Contact Kairomones Research Articles

Does intraspecific variation in Trissolcus japonicus affect its response to non-target hosts?

Host specificity is a key criterion for the implementation of any biological control program. In parasitoid wasps, host specificity is mediated by their response to chemical cues directly and indirectly associated with their hosts during foraging. Although intraspecific variation in foraging behaviour is documented, it is rarely integrated into risk assessment studies of potential biological control agents, as is the case for Trissolcus japonicus (Ashmead) (Hymenoptera: Scelionidae), an Asian egg parasitoid and biological control agent of the invasive Halyomorpha halys (Stål) (Hemiptera: Pentatomidae). In the present study, we compared the behavioural response of T. japonicus females of an adventive line present in Switzerland and an Asian line (Beijing USDA), recently mass released in Italy, to cues of its target host H. halys and non-target host Pentatoma rufipes (L.) (Hemiptera: Pentatomidae). We observed minimal differences in host acceptance, exploitation and suitability in no-choice tests. In contrast, the behavioural response to host contact kairomones left on leaf substrates significantly differed between the two lines. While both lines preferred chemical footprints of H. halys in comparison to P. rufipes, females of the Beijing USDA line responded more strongly to host kairomones. The observed intraspecific variation in behavioural traits highlights the relevance of considering genetically distinct parasitoid lines and their host foraging behaviour in biological risk assessment studies. However, the implications of intraspecific variation in parasitoid foraging behaviour for biological control of a pest and non-target effects in the field remain challenging to predict.

Only Females Oviposit: Chemical Discrimination of Adult Stink Bug Sex by the Egg Parasitoid Trissolcus japonicus

Egg parasitoids foraging for suitable hosts scattered in the environment rely mainly on chemical cues. Elucidating the chemical ecology of natural enemies is important in the development of effective and successful strategies for conservation biological control. In this context, the host cuticular hydrocarbons, which are exploited by several species of egg parasitoids as contact kairomones, could be used to retain them by providing information about the presence and the sex of adults of the target species: sex is important because only females of the host species lay the eggs that can be subsequently utilized for parasitoid reproduction. However, the chemical basis of host sex discrimination in egg parasitoids is not well understood. We carried out behavioral and chemical bioassays to investigate the role played by contact chemical cues left by adults of the brown marmorated stink bug, Halyomorpha halys Stål, in host egg searching behavior and adult host sex discrimination by the egg parasitoid Trissolcus japonicus (Ashmead). A first set of bioassays showed that parasitoids spent more time exploring patches contaminated with chemicals associated with adult H. halys females compared with adult males. Similar responses were displayed by T. japonicus when hexane extracts of H. halys were tested suggesting that non-polar chemical compounds are involved in host sex discrimination. GC-MS analysis of hexane extracts revealed quantitative differences in the cuticular compounds of the two sexes, with 1-hexadecene (more abundant in males) being the most important component in determining these differences. Hexane extracts of H. halys females blended with synthetic 1-hexadecene significantly reduced the wasps’ arrestment responses compared to crude extracts.

The Importance of Methyl-Branched Cuticular Hydrocarbons for Successful Host Recognition by the Larval Ectoparasitoid Holepyris sylvanidis

Cuticular hydrocarbons (CHCs) of host insects are used by many parasitic wasps as contact kairomones for host location and recognition. As the chemical composition of CHCs varies from species to species, the CHC pattern represents a reliable indicator for parasitoids to discriminate host from non-host species. Holepyris sylvanidis is an ectoparasitoid of beetle larvae infesting stored products. Previous studies demonstrated that the larval CHC profile of the confused flour beetle, Tribolium confusum, comprises long chain linear and methyl-branched alkanes (methyl alkanes), which elicit trail following and host recognition in H. sylvanidis. Here we addressed the question, whether different behavioral responses of this parasitoid species to larvae of other beetle species are due to differences in the larval CHC pattern. Our study revealed that H. sylvanidis recognizes and accepts larvae of T. confusum, T. castaneum and T. destructor as hosts, whereas larvae of Oryzaephilus surinamensis were rejected. However, the latter species became attractive after applying a sample of T. confusum larval CHCs to solvent extracted larvae. Chemical analyses of the larval extracts revealed that CHC profiles of the Tribolium species were similar in their composition, while that of O. surinamensis differed qualitatively and quantitatively, i.e. methyl alkanes were present as minor components on the cuticle of all Tribolium larvae, but were absent in the O. surinamensis CHC profile. Furthermore, the parasitoid successfully recognized solvent extracted T. confusum larvae as hosts after they had been treated with a fraction of methyl alkanes. Our results show that methyl alkanes are needed for host recognition by H. sylvanidis.

Host Kairomones Influence Searching Behavior of Trissolcus japonicus (Hymenoptera: Scelionidae), a Parasitoid of Halyomorpha halys (Heteroptera: Pentatomidae).

The brown marmorated stink bug, Halyomorpha halys (Stål), is a highly polyphagous species native to Asia that has become a serious invasive agricultural and nuisance pest across North America and Europe. Classical biological control host range evaluations have revealed egg parasitoid Trissolcus japonicus (Ashmead) to be the primary candidate biocontrol agent for field release against H. halys. However, these evaluations only provide us with the physiological host range of T. japonicus. Other Trissolcus species have demonstrated that contact kairomones from different host species elicit varied responses in the parasitoids' host foraging behaviors. To assess T. japonicus response to host kairomones, mated naive females were exposed to leaf surfaces contaminated with adult kairomones from its preferred host, H. halys, or from a native nontarget host, Podisus maculiventris (Say) (Heteroptera: Pentatomidae). Red maple, apple, and soybean were used as plant substrate treatments. The wasp's residence time on the leaf surface, linear walking velocity, and angular walking velocity were observed and measured using Noldus EthoVision XT tracking software. Within each leaf treatment, T. japonicus displayed stronger behavioral responses on leaves contaminated with contact kairomones from H. halys. The parasitoid resided on H. halys contaminated leaves for approximately twice as a long as it did on P. maculiventris contaminated leaves. Further, both species' kairomones elicited significant decreases in parasitoid walking velocity on all tested substrate types. Overall, our study suggests that kairomone-based behavioral studies can be used to further evaluate the host specificity of T. japonicus and can be an invaluable supplement to classical biocontrol host range testing regimes.

α-Amylase Mediates Host Acceptance in the Braconid Parasitoid Cotesia flavipes

Foraging parasitoids use chemical signals in host recognition and selection processes. Although, the volatiles play a relevant role in the localization by parasitoids of their hosts feeding on plants, the host identification process for acceptance occurs mainly during contact between the parasitoid and its host where host products related to feeding activities, fecal pellets and oral secretions, play a crucial role. The purpose of this study was to identify the nature of the contact kairomone(s) that mediate the acceptance for oviposition of the parasitoid Cotesia flavipes Cameron (Hymenoptera, Braconidae), which was released in Kenya in 1993 to control the invasive crambid Chilo partellus (Swinhoe). Using host and non-hosts of C. flavipes, we showed that it is mainly the oral secretions of the larvae that harbour the active compound(s) that mediate host acceptance for oviposition by C. flavipes. Using an integration of behavioral observations and biochemical approaches, the active compound of the oral secretions was identified as an α-amylase. Using synthetized α-amylases from Drosophila melanogaster (an insect model for which syntheses of active and inactive α-amylases are available), we observed that the conformation of the enzyme rather than its catalytic site as well as its substrate and its degradation product is responsible for host acceptance and oviposition mediation of C. flavipes females. The results suggest that the α-amylase from oral secretions of the caterpillar host is a good candidate for an evolutionary solution to host acceptance for oviposition in C. flavipes.

Cuticular Hydrocarbons of Tribolium confusum Larvae Mediate Trail Following and Host Recognition in the Ectoparasitoid Holepyris sylvanidis.

Parasitic wasps which attack insects infesting processed stored food need to locate their hosts hidden inside these products. Their host search is well-known to be guided by host kairomones, perceived via olfaction or contact. Among contact kairomones, host cuticular hydrocarbons (CHCs) may provide reliable information for a parasitoid. However, the chemistry of CHC profiles of hosts living in processed stored food products is largely unknown. Here we showed that the ectoparasitoid Holepyris sylvanidis uses CHCs of its host Tribolium confusum, a worldwide stored product pest, as kairomones for host location and recognition at short range. Chemical analysis of T. confusum larval extracts by gas chromatography coupled with mass spectrometry revealed a rich blend of long-chain (C25-C30) hydrocarbons, including n-alkanes, mono-, and dimethylalkanes. We further studied whether host larvae leave sufficient CHCs on a substrate where they walk along, thus allowing parasitoids to perceive a CHC trail and follow it to their host larvae. We detected 18 CHCs on a substrate that had been exposed to host larvae. These compounds were also found in crude extracts of host larvae and made up about a fifth of the CHC amount extracted. Behavioral assays showed that trails of host CHCs were followed by the parasitoids and reduced their searching time until successful host recognition. Host CHC trails deposited on different substrates were persistent for about a day. Hence, the parasitoid H. sylvanidis exploits CHCs of T. confusum larvae for host finding by following host CHC trails and for host recognition by direct contact with host larvae.

Pentatomid host suitability and the discovery of introduced populations of Trissolcus japonicus in Oregon

Trissolcus japonicus is an egg parasitoid of Halyomorpha halys, brown marmorated stink bug, a severe agricultural pest in the USA. T. japonicus is being evaluated in quarantine as a classical biological control agent to manage H. halys populations in the USA. To determine T. japonicus’ potential for successful management of the pest, we performed a series of no-choice and paired-host-range tests, evaluating parasitism and host recognition in ten nontarget insects. In laboratory no-choice tests, T. japonicus successfully parasitized egg masses of seven Pentatomidae native to Oregon in addition to H. halys. Mean parasitism proportions of egg masses were greater than 40% in two species, B. dimidiata and H. abbreviatus, and were statistically similar to parasitism of H. halys. However, paired-host tests identified higher proportions of parasitized H. halys egg masses compared to four other pentatomids. T. japonicus was equally attracted to volatiles produced by H. halys and other pentatomids but demonstrated significantly longer arrestment response time on surfaces with H. halys contact kairomones. Although host acceptance patterns were similar between stink bug species, our results suggest a greater potential for parasitoid development in H. halys eggs compared to the native pentatomids. During host-range testing, we detected field populations of T. japonicus at 11 sites in Portland, OR, indicating an unintentional introduction and establishment. Further work is needed to characterize its nontarget activity and dispersal patterns in areas where H. halys causes economic damage in Oregon.

The role of contact chemoreception in the host location process of an egg parasitoid

Taste allows insects to detect palatable or toxic foods, identify a mate, and select appropriate oviposition sites. The gustatory system strongly contributes to the survival and reproductive success of many species, yet it is rarely studied in insect parasitoids. In order to locate and assess a host in which they will lay their eggs, female wasps actively search for chemical cues using their sensory organs present mainly on the antennae. In this paper, we studied the role of antennal taste sensilla chaetica in the perception of contact semiochemicals in Trissolcus brochymenae (Hymenoptera: Platygastridae), an egg parasitoid of the brassicaceae pest Murgantia histrionica (Heteroptera: Pentatomidae). Methanolic extracts obtained from male and female hosts elicited action potentials in taste neurons housed in antennal sensilla chaetica, indicating that these sensilla are involved in the perception of non volatile host kairomones. In behavioural assays, wasp females displayed an intense searching behaviour in open arenas treated with host extracts, thus confirming that these kairomones are soluble in polar solvents. We further investigated the extracts by Gas Chromatography-Mass Spectrometry (GC-MS) and found that they contain several compounds which are good candidates for these contact kairomones. This study contributes to better understanding contact chemoreception in egg parasitoids and identifying gustatory receptor neurons involved in the host location process.

Aphid honeydew: An arrestant and a contact kairomone for Episyrphus balteatus (Diptera: Syrphidae) larvae and adults

Predator searching efficiency increases in response to a variety of environmental cues associated with its prey. The sugary excretion of aphids (honeydew) has been found to act as a prey-associated cue for many aphid natural enemies. In the present study, the honeydew excreted by Acyrthosiphon pisum (Harris) was identified as an arrestant and a contact kairomone for young larvae and adults of a common predatory hoverfly, Episyrphus balteatus (De Geer) (Diptera: Syrphidae). First and second instar larvae increased their foraging behaviour in the honeydew-treated area. When plants were sprayed with crude honeydew, the speed of movement of female E. balteatus was significantly higher than in controls, resulting in a longer period of time spent on treated plants and laying eggs. We conclude that the honeydew excreted by A. pisum induces searching behaviour and acts as and arrestant not only for adults but also for young E. balteatus larvae.

Behavioral and Chemical Investigations of Contact Kairomones Released by the Mud Dauber Wasp Trypoxylon politum, a Host of the Parasitoid Melittobia digitata

Contact kairomones from the host mud dauber wasp Trypoxylon politum Say (Hymenoptera: Crabronidae) that mediate behavioral responses of its ectoparasitoid Melittobia digitata Dahms (Hymenoptera: Eulophidae) were investigated. Chemical residues from host by-products, the cocoon, and the meconium, induced arrestment behavior of macropterous female parasitoids, while those from the host stage attacked, i.e., the prepupa, did not. Melittobia digitata response to polar and apolar extracts of host by-products indicated kairomone(s) solubility mainly in hexane. GC and GC/MS analysis of cocoon and meconium apolar extracts revealed a mixture of linear carboxylic acids from C(6) to C(18), and both extracts contained almost identical compounds. When a reconstructed blend of host by-product carboxylic acids was tested, M. digitata females showed only a weak response, thus suggesting that other unidentified compounds present in small quantities also may be involved. Melittobia digitata's response to contact kairomones was innate and not affected by previous host exposure experience. Our results provide evidence of contact kairomone exploitation in the genus Melittobia. The ecological significance of these findings in the host selection process of M. digitata is discussed.

Host Sex Discrimination by an Egg Parasitoid on Brassica Leaves

Egg parasitoids are able to find their hosts by exploiting their chemical footprints as host location cues. In nature, the apolar epicuticular wax layer of plants that consists of several classes of hydrocarbons serves as the substrate that retains these contact kairomones. However, experiments on chemical footprints generally have used filter paper as substrate to study insect behavior. Here, we explored the ability of Trissolcus basalis (Scelionidae) females to discriminate between footprint cues left by male and female Nezara viridula (Pentatomidae) on leaves of their host plant Brassica oleracea (broccoli). Furthermore, we analyzed the chemical composition of the outermost wax layer of broccoli leaves to evaluate the degree of overlap in insect and plant cuticular hydrocarbons that could lead to masking effects in the detection of footprint cues. Our results showed that B. oleracea epicuticular wax retains the chemical footprints of adult bugs and allows T. basalis females to differentiate hosts of different sex. Traces of female bugs elicited more extensive searching behavior in egg parasitoids than traces of males. The application of n-nonadecane, a compound specific to male N. viridula, on the tarsi of female bugs prevented parasitoid females from distinguishing between host male and host female footprints. Analyses of B. oleracea leaves revealed that epicuticular waxes were mainly composed of linear alkanes, ketones, and secondary alcohols. Alkanes were dominated by n-nonacosane (nC29) and n-hentriacontane (nC31), while male-specific n-nonadecane (nC19) was absent. The ecological significance of these results for parasitoid host location behavior is discussed.

Plant surfaces of vegetable crops mediate interactions between chemical footprints of true bugs and their egg parasitoids

During the host location process, egg parasitoids can eavesdrop on chemical cues released from immature and adult hosts. These indirect host-related cues are highly detectable, but of low reliability because they lead egg parasitoid females to an area where oviposition is likely to occur rather then providing wasps with direct information on the presence of eggs and their location. In the host-parasitoid associations between true bugs and their scelionid egg parasitoids, female wasps perceive the chemical residues left by host adults walking on substrates as contact kairomones, displaying a characteristic arrestment posture. In this study, we demonstrated that epicuticular waxes of leaves of two vegetable crops, broad bean, Vicia faba, and collard greens, Brassica oleracea, mediate the foraging behavior of Trissolcus basalis (Wollaston) by adsorbing contact kairomones from adults of Nezara viridula (L.). Trissolcus basalis females showed no response when released on the adaxial leaf surface of broad bean or collard green plants with intact cuticular wax layers that had not been exposed to bugs, whereas wasps displayed the arrestment posture when intact leaves were contaminated by chemical residues from host females. Adaxial leaf surfaces that were dewaxed with an aqueous solution of gum arabic and afterwards contaminated by N. viridula females elicited no arrestment responses from wasp females. Similarly, leaves contaminated by host females and subsequently dewaxed did not elicit responses from female wasps. These findings reveal the important role of plant waxes in N. viridula - T. basalis semiochemical communication.

The response of Trissolcus basalis to footprint contact kairomones from Nezara viridula females is mediated by leaf epicuticular waxes

Chemical footprints left behind by true bugs are perceived as contact kairomones by scelionid egg parasitoids. Female wasps encountering a contaminated artificial substrate display a characteristic arrestment posture, holding the body motionless and antennating the surface. In the system Nezara viridula (L.) and its egg parasitoid Trissolcus basalis (Wollaston), previous studies have shown that the kairomone mediating such behavior is part of N. viridula's cuticular hydrocarbons (CHC) and furthermore that the wasp's ability to discriminate host male and female footprints is mainly based on the presence/absence of nonadecane (nC(19)). In this study, the effect of epicuticular waxes of leaves of broad bean, Vicia faba, on wasp responses to footprints of N. viridula females were investigated. Approximately 20% of T. basalis females displayed an arrestment posture when released on the adaxial leaf surfaces of broad bean plants with intact wax layer and without host chemical contamination; whereas approximately 70% of wasps displayed the arrestment posture when intact leaves were contaminated by host female footprints. Adaxial leaf surfaces of broad bean plants dewaxed with an aqueous solution of gum arabic and afterwards contaminated by N. viridula females induced arrestment responses in about 10% of female wasps; the same percentage of arrestment (10%) was observed when the wasps were released on leaves contaminated by host females and subsequently dewaxed. The side of the polymer film that was appressed to the leaf surface, peeled from the contaminated leaves, induced an arrestment posture in about 95% of observed wasps. Scanning electron microscopy (SEM) revealed that the epicuticular waxes occurred as a film densely crystallized as irregularly shaped platelets with spherical granules randomly distributed. These findings demonstrated that epicuticular waxes of broad bean leaves can mediate the foraging behavior of T. basalis females by absorbing contact kairomones of the host.

Differential attraction of parasitoids in relation to specificity of kairomones from herbivores and their by‐products

AbstractInfochemicals are used by foraging parasitoids in the host selection process from habitat preference until host recognition. Kairomones from the herbivore host plays a vital role in the attraction of parasitoids, particularly in the micro‐habitat. Parasitoids are specifically attracted to their respective herbivore species even when different herbivores are present on the same plant. Chemicals emitted from different stages of host (eggs, larvae, pupae, adult), host by‐products (e.g., frass, exuviae, mandibular gland secretions, defense secretions etc.), or intra‐specific infochemicals (pheromones) can be main signals for the parasitoids. Parasitoids can differentiate between host and non‐host, between different hosts and host stages by perceiving specific volatile and contact kairomones from the host itself, host along with its by‐product, by‐products alone or intra‐specific infochemicals; of which frass (by‐product) and intra‐specific infochemicals are the most reported ones. Adult and larval parasitoids have been reported to be attracted to kairomones of their target stage or byproduct of their host. Pupal parasitoids have been found to utilize kairomones from the preceding host stage while egg parasitoids are known to exploit a variety of host infochemicals, for example, either from eggs themselves or other non‐target host stages, especially adults and adult‐related by‐products. The kairomonal chemicals identified so far include various groups, but mainly hydrocarbons. A high degree of host specificity and host acceptance is important for the parasitoids as any mistake may result in the loss of fitness.

Effect of Bemisia tabaci honeydew and its carbohydrates on search time and parasitization of Encarsia bimaculata

Abstract: The search time of Encarsia bimaculata Heraty et Polaszek was examined and recorded by using filter paper discs in Petri dish arenas impregnated with Bemisia tabaci (Gennadius) honeydew or its component carbohydrates. Our data show that E. bimaculata used honeydew as a contact kairomone to locate its host. Females responded quantitatively to the honeydew and the search time in the treated patches increased. This increase was concentration‐dependent with regard to the amount of honeydew applied, but it levelled off 0.05 g/ml onwards. The response of E. bimaculata to honeydew decreased with increasing time after application and was significantly reduced 8 days after treatment. The parasitoid females were also arrested in patches treated with glucose, fructose, trehalulose and trehalose; trehalose elicited the highest response in the parasitoids. However, sucrose and low concentrations of melezitose did not elicit a significant effect. Glucose, sucrose, fructose, trehalose and trehalulose increased the longevity of the parasitoid females by a factor of 8.4, 8.1, 6.3, 6.1 and 4.2, respectively. Melezitose and 0.15 g/ml natural honeydew, however, had no effect on parasitoid lifespan. The effects of aqueous spray of honeydew sugars on egg to adult survivorship and parasitization of B. tabaci by E. bimaculata significantly differed when compared with controls only in cases where mixtures of glucose + fructose and trehalose and fructose + trehalose were applied.

Rhythm of honeydew excretion by the tea aphid and its attraction to various natural enemies

The rhythm of honeydew excretion by the tea aphid Toxoptera aurantii (Boyer) and its attraction to following 9 species (or subspecies) of beneficial insects, Aphidius sp., Chrysopa sinica Tjeder, Chrysopa septempunctata Wesmael, Sphaerophoria menthastri L., Coccinella septempunctata L., Leis axyridis (Pallas) ab. bimaculata Hemmelmann, L. axyridis (Pallas) ab. conspicua Faldermaenn, L. axyridis (Pallas) var. spectabilis Faldermaenn and L. axyridis (Pallas) var. novemdecimpunctata Faldermaenn, were investigated. Forty-five wingless virginoparae nymphs, reproduced by the same wingless virginogenia adult within 1 h, were introduced onto tea seedlings with one aphid per seedling. The honeydews excreted from different instars of nymphs and adults were collected under 21°C, 85% RH, 3500 lx and 12 L:12 D photoperiod. It took 32.4 d ±5.8 d for the tested tea aphids to complete the development of their nymph and adult stages, during which 325.6±35.8 droplets (ca. 41.98 μl ±6.14 μl and 45.34 mg ± 8.76 mg) of honeydews were secreted. During the 1st and 4th instars, there was a logistic regression relationship between the amount of honeydews excreted and the time (days). The honeydew secretion during the first two instars was less than that from the later instars. Adult aphids survived for 22.0 d ± 0.0 d, and excreted 176.31 ± 22.38 droplets (ca. 30.38 μl ± 5.32 μl) of honeydews in a rate of 1 drop per ca. 30–50 min for 5–8 h with a pause for 2–5 h before next secretion series. A batch of forty-five virginoparae female adults, reproduced by the same virginoparae female adult within 1 h, were introduced onto the tea seedlings (one aphid/seedling) under 13–21°, 85% RH, 3500 lx and 12 L:12 D photoperiod. Temperature showed a significant effect on the amounts of honeydews excreted within the range of 13–21°. Honeydews excreted by the aphids significantly increased the searching and retention time of the tested 9 species of natural enemies in a positive dose-response fashion. The searching times of Aphidius sp. and S. menthastri were the longest and the shortest, respectively, among all the 9 species, while the searching and retention time of L. axyridis (Pallas) var. spectabilis was the longest among the four varieties of L. axyridis. Tea aphid oneydew is considered as an important contact kairomone for the tested natural enemies.

The use of honeydew in foraging for aphids by larvae of the ladybird beetle, Coccinella septempunctata L. (Coleoptera: Coccinellidae)

Abstract 1. To clarify the use of honeydew in foraging for aphids by larvae of the ladybird beetle, Coccinella septempunctata L., searching behaviour of ladybird larvae for Aphis craccivora Koch and Acyrthosiphon pisum Harris and the abundance of honeydew under aphid colonies were examined in laboratory experiments.2. More larvae responded by climbing the plants with aphids than responded to plants without aphids. When the plants were replaced with sticks, in order to exclude visual and olfactory cues from plants and aphids, more larvae of C. septempunctata climbed sticks above the area that contained honeydew than climbed sticks above the area that did not contain honeydew. Then, ladybird larvae use honeydew as a contact kairomone when foraging for aphids.3. Aphis craccivora deposited a larger number of honeydew droplets beneath the plants than did similar numbers of A. pisum. Thus, C. septempunctata larvae licked more frequently the honeydew of A. craccivora than that of A. pisum and spent longer searching on the area containing honeydew of A. craccivora than that of A. pisum. Consequently, a larger number of larvae climbed a stick above honeydew of A. craccivora than that of A. pisum.4. It may be also considered that C. septempunctata larvae can distinguish honeydew of the two aphid species and respond more strongly to A. craccivora than A. pisum.

The Egg Parasitoid Trissolcus basalis uses n-nonadecane, a Cuticular Hydrocarbon from its Stink Bug Host Nezara viridula, to Discriminate Between Female and Male Hosts

Contact kairomones from adult southern green stink bugs, Nezara viridula (L.) (Heteroptera: Pentatomidae) that elicit foraging behavior of the egg parasitoid Trissolcus basalis (Wollaston) were investigated in laboratory experiments. Chemical residues from tarsi and scutella of N. viridula induced foraging by gravid female T. basalis. Residues from body parts of female N. viridula elicited stronger responses than those from the corresponding body parts of males. Deproteinized tarsi still elicited searching responses from wasps, indicating that the kairomone was not proteinaceous. Hexane extracts of host cuticular lipids induced searching responses from T. basalis, with a strong preference for extracts from female hosts. Extracts consisted primarily of linear alkanes from nC19 to nC34, with quantitative and qualitative differences between the sexes. Extracts of female N. viridula contained more nC23, nC24, and nC25 than the corresponding extracts from males, whereas nC19 was detected only in extracts from males. Direct-contact solid phase microextraction (DC-SPME) of N. viridula cuticle and of residues left by adult bugs walking on a glass plate confirmed gender-specific differences in nC19. Trissolcus basalis females responded weakly to a reconstructed blend of the straight-chain hydrocarbons, suggesting that minor components other than linear alkanes must be part of the kairomone. Addition of nC19 to hexane extracts of female N. viridula significantly reduced the wasps' arrestment responses, similar to wasps' responses to hexane extracts of male hosts. Overall, our results suggest that a contact kairomone that elicits foraging by T. basalis females is present in the cuticular lipids of N. viridula, and that the presence or absence of nC19 allows T. basalis females to distinguish between residues left by male or female hosts. The ecological significance of these results in the host location behavior of scelionid egg parasitoids is discussed.

Prey-specific contact kairomones exploited by adult and larval Teretrius nigrescens: A behavioural comparison across different stored-product pests and different pest substrates

Prostephanus truncatus is a serious exotic beetle pest of stored maize and cassava in Africa. Teretrius nigrescens is a classical biological control predator of this pest that was released into Africa in 1991. In previous work it was shown that adult T. nigrescens are arrested in the presence of dust and frass produced by P. truncatus feeding on maize, and both adult and larval T. nigrescens are arrested by solvent extracts of this dust/frass when presented on filter paper. The current study used these two complementary bioassay approaches to test crude dust/frass and then solvent extracts of the same materials. Results demonstrate that T. nigrescens adults are arrested more strongly by the dust/frass of P. truncatus than the dust/frass of six other beetle pests of stored grain. Similar behaviours are observed for responses of both adults and larvae to solvent extracts. However, extract of dust/frass from Sitophilus zeamais is repellent to adults and larvae of T. nigrescens. Collections of dust/frass from P. truncatus cultured on maize, cassava and an artificial maize substrate previously stripped of all hexane extractable compounds, are all shown to arrest adult T. nigrescens when presented in their crude form and to arrest both adults and larvae when presented as solvent extracts. These experiments demonstrate that P. truncatus produces a species-specific kairomone that is independent of the food or tunnelling substrate. Findings are discussed in the context of prey location, particularly in natural habitats.

Inter-specific variation in the reactive distance of different aphid-parasitoid associations: analysis from automatic tracking of the walking path

The distance from which hosts are perceived by foraging parasitoid females has been repeatedly considered to be an important behavioural trait that is supposed to be related to their efficiency as biological control agents. In the present study, such a ‘reactive distance’ was quantified using the automatic analysis of video-recorded females' walking paths. Inter-specific variation of this trait was analysed by comparison between Aphis gossypii Glover, Myzus persicae Sulzer, and Sitobion avenae Fabricius, and three of their parasitoid species known to present different levels of specificity to these hosts: Aphelinus abdominalis Dalman, Aphidius colemani Viereck, and Aphidius picipes Nees. Responses to two different host instars of different size were also compared in each combination. On average, parasitoid females were able to perceive aphids from a distance of about 3 to 4 mm. The only significant difference observed was between the Aphelinus and the two Aphidius parasitoid species. No difference was detected between the different host instars or species tested. Females seem to be unable to discriminate between host and non-host species from a distance. These results seem to confirm the hypothesis of the use of a cuticular contact kairomone for host recognition at the species level.