Absence of predation eliminates coexistence: experience from the fish–zooplankton interface

Examples from fishless aquatic habitats show that competition among zooplankton for resources instigates rapid exclusion of competitively inferior species in the absence of fish predation, and leads to resource monopolization by the superior competitor. This may be a single species or a few clones with large body size: a cladoceran such as Daphnia pulicaria, or a branchiopod such as Artemia franciscana, each building its population to a density far higher than those found in habitats with fish. The example of zooplankton from two different fish-free habitats demonstrates the overpowering force of fish predation by highlighting the consequences of its absence. Released from the mortality caused by predation, a population of a superior competitor remains at a density equal to the carrying capacity of its habitat, in a steady state with its food resources, consisting of small green flagellate algae, which are successful in compensating high loss rates due to grazing, by fast growth. In such a situation, the high filtering rate of Daphnia or Artemia reduces resources to levels that are sufficient for assimilation to cover the costs of respiration (threshold food concentration) in adults but not in juveniles. This implies long periods of persistence of adults refraining from producing live young, because production of instantly hatching eggs would be maladaptive. Severe competition for limiting resources imposes a strong selective pressure for postponing reproduction or for producing resting eggs until food levels have increased. Offspring can only survive when born in a short time window between such an increase in food levels and its subsequent decline resulting from population growth and intense grazing by juveniles. Such zooplanktons become not only a single-species community, but also form a single cohort with a long-lifespan population. The observations support the notion that diversity may be sustained only where predation keeps densities of coexisting species at levels much below the carrying capacity, as suggested by Hutchinson 50 years ago.     

From inducible defences to population dynamics: modelling refuge use and life history changes in Daphnia

We investigated the relative importance of a behavioural defence (refuge use through diel vertical migration) and a life history change (a reduced size at first reproduction) that are used by daphnids to decrease the risk of predation by visually hunting fish. We used an individual based model of a Daphnia population in a stratified lake to quantify the effects of these inducible defences on Daphnia predation-mortality and the resulting Daphnia population dynamics. Our analysis shows that diel vertical migration (DVM) confers a much stronger protection against fish predation than a reduced size at first reproduction (SFR). DVM allows daphnids to withstand a higher predation pressure in the epilimnion and it decelerates a Daphnia population decline more strongly than a reduced SFR. DVM effectively reduces the (P/B) flow of carbon from daphnids to fish.
Many theoretical studies have only considered the fitness benefits of DVM above 'staying up' in the epilimnion of a lake. Our results suggest that 'staying down' in the hypolimnion would confer an even stronger fitness benefit to Daphnia than DVM at times of peak predation risk. Daphnids that remain in the hypolimnion avoid the predation suffered by migrating daphnids around dusk and dawn. Staying down could prevent a Daphnia population decline, while DVM and a reduced SFR can only decelerate the decrease of Daphnia population densities under heavy fish predation. Staying down at high concentrations of fish infochemicals has in fact been observed within a variety of Daphnia clones and species, both in the laboratory and in stratified lakes.     

Long-term responses of zooplankton to invasion by a planktivorous fish in a subarctic watercourse

1. Introduced or invading predators may have strong impacts on prey populations of the recipient community mediated by direct and indirect interactions. The long-term progression of predation effects, covering the invasion and establishment phase of alien predators, however, has rarely been documented.
2. This paper documents the impact of an invasive, specialized planktivorous fish on its prey in a subarctic watercourse. Potential predation effects on the crustacean plankton, at the community, population and individual levels, were explored in a long-term study following the invasion by vendace ( Coregonus albula ).
3. Over the 12-year period, the density and species richness of zooplankton decreased, smaller species became more abundant and Daphnia longispina , one of the largest cladocerans, was eliminated from the zooplankton community.
4. Within the dominant cladocerans, including Daphnia spp., Bosmina longispina and Bosmina longirostris , the body size of ovigerous females and the size at first reproduction decreased after the arrival of the new predator. The clutch sizes of Daphnia spp. and B. longirostris also increased.
5. Increased predation pressure following the vendace invasion induced many effects on the crustacean zooplankton, and we document comprehensive and strong direct and indirect long-term impacts of an introduced non-native predator on the native prey community.     

Biotic relations affecting species structure in zooplankton communities

Mathematical modelling and laboratory experiments were used to study how exploitative competition and predation influence the species structure in cladoceran community. For five species of Cladocera (Sida crystallina, Daphnia magna, Simocephalus vetulus, Daphnia longispina, and Diaphanosoma brachyurum), representing a gradient of body size, population characteristics were described as functions of food concentration. Abundance dynamics were simulated in mixed species cultures and invasion experiments under different levels of food supply corresponding to oligo-, meso-, and eutrophic conditions. Separate simulations were also run including and excluding (fish) predation. The competitive ability of each species was estimated as the values of the population equilibrium food concentration. Simulation results showed that for the no-predator scenario, increases in the level of food supply promoted species coexistence while under lower food concentrations only one species remained at the end of the simulation runs. When predation was allowed, the number of species that coexisted at the end of the simulations increased up to four species, indicating that predator pressure facilitated species coexistence because it shortened periods of food depletion. Simulation results were verified in laboratory experiments which suggested that population equilibrium food concentration can be used as an estimate of competitive ability. Finally, species structure and relative abundance in Lake Naroch (Belarus) during the summer of 2004 was found to be consistent with our results from computer simulation and laboratory experiments with regard to competition and predation impacts on zooplankton community. Handling editor: S. I. Dodson     

Clutch size and body size at first reproduction in Daphnia pulicaria at different levels of food and predation

Field data from seven alpine lakes in Serra da Estrela. Portugal.show that reproduction in Daphnia may be as efficiently controlledby fish predation and copepod predation on eggs in brood cavitiesas it is by food limitation. Body length and clutch size estimatesin Daphnia pulicaria revealed high inter- and intra-populationvariability in maturation size (body size at first reproduction).and in number of eggs per clutch. Daphnia at first maturationin lakes stocked with rainbow trout were half the size of thosefound in fishless lakes (body length of 0.86–0.95 and1.55–1.81 mm. respectively). The mean number of eggs perclutch was reduced to a similar degree by food limitation, predationby fish and copepod predation on eggs in brood cavities, butthe underlying mechanisms of this reduction were different.Food limitation caused smaller clutch sizes in all individuals,so variation remained the same. Fish predation caused the selectiveremoval of individuals with maximum clutches, so variation decreased.Copepod predation caused removal of eggs from brood cavitiesof randomly infested females, so that variation increased, particularlyat a high food level when non-infested females carried largeclutches of eggs.     

On the different nature of top-down and bottom-up effects in pelagic food webs

SUMMARY 1. Each individual planktonic plant or animal is exposed to the hazards of starvation and risk of predation, and each planktonic population is under the control of resource limitation from the bottom up (growth and reproduction) and by predation from the top down (mortality). While the bottom-up and top-down impacts are traditionally conceived as compatible with each other, field population-density data on two coexisting Daphnia species suggest that the nature of the two impacts is different. Rates of change, such as the rate of individual body growth, rate of reproduction, and each species' population growth rate, are controlled from the bottom up. State variables, such as biomass, individual body size and population density, are controlled from the top down and are fixed at a specific level regardless of the rate at which they are produced.
2. According to the theory of functional responses, carnivorous and herbivorous predators react to prey density rather than to the rate at which prey are produced or reproduced. The predator's feeding rate (and thus the magnitude of its effect on prey density) should hence be regarded as a functional response to increasing resource concentration.
3. The disparity between the bottom-up and top-down effects is also apparent in individual decision making, where a choice must be made between accepting the hazards of hunger and the risks of predation (lost calories versus loss of life).
4. As long as top-down forces are effective, the disparity with bottom-up effects seems evident. In the absence of predation, however, all efforts of an individual become subordinate to the competition for resources. Biomass becomes limited from the bottom up as soon as the density of a superior competitor has increased to the carrying capacity of a given habitat. Such a shift in the importance of bottom-up control can be seen in zooplankton in habitats from which fish have been excluded.     

Larval fish-induced phenotypic plasticity of coexisting Daphnia: an enclosure experiment

1. The indirect effects of predators on lower trophic levels have been studied without much attention to phenotypically plastic traits of key food web components. Phenotypic plasticity among species creates phenotypic diversity over a changing environmental landscape. 2. We measured the indirect effects of planktivorous larval walleye (Stizostedion vitreum) on phytoplankton biomass through their effects on the dominant herbivore species, Daphnia pulicaria and D. mendotae. 3. Fish had no effect on phytoplankton biomass or overall Daphnia density. We observed a compensatory response to predation by functionally comparable species within a trophic level in the form of shifting dominance and coexistence of Daphnia species. We hypothesized that this phenotypically plastic response to predation decoupled a potential trophic cascade in this freshwater pelagic system. Daphnia pulicaria density decreased over time with fish predation, but D. mendotae density increased over time with fish predation. 4. Phenotypically plastic life history trait shifts and reproductive rates differed between species in fishless and fish enclosures, accounting for population trends. Daphnia pulicaria were also proportionally higher in walleye larvae stomachs than in the enclosures, indicating that walleye preferred to feed on D. pulcaria over D. mendotae. The resultant shift in dominance may partially explain the overall benign effect of fish on grazers and supports the hypothesis that trophic level diversity can decouple a trophic cascade.     

Antipredator reaction norms for life history traits in Daphnia pulex: dependence on temperature and food

Synergistic effects between temperature and food level on the vulnerability of Daphnia life histories to predation have previously received little attention, despite their potential significance for summer population dynamics. In this investigation, most traits in the early life history of Daphnia pulex altered in response to increasing concentrations of fish kairomone. Although there was some variation attributable to experimental temperature and food treatments, traits exhibited reaction norms which suggested at least some degree of kairomone dose-dependence. Temperature, food level and their interaction affected the mean values of every trait examined and in some cases also influenced the antipredator response via a three-way interaction with fish kairomone. The insertion of an extra juvenile instar resulted in later maturation at a larger size for most females raised under conditions of low temperature and low food, but this response was largely suppressed in the presence of fish kairomone. Earlier maturation due to deletion of an instar was observed most frequently under high food and temperature conditions with fish kairomone present, suggesting an effect of kairomone on the size threshold for reproductive development. Principal components analysis was used to produce an index reflecting the net potential vulnerability to fish predation of the suite of life history traits. Vulnerability generally declined with increasing kairomone level as a result of the apparently adaptive alterations in most life history traits. Raised temperature and food level also generally reduced potential vulnerability, but a highly significant interaction between these factors was also found. Potentially important implications of these results for optimal vertical migration and summer population dynamics are discussed.     

Ancestral ecological regime shapes reaction to food limitation in the Least Killifish,Heterandria formosa

Populations with different densities often show genetically based differences in life histories. The divergent life histories could be driven by several agents of selection, one of which is variation in per‐capita food levels. Its relationship with population density is complex, as it depends on overall food availability, individual metabolic demand, and food‐independent factors potentially affecting density, such as predation intensity. Here, we present a case study of two populations of a small live‐bearing freshwater fish, one characterized by high density, low predation risk, low overall food availability, and presumably low per‐capita food levels, and the other by low density, high predation risk, high overall food availability, and presumably high per‐capita food levels. Using a laboratory experiment, we examined whether fish from these populations respond differently to food limitation, and whether size at birth, a key trait with respect to density variation in this species, is associated with any such differential responses. While at the lower food level growth was slower, body size smaller, maturation delayed, and survival reduced in both populations, these fitness costs were smaller in fish from the high‐density population. At low food, only 15% of high‐density fish died, compared to 75% of low‐density fish. This difference was much smaller at high food (0% vs. 15% mortality). The increased survival of high‐density fish may, at least partly, be due to their larger size at birth. Moreover, being larger at birth enabled fish to mature relatively early even at the lower food level. We demonstrate that sensitivities to food limitation differ between study populations, consistent with selection for a greater ability to tolerate low per‐capita food availability in the high‐density population. While we cannot preclude other agents of selection from operating in these populations simultaneously, our results suggest that variation in per‐capita food levels is one of those agents.     

Mechanisms regulating zooplankton populations in a high-mountain lake

SUMMARY 1. We studied the seasonal succession of phyto- and zooplankton and the potential impact of predation by salmonids on zooplankton population dynamics in a high-mountain Swiss lake.
2. A comparison of patterns in the abundance, body length, fecundity and age structure in the Daphnia galeata population strongly suggests that trout predation had little impact on the population and was not the cause for a decline in summer.
3. The dominance in the lake of adult trout that feed mainly on benthic prey may buffer the effect of predation on the larger zooplankton. Further, the relatively high amount of phytoplankton after spring thaw could be important for sustaining the Daphnia population under moderate fish predation.
4. Partial correlation analyses proved circumstantial evidence for both exploitative and interference competition between some zooplankton taxa. D. galeata depressed performance of other plankton species through exploitative competition.
5. Our study shows that the impact of fish on zooplankton in high-mountain lakes depends strongly on food web structure and trophic state of the lake. Where fish predation is weak, invertebrate predation combined with competition for food may be responsible for the dominance of large-bodied zooplankton species.     

Enhanced growth at low population density in Daphnia: the absence of crowding effects or relief from visual predation?

1. It has been suggested that chemical information from crowded populations of an animal such as Daphnia carries a cue indicating imminent food limitation, and we suggest that in the presence of fish kairomones, it may also convey a hint of the need to enhance antipredation defences. 2. We performed two‐factorial experiments with Daphnia grown in flow‐through plankton chambers in medium containing high levels of Scenedesmus food plus chemical information on either low or high population density levels and in the presence or absence of fish chemical cues (kairomones) and recorded (i) the effects on Daphnia growth rate and reproduction, and (ii) the effects on Daphnia depth selection. Further depth‐selection experiments were performed to test the reaction of Daphnia to crowding information at different Daphnia concentrations and to test its effect on daytime and night‐time depth selection by different Daphnia instars in the presence of kairomones. 3. The effects of crowding information alone (in the absence of kairomones) were weak and were not significantly strengthened by the addition of kairomones. The effects of kairomones alone (in the absence of crowding information) were much stronger and were increased by the presence of crowding chemicals: Daphnia selected greater depths in daylight (the later the instar and the larger its body size, the greater the depth), their body growth was slower and daily reproductive investment reduced, compared with Daphnia grown in the absence of crowding information. This suggested that crowding chemicals carry a cue indicating the need to invest more into antipredation defences. 4. The adaptive significance of these effects was confirmed by the differential vulnerability to predation of the Daphnia when offered as prey to live roach after being grown for 6 days either in the presence (higher vulnerability) or in the absence (lower vulnerability) of information on high density. 5. The strong interaction between crowding information and fish kairomones may be explained either as the reaction to a cue indicating impending food stress or as the reaction to a signal of increased predation risk. While the former scenario is already known from crowding studies, the latter is a novel idea that stems from the old concept of ‘low‐density anti‐predation refuge’. The two scenarios are not mutually exclusive: each stems from the need to invest in survival rather than in growth and reproduction [Corrections were made to this paragraph after first online publication on 4 April 2012].     

Body size distribution in <Emphasis Type="Italic">Daphnia</Emphasis> populations as an effect of prey selectivity by planktivorous fish

We test the hypothesis that size distribution of a Daphnia population reflects the vulnerability of each size category (instar) to predation by planktivorous fish. We hypothesize that due to the different reaction distances from which separate prey categories can be seen by a foraging fish, each category is preyed upon until its density is reduced and its size-specific apparent density level (number of prey within a hemisphere of radius equal to the reaction distance) or encounter rate (number of prey encountered per time within a tube with a cross section of radius equal to the reaction distance) become equal to those of other size categories. An experiment was performed with populations of Daphnia hyalina and D. pulicaria grown at two Scenedesmus + Chlamydomonas food levels (0.2 and 0.05 mg C per liter) in outdoor mesocosms (1000 l tanks) with predation by invertebrates (phantom midge) prevented by mosquito netting. Once the populations had become established, roach were added to the tanks at dusk each day and allowed to feed for 3 h, while control tanks were kept fish-free. After 20–60 days, while D. pulicaria was at low density level, the densities of D. hyalina in fish tanks were high enough to see that the age structure and size distribution matched those from simulations with the age-structured population model based on size-specific encounter rate. This match, however, remained only up to the point of first reproduction when—in contrast to the size/age distribution predicted by the model—the percentage share of adult instars in the total population decreased rapidly with age. This deviation from the predicted densities of adult instars suggests that neither encounter rates nor apparent densities derived from instar-specific reaction distances are sufficient to explain the instar-specific impact of a visual predator on planktonic prey. This implies that a foraging fish may temporarily change its feeding mode from the typical low-speed harvesting of small but abundant prey from within its visual field volume, to high-speed hunting for more scarce but larger ovigerous females when their abundance allows higher net energy gain. Shifting from one feeding mode to the other may be responsible for damping population density oscillations in Daphnia.     

CHAOBORUS PREDATION AND LIFE-HISTORY EVOLUTION IN DAPHNIA PULEX: TEMPORAL PATTERN OF POPULATION DIVERSITY,FITNESS, AND MEAN LIFE HISTORY

The effect of predation by the aquatic dipteran larva Chaoborus americanus on genetic diversity and life-history evolution in the cladoceran Daphnia pulex was investigated in large replicate laboratory populations. Instantaneous daily loss rates of clonal diversity and genetic variance for fitness indicate that 93–99% of initial genetic diversity can be removed from populations during the 8–12 generations of clonal reproduction that occur each year in natural populations. In the absence of predation, the principal evolved changes in mean population life history were smaller immature body size and increased and earlier fecundity. In the presence of size-selective Chaoborus predation, populations evolved toward larger body size and increased and earlier reproduction. The difference between these two trajectories is an estimate of the direct additive effect of Chaoborus predation. This effect was manifested as evolution toward larger body size with a trend toward earlier and increased reproduction.     

Species-specific population-density thresholds in cladocerans?

The population density of a Daphnia species seems more dependent on properties specific to the species than on those specific to the point in the season, location within a lake basin or the given lake itself. In spite of week-to-week fluctuations, the population density for each of two common European Daphniaspecies was found to be remarkably similar within single lakes (from station to station on a single date, and from date to date at a single station) as well as from lake to lake, regardless of trophic state. All lakes on all dates revealed densities in the range 10–50 ind l^–1 for the smaller-bodied D. cucullata and 1–5 ind l^–1 for the larger-bodied D. hyalina, in spite of different intensity of reproduction resulting from different food levels (chlorophyll a between 0.2 and 4.2 g/l). It can be asserted that the population density of each species remains far below the carrying capacity of the habitat K, and does not depend on food levels, which merely set the rate of population increase, while the population density reflects the species' vulnerability to predation by planktivorous fish. The reactive distance (the distance over which a foraging fish can see its prey) in 1+ roach, a dominant planktivore in the lakes studied, has been found to be twice as great for D. hyalina as for D. cucullata, whatever the light intensity. The relative reactive field volume was therefore an order of magnitude greater for the former than for the latter, showing that the densities of the two prey species would be assessed by a foraging roach as equal when, in reality, they differed by an order of magnitude, as they do in various lakes and in various seasons.The first of the two conclusions is that whatever the growth and reproduction in a population of a cladoceran such as Daphnia, its density would be fixed by mortality induced by fish predation. The second would be, that the difference between the bottom-up and top-down effects in freshwater is more than merely the upward or downward direction along the food web, since the bottom-up effects are about the flow control (the rate of net production, individual growth rate, rate of reproduction) and the top-down about the standing-crop control (biomass, individual body size, population density level).     

Trophic interactions in rockpool food webs: regulation of zooplankton and phytoplankton by Notonecta and Daphnia

We studied trophic interactions in experimental rockpools with three different food web structures: phytoplankton and small-bodied zooplankton; phytoplankton, small-bodied zooplankton and Daphnia ; and phytoplankton, small-bodied zooplankton, Daphnia and Notonecta . Nutrients, primary productivity, chlorophyll a and zooplankton species composition and biomass were measured over eight weeks.
2. Daphnia had a negative impact on other zooplankton and reduced the phytoplankton biomass and primary productivity. In the absence of Daphnia , small-bodied zooplankton species were abundant, in particular cyclopoid copepods. Concentrations of dissolved nutrients were lower and the standing crop of primary producers was higher when Daphnia was absent.
3. The presence of the invertebrate predator Notonecta produced a top-down effect which was similar to that reported for planktivorous fish, i.e. a selective reduction of daphnids followed by an increase of small-bodied zooplankton species and phytoplankton biomass.
4. The study showed that consumer regulation of Daphnia by Notonecta and of algae by Daphnia are important, but also demonstrated that trophic level biomasses were controlled by a combination of predation and resource limitation.     

Refuge availability: a key to understanding the summer disappearance of Daphnia

SUMMARY. 1. The mid-summer declines of Daphnia species in three small lakes were investigated to examine the relative roles of reduced natality and increased mortality. Reduced natality (assessed by quantifying clutch size, lipid index, and available food) could not account for the decline in daphnid abundance in any of the populations examined.
2. The role of increased mortality imposed by zooplanktivorous fish was assessed by estimating the sizes of the mid-water refuge areas where daphnids could escape fish predation. The boundaries of the refuge areas were estimated from field measurements and literature values and were based on gradients of temperature, dissolved oxygen and light. Observed decreases in refuge thicknesses correlated well with the mid-summer declines of large-bodied Daphnia species in all three lakes. Intermediatesized daphnids were less affected as the refuges thinned and small-bodied species increased in abundance.
3. The importance of refuge thickness in modifying zooplanktivore-induced mortality was further tested in large enclosures where refuge thickness was experimentally modified. In the presence of zooplanktivorous fish, large-bodied Daphnia , which used the refuge, persisted when the refuge was thick but disappeared when it thinned. Daphnia galeata mendotae , which did not occupy the refuge zone, was rapidly eliminated regardless of refuge thickness. We conclude that refuge availability plays a major role in Daphnia population dynamics.     

The combined effects of hypoxia and fish kairomones on several physiological and life history traits of Daphnia

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The interaction between predation risk and food ration on behavior and morphology of Eurasian perch

The risk of both predation and food level has been shown to affect phenotypic development of organisms. However, these two factors also influence animal behavior that in turn may influence phenotypic development. Hence, it might be difficult to disentangle the behavioral effect from the predator or resource‐level effects. This is because the presence of predators and high resource levels usually results in a lower activity, which in turn affects energy expenditure that is used for development and growth. It is therefore necessary to study how behavior interacts with changes in body shape with regard to resource density and predators. Here, we use the classic predator‐induced morphological defense in fish to study the interaction between predator cues, resource availability, and behavioral activity with the aim to determine their relative contribution to changes in body shape. We show that all three variables, the presence of a predator, food level, and activity, both additively and interactively, affected the body shape of perch. In general, the presence of predators, lower swimming activity, and higher food levels induced a deep body shape, with predation and behavior having similar effect and food treatment the smallest effect. The shape changes seemed to be mediated by changes in growth rate as body condition showed a similar effect as shape with regard to food‐level and predator treatments. Our results suggests that shape changes in animals to one environmental factor, for example, predation risk, can be context dependent, and depend on food levels or behavioral responses. Theoretical and empirical studies should further explore how this context dependence affects fitness components such as resource gain and mortality and their implications for population dynamics.     

The effects of nymphaeid (Nuphar lutea) density and predation by perch (Perca fluviatilis) on the zooplankton communities in a shallow lake

1. The effects of addition of juvenile perch (Perca fluviatilis) on the microcrustacean and rotifer communities associated with nymphaeid beds were studied, at three different plant densities [high (normal), medium (reduced by a half) and low (reduced to a third of normal)], in eighteen 2 m × 1 m enclosures in a shallow lake. 2. At the low and medium densities of lilies, Daphnia densities were high in the absence of perch but very low in the presence of perch. They increased, even in the presence of perch, to high densities (comparable with those in the absence of perch) at the highest plant density. Body sizes of Daphnia hyalina were consistent with high predation by perch at low and medium plant densities but reduced predation at high plant densities. Patterns of chlorophyll a concentration, in the presence of perch, inversely reflected those of D. hyalina density. 3. At naturally high densities but not at reduced densities, the plants appeared to act as refuges against predation for the Daphnia. Reductions in oxygen concentrations in the plant beds were not responsible for the refuge effect, nor could there be avoidance of the beds by the fish. The mechanism of the refuge effect must therefore lie in frustration of the process of capture of the Daphnia by the fish. 4. Numbers of other small Crustacea and rotifers were mostly unaffected by fish predation. Numbers of Asplanchna sp., Chydorus sphaericus and copepodites were higher in the presence of fish and, although there was no main effect of fish on numbers of Diaphanosoma brachyurum, there was a plant–fish interaction, with this species being less abundant in the presence of fish at low plant densities but more abundant in the presence of fish at medium and high plant densities. Main effects of plants were few, with only Asplanchna sp. and Keratella sp. decreasing in numbers with increasing plant density. Most taxa changed in numbers with time but interaction effects between time and plants, and fish and plants, were few.     

Demographic and life history response of the cladoceranBosmina longirostris to variation in predator abundance

Population dynamics, demography and body size of the cladoceranBosmina longirostris were examined in an experimental study in which the abundance of its predator (the cyprind fishPhoxinus eos) was varied in an unproductive lake. Four densities of fish were used, encompassing the biomass of fish in the lake.Bosmina was most abundant at low and medium fish densities (1.06 and 2.12 g fish biomass · m^-3) and less abundant when fish were either absent or present at high density (3.71 g fish biomass · m^-3). The unimodal response to predator abundance resulted from effects on both birth and death rates.Bosmina birth rates increased as fish biomass increased, in response to increasing food (phytoplankton) biomass. Death rates were highest at high fish biomass (because of fish predation) and in the absence of fish (because of predation by the dipteranChaoborus, which was most abundant in the absence of fish). Size-frequency distributions revealed that fish eliminated the larger size classes ofBosmina, and mean carapace length ofBosmina populations was inversely proportional to fish biomass.Bosmina initiated reproduction at smaller size in the presence of fish than in their absence, and size at maturity was inversely proportional to fish biomass. Size at birth also tended to decrease with increasing fish biomass, but this trend was not as strong as that of size at maturity. Decreased size at maturity apparently allowedBosmina individuals to reproduce before becoming vulnerable to fish predation. Flexibility in size at maturity, together with low abundance of invertebrate predators and large herbivores (which were preyed upon by fish), allowedBosmina to become abundant in low and medium fish treatments. In the high fish treatment, mortality due to fish predation was too severe to be offset by decreased size at maturity, andBosmina population density was low. The net response ofBosmina populations to fish predation results from interactive effects of predation on mortality, natality, and life history traits.