity of certain benthic copepod species, their lipid (FA) response to varying temperatures is understudied. Therefore, the temperate, intertidal copepod Platychelipus littoralis was offered a mono and mixed diatom diet at 4, 15 °C (normal range) and at 24 °C (elevated temperature) to investigate the combined effects of temperature and resource availability on its FA content and composition. P. littoralis showed a flexible thermal acclimation response. Cold exposure increased the degree of FA unsaturation and the EPA%, and induced a shift towards shorter chain FA in the copepod’s membranes. Furthermore, a mixed diet reduced the impact of heat stress on the copepod’s membrane FA composition. Temperature affected the trophic transfer of EPA and DHA differently. While dietary resources could fully compensate for the temperature effects on total lipid and EPA content in the copepods, no such counterweigh was observed for the DHA dynamics. Heat stress lowered the DHA concentration in copepods regardless of the resources available and this implies negative effects for higher trophic levels.     

Are the diatoms Navicula sp. and Thalassiosira fluviatilis suitable to be fed to the benthic harpacticoid copepod Tisbe biminiensis?

The objective of this study was to compare the influence of the diets of two diatoms, Navicula sp. (benthic) and Thalassiosira fluviatilis (planctonic), on the development, fecundity and survival of the harpacticoid copepod Tisbe biminiensis. In order to determine the optimal concentration of food, 35 egg-bearing females were submitted to six algal concentrations and controls (without food). After 24 h, the content of the recipients was fixed with 4% formalin and then fecal pellets produced by each female were counted and measured. The larval development was studied by surveying 50 nauplii on each diet individually until the adult stage, at intervals of 6 h. The cast exoskeletons were removed to count the number of segments and for measurement. The fecundity was obtained counting the naupliar production every 48 h of groups containing 10 females in different algal concentrations in both diets. The diet influence on fecundity was tested by submitting four groups of 10 females fed on optimal algal concentrations based on the fecal pellet experiments. In the Navicula sp. concentration of 0.4 μg Chl-a/ml, considered to be optimal for fecal pellets production, the diatoms were shown toxic, resulting in a low survival rate and inhibiting the egg production of copepods. The optimal concentration considering fecundity was estimated to be 0.1 μg Chl-a/ml for both diets. Copepods fed on Navicula sp. presented a faster development rate and higher naupliar production compared to copepods fed on T. fluviatilis. Size and survival did not vary significantly among diets. The algal concentration interfered significantly in the reproductive success of females. Both very low and very high algal concentrations reduced reproductive success. Concluding, the benthic diatom Navicula sp. was more favorable to the copepod T. biminiensis than the planktonic diatom T. fluviatilis.     

Short-term impacts of polyunsaturated aldehyde-producing diatoms on the harpacticoid copepod, Tisbe holothuriae

Diatoms that produce toxic oxylipins can be detrimental to the reproductive success of aquatic invertebrates. Despite the potential importance of these toxins in shaping aquatic ecosystems, marine studies to date have focused almost exclusively on planktonic calanoid copepods. The current work examines the response of the benthic harpacticoid copepod, Tisbe holothuriae, to direct exposure to diatom-derived oxylipins and the short-term impact of oxylipin-producing diatom diets on reproductive success. The most toxic oxylipin was the polyunsaturated aldehyde (PUA) 2E,4E-decadienal with an LD₅₀ of 9.3 μM for T. holothuriae nauplii. The least tolerant life-stage was the nauplius followed by adult males then adult females. Short-term exposure to PUA-producing diatoms (Skeletonema marinoi and Melosira nummuloides) in maternal diets had no significant impact on reproductive success compared with non-PUA-producing diets (Skeletonema costatum, Navicula hanseni, Phaeodactylum tricornutum and Tetraselmis suecica). The PUA producers had no negative impact on the survival and development of naupliar stages to adulthood. T. holothuriae expresses a higher degree of tolerance to PUA-producing diatoms than many planktonic calanoids, possibly reflecting a degree of adaptation to higher stress levels associated with the benthos. This is the first study to investigate the reproductive responses of harpacticoid copepods feeding on known PUA-producing diatoms.     

The harpacticoid copepod Tisbe holothuriae is resistant to the insidious effects of polyunsaturated aldehyde-producing diatoms

Numerous species of diatoms liberate oxylipins including polyunsaturated aldehydes (PUAs) in response to cellular damage such as may occur during grazing. PUAs are cyto- and genotoxic and negatively disrupt reproductive processes in copepods, their principal grazers, although experimental evidence would suggest that the grazer response may be species specific. The reproduction of the benthic harpacticoid copepod Tisbe holothuriae was compared over two generations. Copepods were reared using four diet treatments: PUA-producing diatom strains Skeletonema marinoi (Adriatic Sea Isolate FE6) and Melosira nummuloides (CCAP 1048/6); and non-PUA-producing diatom strains Phaeodactylum tricornutum (CCAP 1052/A) and S. marinoi (Seasalter (Walney) Ltd). Life tables were generated for each treatment using measured reproductive parameters and the net reproductive rate (R₀) calculated. No significant differences were observed between the individual reproductive parameters of T. holothuriae fed PUA-producing diatoms compared to those fed non-PUA-producing diatoms although diets of P. tricornutum resulted in some decreases in individual reproductive parameters in the second generation. There were no significant differences in the R₀ values between the four tested diets. These observations indicate that T. holothuriae exhibits a tolerance of known PUA-producing diatom diets that has not been similarly demonstrated in pelagic calanoid copepods. Harpacticoid copepods may have a greater capacity to detoxify diatom oxylipins than their planktonic calanoid counterparts.     

The trophic importance of algal turfs for coral reef fishes: the crustacean link

On coral reefs, the epilithic algal matrix (EAM) is widely recognised as an important resource for herbivorous and detritivorous fishes. In comparison, little is known of the interaction between benthic carnivores and the EAM, despite the abundance of Crustacea within the EAM. The trophic importance of the EAM to fishes was investigated in Pioneer Bay, Orpheus Island, Great Barrier Reef. Fish densities were quantified using visual and clove oil censuses, and gut content analyses conducted on abundant fish species. Crustaceans were found to be an important dietary category, contributing between 49.5 and 100 % of the gut contents, with harpacticoid copepods being the dominant component. Of the benthic carnivores, the goby Eviota zebrina was found to consume the most harpacticoids with a mean of 249 copepods m^?2 day^?1. This represents approximately 0.1 % of the available harpacticoid population in the EAM. In a striking comparison, herbivorous parrotfishes were estimated to consume over 12,000 harpacticoids m^?2 day^?1, over 27 times more than all benthic carnivores surveyed, representing approximately 5.3 % of the available harpacticoid copepod population each day. The high consumption of harpacticoid copepods by benthic carnivores and parrotfishes indicates that harpacticoids form an important trophic link between the EAM and higher trophic levels on coral reefs.     

Grazers and Phytoplankton Growth in the Oceans: an Experimental and Evolutionary Perspective

The taxonomic composition of phytoplankton responsible for primary production on continental shelves has changed episodically through Earth history. Geological correlations suggest that major changes in phytoplankton composition correspond in time to changes in grazing and seawater chemistry. Testing hypotheses that arise from these correlations requires experimentation, and so we carried out a series of experiments in which selected phytoplankton species were grown in treatments that differed with respect to the presence or absence of grazers as well as seawater chemistry. Both protistan (Euplotes sp.) and microarthropod (Acartia tonsa) grazers changed the growth dynamics and biochemical composition of the green alga Tetraselmis suecica, the diatom Thalassiosira weissflogii, and the cyanobacterium Synechococcus sp., increasing the specific growth rate and palatability of the eukaryotic algae, while decreasing or leaving unchanged both parameters in the cyanobacteria. Synechococcus (especially) and Thalassiosira produced toxins effective against the copepod, but ciliate growth was unaffected. Acartia induced a 4-6 fold increase of Si cell quota in the diatom, but Euplotes had no similar effect. The differential growth responses of the eukaryotic algae and cyanobacteria to ciliate grazing may help to explain the apparently coeval radiation of eukaryophagic protists and rise of eukaryotes to ecological prominence as primary producers in Neoproterozoic oceans. The experimental results suggest that phytoplankton responses to the later radiation of microarthropod grazers were clade-specific, and included changes in growth dynamics, toxin synthesis, encystment, and (in diatoms) enhanced Si uptake.     

Current velocity influences the facilitation and removal of algae by stream grazers

The modification of flows in lotic ecosystems can have dramatic effects on abiotic and biotic processes and change the structure of basal trophic levels. In high-gradient streams, most of the biota are benthic, and decreased flow may homogenize and reduce benthic current velocity, potentially changing stream ecosystem function. Grazing by macroinvertebrates is an important component of stream function because grazers regulate energy flow from primary producers to higher trophic levels. We conducted an experiment to examine how macroinvertebrate grazers facilitated or removed algal biomass across a gradient of benthic current velocity (0–40 cm s^?1). We chose three grazers (Drunella coloradensis, Cinygmula spp., and Epeorus deceptivus) from a montane stream and conducted our experiment using 24 artificial stream channels that had three treatments: no grazers (control), single-grazer, and combined-grazer treatments. In the absence of grazers, algal biomass increased with benthic current velocity. Grazer treatments differed from the control in that more algal biomass was removed at higher velocities, whereas algal accrual was largely facilitated at low velocities. The transition from facilitation to removal ranged from 4.5 to 5.9 cm s^?1 for individual grazer treatments and occurred at 11.7 cm s^?1 for the combined-grazer treatment. Our data suggest that velocity plays a significant role in the facilitation and removal of algae by macroinvertebrate grazers. Additionally, the patterns revealed here could have general implications for algal accrual in systems where flow is reduced.     

Effects of Grazer Presence on Genetic Structure of a Phenotypically Diverse Diatom Population

Studies of predator–prey systems in both aquatic and terrestrial environments have shown that grazers structure the intraspecific diversity of prey species, given that the prey populations are phenotypically variable. Populations of phytoplankton have traditionally considered comprising only low intraspecific variation, hence selective grazing as a potentially structuring factor of both genetic and phenotypic diversity has not been comprehensively studied. In this study, we compared strain specific growth rates, production of polyunsaturated aldehydes, and chain length of the marine diatom Skeletonema marinoi in both grazer and non-grazer conditions by conducting monoclonal experiments. Additionally, a mesocosm experiment was performed with multiclonal experimental S. marinoi populations exposed to grazers at different levels of copepod concentration to test effects of grazer presence on diatom diversity in close to natural conditions. Our results show that distinct genotypes of a geographically restricted population exhibit variable phenotypic traits relevant to grazing interactions such as chain length and growth rates. Grazer presence affected clonal richness and evenness of multiclonal Skeletonema populations in the mesocosms, likely in conjunction with intrinsic interactions among the diatom strains. Only the production of polyunsaturated aldehydes was not affected by grazer presence. Our findings suggest that grazing can be an important factor structuring diatom population diversity in the sea and emphasize the importance of considering clonal differences when characterizing species and their role in nature.     

Quantifying the trophic base for benthic secondary production in the Nakdong River estuary of Korea using stable C and N isotopes

The overall dependence of benthic secondary production on the main primary producers at three different habitats in the Nakdong River estuarine system, Korea, was estimated. Inventories of macrobenthic invertebrate biomass were combined with multiple-isotope-mixing models to evaluate the trophic base, comparing Scirpus triqueter-dominated and Phragmites australis-dominated marshes and bare intertidal flat. The feasible contributions of four main food sources, marsh macrophytes, the microphytobenthos, and riverine and marine suspended particulate organic matter (RPOM and MPOM), to the consumer biomasses were calculated using the isotopic mixing model. After weighting the feasible contributions of food sources to each taxon by the consumer biomass, the resultant values were summed for all the consumers at each habitat to quantify the trophic base of the benthic invertebrate community. Dual-isotope-mixing model calculations verified the varying dependence on those potential food sources among the functional feeding groups. In addition, the dependence on each source of the same functional group varied between bare intertidal and salt-marsh habitats, shifting from a dominance of benthic and pelagic microalgal sources on the former habitat to a mixed food source at the latter habitat. The biomasses of the species comprising each functional group differed among habitats and sampling dates, so that each functional group made a different contribution to the whole benthic community and its basal food source. Given the calculation of the overall dependence of macrozoobenthic community on each food source, our results indicate that the microphytobenthic source dominates (nearly half) the trophic base in all the intertidal habitats of different vegetational compositions. Marsh-macrophyte-derived organic matter and RPOM served as considerable subsidies only to salt-marsh food webs, reflecting the use of the mixed food source by salt-marsh-bed consumers. Conversely, the dominance of MPOM in the total food base was equal to that of the microphytobenthos in the bare intertidal ecosystem but increased during spring−summer in the salt-marsh systems. Our results also suggest that the river discharge concentrated during the summer monsoon does not lead to any shift in trophic base for estuarine secondary production.     

Copepod grazing during an iron-induced diatom bloom in the Antarctic Circumpolar Current (EisenEx): I. Feeding patterns and grazing impact on prey populations

Feeding activity, selective grazing and the potential grazing impact of two dominant grazers of the Polar Frontal Zone, Calanus simillimus and Rhincalanus gigas, and of copepods < 2 mm were investigated with incubation experiments in the course of an iron fertilized diatom bloom in November 2000. All grazers were already actively feeding in the low chlorophyll waters prior to the onset of the bloom. C. simillimus maintained constant clearance rates and fed predominantly on diatoms. R. gigas and the small copepods strongly increased clearance and ingestion of diatoms in response to their enhanced availability. All grazers preyed on microzooplankton, most steadily on ciliates, confirming the view that pure herbivory appears to be the exception rather than the rule in copepod feeding. The grazers exhibited differences in feeding behavior based on selectivity indices. C. simillimus and R. gigas showed prey switching from dinoflagellates to diatoms in response to the phytoplankton bloom. All grazers most efficiently grazed on large diatoms leading to differences in daily losses for large and small species, e.g. Corethron sp. or Thalassionema nitzschioides. Species-specific diatom mortality rates due to grazing suggest that the high feeding activity of C. simillimus prior to and during the bloom played a role in shaping diatom population dynamics.     

An ecological and economic review of meiofauna as food for fish*

After briefly reviewing the evidence that meiofauna are an integral part of the complex food web of invertebrate infaunal and epifaunal organisms, it is shown that, at least in intertidal and shallow subtidal habitats, meiofauna are an important food for the small (30–60 mm) juveniles of some commercially important fishes (flatfish and salmonids) or for fish which are themselves food for commercial species. The evidence that fish feed preferentially on a few species of harpacticoid copepod rather than on the numerically dominant meiofaunal taxa is presented and discussed. It is argued that, in addition to benthic copepods being energetically efficient as food for fishes, it is differences in the activity and availability between major taxa of meiofauna as they relate to the feeding strategies of the fish which go a long way to explaining this apparent general preference of fish for harpacticoid copepods rather than nematodes. From literature estimates of consumption rates, it is concluded that the supply of meiofaunal food to juvenile fish is unlikely to be limiting although this may not be the case in some shallow subtidal muddy sand habitats. Whilst it is recognized that fish predation may influence the magnitude of the seasonal peaks in numbers, predation is not the principal cause of seasonal fluctuations in harpacticoid copepod species abundances. The effects of predation on other aspects of community structure is unknown. Finally, the possible role of meiofauna in mariculture systems is briefly discussed.