Manoeuvres of female Brachymeria intermedia flying towards host-related odours in a wind tunnel

Abstract. Mature female Brachymeria intermedia (Hymenoptera: Chalcididae) were conditioned to fly towards vanilla odour in a wind tunnel. We analysed the tracks of wasps flying along turbulent plumes of either host odour (pupae of the gypsy moth, Lymantria dispar) or vanilla odour, along either a ribbon plume or a turbulent plume of vanilla odour, and before and after plume removal. Wasps flew in similar shallow zigzagging tracks along the turbulent plume of host and vanilla odours. When the plume was removed while wasps were flying upwind along a turbulent plume of vanilla odour, wasps either maintained an upwind course or drifted sideways, alternating upwind and downwind courses before turning around and flying downwind. No wasp casted upon loss of the plume.     

Larval and adult nutrition effects on blood/nectar choice of Culex nigripalpus mosquitoes

Abstract. The impact of nutritional variables on the development of host-seeking and biting behaviours after emergence by female Culex nigripalpus mosquitoes were studied using air-flow olfactometer and close-range biting assays, respectively. Unfed females failed to develop resting stage ovarian follicles. When offered a bird host in the absence of competing stimuli, sugar-fed mosquitoes were significantly more responsive in both host-seeking and biting than unfed controls. In a choice olfactometer assay using nectar odours (honey scented with artificial apple-blossom oil) versus host odours (a bird), unfed females preferred honey over bird odours except when honey odour was weak. After sucrose feeding, females switched from honey to bird preference. This change in behaviour was accompanied by significant accumulation of lipid and by follicular growth to the resting stage. Elevation of host responsiveness after sugar feeding was reversible; starvation ultimately resulted in females preferring honey over bird odours. When the larval diet was restricted by crowding, the wing-length and total lipid of resultant adult females were reduced. Although differences were subtle, unfed bird-responding females tended to have longer wings and more lipid than their honey-responding counterparts.     

Could learning of pollen odours by honey bees (Apis mellifera) play a role in their foraging behaviour?

Abstract. The role of pollen odour cues in the foraging behaviour of honey bees (Apis mellifera L.) is poorly understood. Using classical conditioning of the proboscis extension response, in which bees learn to associate an odour with a sucrose reward, the present study tests whether odours of bee-collected pollen from the hive environment or odours of fresh pollen on the anthers of flowers could be used in pollen foraging. Honey bees efficiently learn odours from field-bean (Vicia faba) bee-collected pollen and oilseed-rape (Brassica napus) bee-collected pollen, hand-collected pollen, anthers and whole flowers, demonstrating that honey bees can learn pollen odours associatively in biologically realistic concentrations. Honey bees learn pollen odours of oilseed rape better than field bean and, although they generalize these two odours, they easily distinguish between them in discrimination tests, suggesting that pollen odours may be used in species recognition/discrimination. There is little evidence that honey bees can recognize whole flowers based on previous experience of bee-collected pollen odour. However, they generalize the odours of oilseed-rape anthers and whole flowers, suggesting that anther pollen in situ may play a more prominent role than bee-collected pollen in foraging behaviour.     

Carbohydrate bait preferences of wasps (Vespula vulgaris and V. germanica) (Hymenoptera: Vespidae) in New Zealand

Abstract

Carbohydrates are both a source of food and a potential bait for control of common and German wasps. Carbohydrate preferences of wasps were determined in a series of paired trials by counting the number of wasps caught in traps baited with carbohydrate‐based baits versus a standard protein‐based bait. Factors influencing wasp attraction to carbohydrate‐based baits included bait type, season, weather, and location. A 30% sucrose solution was more attractive than honey, jams, gels, or dry (solid) sugars. The sucrose solution was more attractive in spring and autumn than in summer, probably because rainfall washed away natural carbohydrate supplies. There was a significant relationship between rainfall in the previous week and the proportion of wasps caught in traps baited with sucrose solution versus sardine cat‐food. A higher proportion of German wasps than common wasps was caught in sucrose‐baited traps. This may have been related to seasonal differences in abundance and/or behavioural differences between the two species. In some locations, more wasps were caught in traps baited with sucrose solution than in those baited with sardine cat‐food. The proportion of wasps caught in sucrose‐baited traps was higher in locations without beech scale honeydew than with beech scale honey‐dew. Proportionally more non‐target species (mainly honey bees) and fewer wasps were caught in traps baited with sucrose solution than in those baited with sardine cat‐food. All five bee repellents tested also repelled wasps. These results indicate the importance of determining the bait preferences of wasps (carbohydrate or protein) and the presence of non‐target species before attempting a poison‐baiting operation. They also emphasise the need for a wasp‐specific carbohydrate‐based bait.     

Evaluating the Induced-Odour Emission of a Bt Maize and its Attractiveness to Parasitic Wasps

The current discussion on the safety of transgenic crops includes their effects on beneficial insects, such as parasitoids and predators of pest insects. One important plant trait to consider in this context is the emission of volatiles in response to herbivory. Natural enemies use the odours that result from these emissions as cues to locate their herbivorous prey and any significant change in these plant-provided signals may disrupt their search efficiency. There is a need for practical and reliable methods to evaluate transgenic crops for this and other important plant traits. Moreover, it is imperative that such evaluations are done in the context of variability for these traits among conventional genotypes of a crop. For maize and the induction of volatile emissions by caterpillar feeding this variability is known and realistic comparisons can therefore be made. Here we used a six-arm olfactometer that permits the simultaneous collection of volatiles emitted by multiple plants and testing of their attractiveness to insects. With this apparatus we measured the induced odour emissions of Bt maize (Bt11, N4640Bt) and its near-isogenic line (N4640) and the attractiveness of these odours to Cotesia marginiventris and Microplitis rufiventris, two important larval parasitoids of common lepidopteran pests. Both parasitoid species were strongly attracted to induced maize odour and neither wasp distinguished between the odours of the transgenic and the isogenic line. Also wasps that had previously experienced one of the odours during a successful oviposition divided their choices equally between the two odours. However, chemical analyses of collected odours revealed significant quantitative differences. The same 11 compounds dominated the blends of both genotypes, but the isogenic line released a larger amount of most of these. These differences may be due to altered resource allocation in the transgenic line, but it had no measurable effect on the wasps’ behaviour. All compounds identified here had been previously reported for maize and the differential quantities in which they were released fall well within the range of variability observed for other maize genotypes.     

Nutrition and division of labor: Effects on foraging and brain gene expression in the paper wasp Polistes metricus

Deeply conserved molecular mechanisms regulate food-searching behaviour in response to nutritional cues in a wide variety of vertebrates and invertebrates. Studies of the highly eusocial honey bee have shown that nutritional physiology and some conserved nutrient signalling pathways, especially the insulin pathway, also regulate the division of labour between foraging and non-foraging individuals. Typically, lean workers leave the nest to forage for food, and well-nourished workers perform tasks inside the nest. Here we provide the first direct test of whether similar mechanisms operate in a primitively eusocial insect in an independently evolved social lineage, the paper wasp Polistes metricus. We found that food deprivation caused reduced lipid stores and higher levels of colony and individual foraging. Individuals with greatly reduced lipid stores foraged at extremely elevated levels. In addition, brain expression of several foraging-related genes was influenced by food deprivation, including insulin-like peptide 2 (ilp2). Together with previous findings, our results demonstrate that nutrition regulates foraging division of labour in two independently evolved social insect lineages (bees and wasps), despite large differences in social organization. Our results also provide additional support for the idea that nutritional asymmetries among individuals, based on differences in nutritional physiology and expression of conserved nutrient signalling genes in the brain, are important in the division of labour in eusocial societies.     

Influence of different sugars on the longevity of Bathyplectes curculionis (Hym., Ichneumonidae)

Abstract:  Floral nectars, homopteran honeydews and honey are known to increase parasitoid longevity. However, these foods are composed of several sugars which may differentially affect longevity. We tested the effects of individual sugars and mixtures on the longevity of Bathyplectes curculionis Thomson (Hym., Ichenumonidae), a natural enemy of the alfalfa weevil. There was a significant difference in the longevity of female wasps on the various diets. Glucose or fructose alone appeared to have the most benefits while trehalose and melezitose were not as useful for increasing wasp longevity. Suitability of these sugars for provisioning food for parasitoids in the field is discussed.     

Effect of different dietary resources on longevity, carbohydrate metabolism, and ovarian dynamics in two fruit fly parasitoids

A growing body of literature reports the importance of non-prey food sources in boosting fitness of arthropod natural enemies, thus further contributing to their pest control efficacy. Although resources such as nectar, pollen, or honeydew have received a fair amount of research attention, little is known about the role of fruit juices. Under natural conditions, Tephritid fruit fly parasitoids enjoy ample access to fallen or damaged fruits and their saccharide-rich juices, and wasp fitness can potentially benefit in multiple ways from access to these resources. In this study, we compared the effect of fruit juice with other food resources on multiple fitness parameters in parasitoids that commonly forage on fallen, damaged fruits: the braconid Diachasmimorpha longicaudata and figitid Aganaspis pelleranoi. Parasitoids were subject to simple or combined diets of guava juice (Psidium guajava), honey and pollen, and their effect on wasp longevity, ovarian dynamics and (body) carbohydrate levels was assessed. For both species, adult longevity proved lowest on simple diets of water, guava juice, or pollen, while greatest longevity was attained on honey or combined diets. For D. longicaudata, egg load did not differ between the various diets, while A. pelleranoi egg load was higher for individuals that had access to honey or pollen, but did not differ between newly emerged wasps and those fed guava juice. In both parasitoid species, total sugars, fructose, and glycogen levels were highest in wasps fed with honey or combined diets and lowest under (simple) guava juice, pollen, or water diets. In conclusion, D. longicaudata and A. pelleranoi attained superior longevity and body nutrient levels with access to high-sucrose sugar sources, such as honey, but benefited comparatively little from access to guava juice. Our work hints the role of high-sucrose foods such as (extra-) floral nectar or artifical sugar sprays in boosting fitness of fruit fly parasitoids. We further discuss the relevance of these findings for fruit fly biological control, in crops such as guava.     

The importance of location and visual cues during foraging in the German wasp (Vespula germanica F.) (Hymenoptera: Vespidae)

Abstract

The way in which foraging wasps use cues for prey location and choice appears to depend on both the context and on the type of prey. Vespula germanica is an opportunistic, generalist prey forager, and individual wasp foragers often return to hunt at sites of previous hunting success. In this paper, we studied which cues are used by this wasp when relocating a food source. Particularly we analysed the response to a displaced visual cue versus a foraging location at which either honey or cat food had been previously presented. We conclude that location is used over a displaced visual cue for directing wasp hovering, although the landing response is directed differently according to bait type. When wasps are exploiting cat food, location also elicits landing, but if they are exploiting honey, a displaced visual cue elicits landing more frequently than location.     

Development, oviposition, and mortality of Neoseiulus fallacis (Acari: Phytoseiidae) in response to reduced-risk insecticides

Eight reduced-risk insecticides (acetamiprid, thiamethoxam, imidacloprid, thiacloprid, methoxyfenozide, pyriproxyfen, indoxacarb, and spinosad) and three conventional insecticides (azinphosmethyl, fenpropathrin, and esfenvalerate) were tested against Neoseiulus fallacis (Garman) (Acari: Phytoseiidae), the most abundant predacious mite in North Carolina apple (Malus spp.) orchards. To assess the effect of insecticides on development and mortality of N. fallacis immatures, 12-h-old eggs were individually placed on bean leaf disks previously dipped in insecticide solutions. Tetranychus urticae Koch (Acari: Tetranychidae) females were added as a food source. None of the reduced-risk insecticides significantly affected immature N. fallacis compared with the control; however, the pyrethroids esfenvalerate and fenpropathrin were highly toxic to immatures. To evaluate the effect of insecticides on mortality and oviposition of adult N. fallacis, 7- to 8-d-old females were confined on insecticide-treated bean leaves with Malephora crocea (Aizoaceae) pollen added as a food source. Spinosad resulted in the highest mortality, whereas azinphosmethyl, acetamiprid, fenpropathrin, and imidacloprid were moderately toxic, and mortality from esfenvalerate, indoxacarb, thiacloprid, methoxyfenozide, pyriproxyfen, and thiamethoxam did not differ significantly from the control. Oviposition was affected in a similar manner, with the exception of acetamiprid that did not affect oviposition, and thiamethoxam that reduced oviposition.     

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.     

Parasitic wasps avoid ant-protected hemipteran hosts via the detection of ant cuticular hydrocarbons

One of the most studied and best-known mutualistic relationships between insects is that between ants and phloem-feeding insects. Ants feed on honeydew excreted by phloem-feeding insects and, in exchange, attack the phloem feeders'' natural enemies, including parasitic wasps. However, parasitic wasps are under selection to exploit information on hazards and avoid them. Here, we tested whether parasitic wasps detect the previous presence of ants attending colonies of phloem feeders. Behavioural assays demonstrate that wasps left colonies previously attended by ants more frequently than control colonies. This behaviour has a potential cost for the parasitic wasp as females inserted their ovipositor in fewer hosts per colony. In a further bioassay, wasps spent less time on papers impregnated with extracts of the ant cues than on control papers. Gas chromatography coupled with mass spectrometry analyses demonstrated that ants left a blend of cuticular hydrocarbons when they attended colonies of phloem feeders. These cuticular hydrocarbons are deposited passively when ants search for food. Overall, these results suggest, for the first time, that parasitic wasps of honeydew producers detect the previous presence of mutualistic ants through contact infochemicals. We anticipate such interactions to be widespread and to have implications in numerous ecosystems, as phloem feeders are usually tended by ants.     

A novel interference behaviour: invasive wasps remove ants from resources and drop them from a height

This study reports a novel form of interference behaviour between the invasive wasp Vespula vulgaris and the New Zealand native ant Prolasius advenus. By videotaping interactions at bait stations, we found that wasps commonly remove ant competitors from food resources by picking up the workers in their mandibles, flying backward and dropping them unharmed some distance from the food. Both the frequency and the efficiency of the wasp behaviour significantly increased with the abundance of ant competitors. Ant removals were the most common interference events initiated by wasps when ants were numerous, while intraspecific conflicts among wasps were prominent when few ants were present. The 'ant-dropping' behaviour emphasizes how asymmetry in body sizes between competitors can lead to a pronounced form of interference, related to asymmetric locomotion modes.     

Effect of rotational use of insecticides on pyrethroids resistance in Helicoverpa armigera (Lep.: Noctuidae)

Abstract:  To investigate fluctuation in susceptibility to insecticides in the field, natural populations of Helicoverpa armigera were collected from the same field in the region of Multan, Pakistan in 2002 and 2003. The populations were examined against pyrethroids (viz. cypermethrin, esfenvalerate and fenpropathrin) and new insecticides (viz. spinosad, abamectin and indoxacarb). In 2002, the resistance ratio (RR) of cypermethrin and esfenvalerate was significantly higher than fenpropahrin compared with susceptible population. The susceptibility to cypermethrin, esfenvalerate and fenpropathrin increased significantly in 2003; however, the RR for cypermethrin was about half the RR (38) of esfenvalerate (101) and fenpropathrin (89). The toxicity of spinosad, abamectin and indoxacarb was identical in both years. In the field experiments, abamectin was more effective than other compounds tested whereas fenpropathrin, cypermethrin and esfenvalerate had similar toxicity. These results might have important implications in resistance management and suggest that the rotational use of spinosad, abamectin and indoxacarb could help to avoid the development of multiple resistant in H. armigera .     

Host influences on sex ratio, longevity, and egg load of two Metaphycus species parasitic on soft scales: implications for insectary rearing

Metaphycus flavus (Howard) and M. stanleyi Compere (Hymenoptera: Encyrtidae) are currently being screened for use as augmentative biological control agents of citrus-infesting soft scales (Homoptera: Coccidae). Two factors were investigated, host quality-dependent sex allocation and local mate competition, which likely influence these parasitoid's sex allocation strategies and are therefore of interest for their mass-rearing. The results of these studies suggested that, under the mass-rearing protocol that is envisioned for these parasitoids, offspring sex ratios in both M. flavus and M. stanleyi are dominated by host quality (= size) influences, but not by interactions with other females. These results indicated that host size strongly influences offspring sex ratios and brood sizes; larger hosts led to more female offspring and larger broods. In contrast, increasing the number of parental females did not lead to fewer female offspring as expected under local mate competition. Additionally, within-brood sex ratios did not vary with brood size; this result is inconsistent with expected sex ratios due to local mate competition. Other results also indicated that host quality was a dominant influence on M. flavus' and M. stanleyi's sex ratios. Larger hosts led to a larger size in the emerging wasps, and larger wasps had greater egg loads and lived longer than smaller wasps. However, wasp longevity, and the influence of wasp size on longevity were mediated by a wasp's diet. Metaphycus flavus females lived the longest when they had access to hosts, honey, and water, followed by honey and water, and shortest when they had access to water alone; M. stanleyi females lived longest with honey and water, followed by hosts, honey, and water, and shortest with water alone. Greater wasp size led to greater longevity in females only when they had access to food (honey, or hosts and honey). Finally, other results suggested that both M. flavus and M. stanleyi are facultatively gregarious. Wasp size did not decrease with brood size as expected under superparasitism. Overall, the results of these studies suggested that holding newly emerged females of both M. flavus and M. stanleyi for several days in the presence of an appropriate food source before field release could enhance a female's performance as an augmentative biological control agent. It increases their initial life expectancy following release, and maximizes the females' egg load (both Metaphycus species) and resources for replacing oviposited eggs (M. flavus only).     

Competition between Honey-Bees (

Honeydew, the sugary exudate of the scale insect Ultracoelostoma brittini, is an important food source in black beech (Nothofagus solandri var. solandri) forests in the South Island of New Zealand. Two of the most prominent foragers of honeydew are honey bees (Apis mellifera) and wasps (Vespula germanica and V. vulgaris). Observations in the field and using a captive bee hive were used to investigate competition between bees and wasps feeding on honeydew. In laboratory trials, interference competition was often strong, and many cases of aggression were noted. In the forest, there was invariably enough room on the trees for bees and wasps to feed while rarely encountering one another. Over the whole year, environmental variables (especially low temperatures and rain), were found to constrain honey bee foraging to a greater degree than competition with wasps. Because the competition that did occur was primarily exploitation competition, reciprocal effects were likely to be felt. At Coopers Creek, bees may be reducing wasp densities, compared with the situation in Nelson—Marlborough where commercial hives are scarce. It may be possible to reduce wasp densities locally by increasing the number of bee hives in an area.     

Olfactory discrimination in the western lowland gorilla, Gorilla gorilla gorilla

The olfactory abilities of great apes have been subject to little empirical investigation, save for a few observational reports. This study, using an habituation/dishabituation task, provides experimental evidence for a core olfactory ability, namely, olfactory discrimination, in the gorilla. In Experiment 1, six zoo-housed western lowland gorillas were individually presented with the same odour on four trials, and with a novel odour on the fifth trial. Odours (almond and vanilla) were presented on plastic balls, and behavioural responses of sniffing and chewing/licking the balls were recorded. A second experiment presented the same odour on four trials and no odour on the fifth to examine whether any dishabituation was due to the presence of a new odour or the absence of the familiar odour. Gorillas habituated their behaviour with repeated presentation of the same odour, but dishabituated, i.e. increased sniffing and chewing/licking, when presented with the novel odour. No dishabituation was noted when using water as the stimulus across all trials or when used as the novel odour. Overall, results show that gorillas are able to discriminate between odours.     

A six-arm olfactometer permitting simultaneous observation of insect attraction and odour trapping

Abstract.  Behavioural assays to study insect attraction to specific odours are tedious, time consuming and often require large numbers of replications. Olfactometer and flight tunnel tests can usually only be conducted with one or two odour sources at a time. Moreover, chemical information on the odour sources has to be obtained in separate analytical studies. An olfactometer was developed in which six odours can be tested simultaneously for their relative attractiveness while during the assays, part of each test odour can be trapped for further analyses. The effectiveness of this six-arm olfactometer was tested by observing the responses of the solitary endoparasitoid Cotesia marginiventris (Cresson) to host-induced odours from young maize plants. For statistical analyses, we used log-linear models were adapted to account for overdispersion and possible positional biases. Female wasps responded extremely well in tests where they were offered a single odour source, as well as in tests with multiple choices. The responses of wasps released in groups were the same as those released individually and it was found that females did not attract or repel each other, but males preferred arms in which females had been released. Dose–response tests with varying numbers of plants or host larvae on plants revealed that the wasps responded in a dose-related manner, thus showing that the system is well suited to measure relative preference. The clear choices of the insects amongst six possibilities provided substantial statistical power. Gas chromatographic analyses of sampled air revealed clean and effective odour trapping, which largely facilitates the comparison of results from behavioural assays with the actual blends of volatiles that were emitted by the various odour sources. Advantages and disadvantages compared to other methods are discussed.     

The effect of host size on quality attributes of the egg parasitoid,Trichogramma pretiosum

In a study of the quality ofTrichogramma pretiosum Riley (Hymenoptera: Trichogrammidae), we compared female wasps emerging from natural hosts, parasitized in the laboratory or the field with those emerging from factitious hosts used for commercial mass production. Females from the natural hosts were larger, more fecund, and longer lived than those from the factitious hosts. Compared to small females, large female wasps are substantially more fecund when honey (carbohydrate) is available but marginally more fecund when honey is unavailable. The size of a femaleT. pretiosum depends on two factors: the size of the host egg from which it emerges even when the wasp was gregarious, and the number of conspecifics that emerge with it. The similarities in the size distribution of female wasps emerging from natural hosts, in conjunction with the mechanism by whichTrichogramma measure host size and allocate eggs accordingly, suggests the hypothesis that size related components of fitness in femaleT. pretiosum are under strong selection in the field.