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.     

Predation by odonates depresses mosquito abundance in water-filled tree holes in Panama

In the lowland moist forest of Barro Colorado Island (BCI), Panama, larvae of four common species of odonates, a mosquito, and a tadpole are the major predators in water-filled tree holes. Mosquito larvae are their most common prey. Holes colonized naturally by predators and prey had lower densities of mosquitoes if odonates were present than if they were absent. Using artificial tree holes placed in the field, we tested the effects of odonates on their mosquito prey while controlling for the quantity and species of predator, hole volume, and nutrient input. In large and small holes with low nutrient input, odonates depressed the number of mosquitoes present and the number that survived to pupation. Increasing nutrient input (and consequently, mosquito abundance) to abnormally high levels dampened the effect of predation when odonates were relatively small. However, the predators grew faster with higher nutrients, and large larvae in all three genera reduced the number of mosquitoes surviving to pupation, even though the abundance of mosquito larvae remained high. Size-selective predation by the odonates is a likely explanation for this result; large mosquito larvae were less abundant in the predator treatment than in the controls. Because species assemblages were similar between natural and artificial tree holes, our results suggest that odonates are keystone species in tree holes on BCI, where they are the most common large predators. Received: 4 November 1996 / Accepted: 11 April 1997     

Effect of habitat complexity on the predation of Buenoa fuscipennis (Heteroptera: Notonectidae) on mosquito immature stages and alternative prey

Notonectids are well‐known predators in aquatic habitats, where mosquito larvae, chironomids, and cladocerans constitute their main diet. Our purpose was to assess the effect of structural complexity on the predatory ability of Buenoa fuscipennis, a common predator in aquatic habitats of Buenos Aires city (Argentina). Buenoa fuscipennis showed type 2 functional responses in both the presence and absence of prey refuge and no differences in attack rate or handling time between refuge treatments. Regarding mosquito size classes, B. fuscipennis exhibited a significantly higher preference for 2^nd instar larvae and no predation on pupae. In the presence of mosquito larvae and alternative prey, B. fuscipennis preferred mosquitoes over chironomid larvae and adult cladocerans over mosquito larvae. No switching behavior was detected in our experiments. Habitat structure only slightly affected the predator´s consumption rates on mosquito larvae. Overall, preference for prey did not vary with the presence of refuge, except for the preference for mosquitoes over chironomid larvae, which was significantly decreased in the presence of refuge as a consequence of reduced predation on mosquito larvae. The results suggest that B. fuscipennis could efficiently control mosquitoes in structurally simple habitats where chironomids are the most abundant alternative prey but not in temporary pools where cladocerans are abundant.     

Opportunistic foraging by heteropteran mosquito predators

Tropical aquatic environments host a large number of predatory insects including heteropteran water bugs Anisops bouvieri Kirkaldy, 1704 (Heteroptera: Notonectidae), Diplonychus (=Sphaerodema) rusticus Fabricius, 1781 (Heteroptera: Belostomatidae), and Diplonychus (=Sphaerodema) annulatus Fabricius, 1781 (Heteroptera: Belostomatidae) feeding on a range of organisms. In tropical and subtropical wetlands, ponds, and temporary pools these predators play a role in regulation of dipteran populations, particularly mosquitoes and chironomids. Their relative abilities to control mosquitoes depend in part on predator preference for mosquitoes in relation to other natural prey, and the predators’ propensities to switch to mosquitoes as mosquito density increases. The prey electivity and switching dynamics of these predatory water bugs were evaluated in the laboratory under various prey densities, using two instars (II and IV) of chironomid and mosquito larvae as prey. Studies of electivity at relatively high densities (20 prey L⁻¹) in small (5 L) vessels demonstrated that all predators showed opportunistic foraging as the mosquito:chironomid ratio changed, with some evidence that mosquito larvae were positively selected over chironomids. In particular, Anisops showed strong electivity for mosquitoes when presented with any ratio of large mosquito and chironomid prey in the high density experiment, although the preference was not expressed in lower density (2.5 prey L⁻¹) treatments executed in 40 L vessels. In these lower density treatments, D. rusticus demonstrated higher electivity for mosquitoes when the mosquito:chironomid ratio was high, consistent with non-significant trends observed in the higher density experiment. The positive electivity of D. rusticus for mosquitoes was reinforced in an experiment executed over 16 days at varying prey ratios, in which D. rusticus mosquito electivity was high and consistent while D. annulatus showed slight avoidance of mosquito larvae, and Anisops remained largely opportunistic in foraging on prey in proportion with availability. Anisops and D. rusticus are potentially good biocontrol agents for mosquito larvae, in that they preferentially consume mosquitoes under many circumstances but can readily forage on other prey when mosquito density is low.     

Evidence for predatory flatworms as organizers of zooplankton and mosquito community structure in rice fields

I present evidence from two field studies and one laboratory experiment that the predatory flatworm, tentatively indentified as Mesostoma nr. lingua, can be important in organizing invertebrate communities in rice fields. In the first field study, I assessed the importance of various predators including two fishes, in explaining among-plot variation in mosquito abundance during a five week interval. Mosquitofish (Gambusia affinis) abundance was not significantly associated with densities of either mosquito, Culex tarsalis or Anopheles freeborni, except for the final week. Associations between green sunfish (Lepomis cyanellus) juveniles and mosquitoes were demonstrated for only A. freeborni — a positive association. Densities of Mesostoma were negatively associated with C. tarsalis and A. freeborni densities. C. tarsalis, protected from flatworms in small-mesh cages, had a much greater survival rate than those that were exposed in large-mesh cages. C. tarsalis survival, when left unprotected in large-mesh cages, was negatively correlated with field densities of flatworms and with the number of flatworms found inside the cages. Survival was also positively correlated with field densities of mosquitoes.In the second field study, Mesostoma densities were negatively correlated with C. tarsalis, A. freeborni, Cladocera and Ostracoda in rice field enclosures early in the season. Cyclopoid copepod densities were not significantly associated with this predator.In a laboratory experiment, predation by Mesostoma on C. tarsalis larvae was reduced when presented with an alternative prey, Moina micrura (a cladoceran).These results suggest that flatworms, via predation, can have large effects on the structure of mosquito and zooplankton communities in rice fields.     

A parasite's modification of host behavior reduces predation on its host

Parasite modification of host behavior is common, and the literature is dominated by demonstrations of enhanced predation on parasitized prey resulting in transmission of parasites to their next host. We present a case in which predation on parasitized prey is reduced. Despite theoretical modeling suggesting that this phenomenon should be common, it has been reported in only a few host–parasite–predator systems. Using a system of gregarine endosymbionts in host mosquitoes, we designed experiments to compare the vulnerability of parasitized and unparasitized mosquito larvae to predation by obligate predatory mosquito larvae and then compared behavioral features known to change in the presence of predatory cues. We exposed Aedes triseriatus larvae to the parasite Ascogregarina barretti and the predator Toxohrynchites rutilus and assessed larval mortality rate under each treatment condition. Further, we assessed behavioral differences in larvae due to infection and predation stimuli by recording larvae and scoring behaviors and positions within microcosms. Infection with gregarines reduced cohort mortality in the presence of the predator, but the parasite did not affect mortality alone. Further, infection by parasites altered behavior such that infected hosts thrashed less frequently than uninfected hosts and were found more frequently on or in a refuge within the microcosm. By reducing predation on their host, gregarines may be acting as mutualists in the presence of predation on their hosts. These results illustrate a higher‐order interaction, in which a relationship between a species pair (host–endosymbiont or predator–prey) is altered by the presence of a third species.     

Tadpoles of three common anuran species from Thailand do not prey on mosquito larvae

Tadpoles are often considered to be predators of mosquito larvae and are therefore beneficial for the control of certain disease vectors. Nevertheless, only a few species have actually been recorded to prey on mosquito larvae. The mosquito larvae predation rates of tadpoles of three common Thai anuran species (Bufo melanostictus, Kaloula pulchra and Hylarana raniceps) were experimentally tested. Tadpoles in varying developmental stages were used to assess a size/age effect on the predation rate. In addition, different instars of Culex quinquefasciatus were used in order to assess a prey size effect on the predation rates. All three species failed to show any evidence of mosquito larvae predation. Neither small nor large tadpoles fed on mosquito larvae. Prey size also did not affect predation. Although tadpoles do not feed on mosquito larvae, there may be other direct or indirect inter‐specific interactions that adversely impact the development of larvae in shared habitats with tadpoles.     

Effects of predation by the copepod Mesocyclops ogunnus on the sex ratios of mosquito Aedes albopictus

Cyclopoid copepods are important predators in many aquatic ecosystems and have been used as biological agents in successful programs to control mosquito larvae. However, the impacts of this predation on adult mosquito populations are still poorly understood. The present study compared the sex ratios and body sizes (measured as wing length) of Aedes albopictus mosquitoes emerging from recipients containing the copepod predator Mesocyclops ogunnus with control situations without this predator. We found that copepod predation significantly biased mosquito sex ratios toward females, and that both the males and females emerging from copepod-containing recipients were significantly larger than control insects. The ecological and epidemiological consequences of the changes induced by copepod predation on mosquito populations are discussed.     

The role of priority effects in limiting the success of the invasive tiger mosquito,Aedes albopictus

Priority effects (PE), wherein species colonizing a habitat early have a negative impact on later colonizers, can have profound and legacy effects on community organization. In temperate zones, larval mosquito habitats are emptied each year in the winter and recolonized in the spring. There are phenological differences among common species but the role of PE in these communities is largely unexplored. Aedes albopictus, the invasive tiger mosquito, is considered a superior competitor to resident species during the larval phase when conditions are initiated with same-staged heterospecific larvae. However in nature, Ae. albopictus hatches, and resumes activity, later in the spring than other species, suggesting it encounters larger later developed individuals, and denser populations, of species such as Aedes triseriatus. Additionally, despite their competitive inferiority, these species often coexist with Ae. albopictus in larval habitats, with Ae. albopictus often occurring at relatively low abundances in sylvan habitats. Using lab and near field experiments, we tested the hypothesis that PE with early hatching species reduces survivorship and population growth for the invasive Ae. albopictus. When Ae. albopictus larvae encountered larger, later developed heterospecific larvae at greater densities, under controlled lab conditions and in artificial and natural mesocosms, they experienced significant reductions in survival and estimated finite rate of population increase. Additionally, we found that intraguild predation of Ae. triseriatus on Ae. albopictus may be an important mechanism through which PE works. We conclude that PE is a potential mechanism for coexistence between invasive and resident mosquitoes and should be further explored.

     

Experimental analysis of the predation impact of the larvae of pond smelt (<Emphasis Type="Italic">Hypomesus transpacificus nipponensis</Emphasis>) on zooplankton populations established in mesocosms

The predation impact of the larvae of pond smelt Hypomesus transpacificus nipponensis on a zooplankton community was studied using mesocosms. The fish significantly depressed the abundances of copepod nauplii and rotifers, especially Hexarthra mira. The vulnerabilities of these prey might be determined by their swimming behavior and population density, suggesting that larval fish selectively prey on zooplankton that have a high encounter rate with the predator. The larvae did not have a negative effect on the densities of cladocerans, but fish predation altered the cladoceran community structure from the dominance of B. longirostris to that of B. fatalis. This result suggests that larval fish predation is an important factor that shifts the species composition of Bosmina in some lakes, the shift occurring in the season when fish larvae are abundant. Our results have shown that predation by the larval fish would control not only the abundance, but also the community structure of the small-sized zooplankton prey.     

Keystone nonconsumptive effects within a diverse predator community

The number of prey killed by diverse predator communities is determined by complementarity and interference among predators, and by traits of particular predator species. However, it is less clear how predators' nonconsumptive effects (NCEs) scale with increasing predator biodiversity. We examined NCEs exerted on Culex mosquitoes by a diverse community of aquatic predators. In the field, mosquito larvae co‐occurred with differing densities and species compositions of mesopredator insects; top predator dragonfly naiads were present in roughly half of surveyed water bodies. We reproduced these predator community features in artificial ponds, exposing mosquito larvae to predator cues and measuring resulting effects on mosquito traits throughout development. Nonconsumptive effects of various combinations of mesopredator species reduced the survival of mosquito larvae to pupation, and reduced the size and longevity of adult mosquitoes that later emerged from the water. Intriguingly, adding single dragonfly naiads to ponds restored survivorship of larval mosquitoes to levels seen in the absence of predators, and further decreased adult mosquito longevity compared with mosquitoes emerging from mesopredator treatments. Behavioral observations revealed that mosquito larvae regularly deployed “diving” escape behavior in the presence of the mesopredators, but not when a dragonfly naiad was also present. This suggests that dragonflies may have relaxed NCEs of the mesopredators by causing mosquitoes to abandon energetically costly diving. Our study demonstrates that adding one individual of a functionally unique species can substantially alter community‐wide NCEs of predators on prey. For pathogen vectors like mosquitoes, this could in turn influence disease dynamics.     

Typhloplanid flatworms (Mesostoma and related genera): Mechanisms of predation and evidence that they structure aquatic invertebrate communities

Mesostoma species and other typhloplanid flatworms are predators found in a wide variety of habitats. Laboratory observations of some Mesostoma spp found in shallow aquatic habitats (referred to as littoral species) have revealed a wide variety of prey killing mechanisms: 1) mucus trapping, 2) sit-and-wait predation, 3) releasing a toxin into the water, and 4) active searching. We review the existing literature on these mechanisms. We also describe for the first time the predatory behavior of a pelagic Mesostoma sp. found in Brazilian lakes.The existing literature is also reviewed to assess the potential impact of Mesostoma species and related genera on the aquatic invertebrate community. Mesostoma has a high potential for population increase and has been found in high densities in some shallow aquatic habitats. Single prey experiments show that a number of Mesostoma species feed heavily on mosquito larvae, some chironomid larvae and some daphnids but considerably less on most copepods and ostracods. Prey preference experiments reflect the same trends. Hence, these predation studies suggest that the flatworms, at high densities, should reduce populations of certain prey speces and, consequently, alter community structure. Field studies support this prediction. Mesostoma species, at high densities, appear to be important predators of mosquito larvae in shallow aquatic habitats even under conditions where high densities of planktivorous fishes had little impact. They also appear to alter zooplankton community structure. However, much of the field evidence is correlational and not experimental. No studies have assessed the impact of pelagic typhloplanids on zooplankton community structure.It is concluded that many more manipulative field experiments are needed to assess the impact of typhloplanids on aquatic invertebrate communities.     

Behavioural responses to predation may explain shifts in community structure

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Selective predation by larval Agabus (Coleoptera: Dytiscidae) on mosquitoes: support for conservation-based mosquito suppression in constructed wetlands

1. Wetland insect predators can structure aquatic prey communities via selective predation, but receive considerably less attention than vertebrate predators. We conducted laboratory experiments to test selective predation by two species of larval dytiscid beetles ( Agabus ; Coleoptera: Dytiscidae) and the potential contribution of these beetles to suppression of mosquito populations in constructed wetlands.
2.  Agabus consumed copepods, ostracods and mosquito larvae in no-choice tests. When offered a choice, 76% of all prey consumed were mosquito larvae, indicating selective predation. Subsequent experiments revealed this preference was due to ease of capture of mosquito larvae over alternative prey.
3. Cannibalism and intraguild predation were common within and between species of Agabus , which may reduce the overall impact of the observed selective predation.
4.  Agabus larvae selectively preyed on mosquito larvae over alternative prey, which is not characteristic of some fish used as biological control agents for mosquitoes. Predator exclusion or similar experiments in the field could document how these results translate into a natural setting.
5. The findings of this study suggest developing mosquito suppression strategies focused on conservation of native wetland predators. These strategies are preferable to introducing non-native generalist predators, or applying pesticides.     

Effect of water depth on predation frequency by diving beetles on mosquito larvae prey

Diving behavior and its frequency may differ among species of mosquito larvae because of differences in predation pressure. The present study aimed to investigate the relationship between water depth and predation frequency on two mosquito species, Culex tritaeniorhynchus (wetland breeder) and Aedes albopictus (container breeder), by the diving beetle Eretes griseus. Culex tritaeniorhynchus spends more time at the surface than A. albopictus, which spends more time thrashing underwater. When intact mosquito larvae of both species were present, the diving beetles consumed almost all A. albopictus larvae (98.3%). After all the A. albopictus larvae had been consumed, the diving beetles began to prey on C. tritaeniorhynchus. In order to compare the effect of position on the predation preference of the diving beetles, equal numbers of both species were heat‐killed and allowed to settle on the bottom of the container. When all the dead mosquito larvae had sunk to the bottom of a plastic container, the diving beetles caught both mosquito species at random. These results indicate that mosquito larvae near the surface were eaten less frequently by diving beetles than those at the bottom. The low diving frequency of C. tritaeniorhynchus is regarded as a form of anti‐predatory behavior.     

Population regulation of a mosquitoArmigeres theobaldi with description of the animal fauna in zingiberaceous inflorescences

1. Inflorescences of some Curcuma and Zingiber (Zingiberaceae) in tropical Asia provide an unique aquatic habitat being discrete, small, and made of numerous smaller compartments (the bracts).
2. The aquatic community in inflorescences of Curcuma in northern Thailand was composed of immature Diptera, of which the biting midge Dasyhelea and the mosquito Armigeres theobaldi were the commonest. No important competitors, predators or pathogenic parasites for the mosquito were confirmed.
3. Inter-inflorescence distribution of the mosquito was contagious. Within each inflorescence, the fourth-instar larvae or pupae usually occupied bracts singly.
4. The k-value analysis detected density-dependent mortality due to contest competition in the mosquito larvae.
5. Variations in the larval and pupal mosquito size were density-independent and remarkably small as compared with size variations known for other mosquitoes.
6. These population attributes (large density-dependent mortality with density-independent, minimally variable individual size) appear unique among mosquitoes, arising from conspecific killing, efficient foraging (inter-bract movement by crawl and single occupation of bracts), and availability of host plant tissues as supplementary food.
7. A simple population model suggested that a small proportion of adult females lay eggs.

     

Laboratory and field studies of Macrocyclops albidus (Crustacea: Copepoda) for biological control of mosquitoes in artificial containers in a subtropical environment.

The cyclopoid copepod Macrocyclops albidus (Jurine) was tested as a potential biological control agent of mosquitoes in laboratory microcosms, in controlled field conditions, and in a 22-mo field experiment using discarded tires. The predator was highly efficient in controlling mosquitoes in all three settings, reaching close to 90% reduction in larval survival under field conditions and exceeding the recommended predation rates for effective mosquito control in laboratory experiments. The predator was most effective on 1-4-d-old larvae. Alternate food and habitat structure significantly influenced the predation rates on mosquito larvae. Once established, the copepod was able to maintain reproducing populations in the field for the duration of the experiments. However, the predator failed to establish populations at four of the experimental field sites. Two of the failures can be attributed to characteristics of the individual tires, such as leaching chemicals, whereas the other two were probably due to site-specific factors. This copepod species is a promising candidate for control of mosquito larvae because it is a widespread and highly effective predator that is capable of establishing and maintaining populations under a wide variety of field conditions. Additionally, M. albidus is relatively easy to culture, maintain, and deliver to the target areas.     

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.