Effects of spring imidacloprid application for white grub control on parasitism of Japanese beetle (Coleoptera: Scarabaeidae) by Tiphia vernalis (Hymenoptera: Tiphiidae)

Imidacloprid, a relatively long residual neonicotinoid soil insecticide, is often applied to lawns and golf courses in spring for preventive control of root-feeding white grubs. We evaluated effects of such applications on spring parasitism of the overwintered third-instar Japanese beetle, Popillia japonica Newman, by Tiphia vernalis Rohwer, an introduced solitary ectoparasitoid. Natural rates of parasitism on a golf course rough were significantly lower in plots treated with full or one-half label rates of imidacloprid in early May compared with untreated turf. Parasitism also was reduced when female T. vernalis were exposed to imidacloprid residues on turf cores in the laboratory. Such exposures did not affect wasp mortality, longevity, survival, or developmental period of Tiphia larvae feeding on hosts in treated turf. They did, however, reduce wasps' ability to parasitize hosts in nontreated soil for at least 1-2 wk postexposure. In Y-trail choice tests, wasps that previously had been exposed to treated turf failed to respond normally to host frass trails in the soil. Female wasps did not avoid imidacloprid residues, imidacloprid-treated host frass, or host grubs that had previously been exposed to treated soil. This study indicates that applying imidacloprid in early spring can interfere with biological control by T. vernalis, whereas postponing preventive grub treatments until June or July, after the wasps' flight period, will help to conserve T. vernalis populations.     

Use of herbivore‐induced plant volatiles as search cues by Tiphia vernalis and Tiphia popilliavora to locate their below‐ground scarabaeid hosts

Japanese beetle, Popillia japonica Newman, and oriental beetle, Anomala orientalis (Waterhouse) (both Coleoptera: Scarabaeidae) are considered invasive species and have been reported as key pests of urban landscapes in the Northeastern USA. Tiphia vernalis Rohwer and Tiphia popilliavora Rohwer (Hymenoptera: Tiphiidae) were introduced as biocontrol agents against these beetles. These parasitic wasps burrow into the soil and search for grubs. When a host is found, the wasp attaches an egg in a location that is specific for the wasp species. It is unknown if these wasps can detect patches of concealed hosts from a distance above ground and what role, if any, herbivore‐induced plant volatiles play in their host location. This study evaluated the responses of female T. vernalis and T. popilliavora to grub‐infested and healthy plants in Y‐tube olfactometer bioassays. Also the effect of root herbivory on the composition of turfgrass (Poaceae) volatile profiles was investigated by collecting volatiles from healthy and grub‐infested grasses. Tiphia wasps were highly attracted to volatiles emitted by grub‐infested tall fescue (Festuca arundinacea Schreb.) and Kentucky bluegrass (Poa pratensis L.) over healthy grasses. In contrast, wasps did not exhibit a significant preference for grub‐infested perennial ryegrass (Lolium perenne L.) as compared with the control plants. The terpene levels emitted by grub‐infested Kentucky bluegrass and tall fescue were greater than that of control plants. Low levels of terpenes were observed for both test and control perennial ryegrass. The elevated levels of terpenes emitted by grub‐infested Kentucky bluegrass and tall fescue coincided with the attractiveness to the tiphiid wasps. Here, we provide evidence that plant exposure to root‐feeding insects P. japonica and A. orientalis resulted in an increase in terpenoid levels in turfgrasses, which strongly attracts their above‐ground parasitoids.     

Positive and negative impacts of insect frass quality on soil nitrogen availability and plant growth

Frass deposition to soil is an important pathway by which herbivorous insects impact decomposition and soil nutrient availability. However, little is known about how frass quality influences ecosystem properties. Here, we examined the effects of frass quality on the decomposition process, soil nitrogen (N) availability, and plant growth, using frass of Mamestra brassicae (L.) that fed on fertilized or unfertilized Brassica rapa L. var. perviridis Bailey. The frass quality was largely dependent on the host plant quality. Frass excreted by larvae that fed on the fertilized plants had higher N than that of larvae that fed on the unfertilized plants. The decomposition rate of the frass did not differ between N-rich and N-poor frass, except during the early decomposition period. The inorganic N concentration decreased during decomposition in both frass types. However, difference in the initial inorganic N concentration led to different consequences regarding soil N availability. Furthermore, addition of frass to the soil differently influenced the growth of B. rapa plants depending on the frass quality: plant biomass was increased by N-rich frass addition but decreased by N-poor frass addition, compared to the biomass without frass addition. These results indicate that frass quality is an important factor in determining the impact of herbivorous insects on nutrient dynamics, and that frass positively or negatively influences soil N availability and plant growth, depending on its quality.     

Host recognition by the specialist endoparasitoidMicroplitis croceipes (Hymenoptera: Braconidae): Role of host- and plant-related volatiles

The specialist parasitoidMicroplitis croceipes Cresson can parasitize only noctuid larvae in the generaHelicoverpa andHeliothis. To be successful in their search for hosts, the ability to distinguish hosts from nonhosts feeding on the same plant is beneficial. In flight tunnel experiments, we found that prior to landing on the odor sourceM. croceipes were able to distinguish volatiles released from frass of host larvae(Helicoverpa zea Boddie) and nonhost larvae (Spodoptera exigua Hübner andSpodoptera frugiperda J. E. Smith) fed on cotton. However, an initial contact experience with frass of cotton-fed host larvae appeared to be critical for this ability. Wasps that had antennated frass of host larvae fed pinto bean diet were equally attracted to frass of host and nonhost larvae fed on pinto bean diet. In short-range walking experiments, wasps located cotton-fed host larvae faster than diet-fed larvae, regardless of their experience. Wasps that had antennated frass of cotton-fed host larvae were less attracted to cotton-fed nonhost larvae, compared to host larvae, and preferred to sting host larvae. Plant-related volatiles in host frass and larvae appear to play a major role in the successful location of host larvae.     

Specificity of termite trails: Analysis of natural trails of Trinervitermes, macrotermes and Odontotermes from sympatric populations

Laboratory tests for selective trail-following between naturally established trails indicate that termites from neighbouring colonies of Trinervitermes bettonianus (Isoptera: Termitidae) do not show a significant preference for their own colony's trails in a choice situation. However, termites from different genera, T. bettonianus, Macrotermes michaelseni and Odontotermes sp. from neighbouring populations recognize their respective trails with a high degree of precision.

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Résumé Des expériences de laboratoire ont montré que les Trinervitermes bettonianus provenant de deux colonies voisines, quand ils ont le choix, ne préfèrent pas les pistes de leur propre colonie. Par contre, les termites Trinervitermes bettonianus, Macrotermes michaelseni et Odontotermes sp., provenant de populations voisines, reconnaissent avec une grande précision leurs propres pistes.

     

Host Location of Hyssopus pallidus, a Larval Parasitoid of the Codling Moth, Cydia pomonella

The effectiveness of parasitoids as biological control agents depends largely on their host location behavior. In this study we describe the host-searching behavior of Hyssopus pallidus (Askew), a larval parasitoid of the codling moth Cydia pomonella L. We observed parasitoid behavior on mature apples (a potential host patch) and elucidated some of the stimuli which determine host location. Female wasps showed more complex and intensive searching on infested apples than on mechanically damaged or noninfested apples. Wasps were able to enter infested apples through the calyx or the tunnel made by the host larvae and parasitize them. Area-restricted searching was observed on infested apples, in particular on areas contaminated with host frass. In a further bioassay, we confirmed that host frass contains a host location kairomone. The kairomone appears to be produced by the host independently from its diet, even though frass produced by hosts fed on an apple are more attractive than frass produced by hosts fed on a fruit-devoid artificial diet. The capability of H. pallidus to locate its host inside apples makes the foraging strategy of this species potentially useful as a biological control agent.     

Comparative flight behavior of parasitoids Campoletis sonorensis and Microplitis croceipes

Flight behavior of parasitoids Microsplitis croceipes Cresson (Braconidae) and Campoletis sonorensis (Cameron) (Ichneumonidae) was examined in a wind tunnel. Both species exhibited odor directed, oriented flights to cotton, although flight frequency was related to variation in wind velocity or differential cotton volatile emission with changes in wind velocity, or both. Flight frequency at constant wind velocity was affected by illumination. Microplitis croceipes was attracted to the wind borne odor of Heliothis virescens (F.) frass and larvae, whereas C. sonorensis was not. Possible interaction of the wind borne odors from the habitat and host is suggested for parasitoids.

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Résumé Le comportement de vol de M. croceipes Cres. (Hym. Braconidae) et de C. sonorensis Cam. (Hym. Icheumonidae) a été étudié en olfactométrie dans un tunnel à vent. Les femelles des deux espèces, et non les mâles, ont un vol orienté par l'odeur des feuilles fraîches de coton (Gossypium hirsutum L.). La fréquence d'envol est liée aux changements de vitesse du vent, aux variations d'émissions, par le cotonnier, des substances volatiles, en fonction de la vitesse du vent, ou aux deux. La fréquence d'envol à vitesse de vent constante dépend de l'éclairage. La fréquence d'envol de C. sonorensis augmente avec des intensités lumineuses relativement fortes et diminue avec des intensités relativement faibles. L'inverse est observé avec M. croceipes. Le vent chargé d'odeurs de larves ou d'extréments d'Heliothis virescens F., attire M. croceipes et non C. sonorensis. Nous suggérons une éventuelle interaction des odeurs portées par le vent et provenant de l'habitat et de l'hôte.

     

Evaluation of endophytic actinobacteria as antagonists of Gaeumannomyces graminis var. tritici in wheat

Endophytic actinobacteria isolated from healthy cereal plants were assessed for their ability to control fungal root pathogens of cereal crops both in vitro and in planta. Thirty eight strains belonging to the genera Streptomyces, Microbispora, Micromonospora, and Nocardioidies were assayed for their ability to produce antifungal compounds in vitro against Gaeumannomyces graminis var. tritici (Ggt), the causal agent of take-all disease in wheat, Rhizoctonia solani and Pythium spp. Spores of these strains were applied as coatings to wheat seed, with five replicates (25 plants), and assayed for the control of take-all disease in planta in steamed soil. The biocontrol activity of the 17 most active actinobacterial strains was tested further in a field soil naturally infested with take-all and Rhizoctonia. Sixty-four percent of this group of microorganisms exhibited antifungal activity in vitro, which is not unexpected as actinobacteria are recognized as prolific producers of bioactive secondary metabolites. Seventeen of the actinobacteria displayed statistically significant activity in planta against Ggt in the steamed soil bioassay. The active endophytes included a number of Streptomyces, as well as Microbispora and Nocardioides spp. and were also able to control the development of disease symptoms in treated plants exposed to Ggt and Rhizoctonia in the field soil. The results of this study indicate that endophytic actinobacteria may provide an advantage as biological control agents for use in the field, where others have failed, due to their ability to colonize the internal tissues of the host plant.     

Differences in the virulence of Heterorhabditis bacteriophora and Steinernema scarabaei to three white grub species: The relative contribution of the nematodes and their symbiotic bacteria

Because susceptibility of white grub species to entomopathogenic nematodes differs, we compared the virulence of Photorhabdus temperata and Xenorhabdus koppenhoeferi, the symbiotic bacteria of the nematodes Heterorhabditis bacteriophora and Steinernema scarabaei, respectively, to the three white grub species, Popillia japonica, Rhizotrogus majalis, and Cyclocephala borealis. Both bacteria were pathogenic to all three grub species even at 2 cells/grub. However, the median lethal dose at 48 h post injection and median lethal time at 20 cells/grub showed that P. temperata was more virulent than X. koppenhoeferi to C. borealis. Although H. bacteriophora is less pathogenic than S. scarabaei to R. majalis and P. japonica, their symbiotic bacteria did not differ in virulence against these two grub species, and they also showed similar growth patterns both in vitro and inside R. majalis larvae at 20 °C. We then tested the pathogenicity of oral- and intrahemocoel-introduced H. bacteriophora to R. majalis to determine whether nematodes are able to successfully vector the bacteria into the hemolymph. Hemocoel injected H. bacteriophora was pathogenic to R. majalis indicating successful bacterial release, but orally introduced H. bacteriophora were not. Dissection of grubs confirmed that the orally introduced H. bacteriophora were unable to penetrate into the hemolymph through the gut wall. We conclude that the low susceptibility of R. majalis to H. bacteriophora is not due to the symbiotic bacteria but rather to the nematode’s poor ability to penetrate through the gut wall and the cuticle to vector the bacteria into the hemolymph.     

Response of the Carrot Weevil,Listronotus oregonensis(Coleoptera: Curculionidae), to Strains ofBacillus thuringiensis

Mortality and frass production bioassays were used to investigate the toxicity of seven strains ofBacillus thuringiensisagainst the adult carrot weevil,Listronotus oregonensis(Le Conte). A semi-artificial diet of carrot foliage with 4% agar was selected to maximize feeding by the insects.Bacillus thuringiensissubsp.tenebrionis(Krieg, Huger, Langenbruch, and Schnetter) (BTT) and two unidentifiedB. thuringiensisstrains, A30 and A429, gave the lowest LC₅₀values. The frass bioassay supported the conclusions of the mortality assay. Mortality of adults continued after their removal from the insecticidal medium, with the highest mortality being caused by strains A429 and BTT. Survivors from the frass bioassay, initially exposed to strains A30, A429, and BTT, did not resume normal levels of feeding after their removal from the insecticidal medium.     

The parasitoidCotesia glomerata (Hymenoptera: Braconidae) discriminates between first and fifth larval instars of its hostPieris brassicae,on the basis of contact cues from frass,silk, and herbivore-damaged leaf tissue

Adult females of the larval parasitoidCotesia glomerata (L.) respond to chemical cues associated with feeding damage inflicted on cabbage plants by its host,Pieris brassicae (L.). The use of these infochemicals by the parasitoid during selection of the most suitable host instar was investigated. The parasitoid can successfully parasitize first-instar host larvae, while contacts with fifth-instar larvae are very risky since these caterpillars react to parasitization attempts by biting, spitting, and hitting, resulting in a high probability of the parasitoid being seriously injured or killed. Observations of the locomotor behavior of individual wasps on leaves with feeding damage inflicted by the first and the fifth larval instars and on host silk and frass showed that several cues affect the duration of searching by the parasitoids after reaching a leaf: cues on the margin of the feeding damage and cues in the host frass and silk. Whole frass, silk, and hexane extracts of frass obtained from first-instar elicited parasitoid's searching behavior significantly longer than frass, silk, and hexane extract of frass from the fifth instar. The results demonstrate thatC. glomerata can discriminate between first instars, which are more suitable hosts, and fifth instars ofP. brassicae without contacting the caterpillars, by exploiting instar-related cues.     

Recycling of nitrogen in herbivore feces: plant recovery,herbivore assimilation,soil retention,and leaching losses

Herbivores directly and indirectly affect ecosystem functioning in forests. Feces deposition is a direct effect that supplies ephemeral N pulses to soils. Herbivore-mediated changes in plant N allocation and uptake are indirect effects that can also influence soil N availability. These effects may interact if defoliation influences the ability of plants to recover fecal N, and this may affect subsequent generations of herbivores. We added ¹⁵N-enriched insect feces (frass) to a series of replicated red oak, Quercus rubra, mesocosms that had been damaged experimentally and then followed the frass N over the course of 2 years. In the first season, some frass N was mineralized in the soil and leached in organic form from the mesocosms within 1 week of deposition. Within 1 month, frass N had been acquired by the oaks and enriched the foliage; late-season herbivores assimilated the frass N within the same growing season. In the second season, herbivore damage from the previous year lowered total leaf N contents and ¹⁵N recovered in the foliage. A subsequent cohort of early-season herbivores fed on this foliage consequently derived less of their N from the previous year’s frass, and feral leaf rollers colonized fewer of these saplings. The 0- to 5-cm soil fraction was the largest N sink measured, and 42% of the frass N was recovered in the soil. The results demonstrate that: (1) some frass N can be recycled rapidly into foliage and assimilated by successive cohorts of herbivore within the same season; (2) damage can affect N allocation in the following year’s foliage, influencing N availability to and host selection by herbivores; and (3) leaching losses occur soon after deposition but are buffered by soil pools, which are the largest sinks for frass N.     

Electroantennogram (EAG) responses of Microplitis croceipes and Cotesia marginiventris and their lepidopteran hosts to a wide array of odor stimuli: Correlation between EAG response and degree of host specificity?

In order to test whether the electroantennogram (EAG) response spectrum of an insect correlates to its degree of host specificity, we recorded EAG responses of two parasitoid species with different degrees of host specificity, Microplitis croceipes (specialist) and Cotesia marginiventris (generalist), to a wide array of odor stimuli including compounds representing green leaf volatiles (GLVs), herbivore-induced plant volatiles (HIPV), ecologically irrelevant (not used by the parasitoid species and their hosts for host location) plant volatiles, and host-specific odor stimuli (host sex pheromones, and extracts of host caterpillar body and frass). We also tested the EAG responses of female moths of the caterpillar hosts of the parasitoids, Heliothis virescens and Spodoptera exigua, to some of the odor stimuli. We hypothesized that the specialist parasitoid will have a narrower EAG response spectrum than the generalist, and that the two lepidopteran species, which are similar in their host plant use, will show similar EAG response spectra to plant volatiles. As predicted, the specialist parasitoid showed greater EAG responses than the generalist to host-specific odor and one HIPV (cis-3-hexenyl butyrate), whereas the generalist showed relatively greater EAG responses to the GLVs and unrelated plant volatiles. We detected no differences in the EAG responses of H. virescens and S. exigua to any of the tested odor.     

Host-finding behavior, host acceptance, and host suitability of the parasiteXanthopimapla stemmator

Xanthopimpla stemmator (Thunberg)(Hymenoptera: Ichneumonidae), a solitary endoparasite of pupae of Old World lepidopteran stalkborers, was recently imported into Texas as a candidate for biological control of New World stalkborers. Information on host acceptability, host suitability and cues responsible for host finding were necessary to gain an insight into parasite/host interactions, because of the absence of a coevolutionary history.Xanthopimpla stemmator females were exposed to laboratory-reared one-to six-day-oldDiatraea saccharalis (F.) pupae. An average of 62% of host pupae were accepted and all ages of pupae were equally acceptable. Host suitability decreased with host age. One- to five-day-old host pupae averaged 31–37% suitability, whereas only 19% of 6-day-old pupae were suitable. Successful parasitization, defined as the product of the proportion accepted and the proportion suitable, decreased from 22–23% for 1-, 2- and 3-day-old pupae to 13% for 6-day-old pupae. Sex ratio (female:male) of the parasite progeny increased with host age. Females comprised 47% of total parasite progeny of 1-day-old and 84% of 6-day-old pupae. The increase in percent females was a result of a similar number of females in all age classes, coupled with a decrease in the number of males from older hosts.Xanthopimpla stemmator superparasitized 61% of acceptedD. saccharalis pupae in the laboratory. On dissection, 73% of host pupae with multiple probe wounds were found to contain parasite eggs or larvae; these hosts contained up to 10 eggs or 7 first-instar larvae. Increased numbers of probes by the parasites were associated with an increase in successful parasitization. Host seeking activity inX. stemmator was stimulated by the presence of larval frass, host odor and movement of host pupae. Results suggest thatX. stemmator is a good candidate for biological control ofD. saccharalis and possibly other factitious stalkborer hosts.     

Hyperparasitism by Mesochorus spp. (Hymenoptera: Ichneumonidae) in Peristenus sp. (Hymenoptera: Braconidae) and development of PCR primers for hyperparasitoid detection

Peristenus sp. pupae collected from Lygus spp. nymphs in 2001 and 2002 were over-wintered in the laboratory. In both years, more than 30% of adults emerging from over-wintering pupae were identified as ichneumonid hyperparasitoids, Mesochorus curvulus Thomson and Meschorus sp. (Hymenoptera: Ichneumonidae). At the end of the over-wintering period, Peristenus sp. males emerged first followed by Peristenus sp. females and finally Mesochorus spp. The male:female ratio in emerging Peristenus sp. adults was skewed towards males. The Internal Transcribed Spacer (ITS) region and the cytochrome oxidase I (COI) gene from Mesochorus spp. were sequenced. ITS sequences were used to develop PCR primers to detect Mesochorus spp. hyperparasitism in the primary host, Lygus spp. PCR analysis of field-collected Lygus spp. nymphs gave similar estimates of Mesochorus spp. hyperparasitism to the rearing protocols (25–28%). Sequence analysis of COI and ITS regions and subsequent restriction endonuclease analysis of ITS PCR products from Mesochorus spp. indicate the presence of two genotypes in the population. The possibility that these two genotypes represent separate or cyrptic species is discussed.     

Host frass as arrestant chemicals in locating host Mythimna separata by the tachinid fly Exorista japonica

We investigated the response of the tachinid fly, Exorista japonica (Townsend), to host frass or its extracts in order to clarify the host location mechanisms of female flies in a potential host habitat. Host searching time in a patch and the number of patch visits were analyzed by using a frass-containing patch which was excreted by host larvae, Mythimna separata (Walker) (Lepidoptera: Noctuidae), and patches to which host frass extracts were applied. E. japonica females were arrested in response to the host-frass-containing patch after contacting the frass with their front tarsi, thereby spending most of the time to search the patch and to revisit the host-frass patch. While host-searching time in the patch by the females was longest at their first visit of a patch with host frass, searching time decreased with successive visits. The female flies also exhibited area-restricted searching with methanol extracts of the host frass. Area-restricted searching activity increased with the concentration of host-frass extract, i.e., total searching time in the patch and the number of patch visits varied in a dose-dependent manner. E. japonica females likely employ chemicals in host frass as arrestants in host location.     

Synergies between biological and neonicotinoid insecticides for the curative control of the white grubs Amphimallon majale and Popillia japonica

Synergistic combinations of biological and chemical insecticides might yield promising alternatives for soil insect pest management. In turfgrass of the Northeast U.S., control of root-feeding scarab larvae is highly dependent on conventional insecticides. Studies on interactions between entomopathogenic nematodes and neonicotinoid insecticides, however, demonstrate the feasibility of synergies as an approach for reduced-risk curative control. To understand the breadth of potential synergies, we screened numerous combinations of biological control agents with sublethal doses of neonicotinoids against third instars. Interactions were characterized as synergistic, additive or antagonistic. The most promising combinations identified in laboratory bioassays were advanced to greenhouse pot studies and then to field trials featuring microplots with artificially infested populations. To reveal variation across scarab species, trials were conducted on Amphimallon majale and Popillia japonica. Synergies were consistent across trials and specific to white grub species. For A. majale, synergistic combinations of Heterorhabditis bacteriophora with imidacloprid and clothianidin were discernible in laboratory, greenhouse and field trials. For P. japonica, synergistic combinations of Beauveria bassiana and Metarhizium anisopliae with both neonicotinoids were discernible in the laboratory and greenhouse, but not in the field. For both species, antagonistic interactions were discernible between Bt-products and both neonicotinoids. While nematode-neonicotinoid synergies among scarab larvae have been examined before, fungi-neonicotinoid synergies are unreported. In the context of previous studies, however, no patterns emerge to explain variation across target species or control agent. Further study of non-additive interactions will guide how biological and chemical products could be combined to enhance soil insect pest management.     

Effects of different Beauveria bassiana isolates on field populations of Lygus lineolaris in pigweed (Amaranthus spp.)

The tarnished plant bug, Lygus lineolaris (Palisot de Beauvois), is a pest of various fruit, vegetable, fiber, and seed crops; including cotton. Lygus spp. populations often build on alternate host plants before moving to cotton, and in the midsouthern U.S. wild host plants, such as pigweed (Amaranthus spp.), play a major role in L. lineolaris population development. Three isolates of the entomopathogenic fungus Beauveria bassiana (Balsamo) were evaluated for L. lineolaris control in redroot pigweed (Amaranthus retroflexus L.): one from L. lineolaris in Mississippi (TPB3); one from Lygus hesperus (Knight) in California (WTPB2); and one commercial isolate from Mycotrol^® (GHA). Fungal applications resulted in moderate to high mycosis in adults (33 to 80%) and moderate mycosis in nymphs (36 to 53%) that were collected from field plots at 2 days post-treatment and incubated under laboratory conditions. Although TPB3 was previously found to be more pathogenic in laboratory bioassays, there was not a consistent separation of this isolate from the other two isolates in field trials. Where differences in adult mycosis or mortality were observed, TPB3 was the most pathogenic. However, in one field trial 7 day mortality for nymphs treated with GHA was higher than those treated with TPB3 or WTPB2. Infection rates at 2, 7, and 14 days post-treatment from caged and non-caged adults suggested that movement of adults among plots occurred, which could have masked some treatment effects. Fungal treatments did not significantly reduce populations relative to controls. This may have been caused by delayed mortality rates under field conditions and/or difficulties with estimating population change under field conditions characteristic of wild host plant populations (e.g., heterogeneous populations, adult movement, and small plot size). Further work evaluating time–dose–mortality over dynamic temperatures, spring and fall field trials on this and other wild hosts, and improved methods for estimating populations on wild hosts are needed.     

Hunting for moving hosts: Cephalonomia tarsalis, a parasitoid of free-living grain beetles

The parasitic wasp Cephalonomia tarsalis parasitizes larvae of the saw-toothed grain beetle Oryzaephilus surinamensis, which feed on wheat grains. In contrast to most other host–parasitoid systems studied so far, the grain beetles are highly mobile within their habitat, bulk of grains in grain stores. This should increase the wasps’ problem to locate the hosts. To study the host-finding strategy of C. tarsalis females, the reaction of wasps to different grain and host-derived odour sources was tested in a four-chamber-olfactometer. These experiments revealed that wasps were attracted by healthy grains and mechanically damaged grains. In direct comparison, healthy and mechanically damaged grains are equally attractive. Both potential sources of host-derived odours, host faeces and trail-traces of larvae on filter paper were attractive to the wasps. The response to trail-traces vanished 30 min after larvae had been removed from the filter paper. With respect to the specificity of the odours, it turned out that wasps were attracted to odours from the seed–host complexes from O. surinamensis and Oryzaephilus mercator and the non-host complex of larvae of the granary weevil Sitophilus granarius in wheat grains. Odours from the seed–host complex were preferred. From these results, we hypothesize that host habitat location in C. tarsalis is achieved by using grain-derived odours. Within the habitat, wasps search for kairomones from host faeces and host trails. Following these larval trails finally leads wasps to their hosts.     

Host density influences parasitism of the armyworm Pseudaletia unipuncta in agricultural landscapes

Several studies have shown positive responses of parasitism to either host density or landscape complexity. However, no experiments have manipulated host density in landscapes of differing complexity. Here we report the results of a field experiment conducted to determine how host density and agricultural landscape structure jointly affect parasitism and parasitoid diversity of Pseudaletia unipuncta (Haworth) (Lepidoptera: Noctuidae). Parasitism was assessed by experimentally adding P. unipuncta sentinel larvae at low (1 larvae/plant) and high (3 larvae/plant) densities to detect parasitism in commercial cornfields located in a complex and a simple agricultural landscape. The braconid wasps Glyptapanteles militaris (Walsh) and Meteorus spp. accounted for 98.4% of the observed parasitism. Landscape structure did not influence parasitism (80.2% on average) and contrary to expectations, showed a trend towards increased parasitoid richness and diversity in the simple landscape. Increasing host density revealed a trend of increasing parasitoid richness and diversity, and differentially affected parasitism at the parasitoid specific level. G. militaris parasitized a significantly greater proportion of hosts at low host density, while the opposite occurred for Meteorus spp. (primarily M. communis). These offsetting responses of parasitoids resulted in the lack of an overall host density effect on parasitism. The differential response of these parasitoids to host density is discussed in relation to differences in morphological and life history characteristics. Our results suggest that the specific composition of parasitoid assemblages could significantly alter parasitism at different host densities independently of landscape structural complexity.

Zusammenfassung

Verschiedene Untersuchungen haben gezeigt, dass Parasitismus entweder auf die Wirtsdichte oder die Landschaftskomplexität positiv reagiert. Dennoch haben keine Experimente die Wirtsdichte in Landschaften unterschiedlicher Komplexität manipuliert. Hier berichten wir von den Ergebnissen eines Freilandexperiments, das unternommen wurde, um zu bestimmen, wie die Wirtsdichte und die Struktur der Agrarlandschaft gemeinschaftlich die Parasitierung und die Parasitoidendiversität bei Pseudaletia unipuncta (Haworth) (Lepidoptera: Noctuidae) beeinflussen. Die Parasitierung wurde gemessen indem experimentell P. unipuncta Larven in geringen (1 Larve/Pflanze) und hohen (3 Larven/Pflanze) Dichten hinzugefügt wurden, um die Parasitierung in kommerziellen Kornfeldern zu erfassen, die in komplexen und einfachen Agrarlandschaften lagen. 98.4% der Parasitierung entfiel auf die braconiden Wespen Glyptapanteles militaris (Walsh) und Meteorus spp. Die Struktur der Landschaft beeinflusste die Parasitierung nicht (durchschnittlich 80.2%) und zeigte entgegen den Erwartungen einen Trend zu einer erhöhter Parasitoidenartenzahl und -diversität in der einfachen Landschaft. Eine zunehmende Wirtsdichte ließ einen Trend zu einer erhöhten Parasitoidenartenzahl und -diversität erkennen, und sie beeinflusste die Parasitierung auf dem Artenlevel der Parasitoide unterschiedlich. G. militaris parasitierte einen signifikant höheren Anteil der Wirte bei geringen Wirtsdichten, während für Meteorus spp. (vor allem M. communis) das Gegenteil zutraf. Diese sich ausgleichenden Reaktionen der Parasitoide führten zum Fehlen eines Gesamteffekts der Wirtsdichte auf die Parasitierung. Die unterschiedlichen Reaktionen dieser Parasitoide auf die Wirtsdichte werden in Zusammenhang mit Unterschieden in der Morphologie und Lebensweise diskutiert. Unsere Ergebnisse weisen darauf hin, dass die spezifische Zusammensetzung von Ansammlungen von Parasitoiden die Parasitierung bei unterschiedlichen Wirtsdichten unabhängig von der Komplexität der Landschaftsstruktur signifikant verändern könnte.