Predation on Mosquito Larvae by Mesocyclops thermocyclopoides (Copepoda: Cyclopoida) in the Presence of Alternate Prey

The cyclopoid copepod Mesocyclops thermocyclopoides, a dominant invertebrate predator in many shallow ponds and temporary water bodies in northern India, feeds on cladocerans, rotifers, ciliates and when present, on mosquito larvae also. We studied in the laboratory the prey consumption rates of the copepod on first and fourth instar larvae of two species of mosquito (Anopheles stephensi and Culex quinquefasciatus) in relation to their density. We also studied its prey selectivity with mosquito larvae in the presence of an alternate prey (the cladocerans‐either Moina macrocopa or Ceriodaphnia cornuta) in different proportions. With either mosquito species, the copepod actively selected Instar‐I larvae, avoiding the Instar‐IV larvae, and with either instar, selected Anopheles stephensi over Culex quinquefasciatus. When prey choice included the cladoceran as an alternate prey, the copepod selected the cladoceran only when the other prey was Instar‐IV mosquito larvae. Our results point to the potential and promise of M. thermocyclopoides as a biological agent for controlling larval populations of vectorially important mosquito species.     

Lack of prey switching and strong preference for mosquito prey by a temporary pond specialist predator

1. The strengths of trophic interactions within ecosystems can be mediated by complex mechanisms that require elucidation if researchers are to understand and predict population- and community-level stabilities. Where multiple prey types co-occur, prey switching (i.e. frequency-dependent predation) by predators may facilitate low-density prey refuge effects which promote coexistence. On the other hand, lack of switching and strong preferences by predators can strongly suppress prey populations, which is especially important considering vector species such as mosquitoes. 2. The present study quantifies prey switching and preference patterns of the temporary pond specialist copepod Lovenula raynerae towards larvae of the medically important Culex pipiens mosquito complex in the presence of different proportions of alternative Daphnia pulex prey. Further, it examines whether prey switching and preferences are contingent on the sex of the predator. 3. Lovenula raynerae exhibited a lack of prey switching and strong preference for larval mosquito prey overall, irrespective of predator sex. Also, when larval mosquitoes were available in higher proportions over daphniids, the strength of this positive selectivity increased. There was very little low-density refuge for mosquitoes where they were rare. 4. Lack of prey switching and strong preferences towards mosquitoes by predatory paradiaptomid copepods may enhance population-level regulation of disease vector mosquitoes that exploit temporary pond-style habitats. Accordingly, the conservation and promotion of these predators might enable better management of medically important species across landscapes.     

Reciprocity in predator–prey interactions: exposure to defended prey and predation risk affects intermediate predator life history and morphology

A vast body of literature exists documenting the morphological, behavioural and life history changes that predators induce in prey. However, little attention has been paid to how these induced changes feed back and affect the predators’ life history and morphology. Larvae of the phantom midge Chaoborus flavicans are intermediate predators in a food web with Daphnia pulex as the basal resource and planktivorous fish as the top predator. C. flavicans prey on D. pulex and are themselves prey for fish; as D. pulex induce morphological defences in the presence of C. flavicans this is an ideal system in which to evaluate the effects of defended prey and top predators on an intermediate consumer. We assessed the impact on C. flavicans life history and morphology of foraging on defended prey while also being exposed to the non-lethal presence of a top fish predator. We tested the basic hypothesis that the effects of defended prey will depend on the presence or absence of top predator predation risk. Feeding rate was significantly reduced and time to pupation was significantly increased by defended morph prey. Gut size, development time, fecundity, egg size and reproductive effort respond to fish chemical cues directly or significantly alter the relationship between a trait and body size. We found no significant interactions between prey morph and the non-lethal presence of a top predator, suggesting that the effects of these two biological factors were additive or singularly independent. Overall it appears that C. flavicans is able to substantially modify several aspects of its biology, and while some changes appear mere consequences of resource limitation others appear facultative in nature.     

Interactions of multiple predators with different foraging modes in an aquatic food web

Top predators can have different foraging modes that may alter their interactions and effects on food webs. Interactions between predators may be non-additive resulting from facilitation or interference, whereas their combined effects on a shared prey may result in emergent effects that are risk enhanced or risk reduced. To test the importance of multiple predators with different foraging modes, we examined the interaction between a cruising predator (largemouth bass, Micropterus salmoides) and an ambush predator (muskellunge, Esox masquinongy) foraging on a shared prey (bluegill sunfish, Lepomis macrochirus) with strong anti-predator defense behaviors. Additive and substitution designs were used to compare individual to combined predator treatments in experimental ponds. The multiple predator interaction facilitated growth of the cruising predator in the combined predator treatments, whereas predator species had substitutable effects on the growth of the ambush predator. The combined predator treatments created an emergent effect on the prey; however, the direction was dependent on the experimental design. The additive design found a risk-reducing effect, whereas the substitution design found a risk-enhancing effect for prey fish. Indirect effects from the predators weakly extended to lower trophic levels (i.e., zooplankton community). Our results highlight the need to consider differences in foraging mode of top predators, interactions between predators, and emergent effects on prey to understand food webs.     

Retarded growth of cladoceran zooplankton in the presence of a copepod predator

Various instars of four different cladoceran species representing a wide spectrum of body size were grown at high food availability in the presence and in the absence of natural densities of an invertebrate predator, a cyclopoid copepod Acanthocyclops robustus (G.O. Sars). Daily weight increments calculated from individual weights at the end and at the beginning of each 1, 2 or 4 day experiment, showed that individual growth was more or less drastically retarded in the presence of the predator as well as when exposed to water in which the predator had been feeding. The data also showed that the effect of this invertebrate predator was more pronounced in small prey instars and small prey species that were more vulnerable to predation than large prey.     

Prey and predator density‐dependent interactions under different water volumes

Predation is a critical ecological process that directly and indirectly mediates population stabilities, as well as ecosystem structure and function. The strength of interactions between predators and prey may be mediated by multiple density dependences concerning numbers of predators and prey. In temporary wetland ecosystems in particular, fluctuating water volumes may alter predation rates through differing search space and prey encounter rates. Using a functional response approach, we examined the influence of predator and prey densities on interaction strengths of the temporary pond specialist copepod Lovenula raynerae preying on cladoceran prey, Daphnia pulex, under contrasting water volumes. Further, using a population dynamic modeling approach, we quantified multiple predator effects across differences in prey density and water volume. Predators exhibited type II functional responses under both water volumes, with significant antagonistic multiple predator effects (i.e., antagonisms) exhibited overall. The strengths of antagonistic interactions were, however, enhanced under reduced water volumes and at intermediate prey densities. These findings indicate important biotic and abiotic contexts that mediate predator–prey dynamics, whereby multiple predator effects are contingent on both prey density and search area characteristics. In particular, reduced search areas (i.e., water volumes) under intermediate prey densities could enhance antagonisms by heightening predator–predator interference effects.     

Food and feeding of aquatic larvae of the midge chaoborus flavicans (meigen) (diptera : chaoboridae) in the laboratory

Laboratory observations on the feeding of fourth instar larvae of Chaoborus flavicans (Meigen) show that features relating to the prey, such as swimming behaviour, size and shape, play a major role in determining the kind of food eaten by the predator.In tests with two interacting limnetic cyclopoid copepods, Cyclops abyssorum out-competed Mesocyclops leuckarti when Chaoborus was absent but did not do so when Chaoborus was present.     

Effect of the cyclopoid copepod Mesocyclops thermocyclopoides on the interactions between the predatory rotifer Asplanchna intermedia and its prey Brachionus calyciflorus and B. angularis

In many shallow, eutrophic subtropical ponds, brachionid rotifers are common prey of the predatory copepod Mesocyclops thermocyclopoides. The predatory rotifer Asplanchna intermedia, which is itself a potential prey of the copepod, also feeds preferentially on brachionids. We studied in the laboratory the population dynamics of two mutually competing prey species, Brachionus angularis and B. calyciflorus, in the presence of the two predators A. intermedia and M. thermocyclopoides. The experimental design included separate population dynamics studies with one prey–one predator, two prey–one predator, one prey–two predator, and two prey–two predator systems. These combinations were compared with controls, in which both the prey species (B. angularis and B. calyciflorus) were grown separately and in combination with each other. In the absence of any predator, B. angularis generally eliminated the larger B. calyciflorus. Selective predation by the copepod allowed B. calyciflorus to persist longer in competition with B. angularis. Feeding by M. thermocyclopoides on A. intermedia reduced the predation pressure on B. calyciflorus. However, given enough time, the cyclopoid copepod was able to eliminate both the brachionids as well as the predatory Asplanchna.     

Chemical signals and habitat selection by three zooplankters in Andean Patagonian ponds

1. Zooplankton may react differently to chemical signals produced by macrophytes in shallow systems. They may be attracted by macrophytes, as the plants may be used as a refuge against predators, or the plants may have a repellent effect (e.g. when the plants are a habitat for numerous invertebrate predators or fish). In fishless Patagonian ponds, the structural complexity provided by macrophytes modulates the rate of predation on zooplankton by the invertebrate predator Mesostoma ehrenbergii (Turbellaria). 2. We performed a field study to analyse the coexistence of M. ehrenbergii and three of its prey (two copepods, the calanoid Boeckella gracilis and the cyclopoid Acanthocyclops robustus, and the cladoceran Ceriodaphnia dubia) in four ponds. In two of the ponds, we carried out day and night sampling to evaluate the influence of macrophytes on the distribution of these zooplankters. 3. In laboratory experiments, we analysed the response of the zooplankters to the chemical signals produced by macrophytes (the emergent Juncus pallescens and the submerged Myriophyllum quitense), the predator M. ehrenbergii and the ‘alarm signal’ provided by a homogenate of conspecifics. 4. Our field studies demonstrated the coexistence of M. ehrenbergii and the selected prey in different seasons and that A. robustus and C. dubia choose the vegetated area (a mixed bed of J. pallescens and M. quitense) over the non‐vegetated area. The habitat choice experiments indicated that the presence of M. ehrenbergii may directly affect the habitat selection of B. gracilis, because this zooplankter swam away from the predator. In addition, Mesostoma may indirectly affect the habitat selection of the cyclopoid copepod A. robustus and the cladoceran C. dubia as both zooplankters exhibited a negative response to the alarm signal produced by crushed conspecifics. 5. The presence of the submerged M. quitense did not affect the horizontal movements of any of the zooplankters studied. In contrast, the emergent macrophyte J. pallescens elicited a positive response of B. gracilis, suggesting that this aquatic plant may act as a predation refuge. 6. Our results suggest that predator avoidance behaviour can occur in fishless environments in response to a tactile invertebrate predator like Mesostoma. In addition, the refuge effect of emergent macrophytes, enhancing the survival of pelagic zooplankters, may act as a key factor in stabilizing predator–prey interactions in fishless Patagonian ponds, as has been widely recorded in northern temperate lakes with fish.     

Prey size and predator density modify impacts by natural enemies towards mosquitoes

1. Interactions between multiple predators can modify prey risk and profoundly alter ecological community dynamics. Further, ontogenic prey size changes are known to mediate prey risk through refuge effects. Understandings of these biotic factors is important for robust quantifications of natural enemy effects on target species, yet their combined influence lacks investigation. 2. Functional responses were used to quantify the predatory impacts of Notonecta glauca (water boatman; Ng) and Gammarus pulex (river shrimp; Gp) towards four different larval instars of Culex pipiens in container-style habitats. Using conspecific pairs of predators, multiple predator effects (MPEs) of both predator species were examined across larval prey sizes, and prey preference tests were applied to examine prey selectivity across predator–prey body size ratios. 3. Both predators were able to feed on C. pipiens across their larval ontogeny; however, Ng consumed significantly more larvae than Gp. Functional responses of Ng were typically Type IIs, whereas Gp trended towards sigmoidal Type IIIs. Predation by pairs of Ng and Gp showed independent MPEs towards first-, third-, and fourth-instar stages (except predation by Gp at higher densities of fourth-instar) stages, whereas, for second-instar stages, Ng showed synergistic MPEs and Gp showed antagonistic MPEs. Both predators preferred late instar mosquitoes (Ng: fourth instar; Gp: third instar). These preferences reflected predator:prey weight–length ratios, showing that relative sizes of predators and prey are important factors in prey selectivity. 4. The results obtained in the present study demonstrate that MPEs, combined with intraspecific prey preferences, may modulate trophic interactions within ecosystems. Therefore, such effects should be increasingly considered to further the understanding of agent efficacies.     

Modeling the impacts of global warming on predation and biotic resistance: mosquitoes, damselflies and avian malaria in Hawaii

Biotic resistance from native predators can play an important role in regulating or limiting exotic prey. We investigate how global warming potentially alters the strength and spatial extent of these predator–prey interactions in aquatic insect ecosystems. As a simple model system, we use rock pools in streams of rainforests of Hawaii, which contain the beautiful Hawaiian damselfly Megalagrion calliphya as predator and the invasive southern house mosquito Culex quinquefasciatus as prey. This abundant mosquito is the major vector of avian malaria transmission to native forest birds. We use mathematical modeling to evaluate the potential impacts of damselfly predation and temperature on mosquito population dynamics. We model this predator–prey system along an elevational gradient (749-1952 m elevation) and assess the effect of 1°C and 2°C climate warming scenarios as well as the effects of El Niño and La Niña oscillations, on predator–prey dynamics. Our results indicate that the strength of biotic resistance of native predators on invasive prey may decrease with increasing temperature because demographic rates of predator and prey are differentially affected by temperature. Future warming could therefore increase the abundance of invasive species by releasing them from predation pressure. If the invasive species is a disease vector, these shifts could increase the impact of disease on both humans and wildlife.     

Trait‐mediated indirect interactions in a marine intertidal system as quantified by functional responses

Studies of trait‐mediated indirect interactions (TMIIs) typically focus on effects higher predators have on per capita consumption by intermediate consumers of a third, basal prey resource. TMIIs are usually evidenced by changes in feeding rates of intermediate consumers and/or differences in densities of this third species. However, understanding and predicting effects of TMIIs on population stability of such basal species requires examination of the type and magnitude of the functional responses exhibited towards them. Here, in a marine intertidal system consisting of a higher‐order fish predator, the shanny Lipophrys pholis, an intermediate predator, the amphipod Echinogammarus marinus, and a basal prey resource, the isopod Jaera nordmanni, we detected TMIIs, demonstrating the importance of habitat complexity in such interactions, by deriving functional responses and exploring consequences for prey population stability. Echinogammarus marinus reacted to fish predator diet cues by reducing activity, a typical anti‐predator response, but did not alter habitat use. Basal prey, Jaera nordmanni, did not respond to fish diet cues with respect to activity, distribution or aggregation behaviour. Echinogammarus marinus exhibited type II functional responses towards J. nordmanni in simple habitat, but type III functional responses in complex habitat. However, while predator cue decreased the magnitude of the type II functional response in simple habitat, it increased the magnitude of the type III functional response in complex habitat. These findings indicate that, in simple habitats, TMIIs may drive down consumption rates within type II responses, however, this interaction may remain de‐stabilising for prey populations. Conversely, in complex habitats, TMIIs may strengthen regulatory influences of intermediate consumers on prey populations, whilst potentially maintaining prey population stability. We thus highlight that TMIIs can have unexpected and complex ramifications throughout communities, but can be unravelled by considering effects on intermediate predator functional response types and magnitudes. Synthesis Higher‐order predators and habitat complexity can influence behaviour of intermediate species, affecting their consumption of prey through trait‐mediated indirect interactions (TMIIs). However, it is not clear how these factors interact to determine prey population stability. Using functional responses (FRs), relating predator consumption to prey density, we detected TMIIs in a marine system. In simple habitats, TMIIs reduced consumption rates, but FRs remained de‐stabilising for prey populations. In complex habitats, TMIIs strengthened prey regulation with population stabilizing FRs. We thus demonstrate that FRs can assess interactions of environmental and biological cues that result in complex and unexpected outcomes for prey populations.     

Understanding biodiversity effects on prey in multi-enemy systems

Biodiversity–ecosystem functioning theory would predict that increasing natural enemy richness should enhance prey consumption rate due to functional complementarity of enemy species. However, several studies show that ecological interactions among natural enemies may result in complex effects of enemy diversity on prey consumption. Therefore, the challenge in understanding natural enemy diversity effects is to predict consumption rates of multiple enemies taking into account effects arising from patterns of prey use together with species interactions. Here, we show how complementary and redundant prey use patterns result in additive and saturating effects, respectively, and how ecological interactions such as phenotypic niche shifts, synergy and intraguild predation enlarge the range of outcomes to include null, synergistic and antagonistic effects. This study provides a simple theoretical framework that can be applied to experimental studies to infer the biological mechanisms underlying natural enemy diversity effects on prey.     

Predator-prey interactions and community structure: chironomids,mosquitoes and copepods in Heliconia imbricata (Musaceae)

Summary Evidence from both field observations and experimental work indicates that predation by larvae of a midge, Pentaneura n. sp. (Chironomidae), causes the low densities of mosquito larvae (Culicidae) found in the water filled bracts of Heliconia imbricata (Musaceae), microhabitats typically colonized by mosquitoes. This predation affects 2 species of mosquitoes, Wyeomyia pseudopecten, a resident species, and Trichoprosopon digitatum, a non-resident species. Predation keeps resident mosquito densities low while completely excluding the nonresident mosquito from the habitat. Both these effects of predation depend on the presence of an abundant alternative prey, an undescribed species of harpacticoid copepod found in the bracts. These copepod prey sustain chironomids when resident mosquito densities are low, permiting predator densities to remain high enough to exclude the non-resident mosquito. I discuss the evolutionary and ecological implications of predation structuring communities.     

Divergent pheromone-mediated insect behaviour under global atmospheric change

While the effects of global atmospheric changes on vegetation and resulting insect populations(‘bottom‐up interactions’) are being increasingly studied, how these gases modify interactions among insects and their natural enemies (‘top‐down interactions’) is less clear. As natural enemy efficacy is governed largely by behavioural mechanisms, altered prey finding and prey defence may change insect population dynamics. Here we show that pheromone‐mediated escape behaviours, and hence the vulnerability of insects to natural enemies, are divergent under atmospheric conditions associated with global climate change. Chaitophorus stevensis, a common aphid on trembling aspen trees, Populus tremuloides, have diminished escape responses in enriched carbon dioxide (CO₂) environments, while those in enriched ozone (O₃) have augmented escape responses, to alarm pheromone. These results suggest that divergent pheromone‐mediated behaviours could alter predator–prey interactions in future environments.     

The influence of prey behaviour on prey selection of the carnivorous plant Utricularia vulgaris

The mechanisms underlying differential prey selection of two microcrustaceans by the common bladderwort (Utricularia vulgaris) were studied in the laboratory. Functional response experiments with single prey showed that Utricularia had a higher attack rate coefficient and a longer handling time coefficient with the cladoceran Polyphemus pediculus than with the cyclopoid copepod Eucyclops serrulatus. Observation of predation rate, defined as number of prey eaten per unit time, from direct behavioural observation on single prey species, showed a higher predation rate with Polyphemus than on Eucyclops, at low prey densities. The opposite pattern was found at high prey density. When the two prey were presented simultaneously to the predator, Eucyclops was preferred over Polyphemus. Results from the situation with two prey and some of the results from the direct behavioural observations support field data on the diet of Utricularia, which shows that cyclopoid copepods are selected more frequently than Polyphemus.     

Combined impacts of warming and salinisation on trophic interactions and mortality of a specialist ephemeral wetland predator

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Different Predation Impacts of Two Cyclopoid Species on a Small-sized Zooplankton Community: An Experimental Analysis with Mesocosms

We used mesocosms to analyze predation impacts on the prey populations and prey community structures by two cyclopoid copepod species, the larger Mesocyclops pehpeiensis and the smaller Thermocyclops taihokuensis, who coexist with small-sized herbivorous zooplankton species in a fish-abundant lake. The overall predation impact on the prey populations was stronger for Mesocyclops than for Thermocyclops. Mesocyclops had a strong and less selective impact on the rotifer community but a selective impact on the crustaceans. In contrast, Thermocyclops had a selective predation impact on rotifers but a weak and less selective impact on the crustacean community. As a result, the former predator reduced the diversity of the crustacean community but not the rotifer community, while the latter had an opposite impact on the diversities of the two communities. It has been suggested that fish induce development of a zooplankton community dominated by the small-sized zooplankton species in fish-abundant lakes. Our results demonstrated that cyclopoid copepods altered species composition and diversity of the small-sized zooplankton community in such lakes. Thus, the results have given an important suggestion on the role of the invertebrate predator cyclopoid copepods, which often coexist with fish, that they determine population dynamics and community structures of small-sized zooplankton in fish-abundant lakes.     

Selective feeding of Arctodiaptomus salinus (Copepoda, Calanoida) on co-occurring sibling rotifer species

1. Using two‐ and three‐dimensional video recordings, we examined the steps involved in predation that lead to the differential vulnerability of three sympatric rotifer sibling species (Brachionus plicatilis, B. ibericus and B. rotundiformis) to a co‐occurring, predatory, calanoid copepod (Arctodiaptomus salinus). 2. Brachionus rotundiformis, the smallest prey tested, was the most vulnerable with the highest encounter rate, probability of attack, capture and ingestion, and the lowest handling time. 3. Comparison of our results with those of a previous study shows that A. salinus is a more efficient predator than a co‐occurring cyclopoid copepod (Diacyclops bicuspidatus odessanus) feeding on these same rotifer species. However, despite its higher capture rates, A. salinus seems to be less selective than D. b. odessanus based on attack distances and prey handling times. 4. The differential vulnerability to both calanoid and cyclopoid copepod predation can help explain the coexistence and seasonal succession of these co‐occurring rotifer species.     

Evidence for carry-over effects of predator exposure on pathogen transmission potential

Accumulating evidence indicates that species interactions such as competition and predation can indirectly alter interactions with other community members, including parasites. For example, presence of predators can induce behavioural defences in the prey, resulting in a change in susceptibility to parasites. Such predator-induced phenotypic changes may be especially pervasive in prey with discrete larval and adult stages, for which exposure to predators during larval development can have strong carry-over effects on adult phenotypes. To the best of our knowledge, no study to date has examined possible carry-over effects of predator exposure on pathogen transmission. We addressed this question using a natural food web consisting of the human malaria parasite Plasmodium falciparum, the mosquito vector Anopheles coluzzii and a backswimmer, an aquatic predator of mosquito larvae. Although predator exposure did not significantly alter mosquito susceptibility to P. falciparum, it incurred strong fitness costs on other key mosquito life-history traits, including larval development, adult size, fecundity and longevity. Using an epidemiological model, we show that larval predator exposure should overall significantly decrease malaria transmission. These results highlight the importance of taking into account the effect of environmental stressors on disease ecology and epidemiology.