Functional anatomy of the ovipositor clip in the parasitoid leptopilina heterotoma (Thompson) (Hymenoptera : Eucoilidae), a structure to grip escaping host larvae

Observations on the host attack behaviour of the parasitoid Leptopilina heterotoma (Hymenoptera : Eucoilidae) led to the supposition that this wasp should possess a structure on its ovipositor by which it can hold a host larvae in a fixed position until the larva is paralyzed. The ovipositor was studied by light, scanning and transmission electron microscopy and appeared to have a clip with teeth on the unpaired valve of the ovipositor, about 50 μm from the tip. Based on the structure of this ovipositor clip and the oviposition behaviour, the functioning of the ovipositor clip is discussed.     

Sense organs on the ovipositor of Macrocentrus cingulum Brischke (Hymenoptera: Braconidae): their probable role in stinging,oviposition and host selection process

Parasitoid wasps from the insect order Hymenoptera can be deployed successfully as biological control agents for a number of pests, and have previously been introduced for the control of corn pest insect species from the Lepidopteran genus Ostrinia. Organs on the ovipositor of parasitoid wasps have mechanical and tactile senses that coordinate the complex movements of egg laying, and the ovipositor of Hymenopteran insects have evolved associated venom glands as part of their stinging defense. The ovipositor of parasitic wasps has evolved an additional function as a piercing organ that is required for the deposition of eggs within suitable host larvae. The morphology and ultrastructure of sense organs on the ovipositor and sheath of Macrocentrus cingulum Brischke (Hymenoptera: Braconidae) are described using scanning and transmission electron microscopy. Three types of sensilla trichodea were shown to be abundant on the outer sheath of the ovipositor, with types II and III being most distal, and the inner surface of the ovipositor covered with microtrichia, more densely near the apex. Sensilla coeloconica are distributed on both ventral and dorsal valves, while campaniform sensilla and secretory pores are only located on the dorsal valve. The olistheter-like interlocking mechanism, as well as the morphology of the ventral and dorsal valve tips and the ventral valve seal may be important in stinging, oviposition and in the host selection process.     

Predation on the woodwasp Tremex longicollis Konow (Hymenoptera: Siricidae) and its parasitoid Megarhyssa jezoensis (Matsumura) (Hymenoptera: Ichneumonidae) adults during oviposition

Adults of the woodwasp Tremex longicollis Konow (Hymenoptera: Siricidae) oviposit in the wood of dying Celtis sinensis trees using their ovipositors, and adults of the parasitoid Megarhyssa jezoensis (Matsumura) (Hymenoptera: Ichneumonidae) deposit eggs into the woodwasp larvae that live in the wood, also using their long ovipositors. Many T. longicollis ovipositors, both with and without abdominal segments, and a M. jezoensis ovipositor were found stuck in C. sinensis wood at two parks in Osaka Prefecture, central Japan. Field observations showed that ovipositing woodwasps and parasitoids were attacked by predators during the daytime, and their ovipositors were left behind in the wood. Woodwasps and their parasitoids, which have long ovipositors, may be vulnerable to predation during oviposition.     

Host location and oviposition in a basal group of parasitic wasps: the subgenual organ, ovipositor apparatus and associated structures in the Orussidae (Hymenoptera, Insecta)

Anatomical studies and behavioural observations indicate that representatives of the Orussidae use vibrational sounding to detect suitable oviposition sites. During host location, vibrations generated by tapping the tips of the antennae against the wood are picked up by the fore legs through the basitarsal spurs, transmitted along the basitarsi to thin-walled areas on the tibiae and through haemolymph to the subgenual organs, where they are transduced into nerve impulses. The apical antennomeres are distinctly shaped and have the cuticle thickened distally. The fore basitarsi have weakly sclerotised basitarsal lines proximally and membranous basitarsal spurs distally. The external wall of the fore tibiae have thin-walled areas distally on their posterior parts. Internally, large subgenual organs are situated opposite the thin-walled areas and each organ consists of 300–400 scolopidial units suspended between a lateral cuticular spine, a ventral sheet and a median ridge. The ovipositor is several times the length of the body of the wasp. When at rest, it extends all the way into the prothorax, where it is coiled before extending posteriorly to lie between the third valvulae distally. The ovipositor lies in a membranous ovipositor sac attached posteriorly to the proximal parts of the ovipositor apparatus and the posterior margin of sternum 7. In the ovipositor apparatus, the anterior parts of the second valvifers are displaced and expanded anterodorsally, inverting the first valvifers and the base of the ovipositor. When in use, the ovipositor is extended and retracted by median apodemes situated on the anterior margins of abdominal sterna 3–7. Longitudinal muscles between the apodemes allow the latter to grip the ovipositor in troughs between them. The ovipositor extends from the abdomen at the tip of sternum 7, and an internal trough on sternum 7 serves to guide the ovipositor into the wood. Despite the alterations observed in the ovipositor apparatus in the Orussidae, the musculature is almost complete and the mode of operation presumably not much different from that of other representatives of the Hymenoptera. The different ways parasitic wasps with very long ovipositors handle and accommodate these and the implications for the evolutionary history of Hymenoptera are discussed. Accepted: 14 March 2001     

Influence of host behavior and host size on the success of oviposition ofCotesia urabae andDolichogenidea eucalypti (Hymenoptera: Braconidae)

Behavioral interactions among Cotesia urabaeAustin and Allen, Dolichogenidea eucalyptiAustin and Allen (Hymenoptera: Braconidae), and their host Uraba lugensWalker, the gum leaf skeletonizer (Lepidoptera: Noctuidae), were observed at three host sizes over a 20-min period. These sizes were first instar (small, gregarious), fourth-fifth instar (mid, gregarious), sixth-seventh instar (large, solitary) larvae. Unlike C. urabae, D. eucalyptiused its legs to hold small larvae before ovipositor insertion. D. eucalyptialso visited patches of small larvae more frequently, proceeded less often through patches of mid larvae, and made significantly fewer ovipositions in mid and large larvae. Small larvae responded to both parasitoids by dispersing outward, while mid larvae responded to parasitoids by moving inward to form a denser group. Larvae reared or thrashed after each parasitoid visit, especially mid larvae, and some continued to do so for up to 2h after parasitoid departure. Mid and large larvae occasionally injured parasitoids by biting their appendages. By rearing or thrashing immediately prior to an encounter with a parasitoid, mid and large larvae decreased the likelihood of being parasitized by up to 50%.     

Temperature dependent functional response ofEncarsia formosa parasitizing the Poinsettia-strain of the cotton whitefly,bemisia tabaci, on Poinsettia

The control efficiency and performance ofEncarsia formosa Gahan (Hymenoptera: Aphelinidae) as influenced by the density of its host, the Poinsettia-strain ofBemisia tabaci Gennadius (Homoptera: Aleyrodidae), were investigated by laboratory experiments on Poinsettia (Euphorbia pulcherrrima Willd.).E. formosa showed a Type II functional response to fourth instar larvae ofB. tabaci, the response plateau increasing with temperature. A response model for randomly searching parasitoids incorporating temperature-dependent handling time and temperature-independent search rate was in accordance with the results, and gave an estimated search rate of 0.033 leaf·hour⁻¹ and handling times of 1.54, 2.86 and 20.1 h at 28°C, 22°C and 16°C, respectively. From the latter, the maximum number of hosts that can be parasitized at the three temperatures are 10.4, 5.6 and 0.8 larvae per day (provided the light period is 16 h). The number of hosts with ovipositor punctures was higher than the number of parasitized hosts, especially at 22°C and 28°C, implying thatE. formosa refrains from laying eggs in some of the hosts examined with the ovipositor. About 31% of the punctured larvae did not contain any eggs. Superparasitism occured during the experiment presumably originating from young, inexperienced parasitoids. Individual larvae were occasionally punctured several times, also by non-superparasitizingE. formosa. The resulting distribution of ovipositior holes was random, indicating thatE. formosa on the basis of antennal testing is unable to determine if a larva has previously been examined with the ovipositor. Almost fifty percent of the punctures were not followed by egg-depositions. Besides parasitizationE. formosa used hosts as food source. The number of hostfed larvae was independent of density, but varied with temperature being highest at 28°C (0.12 hostfed larvae per parasitoid per day).     

Host discrimination of a larval parasitoid: the quick movement of Microplitis demolitor

Many parasitoids discriminate previously parasitised hosts from unparasitised ones to avoid mortality of offspring. Parasitoids that parasitise aggressive hosts such as lepidopteran larvae are known to attack hosts very quickly to avoid being attacked. However, little is known about host discrimination of such quick attacking parasitoids. We investigated host discrimination of Microplitis demolitor (Wilkinson) (Hymenoptera: Braconidae) a quick attacking parasitoid of Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae). Results showed that ratios of female wasps that rejected the hosts after antennal examination did not differ between parasitised and unparasitised hosts, indicating that M. demolitor did not discriminate hosts by antennal examination. However, 95% of females that inserted ovipositor into unparasitised hosts actually laid eggs, whereas it was only 31% for parasitised hosts, indicating that females discriminated hosts by oviposition insertion. Analyzing video recordings revealed that the ovipositor exploration of the host took 0.3 s. Female wasps that had experienced high-host density of unparasitised hosts readily rejected parasitised hosts, while wasps with experience of low host availability of parasitised hosts tended to accept parasitised hosts. This suggests that host discrimination behaviour of M. demolitor is affected by previous experience of different host availability and host quality.

     

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.      

Host selection by Ibalia leucospoides based on temporal variations of volatiles from the hosts’ fungal symbiont

Insect parasitoids locate hosts via reliable and predictable cues such as volatile emissions from hosts and/or host plants. For insects that depend on mutualistic organisms, such as many wood‐boring insects, symbiont‐derived semiochemicals may represent a source of such cues to be exploited by natural enemies. Ultimately, exploitation of these signals may increase fitness by optimizing foraging efficiency. Female parasitoids of Ibalia leucospoides use volatiles from the fungal symbiont Amylostereum areolatum of their host Sirex noctlio to find concealed host eggs and young larvae within the xylem. We hypothesize that the temporal pattern of fungal emissions may indicate not only the presence of host larvae but also be used as a cue that indicates host suitability and age. Such information would allow female parasitoids to discern more efficiently between hosts within ovipositor reach from those already buried too deep into the xylem and out of reach. In this context, we assessed the behaviour of I. leucospoides females to volatiles of A. areolatum in a ‘Y’‐tube olfactometer at regular intervals over 30 days. We concurrently examined the fungal volatiles by headspace sampling through solid‐phase microextraction (SPME) followed by gas chromatography mass spectrometry (GC‐MS). We observed that females were attracted to volatiles produced by two‐week‐old fungal cultures, a period that matches when older larvae are still within ovipositor reach. Four chemical compounds were detected: ethanol, acetone, acetaldehyde and the sesquiterpene 2,2,8‐trimethyltricyclo[6.2.2.01,6]dodec‐5‐ene, with each compounds’ relative abundance changing over time. Results are discussed in the context of parasitoids fitness. Future studies involving electrophysiology, different collection techniques and further behavioural assays will help in identifying the compounds that convey temporal information to female parasitoids and have the potential for being used in integrated pest management programmes.     

Efficiency of vibrational sounding in parasitoid host location depends on substrate density

Parasitoids of concealed hosts have to drill through a substrate with their ovipositor for successful parasitization. Hymenopteran species in this drill-and-sting guild locate immobile pupal hosts by vibrational sounding, i.e., echolocation on solid substrate. Although this host location strategy is assumed to be common among the Orussidae and Ichneumonidae there is no information yet whether it is adapted to characteristics of the host microhabitat. This study examined the effect of substrate density on responsiveness and host location efficiency in two pupal parasitoids, Pimpla turionellae and Xanthopimpla stemmator (Hymenoptera: Ichneumonidae), with different host-niche specialization and corresponding ovipositor morphology. Location and frequency of ovipositor insertions were scored on cylindrical plant stem models of various densities. Substrate density had a significant negative effect on responsiveness, number of ovipositor insertions, and host location precision in both species. The more niche-specific species X. stemmator showed a higher host location precision and insertion activity. We could show that vibrational sounding is obviously adapted to the host microhabitat of the parasitoid species using this host location strategy. We suggest the attenuation of pulses during vibrational sounding as the energetically costly limiting factor for this adaptation.     

Host discrimination by the endoparasitoid Psyllaephagus pistaciae (Hymenoptera: Encyrtidae): A case of time-dependent ability

Host-discrimination behavior by the koinobiont parasitoid, Psyllaephagus pistaciae Ferrière (Hymenoptera: Encyrtidae), the major biocontrol agent of the common pistachio psylla, Agonoscena pistaciae Burckhardt & Lauterer (Hemiptera: Psyllidae), in Iran, was investigated in the laboratory. The results demonstrated that P. pistaciae quickly detected and avoided freshly parasitized hosts, either after antennating or by probing with their ovipositor. However, this discriminatory ability declined with time, probably influenced by both external and internal markers left by the previous ovipositing female. The results also confirmed that female P. pistaciae responded to changes in host quality associated with the parasitoid's larval development 4 days after the initial parasitization, clearly indicating that the second female could detect the presence of the larvae and adjust her host-selection decision. In addition, psyllid nymphs treated topically with a solution of Dufour's gland were rejected by the parasitoid, showing that Dufour's gland secretion had a significant effect as a host marking chemical. The current study also showed that superparasitism increased the host-mortality and that the rate of encapsulation decreased, suggesting that, when two eggs are laid in the same psyllid nymph, only one parasitoid develops to an adult.     

The evolutionary role and fate of the primary ovipositor in insects

The development of a piercing-sawing ovipositor for introducing eggs into living plant tissues has made its owners independent of the soil characteristics and increased egg protection. This was the most important prerequisite for the appearance of wings and flight which provided the winged insects with tremendous opportunities for finding new niches and led to unparalleled adaptive radiation. The ovipositor has passed several stages of improvement and differentiation. The four principal types of the primary ovipositor are considered: those of Odonata, Diaphanopteroidea (only extinct forms), Cicadina (including Paraneoptera and Hymenoptera), and Orthoptera. A new hypothesis of the gonangulum homologies is put forward, interpreting it as half of sternite IX lateral of the midline plus the paratergite of the same segment. The constructions of the valvae and different homologies of the third valvae in Polyneoptera and Eumetabola are discussed. The primary ovipositor has been repeatedly (i.e., in many lineages) reduced in the evolution of Pterygota. The main circumstances of these reductions are: (1) subterranean (fossorial) life in narrow cavities; (2) aquatic life of the larvae, mostly linked with submerged oviposition; (3) development and perfection of flight to which the heavy and protruding ovipositor was a hindrance. All the holometabolous insects except Hymenoptera lack the primary ovipositor.     

Substrate selection by saprophagous wood‐borer larvae within highly variable hosts

In insects completing their larval development within a single host, oviposition site is seen as a major determinant of offspring performance. However, in previous studies, the saprophagous wood‐borer Anthophylax attenuatus (Haldeman) (Coleoptera: Cerambycidae) showed no strong response to between‐host variations in nutritional factors influencing larval growth and survival. To explain such weak selection in adults, we hypothesized that substrate selection occurs at a smaller scale by larvae within hosts showing high variability in substrate quality. In this study, we described within‐host variability in wood density and determined whether wood‐boring larvae were found more often than expected in specific decay types. We characterized the variability of decay in 24 snags by producing wood density profiles for each. We then collected larvae from the same snags through wood dissection, and associated a wood density value to each by taking a wood sample around each larva found. We then compared ratios of available and used substrate types defined by wood density. We observed substantial within‐snag variation in wood density. Middle decay class (0.275–0.375 g cm^?3) was significantly overused by larvae, whereas more decayed wood was clearly avoided. High within‐host variability in substrate quality and active or passive selection by larvae of specific substrate types suggest that selection pressures on adult behaviour could be lower than expected for a parasitic species, and might be linked with the weak selection observed at a larger scale by ovipositing adults.     

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.     

The ovipositor musculature of Bactrocera depressa Shiraki (Diptera, Tephritidae)

Structure of the ovipositor sclerites and musculature was studied in the tephritid fly Bactrocera depressa Shiraki, whose larvae develop in the fruits of Cucurbita moschata and other Cucurbitaceae. The functioning and adaptations of the ovipositor of this species to laying eggs into soft fruits are discussed with respect to their differences from the ovipositor characteristics in species with different host specializations, e.g., Ceratitis capitata (Wiedemann) (Hanna, 1938), Lenitovena trigona Matsumura (Ovtshinnikova, 2008), Urophora affinis Frauenfeld and U. quadrifasciata (Meigen) (Berube and Zacharuk, 1983) and Campiglossa plantaginis (Haliday, 1833) (Ovtshinnikova, 2010). The ovipositor musculature of Bactrocera depressa is very similar to that of Ceratitis capitata, a species which lays eggs in soft tissues of various fruits. The sclerites and musculature of the ovipositor, especially the basal part of syntergosternite VII, in the fruit infesting Bactrocera (also in Ceratitis) is intermediate between that in Lenitovena laying eggs in the (soft) decaying wood, and Urophora and Campiglossa that deposit eggs in the hard flowerheads of asteraceans. Species of the latter genera have still more complicated structure of the basal part of syntergosternite VII enabling a more intense protruding of the ovipositor and its more closely controlled operating during oviposition in the harder plant tissues. The discussed morpho-functional rearrangements proceed within species of a single lineage of the modern classification (Korneyev, 1999) from the basal Lenitovena to the terminal Urophora and Campiglossa, Bactrocera occupying an intermediate position in this row. In this way, the technology changes (from a muscul lever to a pump) resulting in an increase of the ovipositor mobility during piercing of the substrate.     

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.     

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.     

Spider silk felting—functional morphology of the ovipositor tip of Clistopyga sp. (Ichneumonidae) reveals a novel use of the hymenopteran ovipositor

Apical serrations of the hymenopteran ovipositor have been widely postulated to originally constitute adaptations for cutting through hard substrates. Simplifications of the ovipositor tip have occurred in several ichneumonid wasp genera associated with spiders. Despite such reduction in Clistopyga (Hymenoptera, Ichneumonidae), the ovipositor still possesses some apical serrations. Through the first detailed study, we believe, on the behaviour of an ovipositing Clistopyga species, we show that it can alter its ovipositor for different purposes and that the primary function of the apical serrations is clinging to its spider host as the spider attempts to escape. Intriguingly, we also discover a hitherto undocumented adaptation for the hymenopteran ovipositor. The female wasp seals openings in the silken spider nest by using its ovipositor on the silk in a highly sophisticated way that is comparable to how humans entangle wool by needle felting. By studying the ovipositor morphology through a scanning electron microscope, we elucidate how this works, and we hypothesize that by closing the nest the female wasp protects its developing kin.     

Flexible ovipositor sheaths in parasitoid Hymenoptera (Insecta)

The structure of the ovipositor sheaths is examined in a number of parasitic wasps as well as a few non-parasitic taxa. Parasitic wasps in the families Aulacidae, Braconidae, Ichneumonidae, Megalyridae, and Stephanidae with a very elongate, external ovipositor have the lateral wall of the sheath finely transversely subdivided by narrow furrows. This makes the sheath highly flexible, allowing it to support the ovipositor proper during ovipositing. The taxa having such a flexible sheath all drill into wood to lay their eggs. Support at the tip of the ovipositor is crucial in the initial stages of oviposition. The flexible ovipositor sheath is possibly a very ancient trait, arising prior to the radiation of extant Apocrita. This is corroborated by their presence in fossil Hymenoptera from the Upper Jurassic. The occurrence of short and/or concealed ovipositors with rigid sheaths in many extant apocritan taxa must therefore be assumed to be the result of multiple reversals.     

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.