Habitat complexity reduces prey vulnerability: An experimental analysis using aquatic insect predators and immature dipteran prey

The effects of alternative prey and structural complexity of habitat on the selection of mosquito larvae by aquatic insect predators were evaluated in the laboratory. The water bugs Anisops bouvieri, Diplonychus (= Sphaerodema) rusticus, and D. annulatus, and the odonate nymphs, Ceriagrion coromandelianum and Brachydiplax chalybea chalybea, selected mosquito larvae based on their abundance relative to chironomid larvae and on the levels of habitat complexity. The effect of one prey species on the other was asymmetrical, as indicated through prey selectivity values. Compared to open habitat, the presence of macrophytes reduced the vulnerability of mosquito larvae while the effect was reverse in the presence of sediments. When both sediment and macrophytes were present in habitats, all the predators except D. annulatus consumed more mosquito larvae than chironomid larvae. The clearance rate, an indicator of predatory efficiency, varied among the predator species and habitat types. The results suggest that the outcome of the interactions between insect predators and mosquito immatures was context-dependent and that it was mediated by the presence of alternative controphic species and the habitat complexity.     

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.     

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.     

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.     

Pelagic food web interactions among benthic invertebrates and trout in mountain lakes

1. Benthic chironomid larvae and the amphipod Gammarus lacustris have been observed in the pelagic habitats of many mountain lakes. The main goal of this study was to determine if chironomid larvae and gammarids potentially affect predator–prey and nutrient dynamics in pelagic food webs of mountain lakes. 2. Eighty‐six mountain lakes were surveyed in Alberta and eastern British Columbia during the years 1965–1984, 1991–2004 and 2005–2007. Pelagic chironomid larvae were found in 86% of these lakes, and pelagic gammarids were found in 29% of lakes. Densities of pelagic chironomid larvae were 92% lower in lakes with pelagic gammarids and 76% lower in lakes with trout (P < 0.05). Intraguild predation of trout on gammarids appeared to reduce predation pressure on chironomid larvae. Gammarids consumed in vitro about 1 chironomid per gammarid per day or about 20% of their body mass in chironomid biomass per day. 3. Concentrations of total dissolved P and N, particulate C, and chlorophyll‐a increased with increasing densities of pelagic gammarids and chironomid larvae in situ (R² = 0.14 ± 0.19 SD, P < 0.1) and in vitro (P < 0.001). 4. Our findings suggest that gammarids and chironomid larvae are linked as predators and prey in pelagic food webs, possibly stimulating phytoplankton abundance via nutrient release.     

Relationships between predatory aquatic insects and mosquito larvae in residential areas in northern Thailand

In order to elucidate the poorly understood relationships between mosquito larvae and their predatory aquatic insects in urban and suburban areas of tropical Southeast Asia, where vector‐borne diseases are prevalent, aquatic insects were sampled from 14 aquatic habitats in residential areas of Chiang Mai, northern Thailand, during the rainy season (July to November) in 2016. Correlations among biological variables, densities of major predatory aquatic insect groups (i.e., Odonata, Coleoptera, and Hemiptera: OCH group) in wetlands and artificial lentic habitats, and the density of mosquito larvae were analyzed. Among the sampled mosquito larvae, Culex spp. were the most abundant, and both OCH density and water quality were major determinants of Culex spp. density (r_s = ?0.302 and ?0.396, respectively). Logistic regression analyses indicated that the probability of Culex spp. occurrence was significantly and negatively correlated with OCH density. Furthermore, high macrophyte abundance was associated with higher predator density, potentially reducing mosquito density. Hemipteran predators were most negatively correlated with Culex spp. density, regardless of whether macrophyte abundance was high or low (r_s = ?0.547 and ?0.533, respectively). Therefore, hemipteran predators were the most important aquatic insect predators in the urban and suburban residential areas of Chiang Mai, Thailand, and OCH species, such as the hemipteran Micronecta scutellaris, could be used as biological control agents against mosquitoes in the region.     

Spatial Distribution,Food and Activity of Gomphus pulchellus SELYS 1840 (Insecta; Odonata; Gomphidae) from a Still Water Habitat

Distribution patterns of Gomphus pulchellus larvae in different sediments with different density of prey organisms were studied in the field in a small gravel pit lake in the south of Germany. Larval burrowing behaviour at different temperatures as well as food preference, consumption rates and activity were studied in laboratory experiments. In the study lake G. pulchellus larvae lived exclusively in places where macrophytes were present and in fine sediments (mean grain size <3 mm) with detritus cover. There was a significant positive correlation between larval density and density of food organisms suggesting that abundance of food is one of the determinants of larval distribution. In late autumn larvae migrated to deeper places probably to survive the winter. Low temperatures simulated in laboratory experiments did not induce larvae to burrow deeper. Larvae were always found in a sediment depth of 0.59–0.74 cm. Experiments with mixed prey showed that G. pulchellus larvae preferred tubificid worms and chironomid larvae over gammarids and ephemerid larvae. However, chironomid larvae which stayed in their tubes had a higher survival rate than those outside of tubes. Single-prey experiments showed that G. pulchellus larvae can prey not only on benthic species but also on Daphnia from the open water. Functional-response experiments showed that one G. pulchellus larva consumes a maximum of 2 to 3 tubificid worms or chironomid larvae per day, which corresponds to a maximum biomass (freshweight) of 5 to 30 mg per day. Video recordings of activity showed that G. pulchellus larvae cover long distances of up to 52 m per night on the substrate surface and that activity on the substrate surface started after midnight and ceased before sunrise. Consumption of Zooplankton prey and high activity above the substrate is interpreted as an adaptation of G. pulchellus larvae to the life in still water habitats.     

Food selectivity versus prey availability: a study using the marine fish Pomatoschistus microps

Summary The food selection of the common goby, Pomatoschistus microps Krøyer, was studied in the field and in laboratory experiments on the Swedish west coast. The three most important prey organisms for P. microps in the study area were Corophium volutator, chironomid larvae and Nereis spp. Corophium was consumed more than any other prey, even when it was not the most abundant prey species in the bottom. One reason may be the higher activity of Corophium above the sediment surface, which may increase its visibility and consequently its vulnerability to visual predators. When P. microps was offered Corophium and chironomid larvae with similar exposure in laboratory experiments, it showed no preference for either of the prey items. It always took the closest mobile prey, regardless of species and size.     

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.     

Unsuitability of Cnesterodon decemmaculatus (Jenyns, 1842) for mosquito control in Uruguay: evidence from food‐preference experiments

We analyzed the feeding preference of Cnesterodon decemmaculatus, a small‐bodied poecilid native from the Rio de la Plata and proximate Atlantic Basins in South America. This species has a wide distribution in Uruguayan water bodies but its effectiveness as a predator of mosquito larvae has not been tested. In laboratory trials, five aquatic invertebrates were offered simultaneously as potential prey to fish: Daphnia pulex (Cladocera), copepods, two different instars of mosquito larvae (Culex pipiens), and the 4^th instar of Chironomidae larvae. Preference was measured by the Chesson's electivity index (α). In order to determine differences in prey preference according to fish size, individuals ranging from 9.5 mm to 35.3 mm were classified in three different body size classes: small, medium, and large. Small fish showed preference for copepods, while medium‐sized fish preferred the smallest mosquito larvae instars and Chironomidae larvae. We conclude that C. decemmaculatus is a zooplankton facultative‐feeder fish that prefers large‐bodied zooplankton but is a weak predator of mosquito larvae. Thus, the introduction of C. decemmaculatus as a biological‐control agent in natural environments is not an effective strategy.     

Feeding by the aquatic heteropteran,Diplonychus rusticum (Belostomatidae): an effect of prey density on meal size

The behaviour of Diplonychus rusticum feeding on chironomid larvae has been investigated under laboratory conditions. Changes in the percentage of material extracted from prey indicated that feeding for the first two minutes enabled the predator to obtain approximately 33% of the available food; feeding for 10 minutes resulted in only 60% extraction. Comparing the percentage of each prey consumed by D. rusticum exposed to various prey densities, it was apparent that predators were more wasteful and ate less of each prey as chironomid density increased. Because the rate of food intake declined as a greater proportion of each prey was extracted, predators exposed to high chironomid densities reduced the amount of each prey consumed thereby conforming to a simple optimal feeding model.     

Predator-prey responses in an acarine system

Summary This study examines the responses of the predatory mite,Phytoseiulus persimilis, to the density and distribution of its prey,Tetranychus urticae. It is divided into three parts. Firstly, the functional responses of protonymph, deutonymph and adult females towards different prey stages are displayed. The great majority of the responses are of the type II form, and the variations in the values of attack ratea′ and handling timeT _h are discussed. Experiments are then described in which individual protonymph, deutonymph and adult female predators are presented with varying ratios of two prey age-classes (eggs and deutonymphs, and larvae and deutonymphs). Any observed preference for one of the prey stages is discussed in relation to the predicted preference on the basis of the separate functional response experiments. Finally, the response of different densities of adult female predators to a non-random distribution of deutonymph prey on bean leaflets is examined. The predators show a clear tendency to aggregate on the leaflets of high prey density, counteracted to some extent by interference increasing the probability of dispersal to other leaflets.     

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.     

Comparative Study of Functional Response of Common Hemipteran Bugs of East Calcutta Wetlands,India

The common and abundant hemipteran water bugs Anisops bouvieri, Diplonychus rusticus, D. annulatus, of the wetlands of East Kolkata are known predators of a wide range of aquatic insects including the mosquito larvae. In the laboratory their predation were assessed in respect to short term and long term periods using the larvae of Culex quinquefasciatus to reveal their possible role in regulating the dipteran population in nature. The attack rate (a) and handling time (T_h ) of these predators varied with respect to the prey size. For the backswimmers A. bouvieri the values for a and T_h for the small prey were 5.47 L and 18.72 min respectively, while in case of the belostomatid bugs, the values for the same were 5.37 L and 8.64 min (for D. rusticus), 5.81 L and 20.16 min (for D. annulatus). The predation rate varied with prey and predator densities for both the prey sizes. It was revealed that on an average A. bouvieri can kill and consume 10–82 and 6–44, D. rusticus 10–118 and 10–84 and D. annulatus 10–70 and 10–138 small and large sized prey per day, respectively. However the mutual interference (m) values of the three predators varied with the prey size and ranged between 0.053–0.326 for A. bouvieri, 0.0381–0.066 for D. rusticus and 0.0556–0.115 for D. annulatus, respectively. In the long term experiments A. bouvieri killed between 6–119 small preys and 3–31 large preys, D. rusticus killed 50–94 small preys and 50–96 large preys and D. annulatum were found to kill between 14–74 small prey and 50–131 large prey per day, respectively. The clearance rates were found to be proportional to the predator density as well to the prey size and density, and differed between the predator species significantly. These data are supportive of qualifying the water bugs, A. bouvieri, D. rusticus, and D. annulatus as potential biological resources in regulating the population of mosquito larvae in the wet‐lands. (© 2007 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

The negative effects of pathogen‐infected prey on predators: a meta‐analysis

Intra‐guild predation (IGP) – where a top predator (IG_Pred) consumes both a basal resource and a competitor for that resource (IG_Prey) – has become a fundamental part of understanding species interactions and community dynamics. IGP communities composed of intraguild predators and prey have been well studied; however, we know less about IGP communities composed of predators, pathogens, and resources. Resource quality plays an important role in community dynamics and may influence IGP dynamics as well. We conducted a meta‐analysis on predator–pathogen–resource communities to determine whether resource quality mediated by the pathogen affected predator life‐history traits and if these effects met the theoretical constraints of IGP communities. To do this, we summarized results from studies that investigated the use of predators and pathogens to control insect pests. In these systems, the predators are the IG_Pred and pathogens are the IG_Prey. We found that consumer longevity, fecundity, and survival decreased by 26%, 31% and 13% respectively, when predators consumed pathogen‐infected prey, making the infected prey a low quality resource. Predators also significantly preferred healthy prey over infected prey. When we divided consumers by enemy type, strict predators (e.g. wolf spiders) had no preference while parasitoids preferred healthy prey. Our results suggest that communities containing parasitoids and pathogens may rarely exhibit intraguild predation; whereas, communities composed of strict predators and pathogens are more likely dominated by IGP dynamics. In these latter communities, the consumption of low and high quality resources suggests that IGP communities composed of strict predators, pathogens and prey should naturally persist, supporting IGP theory. Synthesis We investigated how consuming pathogen‐infected prey influence important life‐history parameters of insect predators. Pathogens are used in a variety of biocontrol programs, especially to control crop pests. We found that true predators (i.e. wolf spiders) have no preference for healthy or infected prey and have reduced fecundity, survival and longevity consuming infected prey. However, parasitoids avoided infected prey when possible. In biocontrol programs with multiple control agents, parasitoids and pathogens would do a better job controlling pests as predators would reduce the amount of pathogen available and have reduced fitness from consuming infected prey. However, theory suggests that true predators, prey and pathogens may coexist long term.     

The influence of aquatic predators on mosquito abundance in animal drinking troughs in New Zealand

The occurrence and abundance of mosquito populations may be associated with the abundance of predators. We examined the relationship between aquatic predators and populations of mosquitoes in animal water troughs in Waikanae, New Zealand. We also investigated the effects of water volume and environmental factors (temperature, rainfall, wind speed, humidity, and pressure) in order to further understand factors influencing mosquito and predator populations. Logistic regression indicated that the presence or absence of mosquitoes was primarily affected by three factors: predator abundance, week of observation, and water volume. Pearson's correlation indicated that the presence of predators had a positive correlation with water volume (r²= 0.176, p< 0.05). Otherwise, the presence of mosquito larvae in water troughs was negatively correlated with water volume (r²=?0.159, p=0.022) and wind speed (r²=0.142, p=0.041). We established a translocation experiment in which predators or mosquitoes were moved between troughs in order to examine the prey survival rate after exposure to Anisops wakefieldi predators. The survival rate of mosquitoes was not significantly different, between 0–0.1%, irrespective of the number of predators translocated (1–9) or the initial mosquito density (20–70 larvae). Our results suggested that A. wakefieldi predators may have the potential to be a promising biological control tool for the control of mosquito populations by altering mosquito population dynamics.     

Diet composition,feed preferences and mouth morphology of early stage silver therapon (Leiopotherapon plumbeus,Kner 1864) larvae reared in outdoor tanks

This study examined the diet composition, feeding preferences, and mouth morphology of the silver therapon (Leiopotherapon plumbeus, Kner 1864) larvae under captive conditions. Larvae were reared in outdoor tanks (4 m³) with natural food grown 2 weeks prior to start of larval rearing. Food preference was measured by the Chesson's electivity index (α_i). Gut content analysis of larvae sampled between 5 and 25 days after hatching (DAH) showed the dominance in the diet by zooplankton, mainly copepod nauplii, cladocerans and insect larvae. Small fish larvae (5–9 DAH; 3.32–6.29 mm standard length) preferred cladocerans, ciliates and copepod nauplii; whereas older larvae (12–25 DAH; 5.45–19.26 mm standard length) preferred insect larvae over cladocerans and adult insects. The mouth gape size at 5 DAH was 359 μm and increased to 3.75 mm at 40 DAH when body size grew at an average rate of 0.59 mm d^?1. The standard length (SL) of L. plumbeus larvae was strongly associated with mouth size (r² = 0.98, P < 0.05), indicating a progressive increase of ingested prey size of the fish larvae. These results clarified the early life feeding ecology of this species, which is essential in developing effective hatchery techniques.     

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.