A choice chamber experiment on the selection of algae as food and substrata by Nais elinguis (Oligochaeta: Naididae)

SUMMARY.

• 1 A choice chamber experiment was performed in the laboratory to quantify the selection of benthic algae as food and substrata by Nais elinguis.
• 2 Most worms actively chose filamentous algae and/or diatoms as substrata whereas unicellular or colonial chiorophycean algae were discriminated against. Diatoms were selected in preference to latex beads but the worms could not discriminate between living filaments and nylon monofilaments. A thigmotactic response to filaments and a chemotactic response to diatoms may be implicated.
• 3 The worms ingested unicellular algae at a greater rate than colonial and filamentous algae. The rates at which the algae were ingested were negatively correlated with the lengths of the algae. A filament 200 μm long was the largest item ingested, representing about 63% of the length of the pharynx of the worm.
• 4 Ivlev's electlvity indices were positive when the rates at which the algae were ingested exceeded about 20 individuals h^-1. The index was negatively correlated with the lengths of the algae, but the time periods spent by the worms in association with the algae were unrelated to the electivity index.
• 5 The data confirmed previous observations on the selection of algae as food and substrata by Nais elinguis in a natural habitat.

     

Effects of unicellular and colonial forms of toxic Microcystis aeruginosa from laboratory cultures and natural populations on tropical cladocerans

Three life-table experiments, two growth experiments and one feedinginhibition experiment, were performed to study the effects of the toxiccyanobacterium Microcystis aeruginosa on the cladoceransofa tropical lagoon (Jacarepaguá Lagoon, Rio de Janeiro, Brazil).Differentexperimental designs were used to estimate toxic effects of both field samplesand laboratory cultures of Microcystis aeruginosa oncladoceran life history parameters and juvenile growth rates. Effects ofnutritional deficiency could be distinguished from toxic effects in experimentswhere green algae in high carbon concentration were mixed withMicrocystis. Our results show that natural assemblages ofMicrocystis caused much less pronounced toxic effects thanlaboratory cultures and that unicellular forms were more toxic than colonialforms, even though both contained high concentrations of toxins. One possibleexplanation is that colonies were too large to be ingested by the smallMoina micrura and Ceriodaphniacornuta. Feeding inhibition by single cells and small colonies seemsto be another mechanism that contributes to the harmful effects ofMicrocystis on cladocerans, both in the laboratory and inthe field. Thus, caution is needed in extrapolating results from the laboratoryto the field. We did find, however, that toxic algae in natural seston caninhibit growth and reproduction of native cladocerans populations.     

Testing for resistance of pelagic marine copepods to a toxic dinoflagellate

With few exceptions, the evolutionary consequences of harmful algae to grazers in aquatic systems remain unexplored. To examine both the ecological and evolutionary consequences of harmful algae on marine zooplankton, we used a two-fold approach. In the first approach, we examined the life history responses of two geographically separate Acartia hudsonica (Copepoda Calanoida) populations reared on diets containing the toxic dinoflagellate Alexandrium fundyense . One copepod population was from a region, Casco Bay, Maine, USA, that has experienced recurrent blooms of highly toxic Alexandrium spp. for decades; whereas the other population from Great Bay, New Jersey, USA, has never been exposed to toxic Alexandrium blooms. The life history experiment demonstrated that when the copepod population from New Jersey was reared on a diet containing toxic A. fundyense it exhibited lower somatic growth, size at maturity, egg production and survival than the same population reared on a diet without toxic A. fundyense . In contrast, toxic A. fundyense did not affect the life-history traits of the Maine population. Fitness, finite population growth rate (), was significantly reduced in the New Jersey population, but not in the Maine population. These results are consistent with the hypothesis of local adaptation (resistance) of the historically exposed copepod population to the toxic dinoflagellate. In the second approach, we further tested the resistance hypothesis with a laboratory genetic selection experiment with the naïve New Jersey copepod population exposed to a diet containing toxic A. fundyense. This experiment demonstrated that the ingestion and egg production of adult females of naïve copepods fed A. fundyense improved after three generations of being reared on a diet containing the toxic dinoflagellate. The results of the present study have important implications for understanding how grazer populations may respond to the introduction of toxic algae to their environment, and suggest that grazer resistance may be a feedback mechanism that may lead to bloom control.Co-ordinating editor: Hurst     

Prorocentrum minimum(clone Exuv) is nutritionally insufficient, but not toxic to the copepod Acartia tonsa

This study tested whether the dinoflagellate Prorocentrum minimum is nutritionally insufficient or toxic to the copepod Acartia tonsa. Experiments were carried out with adult female A. tonsa and the P. minimum clone Exuv, both isolated from Long Island Sound. Initially, the functional and numerical responses of A. tonsa feeding on exponentially growing P. minimum cells were characterized. These experiments revealed that A. tonsa readily ingested P. minimum cells, up to the equivalent of 200% of body carbon day⁻¹, but egg production was relatively low, with a maximum egg production rate of 22% of body carbon day⁻¹. Hence, the egg production efficiency (egg carbon produced versus cell carbon ingested) was low (10%). In a separate experiment, ingestion and egg production rates were measured as a function of food concentration with cells in different growth stages (early-exponential, late-exponential/early-stationary, and late-stationary growth phase) to simulate conditions during a bloom. There was no indication that cells in the stationary phase resulted in lower ingestion or egg production rates relative to actively growing cells. Egg hatching success remained high (>80%) and independent of the cell growth phase. In a third experiment specifically designed to test the hypothesis that P. minimum is toxic, ingestion, egg production and egg hatching success were measured when females were fed mixtures of P. minimum and the diatom Thalassiosira weissflogii, but in which total food concentration was held constant and the proportion of P. minimum in the mixed diet varied. A. tonsa readily ingested P. minimum when it was offered in the mixed diet, with no detrimental effects on egg production or egg hatching observed. Supplementing P. minimum with T. weissflogii increased both the egg production rate and the egg production efficiency. It is concluded that P. minimum is nutritionally insufficient, but not toxic to A. tonsa. Finally, it is estimated that in the field grazing by A. tonsa is approximately equivalent to 30% of the maximum daily growth rate of P. minimum. Hence, copepod grazing cannot be ignored in field and modeling studies of the population dynamics of P. minimum.     

Influence of phytoplankton fractions on growth and reproduction of tropical cladocerans

The influence of two seston fractions, < 20 µm (nanoplankton) and 20 µm (microplankton), on growth and reproduction of cladoceran species with different sizes, from Lake Monte Alegre, was evaluated through individual growth and life table experiments. Size, shape and other features of the algae in the fractions were described. The procedure of filtering lake water through a 20 µm net for seston fractionation caused mutual contamination. The smallest cladoceran species, Ceriodaphnia cornuta Sars and Moina micrura Kurz, produced larger clutch sizes and exhibited higher intrinsic rates of population growth (r) in the nanoplankton, despite contamination of their food by inedible algae. The largest species, Simocephalus mixtus Sars, produced larger clutch sizes in the microplankton. There were no differences in juvenile biomass growth between treatments for M. micrura and Daphnia gessneri Herbst, but lower value of the exponential growth rate (g) in the microplankton was significant for M. micrura. Fecundity (eggs/total female) of M. micrura was significantly lower in the microplankton, while D. gessneri did not reproduce in this fraction, at the end of growth experiments. Spines, colonies, cenobium, filaments, hard cell wall, and gelatinous sheaths, represented constraints to cladoceran reproductive performance, besides algae size. Microplankton contamination by nanoplanktonic algae, in the experiments, probably minimized the negative effect of inedible algae. Nanoplankton was more suitable for the smallest species and microplankton for the largest one.     

Selective ingestion of phytoplankton by the bivalvesMytilus edulis L. andCerastoderma edule (L.)

Blue mussles (Mytilus edulis) and cockles (Cerastoderma edule) were collected in the Oosterschelde estuary (SW Netherlands) and exposed to diets consisting of the diatomPhaeodactylum tricornutum and silt. Algal concentrations were kept constant (20.10³ cells.ml^–1), and silt concentrations were varied between 3–80 mg.l^–1 (mussel experiments) and 20–120 mg.l^–1 (cockle experiments). Chlorophyll-a concentrations in the pseudofaeces were reduced compared to the diets, indicating selective ingestion of the algae. It was estimated that the mussels increase the ingestion of chlorophyll-a 2.0 times, and the cockles increase the chlorophyll-a ingestion 2.8 times, compared to the ingestion rate without selection. The selection coefficients were not affected by the SPM concentrations used. Phaeophytin-a concentrations in the faeces showed an increase as a consequence of the digestion of the ingested algae. Digestive efficiencies of chlorophyll-a varied between 36–92% The digestive efficiencies decreased with increasing SPM concentrations.Communication no. 541 of the Delta Institute for Hydrobiological Research.     

Reduction of the filtration rate of Daphnia galeata by dissolved photosynthetic products of edible phytoplankton

The filtration rate of Daphnia galeata was determined in in situ experiments in Bautzen Reservoir and in laboratory experiments, where daphnids were exposed to filtrates that previously contained either natural phytoplankton or cultured eukaryotic algae (Scenedesmus obliquus or Asterionella formosa), respectively. Individual filtration rate (FR) was measured using fluorescent beads, taking into account ingested beads in the gut only. Compared to heated control treatments (100 °C), dissolved compounds released by the nutritious cultured algae during the preconditioning phase or by the natural phytoplankton assemblages from Bautzen Reservoir strongly reduced the filtration rate of D. galeata (down to 60%). Heating deactivated these dissolved compounds. A significant correlation was found between primary production measured in situ and the reduction of FR in the filtrate of reservoir water, indicating that extra-cellular products released during photosynthesis triggered the reduction of the filtration rate. The ratio of ingested to collected beads was used to quantify the proportion of food, which was not only collected but passed the mouth of D. galeata. The ratio of ingestion to collection was compared between filtered and unfiltered reservoir water both media identical with respect to the concentration of dissolved compounds, whereas other factors (e.g. food concentration, temperature, filtration rate) were different. The changes in this ratio between filtered and unfiltered reservoir water suggest that D. galeata is capable of a chemosensory control of the ingestion behaviour by detecting external metabolites.     

Survival of seaweeds ingested by three species of tropical sea urchins from Brazil

Three species of sea urchins, Echinometra lucunter, Lytechinus variegatus, and Arbacia lixula, common on the southeastern coast of Brazil, were selected for experiments on seaweed survival after passage through their digestive tract. L. variegatus and E. lucunter have more than 80% of their diet composed of algae. C. 24% of the species of algae present in the digestive tract survived. A. lixula has only 29% of its diet composed by algae, but the survival rate of the algae is c. 50%.Although Rhodophyta represent 51% of the ingested algae, their survival rate in the faeces is only 19%, while the Chlorophyta, with a survival rate of 63%, comprise only about 26% of the diet. Most species of algae that survived ingestion were r-strategists.     

Carbon budgets for a phytoplanktivorous fish fed three different unialgal populations

Summary The filter feeding blue tilapia, Tilapia aurea, was fed three different algae. Blue tilapia ingestion of two green algae, Chlamydomonas sp. and Ankistrodesmus falcatus and the filamentous blue-green alga, Anabaena flos-aquae, ranged from 21%–89% of the available cells. There were significant differences in the assimilation of algal carbon by the fish depending on the alga fed; A. flos-aquae was the easiest to assimilate (83%). The fish respired significantly less of the Chlamydomonas sp. ingested carbon (15%). The gross growth efficiency of fishes fed either green alga was not significantly different (22%–24%), but these efficiencies were significantly less than the gross growth efficiency of fish fed A. flos-aquae (46%). The carbon budgets for fish feeding on the green algae were similar to that constructed from the literature for a congener fed a mixed algae diet. However, the assimilation component of the budget for blue tilapia fed A. flos-aquae was 2 times greater than that of the literature budget.     

A note on the nutrition of Stylaria lacustris (Oligochaeta: Naididae)

Stylaria lacustris (L.) was offered ¹⁴C-labelled algae. The ingestion rate of diatoms was 0.79 g carbon/hour at 19°C. Diatoms (Nitzschia actinastroides) were well assimilated, green algae (Scenedesmus acuminatus) were practically not assimilated. The less assimilable algae are ingested at higher rates, indicating a regulation mechanism. Biomass doubling time was estimated to be about 3.6 days at 19°C.     

How<Emphasis Type="Italic"> Daphnia</Emphasis> copes with excess carbon in its food

Animals that maintain near homeostatic elemental ratios may get rid of excess ingested elements from their food in different ways. C regulation was studied in juveniles of Daphnia magna feeding on two Selenastrum capricornutum cultures contrasting in P content (400 and 80 C:P atomic ratios). Both cultures were labelled with ¹⁴C in order to measure Daphnia ingestion and assimilation rates. No significant difference in ingestion rates was observed between P-low and P-rich food, whereas the net assimilation of ¹⁴C was higher in the treatment with P-rich algae. Some Daphnia were also homogeneously labelled over 5 days on radioactive algae to estimate respiration rates and excretion rates of dissolved organic C (DOC). The respiration rate for Daphnia fed with high C:P algae (38.7% of body C day^-1) was significantly higher than for those feeding on low C:P algae (25.3% of body C day^-1). The DOC excretion rate was also higher when animals were fed on P-low algae (13.4% of body C day^-1) than on P-rich algae (5.7% of body C day^-1) . When corrected for respiratory losses, total assimilation of C did not differ significantly between treatments (around 60% of body C day^-1). Judging from these experiments, D. magna can maintain its stoichiometric balance when feeding on unbalanced diets (high C:P) primarily by disposing of excess dietary C via respiration and excretion of DOC.     

Low temperature constrains growth rates but not short-term ingestion rates of Antarctic ciliates

Low environmental temperature is a major factor affecting the feeding activities, growth rates, and growth efficiencies of metazooplankton, but these features are poorly characterized for most protistan species. Laboratory experiments were conducted to examine the growth and ingestion rates of cultured herbivorous Antarctic ciliates. Three ciliates fed several algal species individually at 0 °C exhibited uniformly low growth rates (<0.26 day^?1), but the algae varied substantially in their ability to support ciliate growth. Specific ingestion rate (prey biomass consumed per unit ciliate biomass per unit time) was strongly affected by ciliate physiological state (starved vs. actively growing). Starved cells ingested many more prey than cells in balanced growth during short-term (minutes-to-hours) experiment but did not grow faster, indicating temperature compensation of ingestion rate but not growth rate. Field experiments were also conducted in the Ross Sea, Antarctica, to characterize the feeding rates of ciliates in natural plankton assemblages. Specific ingestion rates of two dominant ciliates were an order of magnitude lower than rates reported for temperate ciliates, but estimated rates were strongly affected by prey abundance. Our data indicate that short-term ingestion rates of Antarctic ciliates were not constrained by low environmental temperature although overall growth rates were, indicating the need for caution when designing experiments to measure the ingestion rates of these species at low environmental temperature. We present evidence that artifacts arising from estimating ingestion in short-term experiments may lead to errors in estimating feeding impact and growth efficiencies that are particularly large for polar protists.     

Effects of toxic and non-toxic cyanobacteria on the life history of tropical and temperate cladocerans

1. This study compares the effects of four toxic strains of Microcystis aeruginosa on tropical and temperate Cladocera. Survival was tested in acute toxicity experiments using Microcystis alone or in mixtures with the edible green algae Ankistrodesmus falcatus. The effect of chronic exposure on population growth was estimated in life‐table experiments by varying the proportion of Microcystis and the green alga. Nutritional deficiency was assessed using a non‐toxic cyanobacterium in a zooplankton growth experiment. Feeding inhibition was tested using a C‐labelled green alga as a tracer in mixtures with toxic Microcystis.
2. Toxicity varied consistently between Microcystis strains, while sensitivity varied consistently between cladoceran species. However, no relationship was found between sensitivity and geographical origin or cladoceran body size. Two small‐bodied cladocerans from the same tropical lake, Ceriodaphnia cornuta and Moinodaphnia macleayi, were the least sensitive and most sensitive species, respectively.
3. Surprisingly, two small tropical cladocerans survived longer without food than did three large Daphnia species and a third small tropical species.
4. Each of the three tropical Microcystis strains strongly reduced the population growth rate (little ‘r’) and reproductive output of each cladoceran, this reduction being proportional to the percentage of toxic cells in the diet.
5. As the sole food source, the non‐toxic cyanobacterium, Synechococcus elongatus, supported poor growth in M. macleayi. The nutritional deficiency was overcome when Synechococcus was mixed with either Ankistrodesmus or an emulsion rich in omega‐3 fatty acids.
6. Microcystis inhibited the feeding rate of two cladocerans, even when it comprised only 5% of a mixture with the green algae A. falcatus.
7. Differences in sensitivity to the toxic cyanobacterium appear to be associated with differences in life history between the cladoceran species rather than differences between tropical and temperate taxa. Slow‐growing species that are resistant to starvation appear less sensitive to toxic Microcystis than fast‐growing species, which also tend to die more quickly in the absence of food.     

Ingestion of live filamentous diatoms by the Great Lakes amphipod,Diporeia sp.: a case study of the limited value of gut contents analysis

Individuals of the Great Lakes amphipod, Diporeia sp. (formerly named Pontoporeia hoyi) were collected from a 45-m deep station in southeastern Lake Michigan and isolated in small laboratory feeding vessels at 4 °C, after the animals had voided their guts over a 24-hour period. Over a 20-day period, following introduction of a single ration of live cells of the filamentous diatom, Melosira varians, 9 of 10 animals had ingested this material, and 7 of these 9 individuals had deposited fecal pellets. Subsequent examination of gut contents and fecal pellets showed that although animals had ingested whole algal cells/filaments, little of the material in gut contents or fecal pellets bore any identifiable structural similarity to cells/filaments prior to ingestion. The results suggest that earlier studies of pontoporeiid gut contents may seriously underestimate the importance of algal components in the amphipod's diet and imply that Diporeia sp. growth and production may be more closely linked to primary production than previously thought.     

A novel approach to identifying PST tolerant copepods: An individual ingestion assay

Phenotypic tolerance of individual copepods to paralytic shellfish toxins (PSTs), as determined by fecundity, has recently been reported. In the present study, we tested whether similar phenotypic tolerance related to ingestion also exists. In short-term feeding assays, ingestion rates of individual females of the copepod Acartia hudsonica were measured on a sole diet of toxic Alexandrium fundyense as well as on a sole diet of nontoxic Alexandrium tamarense. When copepods fed on A. fundyense, the frequency distribution of ingestion rates was polymodal, consistent with the hypothesis of levels of phenotypic tolerance associated with toxin ingestion. Four distinct groups of ingestion rates on toxic algae were observed. Mean rates ranged from near zero to more than 100 cells copepod⁻¹ h⁻¹. In contrast, when the same individuals fed on the nontoxic A. tamarense diet, the polymodal distribution of ingestion rates was not apparent. Furthermore, group-specific mean ingestion rates, for two ingestion-defined groups, were always significantly greater on the toxic diet than they were on the nontoxic diet. These results support the hypothesis that discrete groups of grazer ingestion of toxic A. fundyense are related to PST tolerance.     

A comparative approach to study inhibition of grazing and lipid composition of a toxic and non-toxic clone of Chrysochromulina polylepis (Prymnesiophyceae)

Since the massive bloom in 1988 in the North Sea, the prymnesiophyte flagellate Chrysochromulina polylepis Manton et Parke has been known for its ichtyotoxicity. Laboratory experiments using two different clones of C. polylepis were conducted in a comparative approach. Both clones were nearly similar in size and shape, but differed in their toxicity, as demonstrated by the Artemia bioassay. In order to study the effects of toxic C. polylepis on protozooplankton grazers, grazing experiments were performed with the heterotrophic dinoflagellate Oxyrrhis marina Dujardin as grazer. A first experiment was carried out in order to follow batch culture growth and initial grazing of O. marina when fed toxic or non-toxic clones of C. polylepis. Ingestion of the toxic clone was 27% of ingestion when fed with the non-toxic clone. When O. marina was fed with the toxic clone, vacuoles within O. marina contained fewer food particles per cell and the cells attained slower division rate (58% of the division rate estimated for the non-toxic clone). A second experiment was conducted to determine the grazing and growth response of O. marina as a function of algal food concentration. Profound differences in ingestion, clearance, division and gross growth efficiency of O. marina when fed the two clones of C. polylepis again were apparent. However, even at algal concentrations of 400×10³ ml⁻¹, O. marina is not killed by the presence or by the ingestion of toxic C. polylepis, indicating that the toxin deters grazers. In addition to grazing experiments, lipid classes and fatty acids of both algal clones were analysed and compared in order to follow the hypothesis that toxicity of C. polylepis is caused by liposaccharides, lipids, or fatty acids. However, the chemical composition with respect to lipid classes and fatty acids of both clones were quite similar, making an involvement of these substances in the toxicity towards Artemia and O. marina unlikely.     

LIGHT AND ELECTRON MICROSCOPY OF GRAZING BY POTERIOOCHROMONAS MALHAMENSIS (CHRYSOPHYCEAE) ON A RANGE OF PHYTOPLANKTON TAXA1

Grazing of fluorescent latex beads, bacteria, and various species of phytoplankton by Poterioochromonas malhamensis (Pringsheim) Peterfi (about 8.0 μm in diameter) was surveyed. The alga ingested fluorescent beads and various live or killed and nomnotile or motile organisms including bacteria, blue-green algae, green algae, diatoms, and chrysomonads. The size range of grazed prey was from 0.1 to 6.0 μm for latex beads and from 1.0 μm (bacteria) to about 21 μm (Carteria inverse) for organisms. As many as 17 latex beads (2.0 μm) or more than 10 Microcystis cells (5–6 μm) were ingested by a single P. malhamensis cell. Following such grazing, the cell increased in volume by up to about 30-fold. The range of cell volume of ingested prey was from 0.52 μm³ (bacteria) to about 3178 μm³(Carteria inversa). This study demonstrates for the first time that P. malhamensis is capable of grazing algae 2–3 times larger in diameter than its own cell and of grazing intact motile algae. Poterioochromonas malhamensis is an omnivorous grazer. Food vacuole formation and digestion processes were examined. The membrane that was derived from the plasma membrane and surrounded the prey disappeared sometime after ingestion. The food vacuole was then formed by successive fusion of numerous homogeneous vesicles accumulated around the prey. The prey was enclosed in a single membrane-bound food vacuole and then digested.     

Influence of phytoplankton fractions on growth and reproduction of tropical cladocerans

The influence of two seston fractions, < 20 μm (nanoplankton) and ≥ 20 μm (microplankton), on growth and reproduction of cladoceran species with different sizes, from Lake Monte Alegre, was evaluated through individual growth and life table experiments. Size, shape and other features of the algae in the fractions were described. The procedure of filtering lake water through a 20 μm net for seston fractionation caused mutual contamination. The smallest cladoceran species, Ceriodaphnia cornuta Sars and Moina micrura Kurz, produced larger clutch sizes and exhibited higher intrinsic rates of population growth (r) in the nanoplankton, despite contamination of their food by inedible algae. The largest species, Simocephalus mixtus Sars, produced larger clutch sizes in the microplankton. There were no differences in juvenile biomass growth between treatments for M. micrura and Daphnia gessneri Herbst, but lower value of the exponential growth rate (g) in the microplankton was significant for M. micrura. Fecundity (eggs/total female) of M. micrura was significantly lower in the microplankton, while D. gessneri did not reproduce in this fraction, at the end of growth experiments. Spines, colonies, cenobium, filaments, hard cell wall, and gelatinous sheaths, represented constraints to cladoceran reproductive performance, besides algae size. Microplankton contamination by nanoplanktonic algae, in the experiments, probably minimized the negative effect of inedible algae. Nanoplankton was more suitable for the smallest species and microplankton for the largest one.     

The ingestion and digestion of algae by Chloeon dipterum L. (Ephemeroptera)

Summary The ability of larvae of Chloeon dipterum L. to ingest various species of algae was investigated in the laboratory. There was a direct relationship between the size of an alga and the frequency with which it was ingested by different sizes of larva. The frequency of ingestion of Anabaena, Cladophora, and Hydrodictyon appeared to be governed by factors in addition to their size.Several species of algae were thoroughly and rapidly digested by C. dipterum in the laboratory.Cultures of the hind-gut contents of larvae collected in the field, and of those fed upon algae in the laboratory, showed that cells of several species of algae remained viable in the gut for periods of at least 20 hours. These were narrow filamentous forms or small cells with or without a gelatinous sheath. It is suggested that such species pass between the mouthparts without being damaged so that in the absence of a cellulase the digestive enzymes of the gut are unable to penetrate the cell walls.     

Ingestion rate of the burrowing mayfly Hexagenia limbata as determined with14C

The effects of nymph size and temperature on the ingestion rate ofHexagenia limbata were investigated using¹⁴C labelled algae mixed into sediment. Ingestion rate increased with nymph length and temperature. Daily ingestion rates of large 19 mm nymphs burrowing in silt averaged 58 and 192% of the dry body weight at 16° and 21 °C, respectively. Ingestion of seston due to filter feeding in artificial burrows was insignificant compared to that ingested by burrowing nymphs.