Parasitoid Preference for Host-Infested Plants Is Affected by the Risk of Intraguild Predation

Flowering Rorippa indicaplants are attended by ants that collect nectar and, at the same time, prey on herbivorous insects, including larvae of the diamondback moth, Plutella xylostella.Here, we showed that P. xylostellalarvae suffered higher predation on R. indicawhose flowers were accessible by ants than on plants those whose flowers were inaccessible. Ants showed equal predation preference between unparasitized and larvae parasitized by Cotesia plutellae,a dominant specialist parasitic wasp of P. xylostellalarvae. C. plutellaepreferred non-flowering, host-infested R. indicato flowering, host infested R. indica.Based on these results, we infer that the preference of C. plutellaefor non-flowering, host-infested plants is in part explained by the avoidance of intraguild predation by attending ants.     

Intraguild Predation by Anthocoris nemorum (Heteroptera: Anthocoridae) on the Aphid Parasitoid Aphidius colemani (Hymenoptera: Braconidae)

The magnitude of intraguild predation by adult females of the predator Anthocoris nemorum on immature larvae of the aphid parasitoid Aphidius colemani inside mummies of peach-potato aphids Myzus persicae was investigated under laboratory conditions in a preference experiment. Each predator consumed a mean (95% confidence limits) of 2.8 (2.1; 3.8) immature parasitoids within mummies and 3.6 (2.7; 4.6) unparasitised aphid nymphs. Thereby A. nemorum engaged in intraguild predation with A. colemani and did not exhibit prey preference between mummies and unparasitised aphids.     

Effects of gender and mating status on self-directed dispersal by the whitefly parasitoid Eretmocerus eremicus

1. The impacts of gender and mating on short‐range (< 10 km) dispersal by the whitefly parasitoid Eretmocerus eremicus Rose and Zolnerowich (Hymenoptera: Aphelinidae) were examined. A fluorescent dust‐marking technique was also tested. 2. In a vertical flight chamber, female flight duration was significantly longer than that of males, and unmated parasitoids flew for longer than mated parasitoids. The mean flight durations were: unmated females 34 min, mated females 10 min, unmated males 7 min, mated males < 1 min. 3. The dispersal behaviour of E. eremicus was investigated in the field using fluorescent dust. Before doing so, it was determined in the laboratory that this dust did not affect flight behaviour, was retained over the length of the experiments, and allowed large samples to be processed quickly and inexpensively. 4. In the field, traps were placed along annuli at 3, 5, 7, and 10 m from release points. Eighty‐seven per cent of the 4153 parasitoids captured were males. Sex ratios were near parity on release. 5. The difference in dispersal characteristics between males and females may be resource based, suggesting that certain requirements were met within the field plots for males that were not met for females. 6. Locally, males dispersed in a manner consistent with a simple diffusion model while females engaged in wind‐directed flight soon after leaving release sites. The fact that the genders exhibited dissimilar dispersal characteristics, supports the claim that insect flight, even by small species, can be self‐directed.     

Compatibility of Insect Growth Regulators with Eretmocerus eremicus (Hymenoptera: Aphelinidae) for Whitefly (Homoptera: Aleyrodidae) Control on Poinsettias: I. Laboratory Assays

The compatibility of five insect growth regulators (IGRs), buprofezin, pyriproxyfen, fenoxycarb, pymetrozine, and kinoprene, were tested in the laboratory for compatibility with the whitefly parasitoid Eretmocerus eremicus Rose and Zolnerowich (Hymenoptera: Aphelinidae). The survivorship of adult parasitoids foraging on poinsettia leaves with residues 6, 24, and 96 h of age was determined. The toxicity of Bemisia argentifolii Bellows and Perring (Homoptera: Aleyrodidae) patches treated with IGRs presented to female parasitoids 24 and 96 h posttreatment was quantified. Survivorship of immature E. eremicus developing within B. argentifolii nymphs was determined by treating whitefly nymphs with IGRs 5 and 13 days postoviposition by female parasitoids. Finally, behavioral observations of female parasitoids foraging on IGR-treated and untreated B. argentifolii patches presented simultaneously were quantified to determine whether IGR residues had a repellant effect toward E. eremicus. Averaging ranks for IGRs based on their compatibility with E. eremicus and their ability to kill B. argentifolii nymphs produced the following parasitoid compatibility order: buprofezin > fenoxycarb > pymetrozine = pyriproxyfen > kinoprene. Further work in greenhouses assessing the efficacy of buprofezin with E. eremicus for B. argentifolii control on poinsettias is recommended.     

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.     

Compatibility of Insect Growth Regulators with Eretmocerus eremicus (Hymenoptera: Aphelinidae) for Whitefly (Homoptera: Alyerodidae) Control on Poinsettias: II. Trials in Commercial Poinsettia Crops

The efficacy and cost of reduced release rates of the parasitoid Eretmocerus eremicus Rose and Zolnerowich (Hymenoptera: Aphelinidae) when combined with application of the insect growth regulator buprofezin were compared to those of a higher parasitoid release rate used alone for whitefly control (Homoptera: Aleyrodidae) on poinsettia (Euphorbia pulcherrima Willd. ex Koltz.). The trial was conducted in seven greenhouses in Methuen, Massachusetts from August through December 1997 and employed commercial poinsettia production practices. Two whiteflies species, Trialeurodes vaporariorum (Westwood) and Bemisia argentifolii Bellows and Perring (= Bemisia tabaci [Gennadius] strain B), were present. Three treatments were examined: (1) E. eremicus used alone at a release rate of three females per plant per week (two greenhouses); (2) E. eremicus at an intermediate release rate of two females per plant per week, combined with mid-season use of buprofezin (two applications, spaced 1 week apart, applied in weeks 9 and 10) (two greenhouses); and (3) E. eremicus at a low release rate of one female per plant per week, combined with mid-season use of buprofezin, applied as in treatment 2 (two greenhouses). In addition, observations were made in one additional greenhouse at the site, in which the grower used pesticides for whitefly control. Prior to the start of the trial, cuttings used for all treatments experienced some pesticide use, first abamectinduring rooting and later buprofezin at potting to reduce whitefly numbers, which were initially very high. At harvest, densities of live whitefly nymphs were not statistically different among the biological control treatments, indicating that a low parasitoid release rate combined with buprofezin was as effective as a higher release rate of the parasitoid used alone. Nymphal densities in separate market samples (based on smaller sample sizes) showed differences among treatments, but all treatments, including the low parasitoid release rate + buprofezin maintained densities of live nymphs + pupae at or below approximately two per leaf, a level commercially acceptable in local markets. Control costs per single-stemmed poinsettia plant were $1.18 for the high parasitoid release treatment, $0.75 for the treatment of weekly releases of two female parasitoids per plant per week + buprofezin, $0.38 for the treatment of releases of one female parasitoid per plant per week + buprofezin, and $0.14 for the chemical control greenhouse.     

Analysis of Foraging Behaviour of the Whitefly Parasitoid Encarsia formosa on a plant: A Simulation Study

The foraging behaviour of Encarsia formosa was analyzed using a stochastic simulation model of the parasitoid's behaviour. Parasitoids were allowed to search during a day on a tomato plant infested with immatures of the greenhouse whitefly, Trialeurodes vaporariorum. The model simulates searching, host selection, host handling and patch leaving behaviour, and the physiological state of the parasitoid. The outputs of the model are the number of visited leaflets and the number of hosts encountered, parasitized or killed by host feeding. The simulation results agreed well with observations of parasitoids foraging on tomato plants. The number of encounters and ovipositions on the plant increased with host density according to a type II functional response. At a clustered host distribution over leaflets and low host densities, the most important parameters affecting the number of ovipositions were the leaf area, the parasitoid's walking speed and walking activity, the probability of oviposition after encountering a host, the initial egg load and the ratio of search times on both leaf sides. At high densities, the maximum egg load and the giving-up time on a leaflet since latest host encounter were the most essential parameters.     

Predicted Impact of an Exotic Generalist Predator on Monarch Butterfly (Lepidoptera: Nymphalidae) Populations: A Quantitative Risk Assessment

The multicolored Asian lady beetle, Harmonia axyridis (Pallas), was recently identified as a potential hazard to monarch butterflies, Danaus plexippus (L.). A quantitative risk assessment for the potential impact of H. axyridis on D. plexippus in Minnesota corn and soybean fields was developed using risk analysis software. This assessment considered a potential worst-case scenario for the impact of H. axyridis on D. plexippus. Habitat-specific recruitment of D. plexippus eggs was determined empirically. Subsequently, simulated abundance of D. plexippus in each habitat was reduced by two classes of stage-specific mortality: predation by H. axyridis and causes other than H. axyridis. Predation was modeled as a function of D. plexippus exposure to H. axyridis, and predation rate of D. plexippus by H. axyridis. Exposure and subsequent risk varied considerably by habitat, with a low risk of H. axyridis adversely affecting D. plexippus populations developing in corn fields, but a moderate to high risk in soybean fields. Predicted rates of D. plexippus mortality attributable to H. axyridis were greater in soybean compared to corn fields, possibly due to the numerical response of H. axyridis to soybean aphid, Aphis glycines Matsumura. This study demonstrates that H. axyridis has the potential to have a strong adverse impact on D. plexippus populations. However, the likelihood of occurrence for this worst-case scenario remains uncertain. To evaluate the landscape-level risk of H. axyridis impacting D. plexippus, further data on recruitment of D. plexippus in other habitats and exposure estimates specific to additional habitats are needed.     

Effect of Prey and Predator Density on Predation of Rice Leaffolder Eggs by the Cricket Metioche vittaticollis

Cage experiments were conducted to quantify the predation rate of the cricket Metioche vittaticollis (Sta ˚l) on the eggs of rice leaffolder Marasmia patnalis Bradley. Egg predation by adult females was measured in response to changes in egg density, predator density and leaf area per cage. The number of eggs consumed per predator increased with egg density, without reaching a plateau. The predation rate decreased with increasing leaf area. The functional response could be adequately described with a linear Type I model, with the effect of leaf area included. This type of response to leaffolder egg density means that predation was not limited by prey handling time or satiation, but by the search rate. The search rate is here interpreted as the leaf area effectively searched for leaffolder eggs by a single predator in one day. Estimated search rates averaged 0.13 m 2 day -1 for M. vittaticollis females. The search rate of the predators increased with prey density, but a model describing a density dependent search rate explained only 3% of the total variation in search rate. Increasing predator density per cage led to a decrease in the per capita egg predation rate when predator density was more than two per m 2 leaf area. Interference might thus reduce the potential to enhance leaffolder egg predation by conservation or augmentation of predatory cricket populations.     

Behavioral Response to Larval Tracks and the Influence of Tracks on Intraguild Scavenging by Coccinellid Larvae

A paired design was used to determine that Harmonia axyridis 4th instars were not influenced by the presence of conspecific larval tracks, but well-fed H. axyridis 4th instars spent less time on plants that contained tracks left by Coleomegilla maculata 4th instars. To determine if the presence of larval tracks influences intraguild scavenging by H. axyridis 4th instars, dead 4th instars were placed in Petri dishes that contained or did not contain larval tracks. The presence of larval tracks did not influence the feeding frequency or the amount of time before feeding. However, larvae dragged their pygopod on dish surfaces more frequently if the dish contained larval tracks. In addition, starved H. axyridis larvae were more likely to feed on the prey and dragged their pygopod less frequently than well-fed larvae.     

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.     

Dietary Plasticity of Generalist and Specialist Ungulates in the Namibian Desert: A Stable Isotopes Approach

Desert ungulates live in adverse ecosystems that are particularly sensitive to degradation and global climate change. Here, we asked how two ungulate species with contrasting feeding habits, grazing gemsbok (Oryx g. gazella) and browsing springbok (Antidorcas marsupialis), respond to an increase in food availability during a pronounced rain period. We used a stable isotope approach to delineate the feeding habits of these two ungulates in the arid Kunene Region of Namibia. Our nineteen months field investigation included two time periods of drought when food availability for ungulates was lowest and an intermediate period with extreme, unusual rainfalls. We documented thirteen isotopically distinct food sources in the isotopic space of the study area. Our results indicated a relatively high dietary plasticity of gemsbok, which fed on a mixture of plants, including more than 30% of C3 plants during drought periods, but almost exclusively on C4 and CAM plant types when food was plentiful. During drought periods, the inferred gemsbok diets also consisted of up to 25% of Euphorbia damarana; an endemic CAM plant that is rich in toxic secondary plant compounds. In contrast, springbok were generalists, feeding on a higher proportion of C3 than C4/CAM plants, irrespective of environmental conditions. Our results illustrate two dietary strategies in gemsbok and springbok which enable them to survive and coexist in the hostile Kunene arid ecosystem.     

Life History of the Whitefly Predator Nephaspis oculatus (Coleoptera: Coccinellidae) at Six Constant Temperatures

Nephaspis oculatus (Blatchley) is a predator of whiteflies including Bemisia tabaci (Gennadius) and B. argentifolii Bellows and Perring. The usefulness of this and other coccinellid predators of whiteflies could be improved by better information on responses to different temperatures experienced in the field. We reared N. oculatus from egg to adult in incubators at six constant temperatures and observed a linear relationship between developmental time and temperature for all preimaginal lifestages through all but the highest temperatures (33°C). Four larval instars were observed, although a significant proportion of third instar larvae molted directly to the pupal stage at 29°C and above. Complete life table data were obtained at all temperatures except 33°C. Survivorship was greatest over all immature stages at 26°C and finite and intrinsic rates of increase were highest at this temperature. Although 26°C appeared to be the optimum temperature for population growth, estimated rates of increase at all temperatures tested above 20°C were within 75% of the maximum, indicating that the beetle can perform well within the temperature range usually found in greenhouses.     

Intraguild Predation among the Hoverfly Episyrphus balteatus de Geer (Diptera: Syrphidae) and Other Aphidophagous Predators

Aphidophagous predators compete for the same prey species. During their foraging activity they frequently encounter heterospecific aphid predators. These situations can lead to intraguild predation and may disrupt biological control efforts against aphids where more than one predator species is present. We investigated the behavior of larvae of the hoverfly Episyrphus balteatus de Geer and its interaction with three other aphid predators: the ladybird Coccinella septempunctata L., the lacewing Chrysoperla carnea Stephens, and the gall midge Aphidoletes aphidimyza (Rondani). Interspecific interactions between predators were examined in arenas of different sizes and in the presence of extraguild prey. The outcome of interactions between E. balteatus larvae and the other predators depended predominantly on the relative body size of the competitors. Relatively large individuals acted as intraguild predators, while relatively smaller individuals became intraguild prey. Eggs and first- as well as second-instar larvae of E. balteatus were highly susceptible to predation by all other predators, whereas pupae of E. balteatus were preyed upon only by the larvae of C. carnea. Interactions between A. aphidimyza and E. balteatus were asymmetric and always favored the latter. Eggs and first- as well as second-instar larvae of E. balteatus sustained intraguild predation irrespective of the size of the arena or the presence of extraguild prey. However, the frequency of predation on third-instar larvae of E. balteatus was significantly reduced. This study indicated that the same species can be both intraguild predator and intraguild prey. It is suggested that combinations of predators must be carefully chosen for success in biological control of aphids.     

Behavioral and Neurochemical Effects of Fencamfamine on Rats: A Chronobiologic Approach

Biological Rhythms in Clinical and Laboratory Medicine. Edited by Y. Touitou and E. Haus. Berlin, Heidelberg, New York Springer-Verlag, 1992, 730 pp. (347 figures and 54 tables), $210.     

A Behavioral Mechanism for Incorporating an Unpalatable Food in the Diet of a Generalist Herbivore (Orthoptera: Acrididae)

We investigated mechanisms that could lead to incorporation of unpalatable foods into the diet of a generalist grasshopper, Schistocerca americana: nutritional stress, habituation, learning, and attraction to novelty. The model system involved mesquite, a palatable but inferior food, and mulberry, an unpalatable but adequate food. Nutritional stress, due to prolonged intake of the inferior food, mesquite, did not increase the acceptability of mulberry. Habituation to the deterrent compounds in mulberry and associative learning of the nutritional benefits of mulberry also did not occur. However, mulberry became more acceptable after a day of restriction to a single food type other than mulberry, and even deterrent and nutritionally worthless alternatives such as filter paper became acceptable after a day on any one food type. A tendency to feed on novel food types may be a proximate mechanism for the incorporation of relatively unpalatable, but nutritionally valuable foods into the diet. Novelty and the apparent need for diversity of foods are discussed in the context of exploratory foraging behavior by generalist herbivores.     

Behavioral and Neurochemical Effects of Fencamfamine on Rats: A Chronobiologic Approach

Fencamfamine (FCF) is a psychostimulant classified as an indirect dopaminergic agonist. Circadian rhythms of some behavioral and neurochemical parameters were investigated in control rats and in rats which had been treated with a single dose of FCF across the 24-hr span. Rats were entrained to light/dark (LD) 12:12, lights on from 0700 to 1900. In behavioral experiments (performed in March) the rats were injected intraperitoneally with saline or FCF (3.5 mg/kg) at one of six times: 0900, 1300, 1700, 2100, 0100 or 0500. Fifteen minutes after treatment the duration of sniffing, rearing and locomotion was recorded during 120 min. Controls showed circadian rhythms for sniffing and rearing with acrophases at 2255 and 0118, respectively. In animals treated with FCF, only locomotion displayed significant circadian variation with acrophase at 1912. Two-way analysis of variance (ANOVA) showed a statistically significant circadian time-dependent effect of FCF on all behavioral parameters studied; the increase of sniffing, rearing and locomotion induced by FCF was higher in rats treated during the rest phase. In the biochemical studies (performed between March-June), rats were treated (i.p.) with saline or FCF (10 mg/kg) at one of four times: 0900, 1700, 2100 or 0100. The levels of homovanillic acid (HVA) in the striatum and tuberculum olfactorium, 5-hydroxyindolacetic acid (5-HIAA) in the cerebellum and 3-methoxy-4-hydroxypheniglycol (MHPG) in the frontal cortex were determined. Controls showed circadian rhythms for HVA (striatum), MHPG (frontal cortex) and 5-HIAA (cerebellum) with acrophases at 2233, 1955 and 1029, respectively.(ABSTRACT TRUNCATED AT 250 WORDS)     

Bioinsecticide-Predator Interactions: Azadirachtin Behavioral and Reproductive Impairment of the Coconut Mite Predator Neoseiulus baraki

Synthetic pesticide use has been the dominant form of pest control since the 1940s. However, biopesticides are emerging as sustainable pest control alternatives, with prevailing use in organic agricultural production systems. Foremost among botanical biopesticides is the limonoid azadirachtin, whose perceived environmental safety has come under debate and scrutiny in recent years. Coconut production, particularly organic coconut production, is one of the agricultural systems in which azadirachtin is used as a primary method of pest control for the management of the invasive coconut mite, Aceria guerreronis Keifer (Acari: Eriophyidae). The management of this mite species also greatly benefits from predation by Neoseiulus baraki (Athias-Henriot) (Acari: Phytoseiidae). Here, we assessed the potential behavioral impacts of azadirachtin on the coconut mite predator, N. baraki. We explored the effects of this biopesticide on overall predator activity, female searching time, and mating behavior and fecundity. Azadirachtin impairs the overall activity of the predator, reducing it to nearly half; however, female searching was not affected. In contrast, mating behavior was compromised by azadirachtin exposure particularly when male predators were exposed to the biopesticide. Consequently, predator fecundity was also compromised by azadirachtin, furthering doubts about its environmental safety and selectivity towards biological control agents.     

Intraguild predation on Eretmocerus sp. nr. emiratus,a parasitoid of Bemisia tabaci,by three generalist predators with implications for estimating the level and impact of parasitism

Intraguild predation (IGP) is pervasive in many managed and unmanaged ecosystems and may have negative, neutral or positive effects on the biological control of pest insects. Both generalist predators and aphelinid parasitoids attack Bemisia tabaci (Gennadius) Biotype B (=B. argentifolii Perring & Bellows) on cotton in the southwestern USA. Free-choice and no-choice laboratory assays were conducted to quantify prey consumption patterns and preference by three representative generalist predators, Geocoris punctipes (Say), Orius insidiosus (Say), and Hippodamia convergens Guérin-Méneville, offered fourth instar B. tabaci nymphs and nymphs parasitized by Eretmocerus sp. nr. emiratus. All three predators showed a significant preference for larval and pupal stage parasitoids over early fourth instar nymphs, but G. punctipes and O. insidiosus were non-discriminating when offered a choice of larval parasitoids and late fourth instar nymphs. The potential implications of these observed patterns for the field were examined through sensitivity analyses of existing field life table data. First, preference for parasitized hosts alters the methods required for calculating marginal rates of parasitism. Incorporating a preference variable in the estimation procedure had a very small positive effect (0.02–1.13% change) on total generational mortalities observed in previous life table studies. However, further hypothetical analyses suggested that under circumstances of lower generational mortality and higher levels of either apparent parasitism or predation, high levels of predator preference for parasitized prey could alter estimates of total mortality as much as 14%. Second, although intraguild predation was demonstrated, the implications for biological control are unclear. Based on field life table data the rate of IGP ranged from 0.019 to 0.126 depending on predator species and prey comparison, but accounting for these levels of IGP had only small negative effects on total generational mortalities (0.193–1.278% change).