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)     

A comparative study of predation of three aquatic heteropteran bugs on Culex quinquefasciatus larvae

The aquatic bugs Anisops bouvieri Kirkaldy 1704 (Heteroptera: Notonectidae), Diplonychus (=Sphaerodema) rusticus Fabricius 1781, and Diplonychus annulatus Fabricius 1781 (Heteroptera: Belostomatidae) are common members of the freshwater insect communities of the East Calcutta Wetlands along the eastern fringe of Kolkata, India. These insects are established predators of dipteran larvae and other organisms. A comparative account of their predatory efficiency was made using larvae of Culex quinquefasciatus Say 1823 in the laboratory. It was revealed that a single adult of A. bouvieri could consume 28–34 fourth-instar mosquito larvae per day, D. rusticus 11–87 fourth-instar mosquito larvae per day, and D. annulatus 33–122 fourth-instar mosquito larvae per day, depending upon the prey and predator densities. The mean predation rate of A. bouvieri and D. annulatus remained stable over a 6-day feeding period but varied for D. rusticus. The predatory impact (PI) values were 14.77–17.31, 46.9–55.73, and 61.74–72.72 larvae/day for A. bouvieri, D. rusticus, and D. annulatus, respectively. Compared to these, the clearance rate (CR) value range was 9.06–13.25 for A. bouvieri, 13.64–15.99 for D. rusticus, and 13.50–16.52 larvae l/day/predator for D. annulatus. The values of mutual interference, “m,” remained 0.06–0.78 for A. bouvieri, 0.003–0.25 for D. rusticus, and 0.09–0.27 for D. annulatus, and did not vary between the days. The difference in predatory efficiency, CR, and PI values varied significantly among the three predators, indicating the possible difference in the function as predators occupying the same guild. It can be assumed that these predators play an important role in larval population regulation of mosquitoes and thereby impart an effect on species composition and interactions in the aquatic insect communities of the wetlands and other similar habitats where they occur.     

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.     

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.     

Biology of Tupiocoris cucurbitaceus (Hemiptera: Miridae), a predator of the greenhouse whitefly Trialeurodes vaporariorum (Hemiptera: Aleyrodidae) in tomato crops in Argentina

The predator Tupiocoris cucurbitaceus is frequently found attacking Trialeurodes vaporariorum in greenhouses without pesticide applications in Argentina. The objective of these studies was to evaluate some biological characteristics of this species fed on three types of diet (whitefly nymphs, Sitotroga cerealella eggs and a mix of both) and on two host plants (tomato and tobacco), under controlled experimental conditions. Preimaginal developmental time for female and male bugs was shorter in the presence of whiteflies than with only moth eggs. Females lived longer when they ate only whitefly nymphs compared to a mixed diet or only moth eggs. The amount of adult descendants was greater when bugs could eat whiteflies, regardless of the presence of S. cerealella. Embryonic development time, male longevity and sex proportion were not affected by the diet or the host plant. Prey consumption was evaluated for three T. cucurbitaceus life history stages (fourth/fifth instar nymphs, female and male adults) on two types of prey (whitefly nymphs and S. cerealella eggs). On tomato, females were more voracious than males and nymphs. On tobacco, adults and nymphs consumed more S. cerealella than T.vaporariorum nymphs, but again, bug females preyed more than males and nymphs. Results demonstrate that T. cucurbitaceus can survive, develop and reproduce normally using both T. vaporariorum and S. cerealella eggs as prey on tobacco or tomato plants. This information can be useful for managing this predator against T. vaporariorum through conservative or augmentative biological control strategies.     

Intraspecific variation in responses to aposematic prey in a jumping spider (Phidippus regius)

Aposematic signals often allow chemically defended prey to avoid attack from generalist predators, including jumping spiders. However, not all individual predators in a population behave in the same way. Here, in laboratory trials, we document that most individual Phidippus regius jumping spiders attack and reject chemically defended milkweed bugs (Oncopeltus fasciatus), immediately releasing them unharmed. However, a small number of individuals within the population kill and completely consume these presumably toxic prey items. This phenomenon was infrequent with only 14% of our sample (17/122) consuming the milkweed bugs over the course of the study. Individuals that killed and consumed bugs often did so repeatedly; specifically, individuals that consumed a bug in their first test were more likely to kill a bug in their second test and also tended to consume them again. We explored what might drive some (but not all) individuals to consume these bugs and found that neither sex, sexual maturity, body size, laboratory housing type, nor being wild-caught or being laboratory-reared, predicted milkweed bug consumption. Consuming bugs had no negative effects on spider mass or body condition; contrary to expectations, individuals that consumed milkweed bugs actually gained more body mass and increased in body condition. We discuss potential behavioural and physiological variation between individuals that may drive these rare behaviours and the implications for the evolution of prey defences.     

Prey recognition time of praying mantids (Dictyoptera: Mantidae) and consequent survivorship of unpalatable prey (Hemiptera: Lygaeidae)

When juvenile praying mantids (Tenodera sinensis)were exposed to unpalatable prey (the milkweed bug Oncopeltus fasciatus),they attacked, sampled, and then usually rejected the prey. About 70% of the handling time was spent feeding. When offered a second milkweed bug, the mantids usually attacked the prey. However, the overall time required for the mantids to sample, recognize, and then reject the unpalatable prey decreased by half. The proportion of handling time that was spent feeding remained the same as in the first encounter. In contrast, when the second prey individuals encountered by mantids were Drosophila melanogaster,the flies were completely consumed and the proportion of handling time that was spent feeding significantly increased. When praying mantids were exposed to the milkweed bugs for the first time, up to 33% of the bugs survived attack by the mantids. Survival of milkweed bugs increased to 55% when mantids had been previously exposed to the bugs. In contrast, flies that were caught never survived.     

Health food versus fast food: the effects of prey quality and mobility on prey selection by a generalist predator and indirect interactions among prey species

1. In order to understand the relative importance of prey quality and mobility in indirect interactions among alternative prey that are mediated by a shared natural enemy, the nutritional quality of two common prey for a generalist insect predator along with the predator's relative preference for these prey was determined. 2. Eggs of the corn earworm Helicoverpa zea (Lepidoptera: Noctuidae) were nutritionally superior to pea aphids Acyrthosiphum pisum (Homoptera: Aphididae) as prey for big‐eyed bugs Geocoris punctipes (Heteroptera: Geocoridae). Big‐eyed bugs survived four times as long when fed corn earworm eggs than when fed pea aphids. Furthermore, only big‐eyed bugs fed corn earworm eggs completed development and reached adulthood. 3. In two separate choice experiments, however, big‐eyed bugs consistently attacked the nutritionally inferior prey, pea aphids, more frequently than the nutritionally superior prey, corn earworm eggs. 4. Prey mobility, not prey nutritional quality, seems to be the most important criterion used by big‐eyed bugs to select prey. Big‐eyed bugs attacked mobile aphids preferentially when given a choice between mobile and immobilised aphids. 5. Prey behaviour also mediated indirect interactions between these two prey species. The presence of mobile pea aphids as alternative prey benefited corn earworms indirectly by reducing the consumption of corn earworm eggs by big‐eyed bugs. The presence of immobilised pea aphids, however, did not benefit corn earworms indirectly because the consumption of corn earworm eggs by big‐eyed bugs was not reduced when they were present. 6. These results suggest that the prey preferences of generalist insect predators mediate indirect interactions among prey species and ultimately affect the population dynamics of the predator and prey species. Understanding the prey preferences of generalist insect predators is essential to predict accurately the efficacy of these insects as biological control agents.     

Food consumption and diet composition of the web-building spider Agelena limbata in two habitats

Summary Prey capture rate, food consumption, and diet composition of all developmental stages of the funnelweb spider Agelena limbata were estimated in woody and open habitats by a sight-count method. Prey availability was evaluated on the basis of two indices, i.e. the ratios of daily food consumption to dry weight of predator and to daily standard metabolic rate. These indices varied seasonally and between instars in this spider. Comparison of these indices between arthropod predators suggests that A. limbata live under conditions of relatively limited food supply. In the open habitat, the spiders reduced foraging activities to avoid heat stress at midday in summer because the sheet web was exposed to the direct rays of the sun and its temperature exceeded 40°C. The daily food consumption of adult spiders in the open habitat was about half of that in the woody habitat. The lower rate of energy intake of spiders in the open habitat may cause the observed smaller size of adults and lower fecundity. A. limbata captured a great range of prey comprising ten orders of arthropods and ate chemically defended insects, e.g. stink bugs, lady beetles, and ants which were rejected by many spiders. This generalistic foraging may be associated with limited and heterogeneous food supply in this spider.     

Araneophagic behavior of Gardena brevicollis Stål (Heteroptera: Reduviidae): Foraging on pre‐dispersal spiderlings in the spider's web

Araneophagic behavior of an Emesinae assassin bug, Gardena brevicollis Stål (Hemiptera: Reduviidae) was observed in the field. The bug invaded a web of Acusilas coccineus Simon (Araneae: Araneidae) spiderlings and hunted five juvenile spiders. The bug showed stalking tactics to prey on spiders, whereas luring tactics, which have been reported in other Emesinae bugs, were not observed. To the best of my knowledge, the present study is the first report of araneophagy in the genus Gardena and the first report of araneophagic behavior in East Asian assassin bugs.     

Selective and frequency dependent predation of aquatic mosquito predator Diplonychus indicus Venkatesan & Rao (Hemiptera: Belostomatidae) on immature stages of three mosquito species

Frequency dependent mosquito larval size (II and IV instars) and species selection by the water bug Diplonychus indicus against three mosquito species Culex quinquefasciatus, Aedes aegypti and Anopheles stephensi was studied in the laboratory. The different frequencies used for each species selection were 20:30:50, 30:50:20, 50:20:30, 25:35:40, 35:40:25 and 40:25:35 of fourth instars of the respective three prey species. All nymphal water bugs (I–V instars) selected IV instar mosquito larvae and the mean proportion of late (larger) larvae eaten by the predator instars was significantly higher than the mean proportion of early (smaller) larvae eaten (F= 2.28; P < 0.001). In all six ratios used to determine the frequency dependent mosquito species selection, all the stages of the water bug selected Ae. aegypti over the other two species (F= 452.43; P < 0.001). The mean number of mosquito larvae eaten increased as its density increased based on various ratios of larvae offered. The study indicated that the predatory efficiency of D. indicus was high when Ae. aegypti was offered as prey, suggesting the utility of this mosquito predator in the control of dengue vectors.     

Potential population growth and harmful effects on humans from bed bug populations exposed to different feeding regimes

Effects of host availability and feeding period on bed bugs, Cimex lectularius (L.) (Hemiptera: Cimicidae), were measured. Population growth and the potential harmful effect of bed bug populations on human hosts were modelled. Bloodmeal sizes were affected by both feeding length and frequency, with >2‐fold difference between insects fed daily or weekly. Blood consumption increased >2‐fold between bed bugs fed occasionally and often, and 1.5‐fold between occasional and daily feeding. Bed bugs fed more often than once a week, potentially every 2–4 days. Egg production was associated with nutrition, being strongly correlated with blood consumption in the previous week. Bed bug populations can grow under different feeding regimes and are hard to control with <80% mortality. Bed bugs can survive and grow even in locations with a limited blood supply, where bed bug persistence may be important for the continual spread of populations. Persistence in non‐traditional locations and a potential association with human pathogens increase the health risks of bed bugs. Potential blood loss as a result of a bed bug can have serious consequences because uncontrolled populations can reach harmful levels in 3–8 months. The reproduction potential of bed bug populations suggests serious consequences to human health and the need for efficacious control measures.     

The influence of prey size,sediment thickness and fish size on consumption in common sole (Solea solea L.)

This study determined prey consumption in common sole as a function of prey size (0–0.5, 1–1.5, 2–2.5 and 4–5 g), sediment thickness (20 cm and 2 cm) and fish size (50 g, 125 g or 300 g). Prey consumption (in numbers of prey eaten per fish per day) was reduced with increasing prey size and sediment thickness, and was increased with increasing fish size (p < .001 for all factors). All 3 factors showed significant two way interactions (p < .001) when expressed in numbers of prey eaten. Prey consumption decreased with prey size when prey could not escape by burying (2 cm of sediment thickness) irrespective of fish size. We suggest that increasing effort to ingest and handle larger prey played a role. Prey consumption increased with fish size when prey could not bury (2 cm of sediment thickness). However, when prey was able to bury (at 20 cm sediment thickness) prey consumption was similar irrespective of fish size (p < .001 for interaction fish size × sediment). This interaction suggests that with increasing fish size there is an increasing mismatch between foraging adaptation and prey burial depth. This may explain the dominance of crustaceans in the diet of adult common sole in nature, despite the high abundance of polychaetes.     

Differences in prey capture in semiaquatic bugs (Heteroptera: Gerromorpha)

Feeding behavior has been investigated several times in different semiaquatic bugs (Heteroptera: Gerromorpha), but has never been directly compared between species from lentic and lotic habitats and different families. Time required to capture prey was investigated in five gerromorphan species: Aquarius paludum, Gerris lacustris, G. odontogaster (Gerridae), Velia caprai and V. gridellii (Veliidae). This experiment used live and dead prey and intact and artificially blinded bugs. The presence of vibrational stimuli (live prey) significantly decreased latency to capture prey in all species except G. odontogaster. This species was the only one where visual stimuli affected the response significantly (only with dead prey). Latency to capture in Velia spp. was significantly shorter than that in the three gerrid species. Capture efficiency differed significantly between Gerris and Velia spp.; both Velia species carried out unsuccessful attack attempts less frequently than did the Gerris species. Prey capture was quicker in treatments with live prey and/or intact bugs. No difference in capture efficiency was found between sexes. Aquarius and Gerris species from predominantly lentic habitats needed more time to capture prey than did Velia species, which live mostly in streams. This difference in feeding behavior can probably allow coexistence of gerrids and veliids at sites where their occurrence overlaps.     

Variation in the geometry of foreleg claws in sympatric giant water bug species: an adaptive trait for catching prey?

When giant water bugs (Heteroptera: Belostomatidae) encounter prey animals that are larger than they are themselves, they first hook the claw of their raptorial legs onto the animal, and then use all their legs to pin it. The claws of the raptorial legs in giant water bugs play an important role in catching larger prey, but the relationship between the claws, body lengths of predators, and prey size has not been fully investigated. To elucidate the functioning of claws in catching prey, we investigated prey body size relative to predator size in nymphs of two sympatric belostomatid giant water bug species, the vertebrate eater Kirkaldyia (=Lethocerus) deyrolli Vuillefroy and the invertebrate eater Appasus japonicus Vuillefroy, captured in rice fields. The younger nymphs of K. deyrolli caught preys that were larger than themselves, whereas those of A. japonicus caught preys that were smaller. Younger nymphs of K. deyrolli had claws that were curved more sharply than those of A. japonicus. The more curved claws of younger nymphs of K. deyrolli probably hook more easily onto larger vertebrates and thus this shape represents an adaptation for acquiring such prey.     

First observation of Lethocerus cordofanus (Hemiptera: Belostomatidae) preying on Afronycteris nana (Chiroptera: Vespertilionidae)

Giant water bugs (Hemiptera: Belostomatidae) are key predators in freshwater ecosystems and have been reported to feed on several species of vertebrates, including fishes, amphibians and reptiles. Here, we report the opportunistic predation of an adult female vesper bat (Afronycteris nana) by a giant water bug (Lethocerus cordofanus) in a temporary pond in a rice paddy in Guinea-Bissau, West Africa. To our best knowledge, this is the first instance of natural predation upon a mammal by a giant water bug to be documented in a scientific report.     

The role of prey-generated sounds,vision, and echolocation in prey localization by the African batCardioderma cor (Megadermatidae)

Summary Cardioderma cor responded with head movements and flight toward speakers broadcasting calls of frogs and crickets which contained only sonic frequencies. Unlike the frog-eating bat,Trachops cirrhosus, they did not make contact with the speakers. Prey movements that generated sonic and ultrasonic sounds were both sufficient and necessary for the bats to localize and capture prey. Prey dragged across a glass sheet with a thin layer of water did not generate sounds and bats did not attempt to capture these prey, even with the availability of visual and echolocation cues. There was no evidence for the use of visual cues while hunting; bats did not localize prey more readily in light than darkness. Prey were presented such that their movements initially generated sounds, but then the prey moved onto the water layer of the glass sheet and sounds were eliminated. The bats emitted echolocation signals while hunting in this situation; however, the information from these signals was not utilized. The bats landed at the site that prey last made sound. These results demonstrate the importance of passive hearing for prey localization in this bat, and further suggest that when preygenerated sounds and echolocation signals offer conflicting information the bat's behavior is guided by the former.     

Generalist versus specialist strategies of plasticity: snail responses to predators with different foraging modes

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The feeding behaviour of New Zealand Dolomedes species (Araneae: Pisauridae)

Abstract

The prey spectrum and predatory behaviour of Dolomedes sp. (‘D. III’), D. aquaticus, and D. minor are described from a series of field and laboratory investigations, the former made around Nelson and on Banks Peninsula, Canterbury, during summer months. All species are large, robust spiders that capture prey by directly seizing it in their mouthparts, not using silk at any stage of predation. Adult aquatic insects are their main prey, but these are available only irregularly during their activity period (night-time), and the spiders are opportunistic in their feeding habits. They will eat virtually any available small animal, and at least the largest species, D. III, is able to capture and ingest small fishes. Dead as well as live organisms are taken. Furthermore, the spiders are capable of feeding infrequently; when feeding on small prey organisms they may capture several sequentially, to increase the size of the meal. Live prey is caught while it is in flight, or on the ground, or at the water surface (rarely submerged), and is detected primarily by touch and airborne sound; vision is unnecessary for normal predation. Prey is captured very rapidly, even though this may initially involve a dash of up to 40 cm across the water surface to locate the organism. Stages in the behaviour of an active spider, from waiting for prey to grooming after ingestion, are described.     

How Do Predators Learn to Recognize a Mimetic Complex: Experiments with Naive Great Tits and Aposematic Heteroptera

We tested the importance of innate wariness, avoidance learning, memory and generalization for the formation of predatory behaviour in naive great tits (Parus major) towards mimetic complex of four aposematic species of true bugs (Insecta: Hemiptera: Heteroptera): Lygaeus equestris, Spilostethus saxatilis, Pyrrhocoris apterus and Graphosoma lineatum. The birds showed almost no innate wariness against the aposematically coloured bugs, although a hidden wariness elicited by defensive chemicals of some of the bug species is not excluded. Naive birds learned to avoid different species at different rates, which resulted in different prey mortalities. The avoidance learning was faster when the defensive chemicals produced an immediate irritating effect (particularly when squirted into distance – G. lineatum) than when they caused sickness several minutes after the consumption (P. apterus). The experience of birds from learning to avoid a particular species of bug affected their subsequent behaviour to other species – experience with better‐defended species resulted in longer attack latencies, more cautious attacks, broader generalization and lower prey mortality. The least defended species, P. apterus, benefited from the experience of birds with better‐defended species, whereas the birds' experience with P. apterus did not reduce mortality risk of the other species comparably. Judging from the inexperienced young birds, the mimetic relationships are likely to be quasi‐Batesian. However, as wild‐caught great tits avoid all the four species to the same extent, the relationships may become more mutualistic (quasi‐Müllerian) in later phases of learning under natural conditions. The relationships among species in the mimetic complex thus seem to depend on the amount of experience of the bird predators.