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.     

Prey‐mediated changes in the selectivity of the predator Macrolophus pygmaeus (Heteroptera: Miridae)

When foraging in communities with mixed prey, generalist predators may be confronted with prey species that differ in quality, size and mobility and interact with one another. To examine prey selection, predation by Macrolophus pygmaeus (Heteroptera: Miridae) was recorded by providing a diet of either one or two prey species of Myzus persicae (third‐instar nymphs), Aphis gossypii (fourth‐instar nymphs), Trialeurodes vaporariorum (third‐instar nymphs) and Ephestia kuehniella (eggs). In the experiments, prey mobility, prey quality and prey biomass were considered. The biomass consumed by the predator was dependent on the combination of prey species and the quantity of biomass offered. In choice experiments with diets mixed of two prey species at equal densities, the predation to A. gossypii was significantly reduced in the presence of E. kuehniella but the rate of consumption of M. persicae, T. vaporariorum and E.kuehniella was not significantly affected by the coexistence of any other species in the mixed prey diet. When equal amounts of biomass from two prey species were provided in combination, the total consumed biomass was significantly reduced in the mixed prey diets composed of E. kuehniella eggs and aphid nymphs. Thus, under the mixed‐prey situation, prey selection by predators may be affected by interactions among prey species differing in traits such as quality, mobility and size.     

Life‐history parameters and predation capacity of Macrolophus pygmaeus and Nesidiocoris tenuis (Hemiptera: Miridae) on eggs of Plutella xylostella (Lepidoptera: Plutellidae)

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Predation of cotton leafworm first instar larvae,Alabama argillacea [Lep.: Noctuidae]

Natural predation first instar larvae of the cotton leafworm (CLW)A. argillacea was studied in cotton fields in Jaboticabal, Sao Paulo State, Brazil, during 1986. The presence of naturally occurring arthropod predators showed a first instar larvae predation rate of 78.6 and 88.9% after 24 h and 48 h of exposure, respectively. A predator prey ratio of 1∶1 (1 CLW key predator per 1 prey/plant) maintained a level of no more than 1 CLW small larvae per plant. The most evident arthropod predators in the studied fields were: beetles (Coleoptera: Coccinellidae), antsPheidole sp. andConomyrma sp.;Dermaptera Doru lineare (Eschs);Hemiptera Geocoris sp., andOrius insidiosus Say; and the spidersTheridion volubile, Chrysso pulcherrima, Misumenops sp.,Chiracanthium sp., andOxyopes salticus Hentz.      

Toward the Quantification of Predation with Predator Gut Immunoassays: A New Approach Integrating Functional Response Behavior

Immunological methods have been widely used to identify key predator species and qualitatively evaluate predation of target prey. However, despite the quantitative nature of many immunoassays, the translation to number of prey attacked has been problematic because of the many factors that confound interpretation of the strength of the immunoassay response. We developed a new predation model that couples the proportion of predators positive for prey remains determined by enzyme-linked immunosorbent assay (ELISA), predator density, and predator functional response to prey density for estimating total prey attacked. We used single cotton plant arenas in the greenhouse to develop functional response models for two generalist predators, Geocoris punctipes (Say) and Orius insidiosus (Say), preying on Pectinophora gossypiella (Saunders) eggs. The model was validated and compared with other immunologically based predation models in multiple plant/multiple predator arenas. Our predation model was relatively accurate in predicting the total number of prey attacked by both predator species and was a significant improvement over previous models that rely on simple assumptions regarding predator attack rates. The model also improves the predictive capacity of the functional response model alone by correcting for the number of predators actually consuming prey. Sensitivity analyses indicated that model performance was most sensitive to accurate measurement of input variables such as temperature and the proportion of individuals positive for prey antigens by ELISA and less sensitive to changes in estimates of prey density. Accurate estimation of the functional response parameters is also important, especially for the behavioral parameter defining the decline in plant leaf area searched with increases in prey density. Limitations of the model and application to the field are discussed.     

Effects of mummy consumption on fitness and oviposition site selection on Harmonia axyridis

Intraguild predation (IGP) has been commonly reported between predators and parasitoids used as biological control agents as predators consuming parasitoids within their hosts. However, the effect of parasitoid–mummy consumption on the fitness of the predator and subsequent oviposition site selection have not been well studied. In our study, we conducted two laboratory experiments to examine the influence of Aphidius gifuensis Ashmead (Hymenoptera: Braconidae) mummies as prey on fitness and subsequently oviposition site selection of Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae). Results indicate that when H. axyridis was reared on A. gifuensis mummies only, its larval development was prolonged, and body weight of the 4th instar larvae and newly emerged adults, and fecundity decreased. Moreover, H. axyridis did not exhibit oviposition preference on plants infested with unparasitized aphids or aphids parasitized for shorter than 9 days. However, compared with plants with mummies (parasitized ≥9 days), H. axyridis laid more eggs on plants with unparasitized aphids. In contrast, H. axyridis previously fed with A. gifuensis mummies did not show a significant oviposition preference between plants with unparasitized aphids and those with mummies (parasitized ≥9 days). Overall, our results suggest that mummy consumption reduced the fitness of H. axyridis. Although H. axyridis avoided laying eggs on plants with A. gifuensis mummies, prior feeding experience on A. gifuensis mummies could alter the oviposition site preference. Thus, in biological control practice, prior feeding experience of H. axyridis should be carefully considered for reduction of IGP and increase of fitness of H. axyridis on A. gifuensis.     

Stable isotope analyses document intraguild predation in wolf spiders (Araneae: Lycosidae) and underline beneficial effects of alternative prey and microhabitat structure on intraguild prey survival

Intraguild predation (IGP) is common among generalist predators and an important issue in food web theory, because IGP may destabilise communities by increasing extinction of species. Also, IGP may interfere with the effectiveness of generalist predators as biological control agents. In general, occurrence of IGP in laboratory or field studies is inferred from abundance data or direct observations only. We investigated if tracing stable isotopes allows distinction between different types of predation and confirmation of IGP. Wolf spiders were chosen as model organisms for generalist predators; IGP of third instar A. cuneata on second instar P. palustris was investigated in a laboratory experiment. The availability of alternative prey and the complexity of the microhabitat were manipulated, since both factors are thought to facilitate coexistence of predators.
Stable isotope analysis documented predation of A. cuneata on P. palustris and predation on alternative prey by both juveniles. Both the presence of alternative prey and the availability of shelter reduced mortality of juvenile P. palustris during the first week. During the second week mortality increased in complex structure without alternative prey presumably due to enhanced activity and cannibalism among starving P. palustris . Thus, microhabitat complexity and prey abundance may foster coexistence of wolf spiders in the field.
In conclusion, stable isotope analysis was proven a powerful tool to investigate animal behaviour without direct observation. The method allowed disentangling predator feeding behaviour when more than one type of prey was present.     

Arthropods associated with medicinal plants in coastal South Carolina

Arthropods were sampled from feverfew [Tanacetum parthenium （L.） Schultz- Bip], Echinacea purpurea （L.） Moench, Echinacea pallida （Nutt.） Nutt., Valeriana officinalis L., and St. John＇s wort （Hypericum perforatum L.） during 1998-2001. In addition, arthropods were sampled on tansy （Tanacetum vulgare L.） from 2001-2004. In general, 50-60 arthropod species where collected and identified among all of the medicinal plant species. Among the predators, Orius insidiosus （Say） （Hemiptera： Anthocoridae）, Geocoris punctipes （Say） （Hemiptera： Lygaeidae） and spiders were most abundant from 1998-2004. The three-cornered alfalfa hopper, Spissistilus festinus （Say）, was the most abundant herbivore found from 1998 to 2001. Orius insidiosus and G. punctipes were 3-4 times more abundant on T. parthenium than on any other medicinal plant species. Based on the numbers of predatory arthropods found on T. parthenium, this crop could be suitable as a companion or ＂banker＂ plant to attract and maintain populations of predators, especially O. insidiosus and G. punctipes. Whitefly nymphs attacked by predators with piercing/sucking mouthparts are easily identified using a microscope because of the general appearance of the carcass left by the predators. Thus, populations of predators on T. parthenium suppressed Bemisia tabaci populations on E. purpurea when these crops were planted as companion crops.     

Life history characteristics of Orius insidiosus (Say) fed Aphis glycines Matsumura

The soybean aphid, Aphis glycines Matsumura, is a new invasive pest of soybeans throughout most of the soybean production areas of North America. Field studies have demonstrated that the indigenous predator, Orius insidiosus (Say), is an important natural enemy of the soybean aphid early the soybean crop season. Because soybean aphid is newly introduced into North America, the life history characteristics of predators fed this aphid are not known. In laboratory assays, we measured the survival, development, longevity and reproduction of O. insidiosus fed 1, 3, 6 or 12 seconds to third instars of soybean aphid. O. insidiosus nymphal development decreased from 34.0 to 21.4 days as the number of soybean aphid nymphs provided increased from 1 to 6 aphid nymphs daily. Stage-specific mortality was highest at 68% for first instar O. insidiosus nymphs fed 1 soybean aphid nymph per day. Adult longevity (43.9 days) and fecundity (49.7 eggs per female) was highest for O. insidiosus fed 6 soybean aphid nymphs daily, but longevity (23.5 days) and fecundity (10.1 eggs per female) declined for adults fed 1 soybean aphid nymph daily. The intrinsic rate of increase of O. insidiosus ranged from 0.048 to 0.133. Compared to other prey species, soybean aphid is an adequate prey item for O. insidiosus. Our results suggest that O. insidiosus will be most effective in suppressing soybean aphid population growth in the initial phase of the aphid’s colonization of soybeans.     

Evidence for intraguild predation by Podisus maculiventris on a ladybeetle, Coleomegilla maculata: Implications for biological control of Colorado potato beetle, Leptinotarsa decemlineata

The potential for intraguild predation (IGP) between larval and adult life stages of twopolyphagous arthropod predators common in NorthAmerican agroecosystems was studied in thelaboratory. Predators examined were the spinedsoldier bug, Podisus maculiventris Say,and the twelve-spotted ladybeetle, Coleomegilla maculata Lengi. A shared preyitem, eggs of the Colorado potato beetle (CPB),Leptinotarsa decemlineata Say, was alsoprovided to the predators, both to provide analternate food source and to quantify theimpact of IGP on the potential for pestsuppression by these two predators. Experimentswere conducted on single potato leaves inplastic cup arenas, and, subsequently, in cagesenclosing whole potato plants. IGP occurredasymmetrically, with P. maculiventrisadults and nymphs only attacking C. maculata larvae. Even though ladybeetle adultswere generally smaller than soldier bug adults,they were never preyed upon. This appears to bethe first documented case of a coccinellidshowing differential larval and adult immunityto attack by a larger invertebrate predator.The impact of IGP, when it did occur, on CPBegg consumption was equivocal. IGP did notconsistently influence levels of predation onthe eggs. Conversely, even when IGP did notoccur, predation on CPB eggs by both predatorstogether did not increase significantly overlevels inflicted by either predator alone. Theimplications for biological control of CPB bythese predators, which are being considered foraugmentative release in potato and tomato cropsin the United States, are discussed.     

Intraguild predation by the generalist predator Orius majusculus on the parasitoid Encarsia formosa

Intraguild predation of Orius majusculus (Reuter) (Heteroptera: Anthocoridae) on Encarsia formosa (Gahan) (Hymenoptera: Aphelinidae), both natural enemies of Bemisia tabaci (Gennadius) (Homoptera: Aleyrodidae), was studied under laboratory conditions. The experiments quantified prey consumption by 5th instar nymphs and adults of O. majusculus offered unparasitised 3rd, early 4th or 4th instar B. tabaci nymphs or parasitised nymphs containing 2nd or 3rd larval instar or pupal parasitoids. In addition, prey preference of the two stages of O. majusculus for parasitised or unparasitised whitefly nymphs was studied using nine different prey combinations. Both predator stages readily preyed upon on both unparasitised and parasitised B. tabaci. In no-choice experiments, predation on 3rd instar E. formosa by adult predators was the highest, while predator nymphs preyed most on unparasitised 3rd instar B. tabaci and 2nd instar parasitoids. Predation of predator stages was lowest on 4th instar B. tabaci and E. formosa pupae. In all prey combinations, both stages of O. majusculus showed a significant preference for parasitised over unparasitised whitefly nymphs except for the combination of 5th instars of O. majusculus with early 4th instar whiteflies and E. formosa pupae. The results indicate that intraguild interactions between O. majusculus and E. formosa may have negative effects on biological control of B. tabaci.     

Potential of adultOrius insidiosus [Hemiptera: Anthocoridae] as a predator of the fall armyworm,Spodoptera frugiperda [Lepidoptera: Noctuidae]

Orius insidiosus (Say) was observed to successfully prey on eggs and earlier instar larvae of the fall armyworm (FAW),Spodoptera frugiperda (J. E. Smith). Significantly more neonates of FAW were killed per 24 h than either 1, 2 or 3 d old larvae. Female and male predators exhibited a typical type-II functional response when preying on varying densities of eggs or neonates of FAW.      

Predatory ability of generalist predators on eggs,young nymphs and adults of the invasive Halyomorpha halys in southern Europe

Halyomorpha halys (Stål, 1855) is an invasive pest causing serious damage to agricultural crops in Europe and the USA. Very little is known about H. halys predators in Europe. This survey evaluated the potential of generalist predators/omnivorous species by means of predation bioassays in tri-dimensional arenas, where the predator had to locate H. halys prey items on the leaves of a bean plant. Eleven species of different taxa were tested and the prey items consisted in fresh eggmasses, 1st and 2nd instar nymphs. One species was also tested against adults. Some predators were species commercially available as biocontrol agents against plant pests, other predators were wild, captured in habitats shared with H. halys. All tested specimens were starved 24 h before starting the experiment. The survivorship of control prey items in predator-excluding cages was compared to that of predator treatment groups to determine the effect of predator presence. According to the results, the generalist species showed a quite low acceptance of H. halys prey items, since only two species caused 80% mortality on at least one item (Eupholidoptera chabrieri and Rhynocoris iracundus) and mortality due the other species never exceed 60%. Among commercially available species only Adalia bipunctata adults and Chrysoperla carnea larvae were effective, predating the eggs and 1st instar nymphs, respectively. Among the field collected specimens, the orthopteran E. chabrieri and the predatory hemipterans R. iracundus, Nagusta goedelii and Himacerus mirmicoides showed efficacy against 1st instar nymphs, E. chabrieri and R. iracundus showed efficacy against 2nd instar nymphs, whereas only E. chabrieri and N. goedelii predated the eggs. R. iracundus was also tested on the adults and successfully predated them. By identifying some of the species that can exploit H. halys as a suitable prey in southern Europe, the present investigation provides an important contribution for conservation biological control of this pest.

     

Intraguild Predation on the Whitefly Parasitoid Eretmocerus eremicus by the Generalist Predator Geocoris punctipes: A Behavioral Approach

Intraguild predation (IGP) takes place when natural enemies that use similar resources attack each other. The impact of IGP on biological control can be significant if the survival of natural enemy species is disrupted. In the present study, we assessed whether Geocoris punctipes (Hemiptera: Lygaeidae) engages in IGP on Eretmocerus eremicus (Hymenoptera: Aphelinidae) while developing on whitefly nymphs of Trialeurodes vaporariorum (Hemiptera: Aleyrodidae). In choice and non-choice tests, we exposed G. punctipes to parasitized and non-parasitized whitefly nymphs. We found that G. punctipes does practice IGP on E. eremicus. However, choice tests assessing G. punctipes consumption revealed a significant preference for non-parasitized T. vaporariorum nymphs. Subsequently, we investigated whether E. eremicus females modify their foraging behavior when exposed to conditions involving IGP risk. To assess this, we analyzed wasp foraging behavior under the following treatments: i) whitefly nymphs only (control = C), ii) whitefly nymphs previously exposed to a predator ( = PEP) and, iii) whitefly nymphs and presence of a predator ( = PP). In non-choice tests we found that E. eremicus did not significantly modify its number of attacks, attack duration, oviposition duration, or behavior sequences. However, E. eremicus oviposited significantly more eggs in the PEP treatment. In the PP treatment, G. punctipes also preyed upon adult E. eremicus wasps, significantly reducing their number of ovipositions and residence time. When the wasps were studied under choice tests, in which they were exposed simultaneously to all three treatments, the number of attacks and frequency of selection were similar under all treatments. These results indicate that under IGP risk, E. eremicus maintains several behavioral traits, but can also increase its number of ovipositions in the presence of IG-predator cues. We discuss these findings in the context of population dynamics and biological control.     

Differential predation by the generalist predator Orius insidiosus on congeneric species of thrips that vary in size and behavior

We investigated interactions between the generalist predator Orius insidiosus (Say) (Heteroptera: Anthocoridae) and two species of thrips prey, Frankliniella bispinosa (Morgan) and Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae), and interspecific differences in morphology and behavior between these prey species that could contribute to differences in predation by O. insidiosus. Frankliniella occidentalis is significantly larger than F. bispinosa. Frankliniella bispinosa has greater mobility compared with F. occidentalis. When O. insidiosus was offered either F. bispinosa or F. occidentalis as prey in single species trials, there were no significant differences in the number of prey captured. However, O. insidiosus had significantly more encounters with F. bispinosa than with F. occidentalis. In arenas with equal numbers of both species, O. insidiosus encountered and captured F. occidentalis more than F. bispinosa. In large arenas with two pepper plants (Capsicum annuum L.), O. insidiosus preyed on more F. occidentalis than on F. bispinosa. These results indicate that O. insidiosus can prey on both thrips species, but that it preferentially captures F. occidentalis. The greater locomotion and movement of F. bispinosa, perhaps combined with its smaller size, allow it to evade predation by O. insidiosus better than F. occidentalis. Consequently, the observed preference of O. insidiosus for F. occidentalis is not exclusively a function of active selection by the predator but also could arise from inherent differences among prey. We propose this differential predation as a mechanism contributing to observed differences in the temporal dynamics of these species in pepper fields.     

Expression of Bacillus thuringiensis Cry1Ac protein in cotton plants, acquisition by pests and predators: a tritrophic analysis

1. Studies have shown that Cry proteins of the bacterium Bacillus thuringiensis expressed in transgenic plants can be acquired by nontarget herbivores and predators. A series of studies under field and controlled conditions was conducted to investigate the extent to which Cry1Ac protein from Bt transgenic cotton reaches the third trophic level and to measure the amount of protein that herbivores can acquire and expose to predators. 2. Levels of Cry1Ac in Bt cotton leaves decreased over the season. Among herbivores (four species), Cry1Ac was detected in lepidopteran larvae and the amount varied between species. Among predators (seven species), Cry1Ac was detected in Podisus maculiventris and Chrysoperla rufilabris. 3. In the greenhouse, only 14% of the Cry1Ac detected in the prey (Spodoptera exigua larvae) was subsequently found in the predator P. maculiventris. Detection of Cry1Ac protein in Orius insidiosus, Geocoris punctipes and Nabis roseipennis was probably limited by the amount of prey consumed that had fed on Bt cotton. 4. Purified Cry1Ac was acquired by the small predatory bug G. punctipes but at much higher concentration than found in plants or in lepidopteran larvae. 5. Bt protein was shown to move through prey to the third trophic level. Predatory heteropterans acquired Cry1Ac from prey fed Bt cotton, but acquisition was dependent on the concentration of Cry1Ac conveyed by the prey and the amount of prey consumed. The type and availability of prey capable of acquiring the protein, coupled with the generalist feeding behaviour of the most common predators in the cotton ecosystem, probably constrain the flow of Cry1Ac through trophic levels.     

Effect of different diets on reproduction,longevity and predation capacity of Orius insidiosus (Say) (Hemiptera: Anthocoridae)

Orius insidiosus is a generalist predator for which diet influences important biological traits like reproduction and predation. We tested the effects of different diets alone or in combination on reproduction, longevity and predation capacity of this predator. The diets tested were: no food (control); pollen; Frankliniella occidentalis (Pergande) nymphs and adults; F. occidentalis nymphs and adults and pollen; Anagasta kuehniella (Zeller) eggs; A. kuehniella eggs and pollen; A. kuehniella eggs; F. occidentalis nymphs and adults and A. kuehniella eggs, F. occidentalis nymphs and adults and pollen. The pre-oviposition period was shortest when the diet consisted of pollen and prey (A. kuehniella and/or F. occidentalis), of the two prey species together or of only A. kuehniella eggs. The highest values for the length of the oviposition period (50.1, 48.0, 46.3 and 46.1 days), daily fecundity (3.8, 3.9, 4.0 and 4.2 eggs/female/day), total fecundity (190.3, 187.7 185.2 and 193.6 eggs/female) and longevity (52.1, 49.9, 48.7 and 48.0 days) were found with diets consisting of only A. kuehniella eggs. Pollen did not improve any of the performance parameters of the predator when offered exclusively or as a complementary food. The results show that selection of a proper diet can strongly improve reproduction, longevity and predation capacity of O. insidiosus. An important finding is that A. kuehniella eggs, which can easily be produced in large quantities, are an excellent factitious prey for mass production of O. insidiosus.     

Intraguild predation of Serangium parcesetosum (Coleoptera: Coccinellidae), on whitefly Bemisia tabaci (Homoptera: Aleyrodidae) parasitized by Eretmocerus mundus (Hymenoptera: Aphelinidae)

We studied intraguild predation of Serangium parcesetosum on Eretmocerus mundus, both natural enemies of Bemisia tabaci, on eggplant under laboratory conditions. Fourth instar and adult predators attacked parasitized and unparasitized hosts equally if immature parasitoids were larvae, but avoided predating on parasitized hosts containing pupae.     

Comparative studies of predation among feral,commercially-purchased,and laboratory-reared predators

The predatory activities of commercially-purchased adult Hippodamia convergens Guérin-Méneville and two laboratory-reared strains of adult Geocoris punctipes (Say) were compared with their feral counterparts. In single prey choice feeding tests, commercially-purchased and feral H. convergens were provided copious amounts of silverleaf whitefly, Bemisia argentifolii Bellows & Perring adults or eggs of pink bollworm, Pectinophora gossypiella (Saunders). Commercially-purchased H. convergens devoured more pink bollworm eggs and at a faster rate than the feral H. convergens. In contrast, feral H. convergens consumed more adult whiteflies and at a faster rate than commercial H. convergens. In multiple feeding choice tests, two distinctly different laboratory-reared strains and feral G. punctipes were provided a cotton leaf disk containing copious amounts of silverleaf whitefly eggs, nymphs, and adults. Virtually no predation was observed on whitefly eggs, but both laboratory strains of G. punctipes fed on more whitefly nymphs and adults than the feral G. punctipes. Moreover, both of the laboratory strains had a significantly faster feeding rate on adult whiteflies and spent significantly less time feeding on plant tissue than the feral strain. These results suggest that the augmentative biological control candidates retained their ability to prey on these selected prey items, even after being displaced from their overwintering site (H. convergens) or being reared in captivity (G. punctipes) for over 40 generations. Handling editor: Patrick De Clercq. This article reports the results of research only. Mention of a proprietary product does not constitute an endorsement or a recommendation for its use by the USDA.     

The effect of mermithid parasitism on predation of nymphal Baetis bicaudatus (Ephemeroptera) by invertebrates

We investigated how infection by the mermithid nematode Gasteromermis sp. affected predation on its nymphal mayfly host, Baetisbicaudatus, by two invertebrate predators – the stonefly nymphs of Kogotusmodestus and the caddisfly larvae of Rhyacophilahyalinata. Predation trials and behavioral observations were conducted in stream-side, flow-through experimental chambers. When parasitized and unparasitized prey were offered in equal numbers, K. modestus consumed significantly more parasitized than unparasitized nymphs. R. hyalinata consumed equal numbers of both prey types. Behavioral observations of foraging K.␣modestus on parasitized and unparasitized prey suggested that the increased consumption of parasitized nymphs was due to differences in the behavior of infected mayflies in response to the predator. Specifically, parasitized nymphs drifted less often to escape an approaching predator (non-contact encounters) compared to unparasitized nymphs, which increased the number of contact encounters and attacks that occurred between K.␣modestus and parasitized prey. Because all hosts are castrated, these behavioral alterations affect only the fitness of the parasite, which is killed along with its host by invertebrate predation. We present a number of hypotheses to explain why the parasite causes increased predation on its host. These include the large size of the parasite affecting the sensory abilities of the host, the larger energetic costs of escape behavior for parasitized individuals, and natural selection from fish predation against drifting behavior by parasitized individuals. Received: 27 May 1996 / Accepted: 30 September 1996