Zooplankton responses to predation by larval bluegill: an enclosure experiment

SUMMARY 1. Larval fish are gape-limited predators that forage on prey of specific sizes, and thus may be expected to differentially affect members of a zooplankton community, possibly altering the size-structure or species composition.
2. I used an enclosure experiment to look at the effect of predation by larval bluegill on the dynamics of two zooplankton communities, one dominated by large-bodied individuals and the other by small-bodied individuals. Enclosures containing these zooplankton received a zero, low, medium, or high density of larval bluegill predators.
3. Increasing larval density had a negative effect on zooplankton abundance and abundance declined similarly in the large-bodied and small-bodied communities.
4. Zooplankton size-structure, as estimated by the length of the average zooplankton, increased and then decreased during the experiment, decreasing faster at higher larval fish densities. When zooplankton size-structure was estimated as the length of the average cladoceran, size-structure declined in the large-bodied but not in the small-bodied community and the greatest decline in size-structure was seen in the medium and high larval density treatments.
5. Ordination of each community using multidimensional scaling (MDS) indicated that the trajectory of change in species composition differed between the presence and absence of larval fish. In both communities, the degree of response by individual taxa depended on the density of bluegill larvae. This effect on zooplankton abundance, size-structure and community composition suggests that larval fish may make an important contribution to zooplankton dynamics in many lakes and ponds.     

Competition during early ontogeny: Effects of native and invasive planktivores on the growth,survival, and habitat use of bluegill

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Effects of predator-induced thinning and activity changes on life history in a damselfly

We investigated how the lethal and non-lethal presence and absence of a fish predator, perch (Perca fluviatils), influenced behaviour, numbers emerging, size at emergence, and development rate of the damselfly Lestes sponsa. The experiment was carried out in outdoor artificial ponds and spanned from the egg stage to emergence of the damselflies. During the experiment food resources for the damselflies were continuously monitored. Damselflies exposed to a lethal predator showed a significantly lower activity level than those in the absence of predators or subjected to a non-lethal predator. Half-way through the larval stage the reduction in activity level was correlated with the presence of lethal predators, and at the end of the larval stage with higher zooplankton densities. Though larvae decreased activity level, size at emergence was larger and development time faster for individuals in the lethal predator treatment. Since fewer larvae emerged from that treatment we interpret the larger size at emergence to be an effect of a combination of thinning and higher zooplankton densities.     

Influence of gizzard shad on phytoplankton size and primary productivity in mesocosms and earthen ponds in the southeastern U.S.

Gizzard shad (Dorosoma cepedianum), a filter feeding omnivore, can consume phytoplankton, zooplankton and detritus and is a common prey fish in U.S. water bodies. Because of their feeding habits and abundance, shad have the potential to affect primary productivity (and hence water quality) directly through phytoplankton grazing and indirectly through zooplankton grazing and nutrient recycling. To test the ability of shad to influence primary productivity, we conducted a 16-day enclosure study (in 2.36-m³ mesocosms) and a 3-year whole-pond manipulation in 2–5 ha earthen ponds. In the mesocosm experiment, shad reduced zooplankton density and indirectly enhanced chlorophyll a concentration, primary productivity, and photosynthetic efficiency (assimilation number). While shad did not affect total phytoplankton density in the mesocosms, the density of large phytoplankton was directly reduced with shad. Results from the pond study were not consistent as predicted. There were few changes in the zooplankton and phytoplankton communities in ponds with versus ponds without gizzard shad. One apparent difference from systems in which previous work had been conducted was the presence of high densities of a potential competitor (i.e., larval bluegill) in our ponds. We suggest that the presence of these extremely high larval bluegill densities (20–350 larval bluegill m^–3; 3–700 times higher density than that of larval gizzard shad) led to the lack of differences between ponds with versus ponds without gizzard shad. That is, the influence of gizzard shad on zooplankton or phytoplankton was less than the influence of abundant bluegill larvae. Differences in systems across regions must be incorporated into our understanding of factors affecting trophic interactions in aquatic systems if we are to be able to manage these systems for both water quality and fisheries.     

Spatial and temporal niche partitioning in grassland ants

We examined whether the species composition of a community influences the persistence of larval Ambystoma maculatum in assemblages composed of two larger intraguild predators (A. opacum and A. jeffersonianum) and an alternative prey species (tadpoles of Rana sylvatica). We predicted a priori that A. maculatum would have higher survival in more diverse communities containing alternative species of prey and top predators (A. opacum), the latter of which may lower the abundance of intermediate predators (A. jeffersonianum) via intraguild predation. In a factorial experiment, we manipulated the presence of larval A. opacum, A. jeffersonianum, and R. sylvatica in replicated artificial ponds containing larval A. maculatum. The presence of all three species significantly depressed biomass production in A. maculatum: biomass was highest in ponds lacking the other species and was zero in ponds initially containing all four species. Tadpoles severely reduced the growth of filamentous algae in the ponds. This, in turn, may have affected the abundance of some herbivorous prey of larval salamanders, although this possibility was not tested. The presence of congeneric predators severely restricted the presence in the water column of larval A. maculatum, which otherwise exhibited significant diel patterns of activity in the absence of predators. Together, the presence of tadpoles and a predator-mediated reduction in activity patterns may have limited foraging opportunities for A. maculatum, thus exacerbating the direct impact of predation on survival in this species. These results suggest that diverse assemblages consisting of these particular species may actually inhibit, rather than promote, inclusion of A. maculatum in some communities of pond-breeding amphibians.     

Chemical cues from multiple predator-prey interactions induce changes in behavior and growth of anuran larvae

Chemical signals are used as information by prey to assess predation risk in their environment. To evaluate the effects of multiple predators on prey growth, mediated by a change in prey activity, I exposed small and large bullfrog (Rana catesbeiana) larvae (tadpoles) to chemical cues from different combinations of bluegill sunfish (Lepomis macrochirus) and larval dragonfly (Anax junius) predators. Water was regularly transferred from predation trials (outdoor experiment) to aquaria (indoor experiment) in which activity and growth of tadpoles was measured. The highest predation mortality of small bullfrog larvae in the outdoor experiment was due to Anax, and it was slightly lower in the presence of both predators, probably resulting from interactions between predators. There was almost no mortality of prey with bluegill. The activity and growth of small bullfrog larvae was highest in the absence of predators and lowest in the presence of Anax. In the presence of bluegill only, or with both predators, the activity and growth of small bullfrog tadpoles was intermediate. Predators did not affect large tadpole activity and growth. Regressing mortality of small bullfrog tadpoles against activity and growth of bullfrog tadpoles revealed a significant effect for small bullfrog larvae but a non-significant effect for large bullfrog larvae. This shows that the response of bullfrog tadpoles to predators is related to their own body size. The experiment demonstrates that chemical cues are released both as predator odor and as alarm substances and both have the potential to strongly alter the activity and growth of prey. Different mechanisms by which chemical cues may be transmitted to species interactions in the food web are discussed. Received: 28 June 1999 / Accepted: 15 November 1999     

Zooplankton community structure in the kelp beds of the sub-Antarctic Prince Edward Archipelago: are they a refuge for larval stages?

Kelp beds are thought to provide a refuge for a variety of marine organisms by damping waves, changing hydrodynamic flow, offering substrata for epiphytic species, and altering the abundances of predators and prey. At the sub-Antarctic Prince Edward Islands, a typical flow-through system, kelp beds have the potential of acting as a prime habitat for a variety of pelagic species that would otherwise be swept away and lost from the island ecosystem. The current study investigates the effects of kelp beds on the abundance and composition of zooplankton and microphytoplankton, with the main aim of determining whether kelp beds act as a repository for larval stages in general, and for the swimming prawn, Nauticaris marionis, in particular. Salinity gradients created by freshwater run-off indicated that kelp beds at Marion Island retain surface water and restrict water exchange with the open ocean. Both the abundance and diversity of zooplankton were greatest within the kelp beds. This was partially due to the high abundance of kelp-associated species, but also to the apparent retention of naupliar larvae within the kelp canopy. Unlike the larvae of ostracods and copepods, caridean zoeae were rare, indicating that kelp beds are not a prime habitat for larvae of N. marionis. The larval retention mechanism for this species, therefore, remains unknown. Apart from retaining water and certain species of zooplankton, chlorophyll a/phaeopigment ratios confirmed the existence of a substantial detrital pool within the kelp beds. This retention of detritus may explain the observed importance of kelp-derived matter in the diet of many nearshore animals. In contrast to the zooplankton, kelp beds seemed to have little effect on the abundance and composition of microphytoplankton, and chlorophyll a values were low at all stations.     

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.     

Are classical predator-prey models relevant to the real world?

Mathematical models of predator-prey population dynamics are widely used for predicting the effect of predators as biocontrol agents, but the assumptions of the models are more relevant to parasite-host systems. Predator-prey systems, at least in insects, substantially differ from what is assumed by these models. The main differences are: (i) Juveniles and adults have to be considered as two different entities, as the former stay within a patch and do not reproduce, while the latter move between patches of prey and reproduce there. (ii) Because of their high mobility, food availability is likely to be less restrictive for adults than juveniles, which are confined to one patch. Therefore, a functional response to prey abundance may not be important for adults. (iii) Egg and larval cannibalism are common in insect predators. Therefore, the quality of patches of prey for their larvae determines the reproductive strategy of adult predators more than the availability of food for the adults. Here we develop a new model, based on the above considerations, which is suitable for modelling these interactions. We show that selection should favour mechanisms that enable predators to avoid reproducing in patches with insufficient prey and those already occupied by predators.     

The effects of fish on assemblages of amphibians in ponds: a field experiment

1. Bluegill sunfish ( Lepomis macrochirus ) dominate fish assemblages of small lakes and ponds throughout the eastern United States and may play a major role in structuring aquatic communities. We examined the impact of adult bluegill on amphibian density by stocking bluegill at a range of densities into partitions of an experimental pond in which amphibians were free to colonize.
2. Adult bluegill had a major impact on the amphibian assemblage. By the end of the experiment, gray treefrog ( Hyla versicolor ) tadpoles were nine times less abundant, and red-spotted newt ( Notophthalmus viridescens ) adults were three times less abundant in the presence of adult bluegill than in their absence. In contrast, bullfrog ( Rana catesbeiana ) tadpoles tended to increase in the presence of adult bluegill. Adult bluegill also had a negative effect on the abundance of predaceous aquatic insects.
3. There was no indication that interactions among amphibians were significant in determining the above patterns. We suggest that the strong impact of adult bluegill resulted from a combination of direct and indirect effects on amphibian larvae and predaceous aquatic insects.     

Use of prey and non-prey food by the ladybird beetle Eriopis connexa (Coleoptera: Coccinellidae) under laboratory-rearing conditions

Prey for predators can fluctuate in abundance and in quality over time requiring predator strategies to cope with food shortage. Coccinellinae are often associated with sap-sucking pests that exhibit high population unpredictability such as aphids and psyllids. Eriopis connexa (Germar) (Coleoptera: Coccinellidae) is a predator with potential for biological control, especially a well-studied population which is resistant to pyrethroids used to control insect defoliators. Both larvae and adult E. connexa were provided ad libitum prey and non-prey foods (pollen and honey water solution) at increasing intervals from 1 to 10 days. Neonate larvae of E. connexa required eating prey daily to develop into adults. However, non-prey food such as honey water solution did prolong larval and adult survival but neither fulfilled larval development nor adult reproduction. Honey water solution promoted 100% adult survival up to 25 days in the adult stage without prey with oviposition returning after daily feeding on prey. Females subjected to increased feeding intervals over four days reduced oviposition and lived longer, but 10-day feeding intervals correlated with risk to adult survival. These results indicate the importance of non-food sources in E. connexa maintenance and the ability of larvae and adult females to compensate for prey scarcity.     

Separating the effects of intra- and interspecific age-structured interactions in an experimental fish assemblage

We documented patterns of age-structured biotic interactions in four mesocosm experiments with an assemblage of three species of co-occurring fishes from the Florida Everglades, the eastern mosquitofish (Gambusia holbrooki), sailfin molly (Poecilia latipinna), and bluefin killifish (Lucania goodei). These species were chosen based on their high abundance and overlapping diets. Juvenile mosquitofish and sailfin mollies, at a range of densities matching field estimates, were maintained in the presence of adult mosquitofish, sailfin mollies, and bluefin killifish to test for effects of competition and predation on juvenile survival and growth. The mesocosms held 1,200 l of water and all conditions were set to simulate those in Shark River Slough, Everglades National Park (ENP), USA. We placed floating mats of periphyton and bladderwort in each tank in standard volumes that matched field values to provide cover and to introduce invertebrate prey. Of 15 possible intra- and interspecific age-structured interactions, we found 7 to be present at the densities of these fish found in Shark River Slough marshes. Predation by adult mosquitofish on juvenile fish, including conspecifics, was the strongest effect observed. We also observed growth limitation in mosquitofish and sailfin molly juveniles from intra- and interspecific competition. When maintained at high densities, juvenile mosquitofish changed their diets to include more cladocerans and fewer chironomid larvae relative to low densities. We estimated size-specific gape limitation by adult mosquitofish when consuming juvenile mosquitofish and sailfin mollies. At high field densities, intraspecific competition might prolong the time period when juveniles are vulnerable to predation by adult mosquitofish. These results suggest that path analysis, or other techniques used to document food-web interactions, must include age-specific roles of these fishes.     

The effect of invertebrate and vertebrate predators on the foraging movements of Ischnura elegans larvae (Odonata: Zygoptera)

SUMMARY.

• 1 The foraging movements of late instar Ischnura elegans larvae were monitored in laboratory experiments to study the effects of predators on larval feeding behaviour.
• 2 Ischnura larvae are sit-atid-wait, or ambush, foragers, moving occasionally between perches in search of profitable feeding sites. Larval foraging movements, monitored at different densities of Daphnia prey, increased significantly when prey were absent.
• 3 In experiments without prey, larval movement was inhibited by the presence of fish predators, as well as by invertebrate predators (Notonecta glauca), but not by closely related, non-predatory invertebrates (Corixa punctata) or physical disturbance of the water (intermittent air bubbles).
• 4 Further experiments varied Ischnura hunger levels (0–8 days without food) and illumination (light or dark) with and without notonectid predators. Hunger had no consistent effect on penultimate instar behaviour but final instar foraging activity was significantly modified: movements increased after 4 days starvation and decreased again after 8 days. This response was suppressed by the presence of predators. Both larval instars moved significantly less often in the light, even when predators were absent.
• 5 These phenotypically flexible predator-avoidance responses are likely to decrease the risk of predation by both visual and tactile predators. However, predators clearly have an important influence on the feeding niche of Ischnura larvae, and may decrease the overall feeding efficiency, growth rate, and survival of larvae by constraining their movement in search of profitable feeding sites.

     

Propensity towards cannibalism among Hypoaspis aculeifer and H. miles, two soil-dwelling predatory mite species

In biological control programmes, the two predatory soil mites Hypoaspis aculeifer and H. miles are often applied against soil-borne pests like mushroom flies, springtails and mites. Although the mites show high consumption rates on varying prey types in Petri dish experiments and in greenhouses, their overall efficiency is sometimes limited. We hypothesized that intraspecific interactions, like cannibalism, could contribute to this decreased competence. Therefore, experiments were conducted to show the propensity of H. aculeifer and H. miles to cannibalise. Adult mites and nymphs were introduced as predators with conspecific eggs, larvae, nymphs, adult females or males as prey and the number of killed individuals was recorded. Additionally, the oviposition rate on conspecific prey was quantified and the correlation with the number of prey consumed was calculated to assess the influence of cannibalism on egg production. The results illustrate that cannibalism occurs infrequently in both Hypoaspis spp., the only exception being H. aculeifer nymphs, which cannibalised one conspecific egg per day. Moreover, cannibalism never occurred in the presence of alternative prey. Oviposition rate decreased during the experiment in both species but it was positively correlated with the cannibalism rate only for H. aculeifer. The benefit of cannibalism for populations of H. aculeifer and H. miles is discussed.     

Predator evasion in <Emphasis Type="Italic">Daphnia</Emphasis>: the adaptive value of aggregation associated with attack abatement

To evaluate the adaptive value of daphnian aggregation associated with attack abatement, a combined laboratory and field experiment was performed. As the lake investigated does not contain planktivorous fish, only invertebrate predation is important. In the laboratory, newly collected Chaoborus flavicans, among the most important predators to exploit the Daphnia population in the lake, were individually placed in a spherical flask containing different densities of juvenile Daphnia prey. The number of successful attacks of Chaoborus was independent of prey abundance. Thus, in a given density regime, each individual prey gained protection from a dilution effect. In the field, dense aggregates of Daphnia were created by continuously adding food at a localized point in the littoral. Even though the food input produced aggregates of Daphnia which were maintained for 3 h, the number of invertebrate predators did not increase in the input area. It is concluded that each Daphnia in a group reduces the probability of predation through an effect of attack abatement which is approximately proportional to the number of prey in the group. However, when considering the high median density of Daphnia, together with their strong response to food input, it seems more plausible that natural aggregates are driven by a patchy food distribution, whereas attack abatement is only a secondary effect of aggregation.     

The Use of Conspecific Chemical Cues for Cannibal Avoidance in California Newts (Taricha torosa)

Adult California newts, Taricha torosa (Caudata: Salamandridae), prey on both conspecific egg masses and larvae. We used both laboratory and field experiments to determine if newt larvae use conspecific chemical cues to avoid adult intraspecific predators. Larvae showed significant hiding responses when exposed to adult cues in both the field and the laboratory. Larvae did not exhibit significant hiding responses to chemical cues from invertebrate predators.     

Effects of alternative food on cannibalism and herbivore suppression by carabid larvae

1. Predator and alternative food density are important factors influencing herbivore suppression by generalist predators. Herbivore suppression can be reduced if predators forage preferentially on alternative foods. Cannibalism can increase at high predator densities, further reducing herbivore suppression. However, complex interactions are possible, as alternative food can increase predator abundance and survival restoring top‐down effects on herbivores. 2. In two species of carabid larvae (Poecilus chalcites and Anisodactylus ovularis), we studied how alternative foods (fly pupae and grass seeds) and predator density affect predation of black cutworm larvae and how alternative foods affect cannibalism among carabid larvae. 3. Adding alternative food to microcosms generally reduced total predation of cutworms. However, the strength of this effect was dependent on carabid species, larval density, and food type. 4. Increasing larval density from one to three per microcosm reduced per‐capita predation by both species irrespective of alternative food treatment. 5. Alternative food reduced cannibalism in both carabid species and increased survival of carabid larvae in field plots, such that twice as many were captured in plots subsidised with pupae than plots with no alternative food. 6. These results provide new insight into the complex interactions that influence predator survival and herbivore suppression in resource diverse habitats by demonstrating the primacy of intraguild interactions among carabid larvae.     

Threat of predation alters aggressive interactions among spotted salamander (Ambystoma maculatum) larvae

Intraspecific aggression represents a major source of mortality for many animals and is often experienced alongside the threat of predation. The presence of predators can strongly influence ecological systems both directly by consuming prey and indirectly by altering prey behavior or habitat use. As such, the threat of attack by higher level predators may strongly influence agonistic interactions among conspecifics via nonconsumptive (e.g., behaviorally mediated) predator effects. We sought to investigate these interactions experimentally using larval salamanders (Ambystoma maculatum) as prey and dragonfly nymphs (Anax junius) as predators. Specifically, we quantified salamander behavioral responses to perceived predation risk (PPR) from dragonfly nymphs and determined the degree to which PPR influenced intraspecific aggression (i.e., intraspecific biting and cannibalism) among prey. This included examining the effects of predator exposure on the magnitude of intraspecific biting (i.e., extent of tail damage) and the resulting change in performance (i.e., burst swim speed). Salamander larvae responded to PPR by reducing activity and feeding, but did not increase refuge use. Predator exposure did not significantly influence overall survival; however, the pattern of survival differed among treatments. Larvae exposed to PPR experienced less tail damage from conspecifics, and maximum burst swim speed declined as tail damage became more extensive. Thus, escape ability was more strongly compromised by intraspecific aggression occurring in the absence of predation risk. We conclude that multitrophic indirect effects may importantly modulate intraspecific aggression and should be considered when evaluating the effects of intraspecific competition.     

An experimental study of population regulation in the salamander,Notophthalmus viridescens dorsalis (Urodela: Salamandridae)

Summary The roles of density-dependent larval survival and cannibalism of larvae as potential mechanisms of population regulation in the newt (Notophthalmus viridescens dorsalis) were evaluated in laboratory and field experiments. In laboratory containers, adults cannibalized larvae and large larvae cannibalized smaller larvae. In artificial ponds, larval survival did not depend on initial larval density. No cannibalism could be demonstrated in the complex environment, although the experiment was powerful enough to detect an ecologically relevant difference in survival. Adult growth was negatively correlated with the final biomass of larval newts, suggesting that the two life stages competed for resources. Larval growth rates were negatively correlated with final larval density, suggesting that larvae competed with each other. The proportion of larvae that became sexually mature at age 7 months (paedomorphs and adults that skipped the eft stage) varied inversely with larval density. Therefore, the potential regulatory mechanisms identified in this study are competition within and between life stages.     

Priority effects of the early breeding fire salamander on the late breeding banded newt

Early breeding intraguild predators may have advantages over late breeding predators via priority effects; early breeding predators may reduce shared prey resources before late breeders appear and may also prey upon the late breeders. Here we show that predatory larvae of the late-breeding predatory banded newt, Triturus vittatus vittatus, occupy the same temporary pond toward the end of the developmental period of the early-breeding predatory fire salamander, Salamandra salamandra, resulting in a large size disparity between larvae of these two species while they co-occur. We conducted outdoor artificial pool experiments to assess priority effects of large larval Salamandra at the end of their larval development period, on recently hatched larval Triturus. We also assessed how artificial vegetation may influence larval Triturus performance in the presence or absence of Salamandra Salamandra, introduced into the experimental pools two weeks prior to the newt larvae, strongly reduced invertebrate prey abundance shared by these two predatory urodeles and with only a one week period of overlap, strongly reduced abundance of Triturus larvae. The artificial vegetation had only a small ameliorating effect on Triturus survival when Salamandra was present. Triturus size at metamorphosis (snout-tail length) was significantly larger in the Salamandra pools, presumably due to a combination of a strong “thinning effect” and greater vulnerability of smaller Triturus individuals to predation by Salamandra. Time to metamorphosis was not significantly affected by Salamandra. These results have conservation implications as T. v. vittatus is listed as highly endangered and may also explain the largely negative spatial association of the two species. Handling editor: K. Martens