Do Larger Fruits Provide a Partial Refuge for Rose-Hip Flies Against Parasitoids?

The preference of insect herbivores to oviposit into larger structures might be explained by the superior food resource a larger structure offers to herbivore offspring or as an adaptive strategy by the herbivore to protect its offspring against parasitoids, should larger structures provide a partial refuge against parasitoids. We investigated whether larger fruits provide a partial refuge for rose-hip fly (Rhagoletis basiola) offspring against the specialist parasitoid Halticoptera rosae. We tested whether female parasitoids exhibit a preference to land on larger rose-hips into which females of the host fly prefer to oviposit and whether parasitoids are less successful in locating host eggs on large, compared to small, fruits. Female parasitoids showed no clear preference for larger fruits. When searching fruits, they were about equally successful in locating fly eggs on large and small fruits, with a slight, though nonsignificant trend for fly offspring in large fruits to be located more easily. Furthermore, wasps did not require more time to locate host eggs on large fruits than on small fruits, suggesting a nonrandom searching behavior. Consequently, large rose-hips do not provide a partial refuge to fly offspring and the preference of flies to oviposit into larger fruits cannot be explained as an adaptation against parasitoid searching efficiency.     

Exploitation of the Host's Chemical Communication in a Parasitoid Searching for Concealed Host Larvae

We found empirical evidence for the exploitation of the host's chemical communication in a parasitic wasp foraging for a concealed host. Female Halticoptera laevigata wasps (Hym., Pteromalidae) searched longer and probed more frequently on honeysuckle fruit that carried fresh marking pheromone or harbored a first instar larva of its host, the tephritid fruit fly, Myoleja lucida . They further increased their searching efforts on fruits that provided both host-related cues. While the exploitation of host marking pheromones for host location had previously been shown in two parasitoid species attacking the eggs of their tephritid hosts, this is the first evidence for the exploitation of a host's marking pheromone in a parasitoid attacking the larvae of an herbivorous host. Taking into account the time interval between application of host marking pheromones and parasitoid attack plus the fact that these pheromones are generally water soluble and thus might be washed off by rains, we discuss the reliability-detectability problem for the exploitation of those cues for host location in parasitoids attacking a larval host.     

Evidence for a marking pheromone in host discrimination byCephalonomia Stephanoderis (Hym.: Bethylidae)

The bethylidCephalonomia stephanoderis Betrem is an ectoparasitoid that prefers to oviposit on the prepupae and pupae of the coffe berry borerHypothenemus hampei (Ferrari) (Coleoptera: Scolytidae). It has the ability to distinguish unparasitized from parasitized hosts and rarely lays more than one egg per host. The mechanism of this host discrimination byC. stephanoderis was investigated under laboratory conditions. For this, parasitoid eggs that had been deposited on host pupae were removed and pupae were then offered (individually and collectively) to individual female wasps. A total of 92% of individually offered hosts and 93% of collectively offered hosts were not parasitized. It is concluded thatC. stephanoderis recognizes a marking pheromone deposited into or onto the host, preceding, during, or after oviposition which enables female parasitoids to avoid self and conspecific superparasitism.     

Discrimination against previously searched, host-free patches by a parasitoid foraging for concealed hosts

1. The foraging behaviour of the parasitoid wasp Halticoptera laevigata Thomson (Hymenoptera: Pteromalidae) was studied. This wasp attacks the first‐instar larvae of the fruit fly Myoleja lucida Fallén (Diptera: Tephritidae) within honeysuckle Lonicera xylosteum L. (Caprifoliaceae) fruits. It was hypothesised that, to forage efficiently, wasps should be able to maintain a systematic or at least non‐random search pattern on the level of the microhabitat, i.e. branches with fruits, and/or be able to recognise previously searched fruits. 2. Comparisons of observed patch visitation patterns on branches bearing uninfested fruits with expected values for systematic and random visitation patterns indicated that H. laevigata wasps visit patches randomly and do not avoid visiting host‐free patches that they have visited previously. 3. While wasps did not alter their search effort on uninfested fruits searched previously by conspecifics, they searched for much shorter times and probed less frequently for hosts on uninfested fruits that they had searched previously. 4. The results indicate that H. laevigata wasps engage in partially systematic search and increase their search efficiency through the recognition of self‐searched fruits. The reasons why partial rather than fully systematic search should be adaptive in this parasitoid are discussed.     

Resource utilization and sex allocation in response to host size in two ectoparasitoid wasps on subcortical beetles

Patterns of host resource utilization and sex ratio manipulation in relation to host size were investigated for two solitary ectoparasitoid wasps,Atanycolus initiator andSpathius brevicaudis (Hymenoptera Braconidae). Both species parasitize subcortical beetles on the trunks of Japanese pine trees.A. initiator is on average 8 times larger in body weight and has an ovipositor that is 3.7 times longer than that ofS. brevicaudis. In both parasitoids, the size of emerging wasps was positively correlated with host size, but the host/wasp size regressions were linear for all three major host species inA. initiator, whereas inS. brevicaudis the regression was logarithmic for a relatively large host species. The sex ratios (proportion of males) of both parasitoids emerging from different host species decreased with increasing host size, but the overall sex ratio on each host species was male-biased inA. initiator, while female-biased inS. brevicaudis. How the proportion of host consumed changed in response to host size, differed between the two parasitoids for the same host species. In the field survey, the size and sex ratio of the emerging two parasitoids from a dead tree were closely related to host size. However, the spatial distribution of the two parasitoids depended on the bark thickness of the trunk. The data suggest that differences in the relative evaluation of host size and in ovipositor length may enable the coexistence of the two parasitoid wasps.     

Influence of initial egg density and host size on the development of the gregarious parasitoid <Emphasis Type="Italic">Bracon hebetor</Emphasis> on three different host species

Bracon hebetor (Say) (Hymenoptera : Braconidae) is a gregarious parasitoid that attacks a variety of important lepidopterous pests of stored product and in the field. In this study the effect of host species, size and larval competition on parasitoid size, survival and development were investigated. In laboratory studies, wasp eggs at a range of densities, were placed on larvae of different weight of three Lepidoptera host species namely Adoxophyes orana (Fischer von Röslerstamm, Tortricidae), Plodia interpunctella (Hubner, Pyralidae) and, Lobesia botrana(Dennis & Schiffermueller, Tortricidae). On A. orana survival of immature parasitoids was very low at all densities and different host weights. On L. botrana survival progressively reduced as egg density increased at both host weights examined for this host. Survival on P. interpunctella was significantly affected by egg density but not by host weight. Initial egg density had a significant effect on the size of emerging adults from each rearing host. Smaller adult parasitoids emerged as egg density per larva increased. Larval host weight of P. interpunctella and A. orana had a significant effect on the size of emerging adult parasitoids mainly at the higher egg densities used in these experiments. The above results of host quality on fitness of parasitoid are discussed.     

Differential energy allocation as an adaptation to different habitats in the parasitic wasp <Emphasis Type="Italic">Venturia canescens</Emphasis>

Environmental pressures are expected to favour organisms that optimally allocate metabolic resources to reproduction and survival. We studied the resource allocation strategies and the associated tradeoffs in the parasitoid wasp Venturia canescens, and their adaptation to the characteristics of the environment. In this species, individuals of two reproductive modes coexist in the same geographical locations, but they mainly occur in distinct habitats. Thelytokous (asexual) wasps are mostly found in anthropogenic habitats, where hosts tend to aggregate and food is absent. Arrhenotokous (sexual) wasps are exclusively found in natural habitats, where hosts are scattered and food is present. We analysed (1) the quantity of energy stored during ontogeny, (2) the tradeoff between reproduction and survival, by measuring egg load and longevity and (3) the host patch exploitation behaviour of the wasps at emergence. Arrhenotokous wasps emerged with more metabolic resources than thelytokous ones, especially glycogen, a nutrient that could be used for flying in search of hosts and/or food. Thelytokous wasps allocated more energy than arrhenotokous wasps to egg production: this would allow them to parasitize more hosts. The tradeoff between egg production and longevity was not revealed within reproductive modes, but when comparing them. At emergence, arrhenotokous wasps tended to exploit host patches less thoroughly than thelytokous wasps, suggesting that by leaving the host patch, they search for food. The results clearly showed adaptations to the characteristics of habitats preferentially inhabited by the two reproductive modes, and suggested a mechanism that facilitates their coexistence in natural conditions.     

Influence of Host Pheromone on Egg Parasitism by Scelionid Wasps: Comparison of Phoretic and Nonphoretic Parasitoids

The kairomonal activity of the attractant pheromone for the spined soldier bug, Podisus maculiventris (Hemiptera: Pentatomidae), was investigated by exposing fresh pentatomid egg masses in field traps with or without synthetic pheromone. Predominantly two parasitoids were recovered from exposed eggs of P. maculiventris and Euschistus obscurus: Telenomus podisi Ashmead (a generalist pentatomid egg parasitoid) and Telenomus calvus Johnson (a phoretic specialist on Podisus eggs) (Hymenoptera: Scelionidae). The incidences of T. podisi from P. maculiventris and E. obscurus eggs placed in pheromone-baited and nonbaited traps were not significantly different, suggesting that this oophagous wasp does not use the spined soldier bug attractant pheromone as a kairomone. However, T. calvus was reared almost exclusively from egg masses of P. maculiventris placed inside pheromone-baited traps. These results suggest that T. calvus females orient to volatile chemicals emitted by spined soldier bug males as a searching strategy to find areas likely to contain host eggs, in addition to the previously discovered strategy of using the pheromone to guide their phoretic behavior. The data also indicate that T. calvus can distinguish between the egg masses of these pentatomid hosts at close range.     

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.     

Host searching behavior and potential of an aquatic ichneumonid pupal parasitoid of rice caseworm (Parapoynx stagnalis) in an upland rice paddy agro-ecosystem of the Western Ghats,India

Parapoynx stagnalis (Zeller) (=Nymphula depunctalis (Guenée), is a sporadic rice paddy pest of south and southeast Asian rice paddy fields, and widely known as rice caseworm. A recent outbreak of the pest was observed in the upland rice paddy agro-ecosystem of the central Western Ghats, India. No potential natural enemies of this pest have so far been reported, mainly due to the semi-aquatic behavior of the larvae and pupae. However, an undescribed aquatic ichneumonid wasp species (belonging to a genus near Litochila) is reported as a potentially useful early stage pupal parasitoid of this pest in our study. The host searching behavior and potential of the parasitoid wasp was studied in detail using both field and laboratory experiments. The adult female wasps usually dive into the water in search of the host pupae, remaining under water for a maximum of 90.2s, while searching for rice caseworm pupae. In field and laboratory experiments, we found 73% of the parasitized pupae of P. stagnalis kept under water yielded adult parasitic wasps. However, no parasitoids emerged from parasitized host pupae maintained in dry terrariums. Sex ratio was 2.8:1. The present discovery of the semi-aquatic parasitoid wasp could aid in better management of the rice caseworm population in rice paddy fields.     

Differential response of Trichogramma wasps to extreme sex pheromone types of the noctuid moth Heliothis virescens

1. Chemical espionage in nature may occur when predators or parasitoids home in on animal or plant communication signals. Parasitoid wasps are known to use pheromones emitted by adults hosts to locate host eggs, larvae or pupae. The response of Trichogramma egg parasitoids to a synthetic sex pheromone blend of moths has been shown in a number of studies over the past 40 years. 2. Trichogramma pretiosum (Hymenoptera, Trichogrammatidae) is a tiny parasitic wasp, attacking the eggs of the noctuid moth Heliothis virescens (Lepidoptera, Noctuidae). This study investigated whether T. pretiosum homes in on the sex pheromone of H. virescens at close range. The arrestment response of the wasps to sex pheromone gland extracts of two types of female moths, so‐called high and low females, was also tested, referring to two selected extreme pheromone types of H. virescens. The study also investigated whether the wasps would mount females, possibly to hitchhike with them. 3. The wasps were arrested by the common, ‘low’ pheromone, but not by the rare, ‘high’ pheromone or by extracts from male hairpencils. The wasps did not show a preference for separate sex pheromone compounds, but when pre‐exposed to the major sex pheromone component of H. virescens before the tests together with H. virescens eggs, they did show a preference, indicating learning behaviour. In the mounting experiments, mated females were mounted significantly more than virgin females or males, suggesting that hitchhiking is a strategy used by these wasps to locate moth eggs. 4. This represents the first study to show a differential response of parasitoid wasps to two different sex pheromone types in a single host species. The results warrant further investigations into the potential role of parasitic wasps in the evolution of sexual communication in moths.     

Evaluating the impact of a biological control parasitoid on invasive Vespula wasps in a natural forest ecosystem

The overall impact of the parasitoid Sphecophaga vesparum vesparum on invasive Vespula wasps in New Zealand native beech forest was evaluated by assessing the levels of parasitism achieved and the parasitoid’s effect at nest level and population level. The maximum proportion of nests parasitised was 17%, but there was no significant increase with time (r² = 0.139; p = 0.115). However, there was an exponential reduction in the number of parasitoids produced per parasitised nest from a peak of 570 (SE = 143) parasitoids per nest in 1990, declining to only 15 (SE = 6) parasitoids per nest in 2004. Even when parasitoid density was high, the parasitoid had no detectable impact on the number of small cells or the total host nest size, but it halved the number of large (reproductive) cells produced. This may have resulted in fewer queens produced per parasitised nest. Wasp nest density was highly variable from year to year, but there was no evidence that the wasp population density at the parasitised site (Pelorus Bridge) had declined relative to the five sites where the parasitoid had not established. We conclude that the parasitoid is unlikely to have had any significant effect on wasp populations hitherto, nor is it likely to impact host populations in the future. We recommend other biological control programs adopt pre-release assessment of per capita impact as a way of identifying agents that are more likely to be successful and hence minimising economic and potential ecological costs of biocontrol.     

Host-searching behavior ofVenturia canescens (Grav.) (Hymenoptera: Ichneumonidae): Interference—the effect of mature egg load and prior behavior

The foraging behavior ofVenturia canescens, a solitary endoparasitoid of lepidopteran larvae, was investigated in the laboratory. Females with a greater number of mature eggs in their ovarioles and oviducts parasitized a greater number of hosts and won a greater proportion of encounters with other searching females. Wasps which had been exposed to hosts prior to an experimental trial lost a higher proportion ofagonistic encounters with conspecifics than wasps which had no prior exposure to hosts. The behavior of a wasp at the time of the encounter influenced the outcome of the encounter. Wasps involved in active search of the host medium with the ovipositor (probing) were more likely to win encounters than wasps in any other behavioral category. In a situation where the agonistic encounter was between two probing wasps, both contestants were equally likely to win. Results are discussed in the light of the idea that mutual interference arises, in this species, as a result of agonistic encounters between searching females and recent dynamic-programming models which suggest that parasitoid oviposition should be influenced by mature egg load.     

Transfer of a chromosomal <Emphasis Type="Italic">Maverick</Emphasis> to endogenous bracovirus in a parasitoid wasp

Bracoviruses are used by parasitoid wasps to allow development of their progeny within the body of lepidopteran hosts. In parasitoid wasps, the bracovirus exists as a provirus, integrated in a wasp chromosome. Viral replication occurs in wasp ovaries and leads to formation of particles containing dsDNA circles (segments) that are injected into the host body during wasp oviposition. We identified a large DNA transposon Maverick in a parasitoid wasp bracovirus. Closely related elements are present in parasitoid wasp genomes indicating that the element in CcBV corresponds to the insertion of an endogenous wasp Maverick in CcBV provirus. The presence of the Maverick in a bracovirus genome suggests the possibility of transposon transfers from parasitoids to lepidoptera via bracoviruses.     

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.     

Does superparasitism improve host suitability for parasitoid development? A case study in the Microplitis rufiventris-Spodoptera littoralis system

Superparasitism refers to the oviposition behavior of parasitoid females who lay their eggs in an already parasitized host. Recent studies have shown that allocation of additional eggs to an already parasitized host may be beneficial under certain conditions. In the present work, mortality of Microplitis rufiventris wasps was significantly influenced by both host instar of Spodoptera littoralis larvae at parasitism and level of parasitism. In single parasitization, all host instars (first through sixth) were not equally suitable. Percentage of emergence success of wasp larvae was very high in parasitized first through third (highly suitable hosts), fell to 60% in the fourth instar (moderate suitable) and sharply decreased in the penultimate (5th) instars (marginally suitable). Singly parasitized sixth (last) instar hosts produced no wasp larvae (entirely unsuitable), pupated and eclosed to apparently normal adult moths. The scenario was different under superparasitism, whereas supernumerary individuals in the highly suitable hosts were almost always killed as first instars, superparasitization in unsuitable hosts (4th through 6th) had significant increase in number of emergence success of wasp larvae. Also, significantly greater number of parasitoid larvae successfully developed in unsuitable hosts containing three wasp eggs than counterparts containing two wasp eggs. Moreover, the development of surplus wasp larvae was siblicidal in earlier instars and nonsiblicidal gregarious one in the penultimate and last “sixth” instars. It is suggested that the optimal way for M. rufiventris to deal with high quality hosts (early instars) is to lay a single egg, while the optimal way to deal with low quality hosts (late instars) might be to superparasitize these hosts.     

Response of the fruit fly parasitoid Diachasmimorpha longicaudata to host and host‐habitat volatile cues

Chemical information is crucial to insect parasitoids for successful host location. Here, we evaluated the innate response of Diachasmimorpha longicaudata (Ashmead) (Hymenoptera: Braconidae), a fruit fly larval parasitoid, to cues from host and host habitat (i.e., fruit infested with host larvae). We first assessed the preference of female parasitoids between oranges infested with Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) and non‐infested fruit. Females were highly attracted towards infested oranges on the basis of volatile chemical cues. After this initial experiment, we aimed at revealing the potential sources of volatile cues present in an infested fruit. To this end, we considered five potential sources: (1) punctured fruit; (2) fly feeding, frass, or host‐marking pheromone deposited on the orange surface; (3) larval activity inside the fruit; (4) the larvae themselves; and (5) fungi associated with infestation of oranges. Habitat cues associated with host activity and those produced by rotten oranges or oranges colonized by fungi were highly attractive for female wasps, whereas odours associated with the activity of the adults on the surface of the fruit, and those released by the fruit after being damaged (as happens during fruit fly egg‐laying) were not used as cues by female parasitoids. Once the female had landed on the fruit, direct cues associated with larval activity became important although some indirect signals (e.g., products derived from larval activity inside the fruit) also increased host searching activity. Our findings indicate that naïve D. longicaudata uses chemical cues during host habitat searching and that these cues are produced both by the habitat and by the host larvae.     

Vine mealybug sex pheromone increases citrus mealybug parasitism by Anagyrus sp. near pseudococci (Girault)

The present study was aimed at elucidating the role of lavandulyl senecioate (LS), the sex pheromone of Planococcus ficus, in host selection of the parasitoid Anagyrus sp. near pseudococci. Field trials were carried out in Portugal, Italy and Israel. The effect of LS on the parasitism rate of the wasp was determined by exposing sentinel mealybugs combined with pheromone dispensers impregnated with LS, in comparison with other baits: lavandulyl isovalerate (LI); planococcyl acetate (PAc); and unbaited control traps. In addition, in order to study the host location behavior of A. sp. near pseudococci, pheromone dispensers were placed at three different distances: inside the trap, 30 or 60 cm away from the trap. The number of parasitoid females inside the traps, the number of parasitized mealybugs, and the number of days required for the first parasitoid emergence were recorded. The response of A. sp. near pseudococci females to different doses of LS (25–1350 μg) was also evaluated using sticky plate traps. The rate of mealybug parasitism by A. sp. near pseudococci was significantly increased by LS in the three parasitoid populations. PAc and LI had no significant effect on the wasp parasitism rate in most of the trials. However, the Italian population of the parasitoid responded to PAc, showing apparently a different behavioral pattern. The number of parasitoid females trapped did not significantly differ between tested doses of LS. The use of LS as an arrestant in host location by the A. sp. near pseudococci female is discussed.     

Gene Flow in the Exotic Colonizing LadybeetleHarmonia axyridisin North America

The introduced parasitoid,Aphytis melinusDeBach (Hymenoptera: Aphelinidae), is used for augmentative biological control of California red scale,Aonidiella aurantii(Maskell) (Homoptera: Diaspididae). Commercially reared wasps are reared on oleander scale,Aspidiotus neriiBouché (Homoptera: Diaspididae). Oleander scale covers lack the chemical,O-caffeoyltyrosine, a kairomone mediating host selection byA. melinus.Wasps reared on oleander scale but individually exposed, or primed, toO-caffeoyltyrosine more readily accepted California red scale for probing in laboratory bioassays and parasitized a greater proportion of available California red scale in the field than wasps reared similarly but not exposed toO-caffeoyltyrosine. Thus, it may be possible to improve host recognition of commercial, insectary-rearedA. melinusby exposing them toO-caffeoyltyrosine prior to release. The goal of this study was to develop an inexpensive but effective means of priming thousands of wasps simultaneously toO-caffeoyltyrosine. The most effective method, but potentially the most expensive, was simply to spray parasitized oleander scale on their host plant with diluteO-caffeoyltyrosine prior to wasp emergence. In additional experiments, using controlled doses of syntheticO-caffeoyltyrosine applied to scale covers, we showed that primed wasps require both a lower minimum dose ofO-caffeoyltyrosine for recognition and also respond to measuredO-caffeoyltyrosine doses more consistently than unprimed wasps. The ability to mass-prime thousands of wasps prior to release is a crucial step toward realizing the concept of behavioral improvement of host selection of parasitoids on a commercial scale.     

Host Chemical Footprints Induce Host Sex Discrimination Ability in Egg Parasitoids

Trissolcus egg parasitoids, when perceiving the chemical footprints left on a substrate by pentatomid host bugs, adopt a motivated searching behaviour characterized by longer searching time on patches were signals are present. Once in contact with host chemical footprints, Trissolcus wasps search longer on traces left by associated hosts rather than non-associated species, and, in the former case, they search longer on traces left by females than males. Based on these evidences, we hypothesized that only associated hosts induce the ability to discriminate host sex in wasps. To test this hypothesis we investigated the ability of Trissolcus basalis, T. brochymenae, and Trissolcus sp. to distinguish female from male Nezara viridula, Murgantia histrionica, and Graphosoma semipunctatum footprints. These three pentatomid bugs were selected according to variable association levels. Bioassays were conducted on filter paper sheets, and on Brassica oleracea (broccoli) leaves. The results confirmed our hypothesis showing that wasps spent significantly more time on female rather than male traces left by associated hosts on both substrates. No differences were observed in the presence of traces left by non-associated hosts. The ecological consequences for parasitoid host location behaviour are discussed.