Plant—Phytoseiid Interactions Mediated by Herbivore-Induced Plant Volatiles: Variation in Production of Cues and in Responses of Predatory Mites

Phytoseiid mites use herbivore-induced plant volatiles in long-range prey-habitat location and are arrested by these volatiles in a prey patch. The responses of predatory mites to these volatiles are considered to be an important factor in the local extermination of prey populations by phytoseiids such as Phytoseiulus persimilis. Prey-induced plant volatiles are highly detectable and can be reliable indicators of prey presence and prey identity. The composition of herbivore-induced plant volatiles depends on plant species and plant cultivar. Moreover, the composition may also vary with the herbivore species that infests a plant. The responses of phytoseiids to prey-induced plant volatiles from a specific plant-herbivore combination are highly variable. Causal factors include starvation, specific hunger, experience, pathogen infestation and the presence of competitors. Investigating variation in the phytoseiid's behavioural response in relation to these factors is important for understanding how and why behavioural strategies maximize phytoseiid fitness.     

Host plants of the herbivorous insect Mythimna separata (Lepidoptera: Noctuidae) affect its susceptibility to parasitism by the larval parasitoid Cotesia kariyai (Hymenoptera: Braconidae)

Previous studies have shown that the food plant species consumed by herbivorous insects affect their parasitism by parasitoid wasps or flies, but this phenomenon has only been observed in a limited number of systems. Here, we investigate how feeding on different plant species affects the survival of the rice armyworm Mythimna separata (Walker) (Lepidoptera: Noctuidae) and its successful parasitism by the larval endoparasitoid Cotesia kariyai (Watanabe) (Hymenoptera: Braconidae). Survival rate and pupal weight of unparasitised M. separata fed with daikon Raphanus sativus L. var. longipinnatus was lower than with maize Zea mays L. In addition, caterpillars fed with daikon were less susceptible to parasitism by C. kariyai than those fed with maize, and parasitised M. separata larvae fed with daikon had higher survival rates than those fed with maize. These results indicate that while daikon is not an optimal food for unparasitised caterpillars, it may protect the larvae from death by C. kariyai parasitism.     

Short Distance Cues Used by the Adult Parasitoid <Emphasis Type="Italic">Microctonus hyperodae</Emphasis> Loan (Hymenoptera: Braconidae,Euphorinae) for Host Selection of a Novel Host <Emphasis Type="Italic">Listronotus oregonensis</Emphasis> (Coleoptera: Curculionidae)

Insect parasitoids use host kairomone to detect their hosts. However, in parasitoid species that attack adult hosts, the mobility of adult insect may mean that the host can move away for kairomone sources. The effect of Listronotus oregonensis LeConte (Coleoptera: Curculionidae) adult sex, feces and movement on host selection behavior by Microctonus’’ hyperodae Loan (Hymenoptera: Braconidae; Euphorinae) females was evaluated in the laboratory. We hypothesized that, in addition of using host kairomones, parasitoids of adult stage should use host movement for host selection. The sex of L. oregonensis did not affect the host selection behavior of M. hyperodae. However, host feces decreased the number of weevil antennations done by M. hyperodae. Microctonus hyperodae stopped less frequently near immobile L. oregonensis than near walking ones and these latter were frequently pursued by M. hyperodae. Host movement was the stimulus that elicited oviposition by M. hyperodae. The adaptive implications of these results are discussed.     

Behavioural and community ecology of plants that cry for help

Plants respond to insect herbivory with the production of volatiles that attract carnivorous enemies of the herbivores, a phenomenon called indirect defence or 'plants crying for help'. Plants are under selection to maximize Darwinian fitness, and this can be done by making the right 'decisions' (i.e. by responding to environmental stress in ways that maximize seed production). Plant decisions related to the response to herbivory in terms of the emission of herbivore-induced volatiles include 'to respond or not to respond', 'how fast to respond', 'how to respond' and 'when to stop responding'. In this review, the state-of-the-art of the research field is presented in the context of these decisions that plants face. New questions and directions for future research are identified. To understand the consequences of plant responses in a community context, it is important to expand research from individual interactions to multispecies interactions in a community context. To achieve this, detailed information on underlying mechanisms is essential and first steps on this road have been made. This selective review addresses the ecology of herbivore-induced plant volatiles (HIPVs) by integrating information on mechanisms and ecological functions. New questions are identified as well as challenges for extending current information to community ecology.     

Diachasmimorpha longicaudata and D. kraussii (Hymenoptera: Braconidae), potential parasitoids of the olive fruit fly

The olive fruit fly, Bactrocera oleae (Tephritidae), is a significant threat to California's olive industry. As part of a classical biological control program started in 2002, the parasitoids Diachasmimorpha kraussii and D. longicaudata (Hymenoptera: Braconidae) were imported to California from laboratory colonies in Hawaii. Studies on their biology and behavior as parasitoids of the olive fruit fly were conducted in quarantine. Both species tend to oviposit into 2nd and young 3rd instars, with the offspring completing development in the flies’ puparia. Most eggs are deposited in the first two weeks of adult life. Observed lifetime fecundity was low, possibly as a consequence of the relatively poor quality of the harvested olives used as a host substrate. Both pre-imaginal development and adult longevity were limited at constant temperatures above 30°C, which may indicate that these species will have difficulty establishing in the warmest regions of California.     

Host-age discrimination during host location byCotesia glomerata, a larval parasitoid ofPieris brassicae

Some parasitoids are restricted with respect to the host stage that they attack and even to a certain age within a stage. In this paper we investigate whether the parasitoidCotesia glomerata can discriminate between old and young caterpillar instars of its host,Pieris brassicae, before contacting these hosts, since contacts with older instars are very risky with a chance of being killed, due to the aggressive defensive behaviour of the caterpillars. Flight chamber dual choice tests showed that volatile chemicals emitted by Brussels sprouts plants (Brassica oleracea var. gemmifera) after feeding damage by 1st and 5th larval instars are equally attractive to the wasps. Simulated herbivore damage by 2nd and 5th larval instars, obtained by treating mechanically damaged leaves with carterpillar regurgitant, was also equally attractive, even when the wasps were exposed to repeated experience on different larval instars to increase their discriminatory ability. In contrast, single choice contact bioassays showed that the time spent searching on a leaf with feeding damage of 1st instar larvae was significantly longer than the time spent on 5th instar feeding damage or on mechanically damaged leaves. Both flight and contact bioassays did not show any effect of egg-related infochemicals. The results demonstrate thatC. glomerata can discriminate between young and old larval instars ofP. brassicae, without contacting the caterpillars. This is not done through volatile herbivore-induced synomones but through cues that are contacted after arrival at a caterpillar-infested leaf.     

The relative importance of resources and natural enemies in determining herbivore abundance: thistles, tephritids and parasitoids

1. The relative importance of host-plant resources and natural enemies in influencing the abundance of insect herbivores was investigated in potted plant and natural population experiments, using tephritid (Diptera: Tephritidae) flies, their host plant, creeping thistle Cirsium arvense, and their Hymenoptera parasitoids. 2. Experimental manipulation of host-plant quality (i.e. levels of host-plant nutrients) and resource availability (i.e. the number of buds) increased tephritid abundance. There was no evidence that the seed-feeding tephritid fly Xyphosia miliaria preferentially oviposited on fertilized C. arvense. 3. At low thistle densities, X. miliaria showed a constant rate of resource exploitation. At higher thistle densities, a threshold was detected, above which additional buds were not attacked. 4. Parasitism attack was variable across host (tephritid) densities but levels of parasitism were consistently higher on the fertilized thistles. 5. Experimental manipulation of host-plant quality and resource availability (quantity) not only directly affects the tephritid population but also, indirectly, leads to high rates of parasitism. Both chemical and physical characteristics of host plants affect the performance of natural enemies. 6. Both top-down and bottom-up forces act to influence tephritid abundance, with bottom-up influences appearing to be the most important.     

Olfactory response of Anagrus,nigriventrls (HYM.: Mymaridae): Effects of host plant chemical cues mediated by rearing and oviposition experience

The egg parasitoid Anagrus nigriventris Girault is an important natural enemy of the beet leafhopper, Circulifer tenellus (Baker) which has a broad host range that includes cultivated sugar beets and the weed, Russian thistle. When parasitoids were reared on leafhopper eggs deposited in sugar beets, females were attracted to sugar beet volatiles and preferred them over a blank control and over Russian thistle volatiles in Y- tube olfactometer tests. No preference was detected for either plant by female wasps that were reared on Russian thistle. Wasps reared on Russian thistle and allowed an oviposition experience on sugar beet showed a strong attraction to sugar beet volatiles; however, wasps reared on sugar beets and allowed oviposition experience on Russian thistle demonstrated no preference between the two plants. The implications of parasitoid response towards plant volatiles and their importance in biological control are discussed.     

Evaluation of Fopius arisanus as a biological control agent for the olive fruit fly in California

1 The egg‐prepupal parasitoid Fopius arisanus (Hymenoptera: Braconidae) was evaluated in quarantine facilities as a potential biological control agent for the olive fruit fly Bactrocera oleae (Diptera: Tephritidae) in California, U.S.A. 2 Nontarget testing of two weed biological control agents confirmed that F. arisanus will not attack Tephritidae that feed in inflorescences or galls. It may, however, pose risks to native Tephritidae that feed in fruit. 3 Females preferentially oviposited in eggs, although first‐instar B. oleae were also attacked. Low lifetime reproductive potential and high rates of direct mortality inflicted on host eggs indicate that rearing on B. oleae may prove difficult. 4 In multiparasitized B. oleae, F. arisanus prevailed in competition against two species of larval–pupal parasitoids, Diachasmimorpha kraussii and Psyttalia concolor (both Hymenoptera: Braconidae). 5 The broad host‐range of F. arisanus with respect to fruit‐feeding Tephritidae may preclude its introduction to California, as may its low fecundity and its intrinsic competitive superiority over larva l–pupal parasitoids, which include specialists on B. oleae that are currently being introduced to California. High rates of direct mortality, however, point to potential uses in augmentative biological control. Whether or not F. arisanus is released in California, its biology as a parasitoid of B. oleae has been little studied to date and the results herein may be applied in other regions worldwide where B. oleae is a problem.     

Effect of Diachasmimorpha tryoni on two non-target flowerhead-feedingtephritids

Laboratory tests were conducted to evaluate the possible effect of a deliberately introduced fruit fly parasitoid, Diachasmimorpha tryoni, on 2 non-target flowerhead-feeding tephritid flies,Trupanea dubautiae and Ensina sonchi. The former is an nativecomposite endemic Hawaiian tephritid which feeds on flowerheads of the native composite shrub, Dubautia raillar dioides; the latter is an inadvertently introduced tephritid infesting flowerheads of the exotic weed, Sonchus oleraceus. Gravid females of D. tryoniwere confined in test cages with field-collected D.raillardioides and S. oleraceus flowerheads infested with late instars of T. dubautiae and E. sonchi,respectively. D. tryoni showed low levels of visiting and probing responses to D. raillardioides flowerheads and relatively higher responses to S. oleraceus flower heads in both the presence and absence of the parasitoid's normal host, the Mediterranean fruit fly, Ceratitis capitata. With 72-h exposure to D. tryoni, 13.9 and 2.6% of T. dubautiaewere attacked by test parasitoids in the absence and presence of C.capitata in the test cage, respectively; while 56.8 and34.2% of E. sonchi were attacked. In contrast, 94.0 and84.0% of C. capitata larvae presented in screened disheswith diet in test cages were attacked by D. tryoni in testswith T. dubautiae and E. sonchi, respectively. Attackof D. tryoni on flowerhead-feeding T. dubautiae andE. sonchi resulted in significant reduction in the emergence ofadult flies, especially in the absence of the parasitoid's normal host.While 8.8–12.8 adult D. tryoni per test (both males andfemales) successfully emerged from medflies, no adult D. tryoniprogeny emerged from T. dubautiae, and only 3 deformed males ofD. tryoni (much smaller than the normal wasp) emerged fromE. sonchi. The relevance of these findings to the safety offuture biological control programs against tephritid pests is discussed. This revised version was published online in July 2006 with corrections to the Cover Date.