Copepod recruitment and food composition: do diatoms affect hatching success?

Laboratory experiments were conducted to differentiate between factors controlling the hatching success of copepod eggs. Factors that could affect viability of eggs; viz food quality, female condition and external factors were investigated. In a series of experiments the copepod Acartia tonsa Dana was fed several different diets while egg production and hatching success were monitored. The diet was analysed for fatty acid content as an indicator of food quality. Both egg production and hatching were found to be affected by the nutritional quality of the food. Hatching was also highly dependent on female fertility. External effects were tested by exposing eggs to diatom extracts. Negative effects were only evident at high extract concentrations, but disappeared when aeration was supplied to the solution. Oxygen measurements showed that failure to hatch was due to hypoxia in the extracts. No inhibitory or toxic effects of diatom cell components on hatching could be found.     

Fate of domoic acid ingested by the copepod <Emphasis Type="Italic">Acartia clausi</Emphasis>

Two important issues in the studies of harmful algae include ecological role of the toxic compounds and their fate through the food web. The aims of this study were to determine whether the production of domoic acid is a strategy evolved to avoid predation and the role of copepods in the fate of this toxic compound through the food web. The copepod Acartia clausi was fed with single and mixed cultures of the toxic diatom Pseudo-nitzschia multiseries and the non-toxic diatom Pseudo-nitzschia delicatissima. Ingestion rate as a function of diatom abundance was the same for the toxic and non-toxic Pseudo-nitzschia species, indicating no selective feeding behaviour against P. multiseries. The toxins ingested by the copepods did not affect mortality, feeding behaviour, egg production and egg hatching of the copepods. Copepods assimilated the 4.8% of the total domoic acid ingested. Although the amount of toxins daily detoxificated by the copepods was 63.6%, the copepods accumulated domoic acid in their tissues. We conclude that domoic acid is not toxic for copepods and, probably for this reason, this toxin does not act as feeding deterrent for copepods. However, even though the production of domoic acid has apparently not evolved to deter predation, copepods may play an important role on the fate of this toxic compound through the marine food web.     

Measuring RNA:DNA ratios in individual <Emphasis Type="Italic">Acartia tonsa</Emphasis> (Copepoda)

Acartia tonsa Dana is a dominant copepod in coastal waters and is therefore an important link in the food web between microplankton and higher trophic levels. RNA:DNA ratios have been used to describe growth and nutritional condition of field-collected copepods and to show strong correlation between RNA:DNA ratios and group egg production (EP). A method was developed using a sensitive, nucleic acid-specific fluorescent dye, and automated microplate fluorometer to measure DNA, RNA, and the RNA:DNA ratio of individual A. tonsa. DNA, RNA, and RNA:DNA ratios and EP were all significantly higher in copepods fed Thalassiosira sp. compared to starved copepods. There was a general trend toward an increase in RNA:DNA ratios with increase in EP, but due to the high degree of variation in both RNA:DNA ratios and EP among individual copepods there was no significant correlation between RNA:DNA ratios and EP. Significant differences in RNA:DNA ratios between fed (7.2) and starved (3.3) copepods were found after 2 days. This assay may be applied to other species of copepods sampled in the field to provide an index of the health of planktonic food webs.     

The diatom Thalassiosira rotula affects reproductive success in the copepod Acartia clausi

Fecundity, egg viability and fecal pellet production are reported for Acartia clausi females collected in the Bay of Naples, Italy, from April to October 1992 and fed either with a diatom (Thalassiosira rotula) or dinoflagellate (Prorocentrum minimum) diet, at food saturated conditions. The diatom diet significantly reduced both egg and fecal pellet production as well as hatching success. Blockage of egg development occurred with both axenic and non-axenic cultures of T. rotula, suggesting that inhibitors were provided by the diatoms and not by the bacteria associated with diatom cultures. Low hatching success was also artificially induced by exposing newly spawned A. clausi eggs to high concentrations of diatom extracts, indicating the presence of deleterious, inhibitory compounds blocking copepod embryogenesis. Fecundity and hatching success diminished significantly with female age. In contrast, female longevity was not significantly modified by food type. The presence of males did not significantly alter fecundity or egg viability. Females continued to produce viable eggs throughout the period of incubation, with and without males, in both food conditions, indicating that remating is infrequent and not necessary to sustain viable egg production in this species. The succession in low and high population densities may therefore be the outcome of variations in survival rates of eggs, rather than reproductive protential perse; such variations may strongly depend on the adult copepod diet.     

Trophic modification of essential fatty acids by heterotrophic protists and its effects on the fatty acid composition of the copepod Acartia tonsa

To test whether heterotrophic protists modify precursors of long chain n−3 polyunsaturated fatty acids (LCn−3PUFAs) present in the algae they eat, two algae with different fatty acid contents (Rhodomonas salina and Dunaliella tertiolecta) were fed to the heterotrophic protists Oxyrrhis marina Dujardin and Gyrodinium dominans Hulbert. These experiments were conducted in August 2004. Both predators and prey were analyzed for fatty acid composition. To further test the effects of trophic upgrading, the calanoid copepod Acartia tonsa Dana was fed R. salina, D. tertiolecta, or O. marina that had been growing on D. tertiolecta (OM-DT) in March 2005. Our results show that trophic upgrading was species-specific. The presence of eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA) in the heterotrophic protists despite the lack of these fatty acids in the algal prey suggests that protists have the ability to elongate and desaturate 18:3 (n−3), a precursor of LCn−3PUFAs, to EPA and/or DHA. A lower content of these fatty acids was detected in protists that were fed good-quality algae. Feeding experiments with A. tonsa showed that copepods fed D. tertiolecta had a significantly lower content of EPA and DHA than those fed OM-DT. The concentration of EPA was low on both diets, while DHA content was highest in A. tonsa fed R. salina and OM-DT. These results suggest that O. marina was able to trophically upgrade the nutritional quality of the poor-quality alga, and efficiently supplied DHA to the next trophic level. The low amount of EPA in A. tonsa suggests EPA may be catabolized by the copepod.     

Food selection by copepods: discrimination on the basis of food quality

The copepod Acartia tonsa displayed nearly two-fold higher ingestion rates on faster-growing cells of the diatom Thalassiosira weissflogii compared to ingestion rates on slower-growing cells of that species at the same cell concentration. Ingestion rates on slow-growing cells were also enhanced by the addition of cell-free aliquots of algal exudate to the experimental feeding chambers. In addition, the faster-growing algal cells were selectively ingested by the copepod when the two cell types were mixed together in different proportions, indicating that physiological differences between growing cells are a critical factor in the food detection/selection process of zooplankton. Consideration of cell carbon, nitrogen, and protein composition suggests that the copepods are maximizing nitrogenous ingestion (total protein and/or nitrogen). Selectivity for cells with higher protein content results in a higher daily protein ration, even if the selection process results in a decreased rate of ingestion in mixtures of cell types.     

Reproductive toxicity of metals in calanoid copepods

This study investigated the effect of exposure route on metal accumulation, tissue distribution, and toxicity in the marine copepods Acartia hudsonica and A. tonsa. Sublethal toxicity was measured as decreases in egg production, hatching rate, ovarian development and protein (yolk) content of the egg. When algal food, exposed to Hg at 1 nM or Cd at 5 nM resulting in cells containing 34 and 64 nmol metal g^-1 dry weight, respectively, was ingested over a 4-h period by copepods, the total copepod body burden increased nine-fold for Hg and two-fold for Cd over background concentrations, and egg production decreased by 50%. Sublethal concentrations of metals were >2 orders of magnitude lower than LC₅₀ concentrations. Hatching rate, ovarian development and egg protein content all decreased following trophic exposure to metals, implying that the process of yolk accumulation (vitellogenesis) was affected. Exposure to dissolved Cd had no effect, but dissolved Hg at concentrations as low as 0.25 nM did affect egg production. Different toxic effects following different exposure routes were related to different metal distributions in the copepods: exposure to dissolved metal resulted in metal deposition in the exoskeleton, whereas exposure to dietary metal resulted in metal deposition in internal tissues. These findings indicate that enrichment of metal concentrations in internal tissues, which occurs primarily after exposure to dietary metal, affects vitellogenesis. The reproduction rate decreases by about 75% at metal concentrations only moderately higher than levels in coastal waters. Toxicity tests involving aquatic animals need to consider effects following uptake by different pathways, including the trophic transfer of metals.     

A comparative study of the inhibitory effect of diatoms on the reproductive biology of the copepod Temora stylifera

Egg production and viability in the copepod Temora stylifera (collected in the Bay of Naples, Italy in 1992) were strongly dependent on food type. A flagellate (Isochrysis galbana) diet induced the production of good quality eggs that developed to hatching. By contrast, two diatoms (Chaetoceros curvisetum, Phaeodactylum tricornutum) resulted in poor egg quality, with hatching success as low as 20% of total egg production. With the third diatom tested, Skeletonema costatum, females produced eggs for only 3 to 4 d, after which time they either became sterile or died. These results are discussed in relation to previous findings regarding the impact of the dinoflagellate Prorocentrum minimum and the diatom Thalassiosira rotula on the hatching success of T. stylifera eggs. Low egg viability was possibly not due to an absence of remating or a deficiency of some specific essential nutrient required for egg development but to the presence of inhibitory compounds blocking cell division during early copepod embryogenesis. This questions the traditional view that diatoms are an important food item regulating copepod secondary production.     

Effects of suspended sediments on egg production of the calanoid copepod Acartia tonsa

The estuarine copepod Acartia tonsa was collected on several occasions between 4 April and 14 August 1985 from Terrebonne Bay, Louisiana (29°08N; 90°36W) and the effects in its diet of suspended sediments, collected from the same area, were measured at five different concentrations of sediment (100 to 1 000 ppm) and six phytoplankton concentrations (500 to 13 000 cells ml^-1 Thalassiosira weissflogii). Egg production rate was used as an index of diet quality. At low phytoplankton concentrations (500 cells ml^-1), and at intermediate phytoplankton concentrations (2 000 cells ml^-1) for previously starved copepods, egg production was reduced by up to 40% at a sediment concentration of 250 ppm and further reduced at higher sediment concentrations. At higher food concentrations (4 000 to 13 000 cells ml^-1), suspended sediment had no effect on egg production rates at sediment concentrations up to 500 ppm. Rates were reduced only at the highest sediment concentration of 1 000 ppm. Under most natural conditions, suspended sediment would not significantly affect egg production rates in A. tonsa.     

Phytoplankton food quality determines time windows for successful zooplankton reproductive pulses

Recruitment success at the early life stages is a critical process for zooplankton demography. Copepods often dominate the zooplankton in marine coastal zones and are prey of the majority of fish larvae. Hypotheses interpreting variations of copepod recruitment are based on the concepts of "naupliar predation," "nutritional deficiency," and "toxic effect" of diatom diets. Contradictory laboratory and field studies have reached opposite conclusions on the effects of diatoms on copepod reproductive success, blurring our view of marine food-web energy flow from diatoms to higher consumers by means of copepods. Here we report estimates of copepod feeding selectivity and reproduction in response to seasonally changing phytoplankton characteristics measured in a highly productive coastal upwelling area off the coast of central Chile. The variable phytoplankton diversity and changing food quality had a strong and highly significant impact on the feeding selectivity, reproduction, and larval survival of three indigenous copepod species. Seasonal changes in copepod feeding behavior were related to the alternating protozoan-diatom diets, mostly based on dinoflagellates and ciliates during winter and autumn (low highly unsaturated fatty acids [HUFA]/polyunsaturated fatty acids [PUFA] availability), but switched to a diet of centric and chain-forming diatoms (high HUFA/PUFA availability) during the spring/summer upwelling period. Ingestion of diatom cells induced a positive effect on egg production. However, a negative relationship was found between egg hatching success, naupliar survival, and diatom ingestion. Depending on the phytoplankton species, diets had different effects on copepod reproduction and recruitment. In consequence, it seems that the classical marine food web model does not apply to some coastal upwelling systems.     

Hatching and viability of copepod eggs at two stages of embryological development: anoxic/hypoxic effect

The effects of low oxygen concentrations on the hatching and viability of copepod eggs at two stages of embryological development were investigated. Fully developed eggs from Acartia tonsa (Dana) and Labidocera aestiva (Wheeler) collected between July and September 1991 at Turkey Point, Florida, USA, hatched at lower oxygen concentrations than newly spawned eggs given the same incubation periods. Since many of the newly spawned eggs subsequently hatched when exposed to normoxic conditions, it is likely that the exposure to low oxygen delayed embryonic development. At oxygen concentrations where no initial hatching occurred, the subsequent hatching success of fully developed eggs of A. tonsa was higher than that of newly spawned eggs, indicating that newly spawned eggs were more sensitive to low oxygen concentrations. No such difference was observed for the eggs of L. aestiva.     

High reproduction of <Emphasis Type="Italic">Calanus finmarchicus</Emphasis> during a diatom-dominated spring bloom

Feeding, egg production, hatching success and early naupliar development of Calanus finmarchicus were measured in three north Norwegian fjords during a spring bloom dominated by diatoms and the haptophyte Phaeocystis pouchetii. Majority of the copepod diet consisted of diatoms, mainly Thalassiosira spp. and Chaetoceros spp., with clearance rates up to 10 ml ind⁻¹ h⁻¹ for individual algae species/groups. Egg production rates were high, ranging from ca 40 up to 90 eggs f⁻¹ d⁻¹, with a hatching success of 70–85%, and fast naupliar development through the first non-feeding stages. There was no correlation between the egg or nauplii production and diatom abundance, but the hatching success was slightly negatively correlated with diatom biomass. However, the overall high reproductive rates suggested that the main food items were not harmful for C. finmarchicus reproduction in the area, although direct chemical measurements were not conducted. The high population egg production (>1,20,000 eggs m⁻² d⁻¹) indicated that a large part of the annual reproduction took place during the investigation, which stresses the importance of diatom-dominated spring phytoplankton bloom for population recruitment of C. finmarchicus in these northern ecosystems.     

“Good” and “bad” diatoms: development,growth and juvenile mortality of the copepod <Emphasis Type="Italic">Temora longicornis</Emphasis> on diatom diets

We measured development, growth and juvenile mortality of the common copepod Temora longicornis on 11 different monospecific diatom diets in order to estimate (1) how common the negative effects of diatoms are on the development of this copepod and (2) whether the arrested development is connected to deleterious polyunsaturated aldehydes (PUA) or food nutritional quality. Four diatom species (Thalassiosira weissflogii, Thalassiosira rotula CCMP1647, Leptocylindricus danicus CCPM469 and Skeletonema costatum CCMP1281) supported complete development, whereas development failed in or before metamorphosis on seven diatom species/strains (Chaetoceros affinis CCMP158, C. decipiens CCMP173, C. socialis, T. rotula CCMP1018, Thalassiosira pseudonana CCMP1010 and CCMP1335). However, four out of these seven species were not ingested by nauplii, either due to morphology (Chaetoceros spp.) or large size (T. pseudonana CCMP1010). The growth rate did not correlate with the ingestion rate of PUA, neither with ingestion of food mineral (nitrogen) nor with biochemical (polyunsaturated fatty acids, sterols) components. We show that, although some diatoms are of inferior food quality, this is unlikely to be connected to toxicity or due to a direct limitation by a single food nutritional compound.     

Extensive cross-disciplinary analysis of biological and chemical control of Calanus finmarchicus reproduction during an aldehyde forming diatom bloom in mesocosms

Egg and faecal pellet production and egg hatching success of the calanoid copepod Calanus finmarchicus were monitored over a period of 14?days (14?C28 April, 2008) while fed water from 4 differently treated mesocosms and ambient water. Two of the mesocosms used were inoculated with the polyunsaturated aldehyde (PUA)-producing diatom Skeletonema marinoi, while 2 received only nutrient additions with or without silica. The mesocosms developed blooms of S. marinoi, mixed diatoms or the haptophyte Phaeocystis pouchetii, respectively. Faecal pellet production of C. finmarchicus increased with increasing food availability. Egg production increased with time in all mesocosms to a maximum single female production of 232 eggs female^?1?day^?1 (average of 90 eggs female^?1?day^?1) and followed the development of ciliates and P. pouchetii, but was not affected by the observed high (up to 15?nmol?L^?1) PUA production potential of the phytoplankton. The hatching success of the eggs produced on the mesocosm diets was high (78?C96%) and was not affected by either aldehydes in the maternal diet or exposure to the dissolved aldehydes in the water.     

The effect of egg versus seston quality on hatching success,naupliar metabolism and survival of <Emphasis Type="Italic">Calanus finmarchicus</Emphasis> in mesocosms dominated by <Emphasis Type="Italic">Phaeocystis</Emphasis> and diatoms

We studied the effect of a developing Skeletonema marinoi/Phaeocystis spp. bloom on Calanus finmarchicus hatching success, early naupliar survival and metabolism. Our focus was (1) on the development of reproductive rates during a bloom initiation, peak and decline in relation to the production of potentially toxic algal metabolites and (2) on the proportional importance of female nutrition versus naupliar food environment for the production of viable nauplii. Despite polyunsaturated aldehyde (PUA) production by both S. marinoi and Phaeocystis sp., we did not observe any harmful effects on hatching success or naupliar survival and condition in any stages of the short-term (<1 week) algal bloom. Hatching success appeared to be controlled by egg lipid composition, while the beneficial effect of a high food concentration was reflected in naupliar RNA:DNA ratio, protein content and total production of viable nauplii. The egg lipids reflected seston lipids, indicating that the egg fatty acid composition was not modified by the females. Our results suggest that unselective feeding and/or retention of specific lipids can induce qualitative food limitation, although recruitment during the S. marinoi/Phaeocystis sp. bloom was high.     

Effects of water motion and prey behavior on zooplankton capture by two coral reef fishes

Water motion is an important factor affecting planktivory on coral reefs. The feeding behavior of two species of tube-dwelling coral reef fish (Chaenopsidae) was studied in still and turbulent water. One species of blenny, Acanthemblemaria spinosa , lives in holes higher above the reef surface and feeds mainly on calanoid copepods, while a second, A. aspera , lives closer to the reef surface, feeds mainly on harpacticoid copepods, and is exposed to less water motion than the first. In the laboratory, these two blenny species were video recorded attacking a calanoid copepod ( Acartia tonsa, evasive prey) and an anostracan branchiopod (nauplii of Artemia sp., passive prey). Whereas A. spinosa attacked with the same vigor in still and turbulent water, A. aspera modulated its attack with a more deliberate strike under still conditions than turbulent conditions. For both fish species combined, mean capture success when feeding on Artemia sp. was 100% in still water and dropped to 78% in turbulent water. In contrast, when feeding on Acartia tonsa, mean capture success was 21% in still water and rose to 56% in turbulent water. We hypothesize that, although turbulence reduces capture success by adding erratic movement to Artemia sp. (passive prey), it increases capture success of Acartia tonsa (evasive prey) by interfering with the hydrodynamic sensing of the approaching predator. These opposite effects of water motion increase the complexity of the predator-prey relationship as water motion varies spatially and temporally on structurally complex coral reefs. Some observations were consistent with A. aspera living in a lower energy benthic boundary layer as compared with A. spinosa: slower initial approach to prey, attack speeds modulated according to water velocity, and lower proportion of approaches that result in strikes in turbulent water.Communicated by P.W. Sammarco, Chauvin     

Lethal and sublethal effects of naphthalene and 1,2-dimethylnaphthalene on the marine copepod <Emphasis Type="Italic">Paracartia grani</Emphasis>

Here we evaluate the effects of two quantitatively very important components of the water soluble fractions of fuel oils (naphthalene and 1,2-dimethylnaphthalene, hereafter NAPH and C2-NAPH, respectively) on the survival, feeding and egg production rates, and viability of eggs of the coastal copepod Paracartia (Acartia) grani. Acute toxicity responses resulted in lethal concentrations (LC50) of 2,535 and 161 μg l⁻¹ for NAPH and C2-NAPH, respectively, with no evidence of narcotic effects. Hydrocarbon-specific differences in the toxicity response indicate that sublethal effects (EC50) on feeding by C2-NAPH were likely driven by induced mortality, whereas NAPH has direct negative effects on feeding. Sublethal effects on egg production rates followed a similar detrimental pattern to the one exhibited by feeding rates, suggesting that the lower egg production rates were mediated by the decrease in feeding rates. At the exposure time tested (24 h), the 50% reduction effective concentrations (EC50) determined for sublethal effects were relatively high in comparison with hydrocarbons’ concentrations found under natural circumstances. Long exposure (4 days) of P. grani adults to the tested hydrocarbons at concentrations well below the recorded EC50, however, had no significant effects on feeding, egg production and hatching rates. The viability of the eggs was either not affected or only slightly influenced when healthy eggs were incubated under very high concentrations (up to 6,400 and 700 μg l⁻¹ NAPH and C2-NAPH, respectively). The significance of the effects of oil spills on marine zooplankton communities is discussed in light of the results presented in this study.     

Vulnerability of the copepod Acartia tonsa to predation by the scyphomedusa Chrysaora quinquecirrha : effect of prey size and behavior

Although scyphomedusae have received increased attention in recent years as important predators in coastal and estuarine environments, the factors affecting zooplankton prey vulnerability to these jellyfish remain poorly understood. Current models predicting feeding patterns of cruising entangling predators, such as Chrysaora quinquecirrha (Desor, 1948), fail to account for the selection of fast-escaping prey such as copepods. Nevertheless, our analysis of gastric contents of field-collected medusae showed that this scyphomedusa fed selectively on the calanoid copepod Acartia tonsa (Dana, 1846) and preferentially ingested adult over copepodite stages. We measured feeding rates in a planktonkreisel while simultaneously videotaping predator–prey interactions. C. quinquecirrha consumed adult A. tonsa ten times faster than copepodites. Differences in prey behavior, in the form of predator–prey encounter rates or post-encounter escape responses, could not account for the elevated feeding rates on adults. Prey size, however, had a dramatic impact on the vulnerability of copepods. In experiments using heat-killed prey, feeding rates on adults (1.5 times longer than copepodites) were 11 times higher than on copepodites. In comparison, medusae ingested heat-killed prey at only two to three times the rate of live prey. These results suggest that during scyphomedusan–copepod interactions, prey escape ability is important, but ultimately small size is a more effective refuge from predation. Received: 26 September 1997 / Accepted: 20 May 1998     

Measurement of copepod predation on nauplii using qPCR of the cytochrome oxidase I gene

A method to directly measure predation rates by older stage copepods upon copepod nauplii using species-specific primers for the mitochondrial cytochrome oxidase subunit one gene (mtCOI) and real-time quantitative PCR (qPCR) was developed. The general approach is to determine the mtCOI gene copy number of an individual prey organism and the copy number of the same gene in the stomachs of predatory copepods collected in the field. From the knowledge of DNA disappearance rates in the stomachs, ingestion rates can be calculated. In October 2006, laboratory experiments were carried out with Acartia tonsa N1 and N2 as prey and adult female Centropages typicus as predator. The copepods were collected in Narragansett Bay, USA. A. tonsa mtCOI copy numbers copepod⁻¹ were determined for stages N1–C1 and for adults. A. tonsa DNA was detectable in the guts of the predators for as long as 3 h. Exponential rates of decline in prey DNA from the stomachs of the predators are similar to those measured for gut pigments. Because of the very small amount of DNA in an individual N1 or N2 nauplius, procedures were developed to maximize the quantitative extraction and recovery of DNA and to increase the sensitivity of the method. Two quite divergent haplotypes of A. tonsa were found in Narragansett Bay, which required separate qPCR primers; one was present in summer (July) and the other in fall and winter (October and February). With modification, the methods in this study can likely be applied to a range of predator–prey systems.     

Fecundity studies on Acartia tonsa (Copepoda: Calanoida) in standardized culture

Acartia tonsa Dana longevity and egg production data were studied over a 4-year period under standardized culture conditions. Egg-laying patterns and daily rates were evaluated as bioassay tools for measuring the nutritional value of various copepod foods, as well as assessing other aspects of environmental quality. Altogether, 337 females were observed in various aspects of this work. Experiments were performed with newly matured females in loosely covered crystallizing dishes. Fresh water and a standard concentration of algae were supplied daily, and a running daily tally was kept of the eggs produced. The effects of male copepods on egg production were analyzed. Males were either removed after the first eggs were observed or they remained with the female throughout her life span. Daily egg-laying rates of this latter group of females appeared to be more stable, but abnormal embryo development occurred if the males were not removed after fertilization. Likewise, the results of females which were not mated were studied. Life spans were shorter and egg production rates were lower than those of contemporary mated females. Unmated females were subjectively observed to be less active than their mated counterparts. The effects of continuous and temporary starvation on egg-laying were also observed. Under both normal and starved conditions, males appeared to have shorter life spans than either mated or unmated females. During starvation there were no real differences between the life spans or egg productions of mated or unmated females. Non-nutritional particulate matter was evaluated as a food source. It was ingested by the females, but did nothing to prolong the egg-laying period. Finally, the three algal components of the standard ration were evaluated. Females produced equally well with a mixed diet or with Thalassiosira pseudonana (100 m³) or Chroomonas salina (140 m³), the two larger species of this diet, but when Isochrysis galbana (40 m³) was fed, females reacted as if in a starved situation.