Differentiation of innate but not learnt responses to host‐habitat odours contributes to rapid host finding in a parasitoid genotype

Abstract In parasitoids, host‐habitat odour can influence host searching within the habitat. This is the case in Leptopilina sp. (Hymenoptera, Figitidae), a Drosophila parasitoid searching for larvae by ovipositor probes. This a behaviour can be conditioned to fruit odours. In a previous study, the latency of probing to a fruit odour is reported to have a genetic variability within a laboratory strain. This suggests a link between rapidity of host discovery and fitness. In the present study, this hypothesis is tested by comparing responses to host‐habitat odours between two genotypes of Leptopilina heterotoma Thomson, from the Mediterranean coast (Antibes) and from Burgundy (Tailly). The two genotypes present contrasting rhythms and levels of locomotor activity linked to contrasting interspecific competition in their area of origin. The high activity observed in the Mediterranean genotype is interpreted as an adaptive response to a limited time‐window to win against a competitor species absent in Burgundy. The present study finds differentiation in innate but not learnt responses to host‐habitat odours. The more active genotype (Antibes) has a higher probability and a shorter latency of innate probing to the odours than the less active genotype (Tailly); Antibes females also find larvae and complete infestations more rapidly. Learning equalizes the probability and the latency of probing to the odours in both strains, and increases the probing duration. Innate responses to host‐habitat odours would allow time‐limited insects to increase their reproductive rate, when host predictability is high in the habitat. Selection of faster innate responses to host and habitat cues without evolution of learnt responses indicates that the initial host discovery is more crucial to fitness than subsequent ones.     

Host finding and discrimination in Diglyphus isaea,a parasitoid of the chrysanthemum leaf miner,Chromatomyia syngenesiae

Naive and experienced Diglyphus isaea were attracted by host plant odours of lettuce and chrysanthemum to search and probe on infested and uninfested leaves. A slight preference was shown for leaves infested with Chromatomyia syngenesiae. At close range, visual stimuli were unnecessary for oviposition and host‐feeding. Contact with uninfested lettuce and chrysanthemum elicited searching and probing behaviour whereas host frass did not. Host larval movement appeared to aid host detection at close range. The number of ovipositor probes increased with proximity to live hosts but not for stationary, dead hosts. Dead hosts were frequently walked over or missed when D. isaea passed within 0.5 cm of the stationary larvae. Dead hosts were also rejected for oviposition but not for host feeding. Both naive and experienced females discriminated between healthy hosts and those which had been attacked by conspecifics or encountered previously.     

The attractiveness of different odour sources from the fruit–host complex on Leptopilina boulardi, a larval parasitoid of frugivorous Drosophila spp.

In parasitoid insects, successful offspring development depends on the female’s ability to find a suitable host. Specific recognition is often based on responses to olfactory cues, but their source and nature have rarely been determined. –This paper deals with the recognition of odours involved in host location by Leptopilina boulardi[Barbotin, Carton & Kelner-Pillault] (Hymenoptera: Eucoilidae), a larval parasitoid of Drosophila species that develops in mature fruits. The nature and origin of volatile stimuli recognized among odours of the host–fruit complex, and the effect of learning on this recognition, were investigated. Oriented responses to these odours were observed in a four-armed olfactometer and were analysed with the observer software (Noldus Information Technology). Fruit odours alone (banana and pear) were not spontaneously attractive to naive parasitoids, whereas naturally-infested bananas were highly attractive. The attraction was related to the odour that adult Drosophila left on the substrate but not to Drosophila oviposition activity or larval development. A synergism between some fruit odours (banana and pear) and the odour left by adult Drosophila on damp filter paper was observed. However, when testing a non-fruit substrate (mushroom), no synergism was observed. Thus, female L. boulardi may innately recognize host–food substrate odours associated with odours from the adult stage of their host. In addition, an oviposition experience on an infested banana allows L. boulardi females to memorise the fruit odour itself through associative learning. The adaptive significance of this process is discussed.     

Host-handling behavior of the parasitoid,Diglyphus minoeus [Hym.: Eulophidae], parasitizing the honeysuckle leaf-miner,Phytomyza lonicerae [Dipt.: Agromyzidae]

The host-handling behavior of individual female waspsDiglyphys minoeus which attack the larvae of the honeysuckle leaf-miner,Phytomyza lonicerae, include 4 types of behavior: probing, ovipositor insertion, host-feeding, and resting. Ovipositor insertion may either be injection of venom, probing of the host by the sensilla on the ovipositor's tip or egg laying. Three types of attacks were distinguished: oviposition attack for unparasitized larvae, host-feeding attack for unparasitized host larvae and host rejection for previously parasitized host larvae. Oviposition attack was characterized by frequent alternation between probing and ovipositor insertion, long duration of ovipositor insertion and resting, and the long duration of host-handling. Resting behavior is thought to protect the progeny against superparasitism, host-feeding by other wasps or hosts' recovery from paralysis. Host-feeding attack was characterized by frequent alternation between host-feeding and ovipositor insertion and long duration of host-feeding. Host rejection was composed mainly of probing and ovipositor insertion and short handling time.      

Nitrogen fixation by intact colonies of the termite Nasutitermes corniger

Summary Biosteres longicaudatus Ashmead (Hymenoptera: Bracon dae) is a solitary endoparasite of Anastrepha suspensa larvae (Diptera: Tephritidae), which live in fruit tissue. Larvae make andible noises within macerated fruit or larval medium in which they are reared. Parasite females readily located normal, mobile larvae and spent a mean of 16.5±4.7 min/visit to parasitize these hosts. In contrast, females were unable to locate etherized or dead hosts and abandoned them after only 1.9±0.9 and 2.3±0.8 min, respectively. Females of all ages, with and without oviposition experience, exhibited non-random search and ovipositor probe behaviors in response to artifically created vibration. This response was influenced primarily by the number of mature eggs in the ovaries. These findings suggest that 1) an accumulation of mature eggs in the ovaries increase the appetitive drive of females to find and oviposit in hosts and 2) host sound/vibration produced either by movement of hosts through the medium and/or by the rasping mouth hooks during feeding. is used by parasites as a releaser for host finding behavior as well as a cue to the location of the host within the substrate.     

Molecular markers to differentiate two morphologically-close species of the genus Sitobion

The effect of allelochemicals from its host, the larva of the cabbage root fly, Delia radicum (L.) (Diptera: Anthomyiidae), and the host's food plant on the ovipositor probing response of the parasitoid Trybliographa rapae (Westw.) (Hymenoptera: Cynipidae) were investigated. Trybliographa rapae probed both cabbage root fly infested and uninfested swede (Brassica napus var. napobrassica), although significantly more wasps responded to infested swede. Antennal sensilla are likely to be the mediators of this response. The synomones and kairomones involved are extractable in water, diethyl ether and methanol. No response was observed to washed, starved cabbage root fly larvae. Wasps spent significantly longer searching infested swede than uninfested, although probing frequency remained constant. It is suggested that the initiation of probing in T. rapae is dependent on a threshold concentration of general synomones or host related synomones and kairomones, whereas time spent searching a particular area is dependent on the environment perceived by sensilla on the ovipositor.     

Behavioral Attraction of Two Parasitoids,Venturia canescens and Bracon hebetor,to Silk Extracts of a Host Plodia interpunctella

Kairomonal activities of silk extracts of host Plodia interpunctella were determined by measuring the rates of behavioral responses of two parasitic wasps, Venturia canescens and Bracon hebetor. Silk of P. interpunctella larvae attracted both parasitic wasps but the cocoon silk of silkworm, Bombyx mori and the web silk of two-spotted spider mite, Tetranychus urticae did not. Silk components of P. interpunctella were extracted by using either hexane or methanol, and tested the rates of three serial responses of wasps; host location, antennal drumming and ovipositor probing behaviors. The patterns of each behavioral response were similar in two wasps. The rates of each response were increased at the higher concentrations of both extracts. Antennal drumming behavior was much more responsive to lower concentrations of both extracts than ovipositor probing behavior was. Furthermore, the rate of antennal drumming response was higher in hexane-extracts rather than methanol-extract in both wasps; V. canescens and B. hebetor for 20 and 17 times, respectively. However, ovipositor probing response was similar in two different extracts. Both extracts elicited 100% of antennal drumming response but ovipositor probing response was only 60 to 80% of all tested individuals. Our results were shown that silk extracts of host larvae elicited strong behavioral responses of two parasitic wasps and could be applied for practical application of parasitoids attraction in the biological control of agricultural pests.     

Attraction of Drosophila melanogaster males to food-related and fly odours

The fruit fly Drosophila melanogaster has become a model for olfaction and odour-mediated behaviour. In the wild, Drosophila flies aggregate on decaying fruit where they mate and oviposit and a strategy to find mates would be to locate fruit which has already been colonized by other flies. We therefore developed a bioassay to investigate attraction of males to food and fly odours. We showed that upwind flights are initiated by food odours. At shorter distances, males are attracted by volatiles produced by conspecifics. However, only odours produced by copulating flies attract males. This suggests either a synergistic effect of both male and female odours or changes in pheromone release during mating, that indicate the presence of sexually receptive females. Our findings demonstrate the essential role of food odours and pheromones for mate location in D. melanogaster.     

The Role of Chemical Cues in Host and Mate Location in the Pear Psylla Cacopsylla bidens (Homoptera: Psyllidae)

Chemical communication was shown to play a role in the pear psylla, Cacopsylla bidens. Electrophysiological (EAG) and behavioral responses were investigated in males and females pear psylla . Males were found to be attracted to females, and especially to those on host plants, but not to males, uninfested host plants, or plants infested with conspecific larvae. On the other hand, females were not attracted to males or females but displayed some attraction to host plants. Furthermore, females showed a preference for uninfested pear versus plants infested with conspecific larvae. The antennae of males gave highest electroantenographic response to volatiles from pears infested with females but not males, while females, responded also toward the volatiles of pear alone. These results indicate that females of C. bidens emit sex pheromones that are attractive to the males and suggest that, host volatiles may play a role in host selection by pear psylla females.     

Exploitation of the Fecal Shield of the Lily Leaf Beetle, Lilioceris lilii (Coleoptera: Chrysomelidae), by the Specialist Parasitoid Lemophagus pulcher (Hymenoptera: Ichneumonidae)

We investigated signal sources used by the parasitoid Lemophagus pulcher in locating and accepting larvae of its host, the lily leaf beetle, Lilioceris lilii. Olfactometer bioassays revealed that larvae with fecal shields, larvae without shields, the shield alone, and lily leaves damaged by L. lilii were all attractive to female parasitoids. In contact bioassays, L. pulcher females were attracted to shields and showed ovipositor probing independently of whether the larva underneath was L. lilii or a nonhost, suggesting that the shield plays a primary role in short-range host location and host acceptance by L. pulcher. The attractiveness of the shield is at least partly of a chemical nature, since shield extracts applied to dummies increased contact duration and induced ovipositor probing by L. pulcher. Another putative defense system of L. lilii, i.e., oral discharge which is emitted by disturbed larvae, was also attractive to experienced, but not to naive, female parasitoids. In all other tests, naive and experienced female L. pulcher responded to the same signal sources tested, suggesting that the host-selection behavior of this biological control candidate is governed largely by innate responses to host-associated cues.     

Hybrid Lethal Systems in the Drosophila Melanogaster Species Complex. II. the Zygotic Hybrid Rescue (Zhr) Gene of D. Melanogaster

Hybrid females from Drosophila simulans females X Drosophila melanogaster males die as embryos while hybrid males from the reciprocal cross die as larvae. We have recovered a mutation in melanogaster that rescues the former hybrid females. It was located on the X chromosome at a position close to the centromere, and it was a zygotically acting gene, in contrast with mhr (maternal hybrid rescue) in simulans that rescues the same hybrids maternally. We named it Zhr (Zygotic hybrid rescue). The gene also rescues hybrid females from embryonic lethals in crosses of Drosophila mauritiana females X D. melanogaster males and of Drosophila sechellia females X D. melanogaster males. Independence of the hybrid embryonic lethality and the hybrid larval lethality suggested in a companion study was confirmed by employing two rescue genes, Zhr and Hmr (Hybrid male rescue), in doubly lethal hybrids. A model is proposed to explain the genetic mechanisms of hybrid lethalities as well as the evolutionary pathways.     

Influence of pre-emergence experience on response to host and host plant odours in the larval parasitoidEupelmus vuilleti

The response to different host and plant species odours was investigated inEupelmus vuilleti (Crw). This hymenopteran is a solitary ectoparasitoid of several species of bruchids developing inside Leguminosae seeds. The locomotor behaviour of females reared onBruchidius atrolineatus (Pic) larvae developing inVigna unguiculata (Walp) seeds was analysed using a tubular olfactometer. Females showed a specific sensitivity to the semiochemicals emanating from the host and the seed species on which they had developed. Odours fromV. unguiculata seeds were attractive to the parasitoid and stimulated their locomotor activity. Odours fromVigna radiata (Wil) seeds had no effect on the locomotor behaviour. Odours fromB. atrolineatus larvae were attractive to the females whereas odours fromCallosobruchus maculatus (Fab), another bruchid species, had no effect. By isolating the females from the seed and the host at different developmental stages, we found that the specific sensitivity observed resulted from an early adult learning. This learning which occurs before the emergence from the seed while the imago is in the larval chamber of its host is dependent on contact with the seed and the host larval remains.     

Morphological adaptation in host races of Tephritis conura

The present study investigates morphological differentiation among host races of the fruit fly Tephritis conura Loew (Diptera: Tephritidae) for two fitness‐related traits and whether these traits are host induced or genetically determined. Flies were analyzed from independent sympatric regions, and from one syntopic site where parental host plants [Cirsium heterophyllum (L.) Hill. and Cirsium oleraceum (L.) Scop. (Cardueae)] and hybrid plants (C. heterophyllum×C. oleraceum) co‐occur. As both host races may oviposit on hybrid plants and hybrid plants provide an identical environment for larvae of both host races, flies emerging from C. heterophyllum×C. oleraceum hybrids were used to assess whether host‐race morphological differences are genetically determined or due to phenotypic plasticity. No significant size (wing length) differences were found among host races, whereas flies emerging from C. heterophyllum had on average 8.4% longer ovipositors than flies emerging from C. oleraceum. The mean size‐corrected ovipositor length (i.e., the ratio ovipositor/wing length) was 10.3% longer. These proportions were repeated among host races emerging from hybrid plants. Although flies of the C. heterophyllum host race from hybrid plants were smaller than on parental host plants, the ratio ovipositor/wing length was constant. Hybrid flies (which emerged only on hybrid plants) were intermediate in relative and absolute ovipositor length. Thus, ovipositor‐length differences among T. conura host races most likely have a genetic basis. This suggests that host‐related differences in ovipositor length reflect adaptations to the respective host‐plant species, most likely to the host's flower‐head size, whereas both host races experience similar selection regimes on body size.     

Host-marking behaviour as a quantitative signal of competition in the walnut fly Rhagoletis juglandis

Abstract.  1. Walnut-infesting flies in the Rhagoletis suavis species group actively re-use hosts for oviposition despite engaging in a genus-typical host-marking behaviour which, in other Rhagoletis groups, deters oviposition. In a study of the walnut fly, R. juglandis (Cresson), alternative hypotheses for the putative marking behaviour were evaluated.
2. The oviposition site attraction hypothesis proposes that the host mark guides females to oviposition sites on occupied fruit. The competition intensity signal hypothesis proposes that the host mark is an indicator of the level of competition to be incurred if fruit are re-used.
3. In a field cage, females were presented simultaneously with fruit previously exposed to 25 females that were also allowed to oviposit and engage in the putative marking behaviour, and control fruit on which females were allowed only to oviposit. The occurrence of host marking reduced a female's propensity to oviposit from 46% to just over 10%, consistent with the competition intensity signal hypothesis only.
4. In a laboratory assay, the duration of host marking was correlated positively with the size of a female's clutch. This result, also consistent with the competition intensity signal hypothesis, suggests that the amount of marking pheromone on a fruit is a reliable indicator of the number of eggs already deposited within.
5. In a second field-cage experiment, females were allowed to mark on fruit for 0, 10, 20, or 30 min and fruit were presented to test females. Whether or not females alighted on a particular host was not affected by the duration of marking; however, the frequency of both ovipositor probing and egg deposition decreased with increasing duration of marking. Consistent with the competition intensity signal hypothesis, this result suggests that the host mark permits females to assess the level of competition that a clutch will incur within re-used fruit.     

Host location by the fruit fly parasitoid Diachasmimorpha krausii: role of fruit fly species,life stage and host plant

• 1 Diachasmimorpha krausii is a braconid parasitoid of larval tephritid fruit flies, which feed cryptically within host fruit. At the ovipositor probing stage, the wasp cannot discriminate between hosts that are physiologically suitable or unsuitable for offspring development and must use other cues to locate suitable hosts.
• 2 To identify the cues used by the parasitoid to find suitable hosts, we offered, to free flying wasps, different combinations of three fruit fly species (Bactrocera tryoni, Bactrocera cacuminata, Bactrocera cucumis), different life stages of those flies (adults and larvae) and different host plants (Solanum lycopersicon, Solanum mauritianum, Cucurbita pepo). In the laboratory, the wasp will readily oviposit into larvae of all three flies but successfully develops only in B. tryoni. Bactrocera tryoni commonly infests S. lycopersicon (tomato), rarely S. mauritianum (wild tobacco) but never C. pepo (zucchini). The latter two plant species are common hosts for B. cacuminata and B. cucumis, respectively.
• 3 The parasitoid showed little or no response to uninfested plants of any of the test species. The presence of adult B. tryoni, however, increased parasitoid residency time on uninfested tomato.
• 4 When the three fruit types were all infested with larvae, parasitoid response was strongest to tomato, regardless of whether the larvae were physiologically suitable or unsuitable for offspring development. By contrast, zucchini was rarely visited by the wasp, even when infested with B. tryoni larvae.
• 5 Wild tobacco was infrequently visited when infested with B. cacuminata larvae but was more frequently visited, with greater parasitoid residency time and probing, when adult flies (either B. cacuminata or B. tryoni) were also present.
• 6 We conclude that herbivore‐induced, nonspecific host fruit wound volatiles were the major cue used by foraging D. krausii. Although positive orientation to infested host plants is well known from previous studies on opiine braconids, the failure of the wasp to orientate to some plants even when infested with physiologically suitable larvae, and the secondary role played by adult fruit flies in wasp host searching, are newly‐identified mechanisms that may aid parasitoid host location in environments where both physiologically suitable and unsuitable hosts occur.

     

Effect of food deprivation on attractiveness of food sources, containing natural and artificial sugar and protein, to three African fruit flies: Ceratitis cosyra, Ceratitis fasciventris, and Ceratitis capitata

The effects of food deprivation, age, and mating status on the responses of three fruit fly species, Ceratitis cosyra (Walker), Ceratitits fasciventris (Bezzi), and Ceratitits capitata (Wiedemann) (Diptera: Tephritidae) to natural and artificial sugar and protein food sources were investigated. Natural food sources included guava [Psidium guajava L. (Myrtaceae)] juice (a common host fruit for all three fruit fly species) and bird faeces (farm chicken). Artificial food sources included molasses (obtained from a local sugar factory) and a locally produced protein bait (the International Centre of Insect Physiology and Ecology yeast). In all species studied, sugar deprivation of immature (1–2‐day‐old) male and female flies increased their response to food odours, although it did not change their preference for the type of odour (protein or sugar). Protein deprivation of mature (14–17‐day‐old) male and female flies also increased their response to food odours compared to protein‐fed flies. Protein‐deprived females were highly attracted to odours from protein sources in particular. Odours from natural food sources, guava juice, and chicken faeces, were more attractive to food‐deprived flies than were odours from artificial sugar and protein sources. Attraction to food odours increased significantly with increasing age for protein‐deprived females of all species. For males and females of all species, nutritional state was a more important factor than mating status in influencing responses of flies to food odours. Practical implications of these findings are discussed in terms of strategies for fruit fly control using food baits.     

Effect of entomopoxvirus infection of Pseudaletia separata larvae on the oviposition behavior of Cotesia kariyai

Parasitoid behavior and oviposition were monitored to investigate the ability of Cotesia kariyai (Watanabe) (Hymenoptera: Braconidae) to distinguish between entomopoxvirus-infected and noninfected larvae of Pseudaletia separata Walker (Lepidoptera: Noctuidae). Comparison of individual means revealed that searching time was not significantly different between treatments. Ovipositional time, however, differed significantly after day 8 post virus inoculation. The parasitoid adopted ovipositional posture on infected and noninfected host irrespective of the stage of infection and did not distinguish infected from noninfected larvae up to 4 days post infection. Mean number of larvae stung by the parasitoid was not influenced by the length of time interval between exposure to the virus. However, duration of ovipositor insertion was influenced by the time interval between exposure to the virus and subsequent exposure to parasitoid females. Female parasitoids did not completely reject infected larvae as unsuitable for egg deposition. However, 5 days after virus inoculation they rejected significantly more infected than noninfected larvae. Furthermore, significantly more eggs were laid in noninfected than infected larvae from day 5 following virus administration. These observations suggest that probing by Cotesia kariyai may provide information concerning host suitability.     

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.     

Host preference and sex allocation of three hymenopteran parasitoid species (Ichneumonidae and Braconidae) of a longicorn pest, Oemona hirta (Fabr.) (Col., Cerambycidae)

The host preference and sex allocation of two ichneumond parasitoids, Xanthocryptus novozealandicus (Dalla Torre) and Campoplex sp., and one braconid parasitoid, Apsicolpus hudsoni Turner of the longicorn borer, Oemona hirta , were studied in New Zealand. The size of the borer larvae attacked by the parasitoids can be determined by measuring the gallery width of the borers. These parasitoids are idiobionts and their secondary sex ratio is female-biased. Females of these species are significantly larger than males, and the females allocate the sex of offspring depending on the size of host larvae, by laying female-producing eggs on significantly larger host larvae. Xanthocryptus novozealandicus is larger in body size and shorter in ovipositor length than Campoplex sp. and A. hudsoni , and consumes late instar larvae or prepupae in shallow galleries whereas the latter two utilize early to middle instar larvae in deep galleries. Campoplex sp. and A. hudsoni preferred host larvae in significantly larger tree twigs than X. novozealandicus .     

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.