Interclonal variation in diel horizontal migration behaviour of the water flea <Emphasis Type="Italic">Daphnia magna</Emphasis>—searching for a signature of adaptive evolution

In shallow temperate lakes, zooplankton populations may exhibit diel horizontal migration (DHM) and move towards macrophytes during the day to avoid fish. Using a natural Daphnia magna population, we undertook an experimental investigation aimed to describe the genetic variation for DHM and to study whether an adaptive micro-evolutionary response occurred to changes in macrophyte coverage and fish predation pressure through time. Twenty-seven D. magna clones were hatched from ephippia in the sediment of shallow Lake Ring, Denmark. This lake was eutrophied during the 20th century and was subject to restoration measures in the 1970s. The DHM behaviour of the clones was observed both in the presence and absence of fish kairomone. Significant interclonal variation in DHM behaviour occurred in both treatments. To study the micro-evolutionary response of the Lake Ring D. magna population, two approaches were used. First, we compared the DHM behaviour of clones derived from ephippia collected at different depths. A comparison was conducted between clones resurrected from the period of eutrophication (1960–1980) and from the period of recovery (1986–2000). A significant treatment (presence and absence of fish kairomone) × period interaction effect was identified, suggesting a significant micro-evolutionary response for DHM behaviour. The D. magna clones exhibited a significantly stronger horizontal migration response during the period of eutrophication than in the recovery phase. Second, clonal means, representing the influence of the genotype on the trait, were correlated with environmental conditions (macrophyte cover, fish predation pressure and Secchi depth). The results of this analysis also suggest that a micro-evolutionary response by Daphnia has occurred in reaction to changes in fish predation pressure. In periods with high fish predation pressure, Daphnia migrated more strongly towards the plants. Guest editor: Piet Spaak Cladocera: Proceedings of the 7th International Symposium on Cladocera     

Interactive effects of fish kairomone and light on Daphnia escape behavior

We investigated the effects of fish kairomone and light intensity on the inducibility and effectiveness of escape behavior in four clones of the water flea Daphnia from different habitats. To characterize and determine the effectiveness of their escape responses, individuals were observed: (i) escaping from the hydrodynamic disturbances of a simulated predator (a small sphere dropped from above); (ii) being preyed upon by small fish (Poecelia reticulata); (iii) responding to encounters with conspecifics in crowded conditions. The simulated predation experiments revealed that when exposed to fish kairomone for 48 h, two Daphnia pulicaria clones, but not two hybrid clones, became about twice as sensitive to fluid disturbances when tested in the light, relative to no-kairomone and dark treatments. When tested in the dark, kairomone had no effect on sensitivity in any clone. All four clones had an all-or-none escape response, in which the strength of the response, as measured by escape distance and speed, was constant regardless of treatment. In the guppy predation trials, kairomone-treated D.pulicaria escaped significantly more often from guppies, in both bright- and dim-light conditions. In dim light, similar to natural lighting conditions, regardless of kairomone, all but the most weakly escaping clone were able to elude attacking guppies in a significant proportion of attacks. Finally, kairomone had no effect on the number of escapes performed by crowded individuals in response to the comparatively weak, non-threatening signals created by other Daphnia, indicating that the kairomone-induced alertness in the D.pulicaria clones did not make them 'excessively' sensitive. The results suggest that Daphnia escape behavior is under complex and efficient environmental regulation, and may play a significant role in aquatic trophic relationships.      

Use of random amplified polymorphic DNA (RAPD) markers in the discrimination and verification of genotypes in Eucalyptus

We carried out four separate studies using random amplified polymorphic DNA (RAPD) markers to analyse samples of Eucalyptus supplied by several different organisations. The objective was to examine the reproducibility of the RAPD technique and its ability to discriminate between individual genotypes for verification of clonal identities. We found that RAPD profiles that are unique to a genotype can be generated reliably and simply and that even closely related genotypes can be distinguished. In addition, in each of the four studies, we detected cases where the plant material studied had been mis-sampled or mis-labelled (i.e. the RAPD profiles were not consistent with the identification numbers): (1) ramets of a Eucalyptus grandis clone were found to be derived from 2 different clones; (2) ramets labelled as 2 different Eucalyptus hybrid clones were found to be the same clone, owing to a mis-planted clonal hedge; (3) samples supplied as a single progeny of a controlled E. nitens cross were derived from two crosses involving different pairs of parents; (4) mis-labelling was detected for ramets of 4 of a set of 10 clones of E. grandis and E. camaldulensis. For three of the four studies, the detection of genotype mis-identifications was unexpected, suggesting that labelling or sampling errors during the handling of plant material are a frequent occurrence, with potentially serious economic consequences.     

Intraspecific heritable variation in life-history traits can alter the outcome of interspecific competition among insect herbivores

Competition is one of the most important biotic factors determining the structure of ecological communities. In this study, we show that there is variation in competitive ability between two clones of the pea aphid, Acyrthosiphon pisum, both of which out-compete a clone of the vetch aphid, Megoura viciae, in the laboratory. We tested whether this variation in competitive ability would alter the outcome of interspecific competition in the field. While one pea aphid clone followed the pattern set in the laboratory, out-competing the Megoura viciae clone, another showed the reverse effect with Megoura viciae dominating. These differences appear to be the result of variation in early population growth rate between the pea aphid clones, rather than predation, although predation did lead to the eventual extinction of colonies. We also questioned whether intra- and interspecific differences in predator escape behaviour could affect the outcome of competition in the field. All three clones responded similarly to the presence of foraging hoverfly larvae (Episyrphus balteatus), but the Megoura viciae clone dropped from the plant significantly less often in response to the presence of a foraging two-spot ladybird (Adalia bipunctata). This work provides evidence that intraspecific variation in competitive ability can alter the outcome of interspecific competitive interactions in nature and suggests that species–specific behavioural traits may have the potential to modify the outcome of these interactions.     

Formation of morphological defences in response to YOY perch and invertebrate predation in two <Emphasis Type="Italic">Daphnia</Emphasis> species coexisting in a mesotrophic lake

This study examined the formation of morphological defences by two coexisting Daphnia species, the large-sized D. pulicaria (2 mm) and the small-sized D. mendotae (1.4 mm), in response to the presence of young-of-the-year (YOY) yellow perch (Perca flavescens) and invertebrate predators (Chaoborus, Leptodora) during summer in a mesotrophic lake. We hypothesized that due to differential size-selective predation risk by YOY fish and invertebrates, the large-sized and the small-sized Daphnia species would show different morphological responses to predation threats. We followed changes in two morphological traits (relative length of the tail spine in D. pulicaria and of the helmet in D. mendotae) among different periods during summer according to YOY fish and invertebrate predation. We defined four YOY fish predation periods based on the presence of YOY perch in the pelagic zone of the lake and the relative abundance of Daphnia preys in their gut contents, and two invertebrate predation periods based on exclusive or mutual occurrence of the invertebrate predators. The large-sized (D. pulicaria) and the small-sized (D. mendotae) species showed different morphological responses to YOY fish and invertebrate predators, respectively. The tail spine ratio of the juveniles and adults of D. pulicaria did not change in response to YOY fish predation or to invertebrate predation. A gradual increase in the helmet ratio was observed in the small-sized D. mendotae over the summer period. This change was related to the co-occurrence of the invertebrate predators (Chaoborus and Leptodora) and to YOY fish predation. The warmer temperature cannot be accounted for helmet elongation since it was constant across depths, and not related with the co-occurrence of D. mendotae and YOY perch. Guest editor: Piet Spaak Cladocera: Proceedings of the 7th International Symposium on Cladocera     

Optimising survival under predation: chemical cues modify curvature in Daphnia galeata

Morphological responses to the presence of predator info-chemicals havebeen described for many Daphnia (Cladocera) species, butD. galeata is generally considered to exhibit almost nomorphological changes that could increase its fitness under predation.Therefore, the aim of our study was to examine the nature and magnitude ofmorphological responses of D. galeata to their predatorsindetail and assess their potential role in decreasing the predation threat. Twoclones of Daphnia were exposed to predator info-chemicals(kairomones) from perch, a fish (Perca), and a phantommidge larvae (Chaoborus) an invertebrate, and a kairomone mixture fromboththese organisms. Laboratory life-table experiments were carried out and fiveparameters characterising the body shape of the daphnids were measured: helmetlength, head- and carapace width, eye diameter and body size. The last-namedthree parameters did not differ significantly between the clones or thetreatments. The differences found between the clones were significant for headwidth and helmet length, but only in combination with the treatment effects.Ourresults on genotype-dependent phenotypic plasticity indicated that, althoughphenotypic plasticity is present, the clonal composition of aDaphnia population can be altered by selection on themorphotype. This potential for a change in clonal frequencies is given by thedifferences measured between the two clones in head width and helmet length,altering the curvature of the Daphina body in response tokairomone presence.     

Trophic interactions and habitat segregation between competing Daphnia species

Summary Two commonly coexisting species of Daphnia segregate by habitat in many stratified lakes. Daphnia pulicaria is mostly found in the hypolimnion whereas D. galeata mendotae undergoes diel vertical migration between the hypolimnion and the epilimnion. I examined how habitat segregation between these two potentially competing species might be affected by trophic interactions with their resources and predators by performing a field experiment in deep enclosures in which I manipulated fish predation, nutrient levels, and the density of epilimnetic Daphnia. The results of the experiment indicate that habitat use by D. pulicaria can be jointly regulated by competition for food from epilimnetic Daphnia and predation by fishes. Patterns of habitat segregation between the two Daphnia species were determined by predation by fish but not by nutrient levels: The removal of epilimnetic fish predators resulted in higher zooplankton and lower epilimnetic phytoplankton densities and allowed D. pulicaria to expand its habitat distribution into the epilimnion. In contrast, increased resource productivity resulted in higher densities of both Daphnia species but did not affect phytoplankton levels or habitat use by Daphnia. The two species exhibit a trade-off in their ability to exploit resources and their susceptibility to predation by fish. D. g. mendotae (the less susceptible species) may thus restrict D. pulicaria (the better resource exploiter) from the epilimnion when fish are common due to lower minimum resource requirements than those needed by D. pulicaria to offset the higher mortality rate imposed by selective epilimnetic fish predators. D. g. mendotae does not appear to have this effect in the absence of fish.     

Antennule shape and body size of <Emphasis Type="Italic">Bosmina</Emphasis>: key factors determining its vulnerability to predacious Copepoda

Cyclopoid copepods are common in lakes and ponds, and they have a significant predation impact on the communities of the small zooplankton species. To reduce the predation risk, some cladoceran zooplankters develop protuberant (defensive) morphologies in the presence of the copepods. In the case of the small cladoceran Bosmina, they elongate their appendages (antennule and mucrone) and change the antennule morphotype. However, information about the effectiveness of these defensive devices against copepod predation is still insufficient. In our study, to find the compositive effects of these appendages on the vulnerability of Bosmina, we exposed two bosminid species (B. longirostris and B. fatalis) of different body sizes and with appendages of different lengths and shapes to copepod (Mesocyclops) predation. The experiment revealed that the shape of the antennule is a main factor determining the bosminid’s vulnerability to copepod predation and indicated that the protection of the opened ventral carapace must be a key strategy by which Bosmina avoids copepod predation.     

Divergent ecology of sympatric clones of the asexual gecko,Lepidodactylus lugubris

We report differences in the thermal biology, elevational, temporal and geographic distributions of sympatric clones of the widespread asexual house gecko, Lepidodactylus lugubris. The two most common L. lugubris clones in Fiji, clones 2NA and 2NB, differ significantly in preferred temperature as measured in a laboratory heat gradient, but were similar in critical thermal maximum and minimum. Significant differences were found in the relative frequency of clones 2NA, 2NB, and a third Fijian clone, clone 3NB, at seven sites along an elevational gradient in Fiji. Clone 2NB was not collected at sites above 235 m, consistent with its higher preferred temperature, whereas clone 2NA was captured as high as 835 m. Clone 3NB was extremely rare at sealevel (1% of all individuals at three sites below 100 m), but predominated at the two highest-elevation sites (42% and 100%). Clones 2NA and 2NB did not differ significantly in their activity time or ambient activity temperature at low-elevation sites. Clone 3NB however, was active on significantly cooler nights at two of those sites. These significant inter-clonal differences in spatial and temporal distribution should allow a more complete utilization of resources by the assemblage of clones than by any single clonal genotype, and may promote coexistence of clones at a within-island and within-site scale. Clone 2NA, which is the most common clone in Fiji and has the broadest elevational distribution, also has the widest geographic distribution. It was the predominant clone at 27 of 34 sites surveyed in nine Pacific archipelagoes. This suggests that the ecological attributes that favor this clone in Fiji also favor it elsewhere in the Pacific despite differing environmental conditions and clonal composition in those areas.     

More than one `fish kairomone'? Perch and stickleback kairomones affect Daphnia life history traits differently

Life history traits of Daphnia galeata clones were affected differently by kairomones exuded by two species of fish. The exposure to Gasterosteus kairomone resulted in an increase in population intrinsic rate of increase and body growth rate relative to the exposure to Perca kairomone. These differences between fish treatments have not been reported earlier. The results suggest differences between fish kairomones that are not due to concentration effects alone. This raises the possibility that kairomones are cocktails of chemicals rather than a single `fish substance'. It also supports the view the effects of size-selective predation vary depending on the local predator–prey assemblage examined. This finding leads to an additional, previously underestimated but potentially complex aspect in zooplankton ecology.     

Intraspecific variability in response to stimuli for male and ephippia formation in Daphnia pulex

Natural clones and clones from laboratory crossings of Daphnia pulex have been tested in their response to stimuli for male and ephippia formation. The clones are from ponds in Illinois, USA, an area with both obligate and cyclical clones of D. pulex. The progeny from two types of crosses were studied: 1) crosses between an obligate clone producing males and a cyclical clone producing only females, and 2) self-fertilization within a cyclical clone producing both sexes. Both the natural and the artificial clones showed great variation in response to environmental stimuli for males. Ephippia response seems more equal among the clones, but two of the artificial clones could hardly be stimulated to produce ephippia in the induction experiments. In crowded cultures, however, they produced a few. Crossings between cyclicals and obligates yielded mostly cyclical progeny. Progeny from the selfed clone had low survival and fecundity.     

Predator detection and avoidance by lotic mayfly nymphs of different size

We studied antipredatory responses of lotic mayfly (Baetis) nymphs in a factorial experiment with four levels of fish presence: (1) a freely foraging fish (the European minnow,Phoxinus phoxinus), (2) a constrained fish, (3) water from a fish stream, (4) water from a fishless stream. LargeBaetis nymphs drifted mainly during night-time in treatments involving either the chemical or actual presence of fish, whereas no diel periodicity was observed when the water was not conditioned with fish odour. The response was strongest when the fish was uncaged, which suggests that visual or hydrodynamic cues are needed in addition to chemical ones for an accurate assessment of predation risk. Fish presence had no effect on the drift rates of small nymphs. Instead, they increased their refuge use in the presence of a live fish. Chemical cues alone did not have any effect on the refuge use of any of theBaetis size classes. Our results indicate active drift entry by mayfly nymphs. Because predation pressure is spatially and temporally variable, nymphs must sample the environment in order to locate predator-free areas or areas with low predation risk. Drifting should be the most energy-saving way to do this. To avoid the risk from visually feeding fish, large individuals can sample safely (i.e. enter drift) only at night-time, while the small ones can also do this safely during the day. We suggest that, contrary to some earlier assumptions, mayfly drift is not a fixed prey response. Instead,Baetis nymphs are able to assess the prevailing predation pressure, and they adjust their foraging behaviour accordingly.     

Phosphorus availability mediates plasticity in life-history traits and predator–prey interactions in Daphnia

We analysed growth plasticity of two Daphnia pulex clones under low‐phosphorus (LP) and high phosphorus (HP) conditions, in the presence of Chaoborus kairomones to examine how food quality (P‐availability) might impact life‐history responses and vulnerability to predation. Overall, clone 1 grew faster, and was larger at maturity. Under HP, both clones responded to kairomones by increasing growth, age and size at maturity, and decreasing fecundity. Under LP, both clones suffered reduced growth, and fecundity. However, the magnitude of response to kairomones depended on a clone by P‐availability interaction. Chaoborus presented a 1 : 1 clonal mixture under HP or LP, consumed more individuals under LP. Moreover, fewer clone 1 individuals were consumed. Studying the effects of P‐availability on life histories, and predator–prey interactions may help us understand the mechanisms generating and maintaining plasticity, as well as influencing genotypic diversity and microevolutionary processes in natural populations.     

Developmental Polymorphism Induced by Intraspecific Predation in the Ciliated Protozoon Onychodromus quadricornutus1

ABSTRACT. The hypotrich ciliate Onychodromus quadricornutus is remarkable in its potential for voluminous size (up to 900 μm in length), its possession of a unique set of four dorsal spines or horns, and its capability to express two kinds of developmental polymorphism induced by intraspecific predation. Cell length frequencies in replicates of a well-fed clone show normal distributions; starvation followed by intraspecific predation, however, induces cells within a clone to transform into two size classes: small lanceolate cells and cannibal giants. Induction experiments indicate that a substance released by cannibal giants stimulates defensive spine growth in clonemates within 24 h. Giants can also induce spine growth in non-clonemates. Furthermore, O. quadricornutus cells exposed to the predacious ciliate Lembadion magnum also develop hypertrophied spines. Selection experiments show that conspecific giants prey on cells with undeveloped spines (< 20 μm in length) to a much greater extent than on cells with developed spines (>40 μm in length). Transformation of a population of similarly sized O. quadricornutus cells into two different size classes may function to increase the range of potential prey sizes available to the O. quadricornutus population; hypertrophied spines appear to function as an inducible defense against intermittent predators appearing in the system including conspecific giants. This is the first reported case of a defensive developmental polymorphism induced by intraspecific predation.     

High variation in host adaptation among clones of the pea aphid,Acyrthosiphon pisum on peas,Pisum sativum

The performance of one clone of the pea aphid,Acyrthosiphon pisum (Harris), was assessed on 37 different cultivars and species ofPisum L. In addition, random samples of 36 pea aphid clones collected on alfalfa and clover were tested on a selection of fivePisum sativum L. cultivars. Aphid performance was evaluated in terms of the mean relative growth rate (MRGR) during the first five days of life or other life history variables. The MRGR of the first-mentioned pea aphid clone differed little between cultivars. No significant differences in MRGR were found between wild and cultivatedPisum species or between modern and oldP. sativum cultivars. There was considerable variation in host adaptation among the 36 pea aphid clones within each sampled field. The pea aphid clones showed no consistent pattern in performance on four of the five pea cultivars i.e. there was a significant pea aphid genotype —pea genotype interaction. On one of the cultivars all clones performed well. Pea aphid clones collected from red clover generally performed relatively poorly on pea cultivars, in contrast to the pea aphid clones collected on alfalfa. There was no difference in performance between the two pea aphid colour forms tested. Possible reasons for the high variation and the observed adaptation patterns are discussed. The fact that all clones were collected in two adjacent fields indicates thatA. pisum shows high local intraspecific variability in terms of host adaptation.     

Differences in growth,total lipid content and fatty acid composition among 60 clones of <Emphasis Type="Italic">Cylindrotheca fusiformis</Emphasis>

Differences between clones of the diatom Cylindrotheca fusiformi were studied with respect to growth rate, total lipid content and fatty acid composition. Sixty clones were isolated and cultivated under batch conditions. All clones were grown under identical conditions (temperature 22±1°C, light intensity 100 μmol photon m⁻² s⁻¹, salinity 28, F/2 medium) and were harvested in the late exponential growth phase for lipid and fatty acid analysis. The results show a wide variation in growth, total lipid content and fatty acid profiles among clones (p<0.05). The major fatty acids in the 60 clones were 14:0 (4.6–9.1%), 16:0 (18.2–32.0%), 16:1n-7 (21.6–33.1%), 20:4n-6 (4.1–13.5%) and 20:5n-3 (6.2–17.2%), with the highest proportion of 20:4n-6 in clone CF13 (13.5%), and the highest proportion of 20:5n-3 in clone CF5 (17.2%). The results support the view that some microalgal fatty acid variation is not restricted to interspecific variation and external factors, but also varies from clone to clone within the same species.     

Differences in demographic parameters among electrophoretic clones of Daphnia obtusa Kurz (Crustacea: Cladocera)

In an 18-month electrophoretic study, a Daphnia obtusa population that recently colonized Lake Orta (a large, severely polluted lake in Northern Italy) showed no genetic variation.Populations from other localities were sampled. From clones established by single females, genotype differences were determined by two life table experiments (using Lake Orta and Lake Maggiore water as a culture medium).As expected, in the Lake Orta water the colonizing clone (LO) had longer survival, produced more newborns and had a larger rate of increase than the other clones. In the Lake Maggiore water the LO clone still fared better than the other clones.     

THE SIGNIFICANCE OF OUTCROSSING IN AN INTIMATE PLANT-HERBIVORE RELATIONSHIP. I. DOES OUTCROSSING PROVIDE AN ESCAPE FROM HERBIVORES ADAPTED TO THE PARENT PLANT?

Sexual reproduction may be advantageous for hosts that are preyed on or parasitized by enemies that are highly adapted to them. Sexual reproduction can create rare genotypes that may escape predation by virtue of rarity and can create variable progeny that may escape predation if enemies are specialized to only one genotype of host. Populations of the herbivorous thrips, Apterothrips apteris, have been shown to be adapted to individual Erigeron glaucus clones. Here, we show that thrips adapted to the parental clone could better use plant progeny of the “home” clone produced through selfing than progeny derived from selfing of other clones. Thus, despite recombination, progeny produced by selfing presented a resource that was similar to the parental phenotype with respect to use by adapted thrips. We also show that E. glaucus susceptibility to thrips has a genetic basis and then ask whether outcrossing provides a means for E. glaucus clones to escape attack by adapted thrips. When we compared the success of thrips on progeny produced by selfing or outcrossing of the home clone, we found that the merits or disadvantages associated with outcrossing were dependent on the susceptibility to infestation of the parental clones. Selfing by clones characterized by low infestations of thrips appeared to preserve resistant genotypes; all outcrossed progeny had, on average, higher infestation levels than selfed progeny. In contrast, outcrossed progeny of clones characterized by high infestations of thrips had either the same thrips density as progeny from selfing, when the pollen donor was a highly infested clone, or lower density, when the pollen donor was a low infestation clone. The advantages of outcrossing were caused by the alleles contributed to progeny rather than to progeny variability or rarity.     

Impact of whitefish on an enclosure ecosystem in a shallow eutrophic lake: selective feeding of fish and predation effects on the zooplankton communities

Bag-type enclosures (75 m³) with bottom sheets and tube-type enclosures (105 m³) open to the bottom sediment were stocked with exotic whitefish (Coregonus lavaretus maraena) to study their predation effects on the plankton community. The fish fed mainly on adult chironomids during the period of their emergence (earlier part of the experimental period). Thereafter, the food preference was shifted to larvae of chironomids and crustacean zooplankters. The predation effects on the plankton community were not evident in the bag-type enclosures where zooplankton densities were consistently low. The fish reduced the crustacean populations composed ofBosmina fatalis, B. longirostris andCyclops vicinus in the tube-type enclosures where the prey density was high (above ca. 50 individuals 1⁻¹). The results suggested that the intensity of predation depended on the prey density. Rotifers increased in the fish enclosure, probably becauseCoregonus reduced the predation pressure byCyclops vicinus on rotifers and allowed the latter to increase. In the fish enclosures, no marked changes in species composition were observed. Zooplankton predated by the fish seemed to be distributed near the walls of the enclosures. Problems of enclosure experiments for examining the effects of fish predation on pelagic zooplankton communities are discussed.     

Impact of Holoshestes heterodon Eigenmann (Pisces: Characidae) on the plankton community of a subtropical reservoir: the importance of predation by Chaoborus larvae

We analyzed the effects of planktivorous Holeshestes heterodon Eigenmann (Characidae) predation on the plankton community of a small subtropical reservoir, using four enclosures (volume about 17.5 m³), open to the sediment, established in the littoral zone. Two enclosures were stocked with fish (mean TL 5.7 cm), at a density of about 4–5 fish m^–3 (approx. 8 g m^–3), whereas two remained fishless. The experiment lasted a little longer than one month. In the fish enclosures, most Crustacea and Chaoborus larvae remained scarce, probably as a result of visually selective fish predation. In both fishless enclosures, Chaoborus larvae became abundant. However, in only one of these did large individuals become relatively numerous; this discrepancy in the demographic structure of the Chaoborus populations between the two fishless enclosures is unexplained. Only in the fishless enclosure without appreciable numbers of large Chaoborus did densities of Crustacea increase greatly. It is suggested that in the enclosure containing large Chaoborus individuals, crustacean populations were prevented from developing due to predation pressure, while the small Chaoborus larvae of the other enclosure could not readily consume these prey. Rotifers were low in abundance in the absence of fish, probably as a consequence of Chaoborus predation. Phytoplankton density increased in all four enclosures, due probably to the lack of water flow. Only in the fishless enclosure with high densities of crustaceans did phytoplankton abundance decrease markedly at the end of the experiment, perhaps because of grazing losses.