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Abstract A large body of evidence shows that plants release volatile chemicals upon attack by herbivores. These volatiles influence the performance of natural enemies. Nearly all the evidence on the effect of plant volatiles on natural enemies of herbivores concerns predators, parasitoids, and entomophagous nematodes. However, other entomopathogens, such as fungi, have not been studied yet for the way they exploit the chemical information that the plant conveys on the presence of herbivores. We tested the hypothesis that volatiles emanating from cassava plants infested by green mites (Mononychellus tanajoa) trigger sporulation in three isolates of the acaropathogenic fungus Neozygites tanajoae. Tests were conducted under climatic conditions optimal to fungal conidiation, such that the influence of the plant volatiles could only alter the quantity of conidia produced. For two isolates (Altal.brz and Colal.brz), it was found that, compared with clean air, the presence of volatiles from clean, excised leaf discs suppressed conidia production. This suppressive effect disappeared in the presence of herbivore-damaged leaves for the isolate Colal.brz. For the third isolate, no significant effects were observed. Another experiment differing mainly in the amount of volatiles showed that two isolates produced more conidia when exposed to herbivore-damaged leaves compared with clean air. Taken together, the results show that volatiles from clean plants suppress conidiation, whereas herbivore-induced plant volatiles promote conidiation of N. tanajoae. These opposing effects suggest that the entomopathogenic fungus tunes the release of spores to herbivore-induced plant signals indicating the presence of hosts.

The most common post–zygotic isolation mechanism between populations of the phytophagous mite Tetranychus urticaeis ‘hybrid breakdown’, i.e. when individuals from two different populations are crossed, F1hybrid females are produced, but F2recombinant male offspring suffer increased mortality. Two spotted spider mites collected from two populations, one on rose and the other on cucumber plants, were infected with Wolbachiabacteria. These bacteria may induce cytoplasmic incompatibility in their hosts: uninfected (U) females become reproductively incompatible with infected (W) males. We report on the effect of Wolbachiainfections in intra–and interstrain crosses on (i) F1mortality and sex ratios (a test for cytoplasmic incompatibility), and (ii) the number of haploid offspring and mortality in clutches of F1virgins (a test for hybrid breakdown). U x W crosses within the rose strain exhibited partial cytoplasmic incompatibility. More interestingly, F2males suffered increased mortality, a result identical to the hybrid breakdown phenomenon. The experiments were repeated using females from the cucumber strain. In interstrain U x W and U x U crosses, hybrid breakdown was much stronger in the former (80 versus 26).This is the first report of a Wolbachiainfection causing a hybrid breakdown phenotype. Our results show that Wolbachiainfections can contribute to reproductive incompatibility between populations of T.urticae.

Arthropods use odours associated with the presence of their food, enemies and competitors when searching for patches. Responses to these odours therefore determine the spatial distribution of animals, and are decisive for the occurrence and strength of interactions among species. Therefore, a logical first step in studying food web interactions is the analysis of behaviour of individuals that are searching for patches of food. We followed this approach when studying interactions in an artificial food web occurring on greenhouse cucumber in the Netherlands. In an earlier paper we found that one of the predators of the food web, the predatory mite Phytoseiulus persimilisAthias‐Henriot, used to control spider mites, discriminates between odours from plants with spider mites, Tetranychus urticaeKoch, and plants with spider mites plus conspecific predators. The odours used for discrimination are produced by adult prey in response to the presence of predators, and probably serve as an alarm pheromone to warn related spider mites. Other predator species may also trigger production of this alarm pheromone, which P. persimiliscould use in turn to avoid plants with heterospecific predators. We therefore studied the response of the latter to odours from plants with spider mites and 3 other predator species, i.e. the generalist predatory bug Orius laevigatus(Fieber), the polyphagous thrips Frankliniella occidentalisand the spider‐mite predator Neoseiulus californicus(McGregor). Both olfactometer and greenhouse release experiments yielded no evidence that P. persimilisavoids plants with any of the 3 heterospecific predators. This suggests that these predators do not elicit production of alarm pheromones in spider mites, and we argue that this is caused by a lack of coevolutionary history. The consequences of the lack of avoidance of heterospecific predators for interactions in food webs and biological control are discussed.

The occurrence and strength of interactions among natural enemies and herbivores depend on their foraging decisions, and several of these decisions are based on odours. To investigate interactions among arthropods in a greenhouse cropping system, we studied the behavioural response of the predatory bug Orius laevigatus(Fieber) (Hemiptera: Anthocoridae) towards cucumber plants infested either with thrips (Frankliniella occidentalis(Pergande) (Thysanoptera: Thripidae)) or with spider mites (Tetranychus urticaeKoch (Acari: Tetranychidae)). In greenhouse release‐recapture experiments, the predatory bug showed a significant preference for both thrips‐infested plants and spider mite‐infested plants over clean plants. Predatory bugs preferred plants infested with spider mites to plants with thrips. Experience with spider mites on cucumber leaves prior to their release in the greenhouse had no effect on the preference of the predatory bugs. However, this experience did increase the percentage of predators recaptured. Y‐tube olfactometer experiments showed that O. laevigatuswas more attracted to odours from plants infested with spider mites than to odours from clean plants. Thus, O. laevigatusis able to perceive odours and may use them to find plants with prey in more natural conditions. The consequences of the searching behaviour for pest control are discussed.

To understand the role of allelochemicals in predator-prey interactions it is not sufficient to study the behavioral responses of predator and prey. One should elucidate the origin of the allelochemicals and be aware that it may be located at another trophic level. These aspects are reviewed for predator-prey interactions in general and illustrated in detail for interactions between predatory mites and herbivorous mites. In the latter system there is behavioral and chemical evidence for the involvement of the host plant in production of volatile allelochemicals upon damage by the herbivores with the consequence of attracting predators. These volatiles not only influence predator behavior, but also prey behavior and even the attractiveness of nearby plants to predators. Herbivorous mites disperse away from places with high concentrations of the volatiles, and undamaged plants attract more predators when previously exposed to volatiles from infested conspecific plants rather than from uninfested plants. The latter phenomenon may well be an example of plant-to-plant communication. The involvement of the host plant is probably not unique to the predator-herbivore-plant system under study. It may well be widespread since it makes sense from an evolutionary point of view. If so, prospects for application in pest control are wide open. These are discussed, and it is concluded that crop protection in the future should include tactics whereby man becomes an ally to plants in their strategies to manipulate predator-prey interactions through allelochemicals.

Abstract: Plants under herbivore attack produce volatiles, thus attracting natural enemies of the herbivores. However, in doing so, the plant becomes more conspicuous to other herbivores. Herbivores may use the odours as a cue to refrain from visiting plants that are already infested, thereby avoiding competition for food, or, alternatively, to visit plants with defences weakened by earlier attacks. We investigated the response of one species of herbivore (the spider mite Tetranychus urticae) to odours emanating from cucumber plants infested by conspecific or heterospecific (the western flower thrips, Frankliniella occidentalis) herbivores. Olfactometer experiments in the laboratory showed that spider mites have a slight, but significant, preference for plants infested with conspecifics, but strongly avoid plants with thrips. These results were substantiated with greenhouse experiments. We released spider mites on the soil in the centre of a circle of six plants, half of which were infested with either conspecifics or heterospecifics (thrips), whereas the other half were uninfested. It was found that 60–70% of the mites were recaptured on the plants within 5 h after release. Results of these experiments were in agreement with results of the olfactometer experiments: (1) significantly fewer spider mites were found on plants infested with thrips than on uninfested plants and (2) more mites were found on plants with conspecifics than on clean plants (although this difference was not significant). From a functional point of view it makes sense that spider mites prefer clean plants over thrips-infested plants, since thrips are not only competitors, but are also known as intraguild predators of spider mites. Possible reasons for the slight attraction of spider mites to plants infested with conspecifics are discussed.

In his mathematical treatment of Fisher's ideas on sexual selection (so-called runaway selection) Lande (1981) predicted that males may evolve increasingly elaborate sexual characters despite opposing viability selection as a consequence of the associated costs.Lande thereby assumed that female mate preferences are not subject to selection since (1) females are all inseminated and (2) the quantity and quality of their offspring are independent of the female's mate preferences. Kirkpatrick (1985) removed the latter assumption and investigated the consequences for the mean phenotype with respect to both female and male traits. He also explored the dynamics of the (co)-variance matrix by numerical methods.In this paper we consider a simpler model with just two multi-allelic loci. This enables us to derive explicit expressions for (co)-variances under steady state conditions. Rather than assume natural selection through differential fertility (as in Kirkpatrick, 1985), we take sexual selection on females into account by modelling the preference-dependent risk that females remain unmated.We argue that this wallflower effect is a realistic feature of any mating system, since it merely depends on the existence of (1) variation in mating preferences and (2) a finite mating season. Our approach provided an insight into the dynamic behaviour of the means of the phenotypes. This is because the dynamics of the means depend on the steady state (co)-variance matrix. Thus, an insight into the former requires explicit expressions for the latter.Whereas Lande and Kirkpatrick predicted runaway processes, despiteopposing viability selection, our model predicts a globally stable steady state, i.e. no runaway, even withoutopposing viability selection (under the assumption of an asymptotically stable steady state of the (co)-variances. Admittedly, we have no analytic proof of this stability but only support for it, based on simulations.) The absence of the runaway processes in our model is caused by the wallflower effect, since it imposes constraints on the steady state of the (co)-variance matrix.When mutational input applies to female traits but not to male traits, explicit expressions for the (co)-variances under steady state conditions can be derived, and these show that: (1) both the genetic covariance and the variance of male traits are equal to zero, but (2) the variance of the female trait exceeds to zero. Should there be mutational input influencing the male trait, then these results would suggest that the male-to-female ratio of variances is much smaller than unity. This prediction is of tremendous importance for speciation through founding events.

Prey-selection behaviour of the phytoseiid mite Typhlodromus pyri Scheuten was analysed with a Markovtype model of feeding-state dynamics and feeding-state dependent searching behaviour (Sabelis 1981, 1986, 1989; Metz and Van Batenburg 1985a, b). All behavioural characteristics of the predator which are independent of the feeding state were represented by one parameter. The remaining feeding-state dependent characteristics were represented by a function of the feeding state, with one parameter. The best parameter values to describe a predator-prey interaction were determined by fitting the model to the predation rates in monocultures. Under the assumption that the parameter values are not dependent on the composition of prey species supply, the diet of the predators in mixed cultures was predicted from parameters estimated in monoculture experiments.

Prey preference of three phytoseiid species,Typhlodromus pyriScheuten,Amblyseius potentillae(Garman) andA. finlandicus(Oudemans) which occur in Dutch orchards, was analysed with respect to two economically important phytophagous mites, the European red spider mitePanonychus ulmi(Koch), and the apple rust miteAculus schlechtendali(Nalepa). Two types of laboratory experiments were carried out: (1) olfactometer tests to study the response when volatile kairomones of both prey species were offered simultaneously; and (2) predation tests in mixtures of the two prey species and comparison with calculated predation rates, using a model provided with parameters estimated from experiments with each prey species alone. In addition, the diet of field-collected predators was analysed using electrophoresis.

Apart from interacting, prey and predators may also avoid each other by moving into refuges where they lack food, yet survive by switching to an energy-saving physiological state. Lotka–Volterra models of predator–prey interactions ignore this option. Therefore, we have modelled this game of ‘joining versus opting out’ by extending Lotka–Volterra models to include portions of populations not in interaction and with different energy dynamics. Given this setting, the prey's decisions to join or to opt out influence those of the predator and vice versa, causing the set of possible strategies to be complex and large. However, using game theory, we analysed and published two models showing (i) which strategies are best for the prey population given the predator's strategy, and (ii) which are best for prey and predator populations simultaneously. The predicted best strategies appear to match empirical observations on plant-inhabiting predator and prey mites. Here, we consider a plausible third model that does not take energy dynamics into account, but appears to yield contrasting predictions. This supports our assumption to extend Lotka–Volterra models with ‘interaction-dependent’ energy dynamics, but more work is required to prove that it is essential and that what is best for the population is also best for the individual.

Most studies on ecological impact of solar ultraviolet (UV) radiation generally focus on plants. However, UV radiation can also affect organisms at other trophic levels. Protection against mortality induced by solar UV has, therefore, been hypothesized as one of the reasons why Typhlodromalus aripo hides in the apex of cassava plants during the day and comes out at night to prey on spider mites on leaves. In laboratory experiments using UV lamps, we determined the impact of UVA and UVB radiation on survival and oviposition of two leaf-inhabiting mites (Amblydromalus manihoti, Euseius fustis) and the apex-inhabiting mite (T. aripo), all three species being predators used for controlling the cassava green mite Mononychellus tanajoa in Africa. Whereas on leaf discs UVA has no negative impact on survival of the three predators, UVB is lethal to all of them. In contrast, nearly 85% of T. aripo survived after exposure to UVB inside apex of cassava plants. Exposure of A. manihoti and E. fustis to UVB radiation on the lower surface of a cassava leaf resulted in 36% survival. Oviposition and hatching of eggs laid after exposure to UVB were not affected, but eggs directly exposed to UVB did not hatch. Although caution should be exercised to extrapolate laboratory studies to the field, our results support the hypothesis that lower side of leaves, but especially plant apices, represent refuges that protect predatory mites from UVB. This might explain why T. aripo moves out of the apex to forage on leaves only during the night.