Relative significance of direct and indirect effects of predation by planktivorous fish on zooplankton

One of the most obvious features of tropical lakes and reservoirs is the small body size of their zooplankton taxa. It is believed that this is the result of high and persistent predation by abundant planktivorous fish, which select large-bodied zooplankton prey thus making them more vulnerable to extinction in tropical as compared to temperate habitats. Do these extinctions result directly from fish predation? Could the high predation-induced mortality alone be responsible for an extermination of the population from a habitat? Or could indirect effects of predation be responsible? Some important indirect effects can be seen at the demographic level; these include reduced reproduction in the population resulting from higher vulnerability of ovigerous females to predation by visually oriented planktivores. Other important indirect effects can be observed at the individual level; these include shifts in behavior (from foraging to predator avoidance) and adjustments in physiology (from high to low feeding rate) in those planktonic animals which detect danger from their predators by sensing either the ‘predator odor’ or an ‘alarm substance’ originating from injured conspecific prey. Although a zooplankton species density may mostly result from the brutal force of direct predator impact on the population (mortality), it is more likely that its distribution in time and space could be attributed to a combination of indirect effects of predation on individual behavior and physiology. An example of periodicity in density and depth distribution patterns of Cahora Bassa zooplankton species and their periodic exterminations seems to confirm the role of indirect effects of predation by planktivorous fish.     

Feeding behavior of the cichlid,Sarotherodon galilaeum: selective predation on Lake Kinneret zooplankton

In laboratory feeding trials, we analyzed the feeding behavior and selectivity of the cichlid, Sarotherodon galilaeum, for zooplankton prey from Lake Kinneret, Israel. The feeding behavior was dependent on fish size. Fish less than 20 mm SL fed on zooplankton as obligate particulate feeders. Fish from 20 to 42 mm SL fed either as particulate feeders or as filter feeders. Fish larger than 62 mm SL fed as obligate filter feeders. Particulate-feeding fish were size selective and had highest feeding electivities for large-sized zooplankton species. Filter-feeding fish had highest feeding electivities for zooplankton species with poor escape ability. In general, S. galilaeum predation pressure would be greatest on Ceriodaphnia reticulata, a large-bodied and easily captured species which is selected by both particulate-feeding and filter-feeding fish.     

Differential zooplankton feeding behaviors,selectivities, and community impacts of two planktivorous fishes

Synopsis We examined the feeding behaviors and selectives of two common planktivorous fishes, pumpkinseeds Lepomis gibbosa and fathead minnows Pimephales promelas in the laboratory. Ingestion rates for both pumpkinseeds and fathead minnows feeding on zooplankton increased as a function of fish length. Pumpkinseeds fed on zooplankton strictly as particulate feeders, with preferences increasing as a function of zooplankton body size regardless of taxonomic identity. Preferences were highest for large Daphnia, intermediate for intermediate-sized copepods, and lowest for small Ceriodaphnia. Fathead minnows displayed the ability to use both particulate-feeding and filter-feeding behaviors. Differential preferences tended to reflect both zooplankton size and taxon, being highest for large, slow-swimming Daphnia, intermediate for small Ceriodaphnia, and lowest for faster-swimming copepods. These differences in prey capture behaviors and preferences of the two fishes are reflected in the zooplankton taxonomic composition of small ponds containing each fish type. The crustacean zooplankton assemblages in ponds containing both pumpkinseeds and fathead minnows were dominated by copepods. Cladocerans were rare. In ponds containing pumpkinseeds, but no fathead minnows, cladocerans were abundant, generally accounting for up to 80% of total crustacean zooplankton biomass. These results suggest that the type of planktivore, and not simply the presence or abundance of planktivores in a system, can determine zooplankton community structure.     

Feeding convergence in South American and African zooplanktivorous cichlids Geophagus brasiliensis and Tilapia rendalli

Synopsis Acará, Geophagus brasiliensis, and red-breasted bream, Tilapia rendalli, are important planktivorous cichlids in southern Brazilian lakes and reservoirs. In laboratory experiments, I quantified behavior and selectivity of different sizes of these two fish feeding on lake zooplankton. Feeding behavior depended on fish size. Fish < 30 mm were visual feeders. Fish 30–50 mm either visually fed or pump-filter fed depending on zooplankton size. Fish > 70 mm were pump-filter feeders. Replicate 1 h feeding trials revealed that, as the relative proportions of prey changed during an experiment, acará (30–42 mm, standard length) and tilapia (29–42 mm) shifted from visual feeding on large evasive copepods to filter feeding on small cladocerans and rotifers. Electivity and feeding rate increased with prey length, but were distinct for similar-sized cladocerans and copepods. Visual/filter-feeding fish had lowest electivities for small and poorly evasive rotifers and cyclopoid nauplii. They fed non-selectively on cyclopoid copepodites, had intermediate electivities for calanoid nauplii and small cladocerans, and had highest electivities for large cladocerans, cyclopoid adults, and calanoid copepodites and adults. Although belonging to different cichlid genera and native to South America and Africa, respectively, acará and red-breasted bream (= congo tilapia) exhibited similar selectivity for zooplankton. Apparently, few stereotyped feeding behaviors have evolved during the acquisition of microphagy in fish. Shift in feeding modes allows these two species to optimally exploit the variable and dynamic patchy distribution of planktonic resources.     

Cladoceran Densities,Day-to-Day Variability in Food Selection by Smelt,and the Birth-Rate-Compensation Hypothesis

Late-evening gut inspection of a dominant planktivore (smelt) and evaluation of densities, fecundities, and body-size distributions in dominant zooplankton prey (cladocerans) were made in day-to-day sequences in June–July (24 days in 1999 and 24 days in 2000). This was conducted as a field test of the hypothesis that species-specific population densities in cladocerans result from size-selective predation by a dominant fish assumed to be a general predator, switching from one prey to another as relative abundance changes. Little of the expected coincidence has been revealed between population density declines and increased numbers of a given prey in smelt diet. However, the data were consistent with the notion that fish would switch from one prey to another depending on the prey relative abundance (the number of prey a fish would see in its reaction field volume). Each cladoceran population fluctuated around its species-specific density level, lower or higher, depending on individual susceptibility to smelt predation, from 0.2 ind. l^–1 in large-bodied Daphnia hyalina and Leptodora kindtii, to 30.0 ind. l^–1 in small-bodied Daphnia cucullata andBosmina thersites. In spite of high fish-to-fish and day-to-day variability in both smelt diet and smelt selectivity for different prey, all cladocerans (also copepods and midge larvae) were equally persistent in smelt diet, and smelt selectivity was similar for small- and large-bodied prey categories, but lower for elongated-(Daphnia, Diaphanosoma) than for compact-body (Bosmina, Chydorus) species, when integrated for the entire sampling time. Closer examination of D. cucullata and B. thersites revealed strong smelt selection for later instars and females with greater clutches, showing that size distribution in a cladoceran population might be structured by fish predation in a similar way to that a cladoceran community (species relative abundance) is structured in a lake habitat. The birth-rate-compensation hypothesis is offered to explain why the value of food selectivity index in a planktivorous fish would remain the same for alternate prey categories with similar life-history traits, unless they differ in susceptibility to predation before the time of first reproduction.     

Nonvisual feeding in a visual planktivore,Xenomelaniris venezuelae

Summary Studies of the diel feeding patterns of the planktivorous fish, Xenomelaniris venezuelae, in Lake Valencia, Venezuela, revealed that, although the fish is primarily a diurnal feeder, it consumes substantial numbers of Chaoborus larvae and pupae at night. A number of fish species are known which feed on plankton at night, but these fish are filter feeders and their diets largely consist of relatively small, nonevasive prey. Chaoborus, however, is large and agile. Predation by Xenomelaniris in the dark was also studied experimentally. Captured fish were placed in completely darkened aquaria with zooplankton from Lake Valencia. After several hours the plankton was removed and examined for evidence of feeding. The fish were found to consume Chaoborus pupae and fourth instar larvae but not other types of prey. The mode of feeding by Xenomelaniris in the dark is unknown.     

A method for in situ estimation of prey selectivity and predation rate in large plankton, exemplified with the jellyfish Aurelia aurita (L.)

A method to estimate predation rates of large predatory zooplankton, such as jellyfish and ctenophores, is outlined. Large plankton size allows direct visual tracking of the predator during the process of foraging. The presented method is novel in the sense that it measures predation rate of a specific individual plankton predator in situ.After prey has been evacuated from the gut of an individual predator, the predator is incubated in situ, and observed by SCUBA-divers who recapture the individual after a defined time. Given that this incubation time is shorter than prey digestion time, predation rate can be calculated as increase in gut content over time. Clearance rates for different prey can be calculated from predation rates and prey concentrations in the water, allowing accurate estimates of prey selectivity. Thus, the problem of unknown feeding history and feeding environment, which can otherwise be a problem in prey selectivity studies of in situ-captured predators, is circumvented. Benefits and limitations of the method are discussed.The method was applied to adult medusae of the common jellyfish Aurelia aurita. A large variation in number of captured prey was detected both among individual jellyfish and among the various oral arms and gastric pouches within individuals. Clearance rates varied strongly with prey type. The medusae selected large crustacean prey (cladocerans and copepods/copepodites) over echinoderm larvae and copepod nauplii. Prey distribution within the medusae indicates that both tentacles and oral arms were used as prey capturing sites. Food passage time from prey capturing organs to gastric pouches was estimated.     

Size‐based interactions and trophic transfer efficiency are modified by fish predation and cyanobacteria blooms in Lake Mývatn,Iceland

相似文献   

Modifying the PEG model for Mediterranean lakes – no biological winter and strong fish predation

相似文献   

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.     

The Influence of Structural Complexity on Fish–zooplankton Interactions: A Study Using Artificial Submerged Macrophytes

Aquatic macrophytes produce considerable structural variation within the littoral zone and as a result the vegetation provides refuge to prey communities by hindering predator foraging activities. The behavior of planktivorous fish Pseudorasbora parva (Cyprinidae) and their zooplankton prey Daphnia pulex were quantified in a series of laboratory experiments with artificial vegetation at densities of 0, 350, 700, 1400, 2100 and 2800 stemsm^–2. Swimming speeds and foraging rates of the fish were recorded at different prey densities for all stem densities. The foraging efficiency of P. parva decreased significantly with increasing habitat complexity. This decline in feeding efficiency was related to two factors: submerged vegetation impeded swimming behavior and obstructed sight while foraging. This study separated the effects of swimming speed variation and of visual impairment, both due to stems, that led to reduced prey–predator encounters and examined how the reduction of the visual field volume may be predicted using a random encounter model.     

Predictability and possible mechanisms of plankton response to reduction of planktivorous fish

The predictability of plankton response to reductions of planktivorous fish was investigated by comparing the plankton community in three biomanipulated lakes and ten unmanipulated lakes differing in intensity of fish predation. Data collected on total phosphorus, phytoplankton and zooplankton biomass and share of cyanobacteria and large grazers, as well as specific growth rate of phytoplankton, were further used to test some of the proposed underlying response-mechanisms. In the biomanipulated lakes the algal biomass and share of cyanobacteria decreased, specific growth rate of phytoplankton increased, and zooplankton biomass and share of large grazers increased or remained unchanged. This pattern was largely reflected in the differences in food-chain structure between the unmanipulated lakes with highversus those with low fish predation. The qualitative response to planktivorous fish reduction thus seems largely predictable. The biomanipulated lakes differed, however, in magnitude of response: the smallest hypertrophic, rotenone-treated lake (Helgetjern) showed the most dramatic response, whereas the large, deep mesotrophic lake (Gjersjøen), which was stocked with piscivorous fish, showed more moderate response, probably approaching a new steady state. These differences in response magnitude may be related to different perturbation intensity (rotenone-treatmentversus stocking with piscivores), food-chain complexity and trophic state. Both decreased phosphorus concentration and increased zooplankton grazing are probably important mechanisms underlying plankton response to biomanipulation in many lakes. The results provide tentative support to the hypothesis that under conditions of phosphorus limitation, increased zooplankton grazing can decrease algal biomassvia two separate mechanisms: reduction of the phosphorus pool in the phytoplankton, and reduction of the internal C:P-ratio in the phytoplankton cells.

     

Feeding patterns of the dominant benthic and demersal fish community in a temperate estuary

The diet and feeding relationships of the 12 most abundant benthic and demersal fish species of the Mondego Estuary, Portugal, were studied between June 2003 and May 2006. Fishes were caught during the night using a 2 m beam trawl. The stomach contents were analysed for prey identification, counting and weighing. According to prey importance in diets, three main feeding guilds were identified: (1) invertebrate feeders, (2) invertebrate and fish feeders and (3) plankton and invertebrate feeders. Besides these main feeding guilds, some fishes also presented fractions of algae and zooplankton in their stomach contents. The most abundant prey items were macroinvertebrates, with several polychaetes ( Nephtys spp., Capitellidae, Spionidae and Eunicidae), Corophium spp. and Crangon crangon among the dominant prey. Pomatoschistus spp. were the most preyed on fishes. Several fish species showed a tendency for a specialized diet, but almost all also showed some degree of opportunistic feeding by preying on other food resources. High diet overlap was found between some fish species, yet exploitative competition could not be concluded.     

Strategies of zooplanktivory shape the dynamics and diversity of littoral plankton communities: a mesocosm approach

Planktivorous fish can exert strong top‐down control on zooplankton communities. By incorporating different feeding strategies, from selective particulate feeding to cruising filter feeding, fish species target distinct prey. In this study, we investigated the effects of two species with different feeding strategies, the three‐spined stickleback (Gasterosteus aculeatus (L.)) and roach (Rutilus rutilus (L.)), on a low‐diversity brackish water zooplankton community using a 16‐day mesocosm experiment. The experiment was conducted on a small‐bodied spring zooplankton community in high‐nutrient conditions, as well as a large‐bodied summer community in low‐nutrient conditions. Effects were highly dependent on the initial zooplankton community structure and hence seasonal variation. In a small‐bodied community with high predation pressure and no dispersal or migration, the selective particulate‐feeding stickleback depleted the zooplankton community and decreased its diversity more radically than the cruising filter‐feeding roach. Cladocerans rather than copepods were efficiently removed by predation, and their removal caused altered patterns in rotifer abundance. In a large‐bodied summer community with initial high taxonomic and functional diversity, predation pressure was lower and resource availability was high for omnivorous crustaceans preying on other zooplankton. In this community, predation maintained diversity, regardless of predator species. During both experimental periods, predation influenced the competitive relationship between the dominant calanoid copepods, and altered species composition and size structure of the zooplankton community. Changes also occurred to an extent at the level of nontarget prey, such as microzooplankton and rotifers, emphasizing the importance of subtle predation effects. We discuss our results in the context of the adaptive foraging mechanism and relate them to the natural littoral community.     

Near-bottom depletion of zooplankton over coral reefs: III: vertical gradient of predation pressure

Many diurnal planktivorous fish in coral reefs efficiently consume zooplankton drifting in the overlying water column. Our survey, carried out at two coral reefs in the Red Sea, showed that most of the diurnal planktivorous fish foraged near the bottom, close to the shelters from piscivores. The planktivorous fish were order of magnitude more abundant near (<1.5 m) the bottom than higher in the water column. The predation pressure exerted by these fish was assessed by measuring the consumption of brine shrimps tethered at different heights above the bottom on a vertical line which was pulled over the reef. Below 1.5 m above bottom, the shrimps survival probability sharply decreased toward the bottom. Higher in the water column, survivorship was nearly 100% with little vertical variation. Our results indicate that near-bottom depletion of zooplankton in coral reefs is likely due to intense predation at that boundary layer. Risk of predation by piscivorous fish apparently restricts planktivorous fish to forage near the bottom, with a distribution pattern greatly deviating from ideal-free distribution.     

Do the effects of piscivorous largemouth bass cascade to the plankton?

Ecologists have hypothesized that an increase in the biomass of piscivorous fish in lakes will cause a decrease in populations of planktivorous fish, an increase in the size of herbivorous zooplankton and a decrease in the biomass of phytoplankton. Here we present an experimental test of whether the effects of largemouth bass (Micropterus salmoides) cascade to the planktivorous fish, zooplankton and phytoplankton of a 15-ha water storage reservoir. A pilot study indicated that the reservoir was eutrophic with dense populations of planktivorous fish dominated by threadfin shad (Dorosoma petenense). No piscovorous fish were present in the reservoir. We conducted a one-month mesocosm experiment using water and plankton from the reservoir showing that the presence of threadfin shad reduced large-sized zooplankton and increased the productivity and biomass of phytoplankton. To test whether the effects of piscivorous fish could cascade to the plankton, we assessed the effects of the addition of piscivorous largemouth bass on the planktivorous fish, zooplankton and biomass of phytoplankton of the reservoir by monitoring the reservoir during the year before and the two years after largemouth bass were stocked. In the second year after the addition of largemouth bass, the number of planktivorous fish decreased and the relative abundance of threadfin shad declined. Although the abundance of cladocerans increased after the addition of largemouth bass, the average size of zooplankton did not change. We did not detect changes in chlorophyll a, Secchi depth, or concentrations of total phosphorus and total nitrogen as a result of the addition of largemouth bass.     

Omnivory by Planktivores Stabilizes Plankton Dynamics, but May Either Promote or Reduce Algal Biomass

Classical models of phytoplankton–zooplankton interaction show that with nutrient enrichment such systems may abruptly shift from limit cycles to stable phytoplankton domination due to zooplankton predation by planktivorous fish. Such models assume that planktivorous fish eat only zooplankton, but there are various species of filter-feeding fish that may also feed on phytoplankton. Here, we extend these classical models to systematically explore the effects of omnivory by planktivorous fish. Our analysis indicates that if fish forage on phytoplankton in addition to zooplankton, the alternative attractors predicted by the classical models disappear for all realistic parameter settings, even if omnivorous fish have a strong preference for zooplankton. Our model also shows that the level of fish biomass above which zooplankton collapse should be higher when fish are omnivorous than when fish are zooplanktivorous. We also used the model to explore the potential effects of the now increasingly common practice of stocking lakes with filter-feeding fish to control cyanobacteria. Because omnivorous filter-feeding fish forage on phytoplankton as well as on the main grazers of phytoplankton, the net effect of such fish on the phytoplankton biomass is not obvious. Our model suggests that there may be a unimodal relationship between the biomass of omnivorous filter-feeding fish and the biomass of phytoplankton. This implies that to manage for reductions in phytoplankton biomass, heavy stocking or strong reduction of such fish is best.     

Cascading effects of generalist fish introduction in oligotrophic lakes

Introduction of strictly planktivorous fish to lakes can alter plankton communities via cascading interactions in food webs. Less is known about the large-scale and long-term effects resulting from the introduction of fish with more generalist feeding habits, and the extent to which these effects depend on lake trophic status. Paleolimnological records of three oligotrophic lakes in Maine, USA were used to analyze the response of plankton communities to the introduction of white perch (Morone americana), a fish that often numerically dominates fish assemblages and switches from strict planktivory to a more generalist diet during ontogeny. After white perch introduction, cladoceran ephippia size increased up to 50 %, suggesting that the most important role of this generalist fish, with respect to water quality, is as a piscivorous trophic link. Algal standing crop declined by a quarter to over a half of pre-introduction levels, suggesting that top-down effects of white perch reduced phytoplankton biomass. In comparing these oligotrophic lakes to prior work in a eutrophic system, white perch introduction had similar effects on zooplankton body size; however, cascading effects to phytoplankton were only observed in low productivity lakes.     

The influence of turbidity on growth and survival of fish larvae: a numerical analysis

Growth and survival through the early larval phase probably limit the production potential of many commercially important fish stocks. Attempts to increase the production of these stocks by restocking of juveniles have generally failed. Here, we analyse how enhanced concentrations of phytoplankton and zooplankton affect the survival of fish larvae during their early life stages. The analysis is developed for larvae feeding on copepod eggs and nauplii, with fish and invertebrates as major predators. A model of feeding and growth of fish larvae is applied to assess the benefit of enhanced phytoplankton and zooplankton abundance. The analysis shows that the shading effect of higher phytoplankton concentration may reduce predation rates on fish larvae substantially. This `top-down' effect may be more important for the cohort survivorship than the `bottom-up' mechanism in situations when larval food is sufficiently abundant. However, while increased algal biomass will improve recruitment at high zooplankton concentration, it may also reduce recruitment at low zooplankton concentrations and shallow mixing depths. Both the larvae and their vertebrate predators are dependent on light to detect their prey, and the longer reactive distance of the predators make them more susceptible than the larvae to reduced light levels and increased turbidity. We discuss the implications of reduced predation and increased zooplankton abundance on recruitment and production of fish larvae, and point at environmental conditions where changing algal biomass is likely to affect recruitment success.     

A comparison of methods for quantitative analysis of feeding selection of fishes

Synopsis Predator-prey data were obtained in a study of feeding ecology of land-locked alewife,Alosa pseudoharengus, in Claytor Lake, Virginia, and subjected to two indices of electivity as well as a nonparametric paired-comparisons statistical test. Values obtained from the two indices were often contradictory with each other and the statistical test. Attributes of the statistical procedure render it more appropriate than electivity indices to describe feeding selectivity in fish populations. Results of the study also demonstrated the importance of considering the size composition of zooplankton populations in prey preference studies; measures of selection were often positive for zooplankters 1.0 mm while negative for those of the same prey group < 1.0 mm length. Because numerous problems are inherent with in situ feeding selectivity studies of planktivorous fishes, selectivity determinations, regardless of methodology, should be construed in a relative rather than absolute context.