A genetic reconstruction of the invasion of the calanoid copepod <Emphasis Type="Italic">Pseudodiaptomus inopinus</Emphasis> across the North American Pacific Coast

The rate of aquatic invasions by planktonic organisms has increased considerably in recent decades. In order to effectively direct funding and resources to control the spread of such invasions, a methodological framework for identifying high-risk transport vectors, as well as ruling out vectors of lesser concern will be necessary. A number of estuarine ecosystems on the North American Pacific Northwest coast have experienced a series of high impact planktonic invasions that have slowly unfolded across the region in recent decades, most notably, that of the planktonic copepod crustacean Pseudodiaptomus inopinus. Although introduction of P. inopinus to the United States almost certainly occurred through the discharge of ballast water from commercial vessels originating in Asia (the species’ native range), the mechanisms and patterns of subsequent spread remain unknown. In order to elucidate the migration events shaping this invasion, we sampled the genomes of copepods from seven invasive and two native populations using restriction-site associated DNA sequencing. This genetic data was evaluated against spatially-explicit genetic simulation models to evaluate competing scenarios of invasion spread. Our results indicate that invasive populations of P. inopinus exhibit a geographically unstructured genetic composition, likely arising from infrequent and large migration events. This pattern of genetic patchiness was unexpected given the linear geographic structure of the sampled populations, and strongly contrasts with the clear invasion corridors observed in many aquatic systems.     

Effects of UV-B radiation on ingestion, fecundity, population dynamics and antioxidant enzyme activities of Schmackeria inopinus (Copepoda Calanoida)

Effects of enhanced ultraviolet B (UV-B, 280-320 nm) on copepods have gained particular attention in recent years. In this study, we investigated the effects of UV-B radiation on ingestion, fecundity, population dynamics and antioxidant enzyme activities of copepod Schmackeria inopinus exposed to varying doses of UV-B irradiance. Artificial UV-B radiation resulted in an increased mortality of nauplii, copepodites and adults with increasing UV-B doses. Nauplii and copepodites were more sensitive to UV-B radiation than adults, and adult males had a higher UV-B radiation susceptivity in comparison with adult females. Both ingestion rates and proportion of gravid females decreased with the increase of UV-B doses; and at the same time, we also observed that adult females had higher ingestion rates as compared with adult males. In comparison with the control, the abundance of the treatment significantly decreased. Antioxidant enzyme (GPx and GR) activities attained a significant increase at lower UV-B radiation doses when compared to the control, but declined at higher UV-B doses. These results suggested that enhanced UV-B radiation might change the species composition of copepods. Our study also showed that antioxidant enzymes might protect S. inopinus against UV-induced oxidative damage.     

Modelling physico-chemical factors affecting occurrences of a non-indigenous planktonic copepod in northeast Pacific estuaries

Estuarine ecosystems along the Pacific coast of North America are vulnerable to invasions by non-indigenous planktonic copepods, with documented invasions by at least nine species introduced via ship’s ballast. One of these, the calanoid copepod Pseudodiaptomus inopinus, now occurs in a relatively wide geographical area in coastal estuaries of Washington and Oregon States. Although it appears to be well established in the region, plankton surveys conducted in 1992, 1996, 2000, and 2004 in estuaries from southern Vancouver Island in British Columbia, Canada, to northern California, United States indicate that it has not expanded its range. This static distribution suggests that P. inopinus has reached a distributional limit, and it may thus be a good organism for applying models for predicting planktonic invasions, by characterizing estuaries with and without populations of the copepod. In this study, we applied both parametric, linear (discriminant function analysis, logistic regression) and nonparametric, non-linear (classification trees) techniques to develop models for occurrence of P. inopinus, to identify parameters that may lead to successful invasions and to identify specific estuaries or regions that might be at risk of invasion by this species. Both model types had similar results, identifying relatively simple salinity- and stratification-based models as good predictors of P. inopinus. While different models selected slightly different sets of variables and thresholds, all models identified relatively low salinity and stratification of water column temperature and salinity as important predictors of P. inopinus presence. The models also identified several “false positives” that mainly occurred in more inland waters of Puget Sound—estuaries that did not have P. inopinus, but had the conditions that support it, and which may be at risk for future invasions by this species.     

Predation on the Invasive Copepod,Pseudodiaptomus forbesi,and Native Zooplankton in the Lower Columbia River: An Experimental Approach to Quantify Differences in Prey-Specific Feeding Rates

Invasive planktonic crustaceans have become a prominent feature of aquatic communities worldwide, yet their effects on food webs are not well known. The Asian calanoid copepod, Pseudodiaptomus forbesi, introduced to the Columbia River Estuary approximately 15 years ago, now dominates the late-summer zooplankton community, but its use by native aquatic predators is unknown. We investigated whether three species of planktivorous fishes (chinook salmon, three-spined stickleback, and northern pikeminnow) and one species of mysid exhibited higher feeding rates on native copepods and cladocerans relative to P. forbesi by conducting `single-prey’ feeding experiments and, additionally, examined selectivity for prey types with `two-prey’ feeding experiments. In single-prey experiments individual predator species showed no difference in feeding rates on native cyclopoid copepods (Cyclopidae spp.) relative to invasive P. forbesi, though wild-collected predators exhibited higher feeding rates on cyclopoids when considered in aggregate. In two-prey experiments, chinook salmon and northern pikeminnow both strongly selected native cladocerans (Daphnia retrocurva) over P. forbesi, and moreover, northern pikeminnow selected native Cyclopidae spp. over P. forbesi. On the other hand, in two-prey experiments, chinook salmon, three-spined stickleback and mysids were non- selective with respect to feeding on native cyclopoid copepods versus P. forbesi. Our results indicate that all four native predators in the Columbia River Estuary can consume the invasive copepod, P. forbesi, but that some predators select for native zooplankton over P. forbesi, most likely due to one (or both) of two possible underlying casual mechanisms: 1) differential taxon-specific prey motility and escape responses (calanoids > cyclopoids > daphnids) or 2) the invasive status of the zooplankton prey resulting in naivety, and thus lower feeding rates, of native predators feeding on invasive prey.     

Bacteria Associated with the Surface and Gut of Marine Copepods

Little is known about the nature of bacteria associated with the surface and gut of marine copepods, either in laboratory-reared animals or in the natural environment. Nor is it known whether such animals possess a gut flora. The present report deals with studies of microorganisms isolated from healthy, laboratory-reared copepods of the species Acartia tonsa Dana, from several species of wild copepods collected from a marine or estuarine environment, and from laboratory dishes containing moribund copepods. Evidence for a unique gut flora in laboratory-reared animals is presented; the predominant bacteria were represented by the genus Vibrio. Other organisms such as Pseudomonas and Cytophaga were found less abundantly associated with the copepods and not specifically associated with the gut.     

Predation by Acartia tonsa Dana on planktonic ciliates and rotifers

In laboratory experiments, adults and nauplii of the calanoid copepod Acartia tonsa Dana feed on planktonic ciliates and rotifers. Adults have a higher clearing rate for planktonic ciliates and rotifers than for phytoplankton. Adult copepods have a maximum clearing rate of ≈200 ml copepod⁻¹ · d⁻¹ for large ciliates, with lower clearing rates for small ciliates. Nauplii have higher clearance for small than for large ciliates. Addition of ciliates or rotifers to the diet of A. tonsa enhances egg production; this effect is due to improved food quality. Microzooplankton may be an important component of the diet of A. tonsa even when phytoplankton are plentiful. Selective predation by copepods probably influences the population dynamics of planktonic ciliates and rotifers in coastal waters.     

Possible mechanisms underlying copepod grazing responses to levels of toxicity in red tide dinoflagellates

A previous study indicated that rates of ingestion exhibited by adult female copepods of Acartia hudsonica (Pinhey) and Pseudocalanus spp. were lowered by increasing levels of toxicity in clonal cultures of the bloom-causing dinoflagellate Protogonyaulax tamarensis (Taylor). In the present study, three types of laboratory grazing experiments were performed to determine which of two contending hypotheses —behavioural rejection or physiological incapacitation - could explain the observed relationship best. The experimental results consistently supported the postulated mechanism of physiological incapacitation, and not the mechanism of behavioural rejection, as the reason for the lowered rates of ingestion on the more toxic dinoflagellate clones.     

Toxin content and toxic effects of the dinoflagellate Gyrodinium corsicum (Paulmier) on the ingestion and survival rates of the copepods Acartia grani and Euterpina acutifrons

Short-term experiments showed that ingestion rates of the copepods Acartia grani and Euterpina acutifrons on different concentrations of the dinoflagellate Gyrodinium corsicum were not usually different from those obtained with the non-toxic and similar sized Prorocentrum triestinum. Long-term experiments showed that no A. grani survived for more than 288 h on concentrated cultures (1500 μg C l⁻¹) of G. corsicum compared with survival rates of 86.7% on the non-toxic P. triestinum. In contrast high and similar survival rates were found for E. acutifrons exposed for the same time to similar concentrations of both dinoflagellates. These results demonstrated that G. corsicum produce toxins which have significant adverse effects on the long-term survival rates of some copepods like A. grani but not on other copepods like E. acutifrons. The possibility of PSP-toxin production by G. corsicum was analysed by chromatographic analysis (HPLC-FD) and mouse bioassay of extracts obtained from cultures maintained in exponential growth phase in f/2-Si and L1 culture mediums. These analyses and the bioassay ruled out the presence of PSP toxins in this dinoflagellate. However, mouse tests provided the first evidence for the presence of some type of NSP toxin in G. corsicum. Haemolytic assays also gave the first positive results for the methanol extract of this dinoflagellate species.     

Hydrostatic Pressure and Temperature Effects on the Membranes of a Seasonally Migrating Marine Copepod

Marine planktonic copepods of the order Calanoida are central to the ecology and productivity of high latitude ecosystems, representing the interface between primary producers and fish. These animals typically undertake a seasonal vertical migration into the deep sea, where they remain dormant for periods of between three and nine months. Descending copepods are subject to low temperatures and increased hydrostatic pressures. Nothing is known about how these organisms adapt their membranes to these environmental stressors. We collected copepods (Calanoides acutus) from the Southern Ocean at depth horizons ranging from surface waters down to 1000 m. Temperature and/or pressure both had significant, additive effects on the overall composition of the membrane phospholipid fatty acids (PLFAs) in C. acutus. The most prominent constituent of the PLFAs, the polyunsaturated fatty acid docosahexanoic acid [DHA – 22:6(n-3)], was affected by a significant interaction between temperature and pressure. This moiety increased with pressure, with the rate of increase being greater at colder temperatures. We suggest that DHA is key to the physiological adaptations of vertically migrating zooplankton, most likely because the biophysical properties of this compound are suited to maintaining membrane order in the cold, high pressure conditions that persist in the deep sea. As copepods cannot synthesise DHA and do not feed during dormancy, sufficient DHA must be accumulated through ingestion before migration is initiated. Climate-driven changes in the timing and abundance of the flagellated microplankton that supply DHA to copepods have major implications for the capacity of these animals to undertake their seasonal life cycle successfully.     

Observations on the existence of lower threshold and upper critical food concentrations for the copepod Acartia tons a Dana

Acartia tonsa Dana represents a genus of temperate-tropical inshore copepods which, by virtue of its high biomass and rapid generation times, may be assumed to be an important planktonic primary consumer in terms of total production world-wide; its grazing response to different food concentrations is little known. Using wide ranges of concentrations of phytoplankton cultures, we have found that A. tonsa has a maximum grazing rate of ≈ 10.0 μg chl $\text{a}\text{1}$, decreasing to zero below 1.0 μg chl $\text{a}\text{1}$. This was confirmed using the more limited range of naturally-occurring particulate material. Although grazing rate became progressively reduced above 10 μg chl $\text{a}\text{1}$, ingestion rates continued to increase over the next order of magnitude of food concentration. It appears that A. tonsa is rarely exposed to food concentrations in its natural environment (as measured by conventional techniques) high enough to stimulate the maximum grazing effort. On the other hand, it is suggested that the continued increase of ingestion rates in the laboratory at much higher concentrations may indicate an adaptive mechanism associated with the encountering of ephemeral micro-patches of such concentrations, which would permit rapid filling of an empty gut in an energy-efficient manner. Fecal pellet production was also measured and shown to be a good indicator of ingestion rate.     

Trophic modification of food quality by heterotrophic protists: species-specific effects on copepod egg production and egg hatching

Recent literature suggests that heterotrophic protists could improve the biochemical constituents of poor quality algae for subsequent use by higher trophic levels, a phenomenon dubbed “trophic upgrading”. We conducted experiments to test trophic upgrading effects on the omnivorous calanoid copepod Acartia tonsa. Two heterotrophic protists, Oxyrrhis marina and Gymnodinium dominans, were grown on three algal species of different nutritional qualities: Dunaliella tertiolecta, Isochrysis galbana and Rhodomonas salina. In parallel incubations, the copepods were fed the heterotrophic protists or the algae that the protists grew on. We measured the ingestion rate, egg production rate and egg hatching success of the copepods in each treatment. Comparison of the egg production efficiency (EPE; egg carbon production/carbon ingestion×100%) between parallel diet treatments allowed us to calculate the trophic upgrading index (TUI; ratio of EPE-heterotrophic protist to EPE-alga). The copepods ingested similar amount, or 20-30% more algae than heterotrophic protists. Daily rations ranged 50-122% of body C per day. Copepods feeding on O. marina that grew on D. tertiolecta had higher egg production rate and egg hatching success than copepods feeding on D. tertiolecta directly, and the resultant TUI between the two treatments was 19.7. Similarly, the copepod egg production was improved by G. dominans that grew on D. tertiolecta and the resultant TUI was 23.3. TUI was near 1 between I. galbana treatment and the parallel heterotrophic protist treatments, indicating no or little trophic upgrading effects on the copepod's egg production. However, egg hatching success was significantly lower with G. dominans growing on I. galbana. O. marina that grew on R. salina induced a lower EPE relative to the alga itself, yielding a TUI of less than 1. Biochemical data showed that the heterotrophic protists contained eicosapentanoic acid (EPA) and docosahexanoic acid (DHA) even when they fed on alga that lacked these essential fatty acids, which may explain some of the observed trophic upgrading effects and species-specific differences in our experiments. However, our data also suggested that dietary EPA and DHA could not be the sole factors regulating the copepod's reproductive output, and that egg production and egg hatching appeared to have different nutritional requirements and respond differently to the copepod's diets.     

A model study for an Oceania watershed: spatio-temporal changes of mesozooplankton in riverine and estuarine parts of the Lanyang River in Taiwan

The mesozooplankton of a river tributary in Oceania is evaluated and is correlated against environmental, abiotic, and biological attributes of this lotic system. Abundance, distribution, and the diversity of mesozooplankton was studied at nine stations including one estuarine station during ten sampling campaigns from June 2004 to December 2005 along the Lanyang River, the largest river and estuarine ecosystem in northeastern Taiwan. Mesozooplankton was dominated by copepods, cladocerans, and fish larvae. Among all samples, the highest abundances of mesozooplankton (5,049.36 individuals m^?3) occurred in the estuary station in August 2004, which also corresponds to the highest salinities (37.0), indicating the marine role in shaping the estuarine planktonic assemblages. The abundance of mesozooplankton and number of mesozooplankton taxa were significantly higher in samples of the estuarine station than in the riverine stations (p < 0.05, one-way ANOVA). The number of mesozooplankton taxa number was affected by water temperature (r = 0.697; p = 0.025, Pearson’s correlation) that was primarily influenced by the weather that was in turn affected by seasonal monsoons.     

Selection and viability after ingestion of vegetative cells, resting spores and resting cells of the marine diatom, Chaetoceros pseudocurvisetus, by two copepods

Feeding response of two copepods Neocalanus flemingeri and Calanus sinicus on cultures of three life-forms; vegetative cells, resting spores and resting cells, of Chaetoceros pseudocurvisetus was investigated. N. flemingeri fed heavily on the vegetative cells but scarcely responded to feed on the resting spores. C. sinicus showed significantly higher filtering rate on the vegetative cells and resting cells than on the resting spores. Survival of the three life-forms of C. pseudocurvisetus after gut passage of the copepods was also studied. The resting spores could germinate from fecal pellets of both N. flemingeri and C. sinicus; however, both the vegetative cells and the resting cells could not survive ingestion by the copepods. These results suggest that resting spore forming diatoms, such as C. pseudocurvisetus form spores which have a low nutritional value and during gut passage are largely indigestible due to the heavily silicified frustules and thus minimize the effects of grazing by copepods.     

Trophic Level Stability-Inducing Effects of Predaceous Early Juvenile Fish in an Estuarine Mesocosm Study

Background

Classically, estuarine planktonic research has focussed largely on the physico-chemical drivers of community assemblages leaving a paucity of information on important biological interactions.

Methodology/Principal Findings

Within the context of trophic cascades, various treatments using in situ mesocosms were established in a closed estuary to highlight the importance of predation in stabilizing estuarine plankton abundances. Through either the removal (filtration) or addition of certain planktonic groups, five different trophic systems were established. These treatments contained varied numbers of trophic levels and thus different “predators” at the top of the food chain. The abundances of zooplankton (copepod and polychaete), ciliate, micro-flagellate, nano-flagellate and bacteria were investigated in each treatment, over time. The reference treatment containing apex zooplanktivores (early juvenile mullet) and plankton at natural densities mimicked a natural, stable state of an estuary. Proportional variability (PV) and coefficient of variation (CV) of temporal abundances were calculated for each taxon and showed that apex predators in this experimental ecosystem, when compared to the other systems, induced stability. The presence of these predators therefore had consequences for multiple trophic levels, consistent with trophic cascade theory.

Conclusions/Significance

PV and CV proved useful indices for comparing stability. Apex predators exerted a stabilizing pressure through feeding on copepods and polychaetes which cascaded through the ciliates, micro-flagellates, nano-flagellates and bacteria. When compared with treatments without apex predators, the role of predation in structuring planktonic communities in closed estuaries was highlighted.     

The dominant copepods <Emphasis Type="Italic">Senecella siberica</Emphasis> and <Emphasis Type="Italic">Limnocalanus macrurus</Emphasis> in the Ob Estuary: ecology in a high-gradient environment

Intensive transformation and sedimentation of suspended matter from riverine runoff occur in estuarine frontal zones. The mesozooplankton community plays an important role in these processes. In the Ob Estuary, the dominant copepods Limnocalanus macrurus and Senecella siberica form dense local aggregations, but only scarce data on the ecology of these species in the estuarine environment are available. We aimed at analyzing the main aspects of the ecology of the two species including their grazing impact on phytoplankton. The distribution (net tows), ingestion rates (gut fluorescence analysis), respiration and excretion rates (incubation experiments), diet composition, gonad development and size of the lipid sacs of these copepods in a high-gradient area of the Ob Estuary were studied during a cruise of the R/V Professor Stockman in September 2013. S. siberica predominantly inhabited the freshwater zone; L. macrurus was more abundant in the estuarine frontal zone. In L. macrurus, adult females and males dominated the population, the herbivorous feeding hardly met the metabolic demands, the specific lipid content was high, and the gonads were developed. In S. siberica, the fifth copepodite stage (CV) dominated. The feeding rate considerably exceeded the metabolic requirements, and the lipid content was variable. The gonads were undeveloped. The two species grazed one-fifth of the phytoplankton biomass and more than 100 % of primary production, with S. siberica responsible for the main part of the total grazing impact (up to 90 %). These results are discussed in connection with the hydrophysical parameters and phase of the population’s life cycle. The obtained results contribute to the knowledge about zooplankton ecology and the transformation of suspended matter in an estuarine high-gradient environment.     

Predators Exacerbate Competitive Interactions and Dominance Hierarchies between Two Coral Reef Fishes

Predation and competition are critical processes influencing the ecology of organisms, and can play an integral role in shaping coral reef fish communities. This study compared the relative and interacting effects of competition and predation on two competing species of coral reef fish, Pomacentrus amboinensis and P. moluccensis (Pomacentridae), using a multifactorial experiment. Fish were subjected to the sight and smell of a known predator (Pseudochromis fuscus), the presence of the heterospecific competitor (i.e., P. amboinensis vs. P. moluccensis), or a combination of the two for a period of 19 days. The sub-lethal effects of predator/competitor treatments were compared with controls; a combination of otolith microstructure analysis and observations were used to determine otolith growth patterns and behaviour. We predicted that the stress of competition and/or predation would result in strong sub-lethal impacts, and act synergistically on growth and behavioural patterns. We found strong evidence to support this prediction, but only for P. amboinensis, which suffered reductions in growth in both predator and competitor treatments, with the largest reductions occurring when subjected to both predation and competition concurrently. There was strong evidence of asymmetrical competition between the two damselfish species, with P. moluccensis as the dominant competitor, displaying strong aggressive behaviour towards P. amboinensis. Growth reductions for P. amboinensis in predator/competitor treatments appeared to come about primarily due to increases in shelter seeking behaviour, which significantly reduced the foraging rates of individuals compared with controls. These data highlight the importance of predator/competitor synergisms in influencing key behaviours and demographic parameters for juvenile coral reef fishes.     

Feeding by an omnivorous planktonic copepod aetideus divergens Bradford

Aetideus divergens Bradford is representative of a large group of marine planktonic calanoid copepods which are omnivores or mixed feeders. Because very little quantitative information is available on the feeding behavior of these copepods, laboratory feeding experiments have been carried out with adult female A. divergens presented with various sizes and concentrations of diatoms and freshly hatched nauplii of Artemia. The copepod fed most efficiently on the largest size of diatom and on Artemia nauplii, but was peculiarly inefficient at feeding on small diatoms, even when they were available at very high concentrations. In this respect, the copepod differs from filterfeeding copepods, such as Calanus pacificus Brodsky. A possible explanation of this difference depends upon Aetideus being less capable of handling very small food particles than Calanus. Aetideus divergens and its congeners usually occur at subsurface depths not far below the mixed layer and seem to be adapted for feeding on large particles, possibly large phytoplanktonic organisms and fecal pellets, which sink out of the mixed layer.     

埃玛菌素对小菜蛾幼虫的毒力及其对子代种群增长的影响

研究了不同温度下埃玛菌素（emamectin）对小菜蛾Plutella xylostella幼虫的毒力,并用生命表法研究了25℃下埃玛菌素亚致死剂量LC₃₀）处理小菜蛾3龄初期幼虫后对其子代的影响。结果表明,埃玛菌素为典型的正温度系数药剂,对小菜蛾幼虫的触杀毒力在温度16～31℃间提高了10倍左右,口服毒力在相同温度范围内提高1 000倍左右,显示了埃玛菌素对该虫极高的口服毒力。25℃下埃玛菌素亚致死剂量处理当代幼虫后,对其子代的存活和繁殖均有明显影响,处理组和对照组子代的世代存活率分别为3.7%和35.5%,平均产卵量分别为92.4和148.3,种群趋势指数分别为1.8和27.4,净增殖率（R₀）分别为3.66和33.81,内禀增长力（r_m）分别为0.0665和0.0978。     

Zooplankton invasions: a brief review,plus two case studies from the northeast Pacific Ocean

Invasions of aquatic habitats by non-indigenous species (NIS), including zooplankton, are occurring at an alarming rate and are causing global concern. Although hundreds of such invasions have now been documented, surprisingly little is known about the basic biology and ecology of these invaders in their new habitats. Here we provide an overview of the published literature on NIS zooplankton, separated by life history (holoplankton vs. meroplankton), habitat (marine, estuarine, freshwater), and biological level of organization or topic (e.g. distribution and range extension, physiology, behavior, feeding, community impacts, ecosystem dynamics, etc). Amongst the many findings generated by our literature search, perhaps the most striking is the paucity of studies on community and ecosystem level impacts of NIS zooplankton, especially in marine and estuarine systems. We also present some results from two ongoing studies of invasive zooplankton in the northeast Pacific Ocean – Pseudodiaptomus inopinus in Washington and Oregon coastal estuaries, and Tortanus dextrilobatus in San Francisco Bay. Both of these Asian copepods have recently expanded their range and can at times be extremely abundant (10³ m^–3). We also examine some aspects of the trophic (predator–prey) ecology of these two invasive copepods, and find that they are likely to be important in the flow of material and energy in the systems in which they now pervade, although their impacts at the ecosystem level remain to be quantified. Finally, the findings of both our literature search and our two case studies of invasive zooplankton lead us to make several recommendations for future research.     

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.