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Cucurbitaceae plants produce cucurbitacins, bitter triterpenoids, to protect themselves against various insects and pathogens. Adult banded cucumber beetles (Diabrotica balteata), a common pest of maize and cucurbits, sequester cucurbitacins, presumably as a defensive mechanism against their natural enemies, which might reduce the efficacy of biological control agents. Whether the larvae also sequester and are protected by cucurbitacins is unclear. We profiled cucurbitacin levels in four varieties of cucumber, Cucumis sativus, and in larvae fed on these varieties. Then, we evaluated larval growth and resistance against common biocontrol organisms including insect predators, entomopathogenic nematodes, fungi and bacteria. We found considerable qualitative and quantitative differences in the cucurbitacin levels of the four cucumber varieties. While two varieties were fully impaired in their production, the other two accumulated high levels of cucurbitacins. We also observed that D. balteata larvae sequester and metabolize cucurbitacins, and although the larvae fed extensively on both belowground and aboveground tissues, the sequestered cucurbitacins were mainly derived from belowground tissues. Cucurbitacins had no detrimental effects on larval performance and, surprisingly, did not provide protection against any of the natural enemies evaluated. Our results show that D. balteata larvae can indeed sequester and transform cucurbitacins, but sequestered cucurbitacins do not impact the biocontrol potential of common natural enemies used in biocontrol. Hence, this plant trait should be conserved in plant breeding programs, as it has been demonstrated in previous studies that it can provide protection against plant pathogens and generalist insects., Journal of Pest Science, 96 (3), ISSN:1612-4758, ISSN:1612-4766

Plants and herbivores have been engaged in a co-evolutionary arms race for millions of years, during which plants evolved various defenses and other traits to cope with herbivores, whereas herbivores evolved traits to overcome the plants' resistance strategies. Herbivores may also avoid certain plants merely because these lack suitable nutrients for their development. Interestingly, the number of herbivores that attack individual early land plants like mosses and ferns is quite low. Among others, poor nutrient quality has been hypothesized to explain the apparent low herbivory pressure on such plants but still waits for scientific evidences. Here, the nutritive suitability of representative mosses and liverworts (bryophytes) and ferns (pteridophytes) for herbivores was investigated using feeding assays combined with quantifications of nutrients (proteins, amino acids, and sugars). Growth and survival of two polyphagous herbivores, a caterpillar and a snail, were monitored when fed on 15 species of bryophytes and pteridophytes, as well as on maize (

Previous studies have frequently reported effects of abiotic factors on herbivore‐induced plant defences based on effects on single plant traits. However, plants commonly express multiple defences simultaneously and these traits are often correlated. Thus, a fuller understanding of abiotic‐context dependency in plant defence requires measuring multiple traits and addressing their patterns of correlated expression. We evaluated the effects of soil salinity on the expression of direct (phenolic compounds, gossypol gland density) and indirect (volatile organic compounds, extrafloral nectar) defensive traits in wild cotton Gossypium hirsutum. Specifically, we asked whether soil salinity affects the induction of these traits, and whether it shapes trait correlations potentially underlying altered patterns of trait induction. We conducted a factorial experiment with 16 cotton genotypes where we manipulated soil salinity and defence induction by applying artificial leaf damage (25% mechanical damage and caterpillar oral secretions) and measured defence levels at different time points post damage. Leaf damage induced most traits except gossypol gland density, whereas salinity did not have a mean effect (across constitutive and induced levels) on any of the measured traits. Nonetheless, salinity prevented the induction of phenolic compounds (condensed and hydrolysable tannins), and also affected trait correlations. Specifically, phenolic compounds were negatively associated with nectar production only under salinized conditions, an apparent trade‐off that could affect the induction of phenolic compounds. In addition, positive correlations between phenolic compounds and gland density and root biomass observed under control conditions were lost under salinized conditions. Synthesis. By investigating the effects of soil salinity on the expression of multiple direct and indirect defensive traits and their underlying correlations, these findings build towards a better understanding of how abiotic context dependency shapes plant allocation to and expression of multiple defensive traits.

Significance Plants respond to herbivory and pathogenic infection with the synthesis of various defense compounds, including volatile compounds that are emitted into the environment. These volatiles can be perceived by neighboring plants and “prime” them for an attack by the specific attacker. We found that whitefly-infested tomato plants release volatiles that prime a defense against pathogens in neighboring plants, which goes at the cost of defenses against insect herbivores, making the neighboring plants more suitable for whitefly development. This apparent ability of whiteflies to manipulate plant defense responses through induced volatile emissions could explain the rapid spread of this invasive pest, and a good understanding of the mechanisms that are involved may lead to novel pest control strategies., The whitefly Bemisia tabaci is one of the world’s most important invasive crop pests, possibly because it manipulates plant defense signaling. Upon infestation by whiteflies, plants mobilize salicylic acid (SA)-dependent defenses, which mainly target pathogens. In contrast, jasmonic acid (JA)-dependent defenses are gradually suppressed in whitefly-infested plants. The down-regulation of JA defenses make plants more susceptible to insects, including whiteflies. Here, we report that this host–plant manipulation extends to neighboring plants via airborne signals. Plants respond to insect attack with the release of a blend of inducible volatiles. Perception of these volatiles by neighboring plants usually primes them to prepare for an imminent attack. Here, however, we show that whitefly-induced tomato plant volatiles prime SA-dependent defenses and suppress JA-dependent defenses, thus rendering neighboring tomato plants more susceptible to whiteflies. Experiments with volatiles from caterpillar-damaged and pathogen-infected plants, as well as with synthetic volatiles, confirm that whiteflies modify the quality of neighboring plants for their offspring via whitefly-inducible plant volatiles.

It is increasingly evident that plants interact with their outside world through the production of volatile organic compounds,1-5 but whether the volatiles have evolved to serve in plant defense is still a topic of considerable debate.3,6-8 Unharmed leaves constitutively release small amounts of volatiles, but when the leaves are damaged by herbivorous arthropods, they emit substantially more volatiles. These herbivore-induced plant volatiles (HIPVs) attract parasitoids and predators that kill insect herbivores,9-12 and this can benefit the plants.13,14 As yet, however, there is no tangible evolutionary evidence that this tritrophic interplay contributes to the selection forces that have shaped the volatile emissions of plants.2,3,5-8,15 With this in mind, we investigated the evolutionary changes in volatile emissions in invasive common ragwort and the respective defensive roles of its constitutive and inducible volatiles. This Eurasian plant has invaded other continents, where it evolved for many generations in the absence of specialized herbivores and their natural enemies. We found that, compared to native ragworts, invasive plants release higher levels of constitutive volatiles but considerably lower levels of herbivore-induced volatiles. As a consequence, invasive ragwort is more attractive to a specialist moth but avoided by an unadapted generalist moth. Importantly, conforming to the indirect defense hypothesis, a specialist parasitoid was much more attracted to caterpillar-damaged native ragwort, which was reflected in higher parasitism rates in a field trial. The evolution of foliar volatile emissions appears to be indeed driven by their direct and indirect roles in defenses against insects.

Plants typically release large quantities of volatiles in response to herbivory by insects. This benefits the plants by, for instance, attracting the natural enemies of the herbivores. We show that the brown planthopper (BPH) has cleverly turned this around by exploiting herbivore-induced plant volatiles (HIPVs) that provide safe havens for its offspring. BPH females preferentially oviposit on rice plants already infested by the rice striped stem borer (SSB), which are avoided by the egg parasitoid Anagrus nilaparvatae, the most important natural enemy of BPH. Using synthetic versions of volatiles identified from plants infested by BPH and/or SSB, we demonstrate the role of HIPVs in these interactions. Moreover, greenhouse and field cage experiments confirm the adaptiveness of the BPH oviposition strategy, resulting in 80% lower parasitism rates of its eggs. Besides revealing a novel exploitation of HIPVs, these findings may lead to novel control strategies against an exceedingly important rice pest.

Tritrophic interactions between plants, herbivores, and their natural enemies are an integral part of all terrestrial ecosystems. Herbivore-induced plant volatiles (HIPVs) play a key role in these interactions, as they can attract predators and parasitoids to herbivore-attacked plants. Thirty years after this discovery, the ecological importance of the phenomena is widely recognized. However, the primary function of HIPVs is still subject to much debate, as is the possibility of using these plant-produced cues in crop protection. In this review, we summarize the current knowledge on the role of HIPVs in tritrophic interactions from an ecological as well as a mechanistic perspective. This overview focuses on the main gaps in our knowledge of tritrophic interactions, and we argue that filling these gaps will greatly facilitate efforts to exploit HIPVs for pest control.

Global climatic changes may lead to the arrival of multiple range-expanding species from different trophic levels into new habitats, either simultaneously or in quick succession, potentially causing the introduction of manifold novel interactions into native food webs. Unraveling the complex biotic interactions between native and range-expanding species is critical to understand the impact of climate change on community ecology, but experimental evidence is lacking. In a series of laboratory experiments that simulated direct and indirect species interactions, we investigated the effects of the concurrent arrival of a range-expanding insect herbivore in Europe, Spodoptera littoralis, and its associated parasitoid Microplitis rufiventris, on the native herbivore Mamestra brassicae, and its associated parasitoid Microplitis mediator, when co-occurring on a native plant, Brassica rapa. Overall, direct interactions between the herbivores were beneficial for the exotic herbivore (higher pupal weight than the native herbivore), and negative for the native herbivore (higher mortality than the exotic herbivore). At the third trophic level, both parasitoids were unable to parasitize the herbivore they did not coexist with, but the presence of the exotic parasitoid still negatively affected the native herbivore (increased mortality) and the native parasitoid (decreased parasitism rate), through failed parasitism attempts and interference effects. Our results suggest different interaction scenarios depending on whether S. littoralis and its parasitoid arrive to the native tritrophic system separately or concurrently, as the negative effects associated with the presence of the parasitoid were dependent on the presence of the exotic herbivore. These findings illustrate the complexity and interconnectedness of multitrophic changes resulting from concurrent species arrival to new environments, and the need for integrating the ecological effects of such arrivals into the general theoretical framework of global invasion patterns driven by climatic change.

In the arms race between plants, herbivores, and their natural enemies, specialized herbivores may use plant defenses for their own benefit, and variation in plant traits may affect the benefits that herbivores derive from these defenses. Pieris brassicae is a specialist herbivore of plants containing glucosinolates, a specific class of defensive secondary metabolites. Caterpillars of P. brassicae are known to actively spit on attacking natural enemies, including their main parasitoid, the braconid wasp Cotesia glomerata. Here, we tested the hypothesis that variation in the secondary metabolites of host plants affects the efficacy of caterpillar regurgitant as an anti‐predator defense. Using a total of 10 host plants with different glucosinolate profiles, we first studied natural regurgitation events of caterpillars on parasitoids. We then studied manual applications of water or regurgitant on parasitoids during parasitization events. Results from natural regurgitation events revealed that parasitoids spent more time grooming after attack when foraging on radish and nasturtium than on Brassica spp., and when the regurgitant came in contact with the wings rather than any other body part. Results from manual applications of regurgitant showed that all parameters of parasitoid behavior (initial attack duration, attack interruption, grooming time, and likelihood of a second attack) were more affected when regurgitant was applied rather than water. The proportion of parasitoids re‐attacking a caterpillar within 15 min was the lowest when regurgitant originated from radish‐fed caterpillars. However, we found no correlation between glucosinolate content and regurgitant effects, and parasitoid behavior was equally affected when regurgitant originated from a glucosinolate‐deficient Arabidopsis thaliana mutant line. In conclusion, host plant affects to a certain extent the efficacy of spit from P. brassicae caterpillars as a defense against parasitoids, but this is not due to glucosinolate content. The nature of the defensive compounds present in the spit remains to be determined, and the ecological relevance of this anti‐predator defense needs to be further evaluated in the field.

Herbivore-induced plant volatiles (HIPVs) are important cues for female parasitic wasps to find hosts. Here, we investigated the possibility that HIPVs may also serve parasitoids as cues to locate mates. To test this, the odour preferences of four braconid wasps - the gregarious parasitoid Cotesia glomerata (L.) and the solitary parasitoids Cotesia marginiventris (Cresson), Microplitis rufiventris Kokujev and Microplitis mediator (Haliday) - were studied in olfactometers. Each species showed attraction to pheromones but in somewhat different ways. Males of the two Cotesia species were attracted to virgin females, whereas females of M. rufiventris were attracted to virgin males. Male and female M. mediator exhibited attraction to both sexes. Importantly, female and male wasps of all four species were strongly attracted by HIPVs, independent of mating status. In most cases, male wasps were also attracted to intact plants. The wasps preferred the combination of HIPVs and pheromones over plant odours alone, except M. mediator, which appears to mainly use HIPVs for mate location. We discuss the ecological contexts in which the combined use of pheromones and HIPVs by parasitoids can be expected. To our knowledge, this is the first study to show that braconid parasitoids use HIPVs and pheromones in combination to locate mates.

Interactions between plants and aquatic insects are poorly documented, especially for turbid freshwater ecosystems. Many Swiss lakes offer such habitats, several of which are inhabited by the leaf beetle Macroplea appendiculata (Panzer 1794). This donaciid beetle is the only coleopteran species known to complete its life cycle entirely under water, where it lives primarily on perfoliate pondweed (Potamogeton perfoliatus L.), with Eurasian watermilfoil (Myriophyllum spicatum L.) as an alternative host plant. Direct observations during diving trips, aquatic olfactometer bioassays, and stir bar sorptive extractions (SBSE) coupled with GC–MS analysis were used to understand how these beetles locate their patchily distributed host plants and congeners in a harsh, often swirling environment. In olfactometer assays we observed that the aquatic beetle is strongly attracted to water extracts of pondweed, whereas neither mature males nor females beetles seem to produce attractive cues. The chemical analyses revealed that perfoliate pondweed releases one dominating compound, eucalyptol. Olfactometer assays confirmed that this is a potent attractant for the beetle. We also observed attraction to phytol, which is released by the main, as well as the alternative host plant. These finding are somewhat surprising as eucalyptol has never been reported for aquatic plants and phytol is poorly soluble in water. In addition, both are frequently described as insect repellents in terrestrial ecosystems. We suggest that these terpenoids normally have a defensive function against herbivores and pathogens, but that the highly specialized leaf beetle has evolved to exploit its host’s defence chemistry for optimal foraging.

Plants release herbivore-induced volatiles (HIPVs), which can be used as cues by plants, herbivores and natural enemies. Theory predicts that HIPVs may initially have evolved because of their direct benefits for the emitter and were subsequently adopted as infochemicals. Here, we investigated the potential direct benefits of indole, a major HIPV constituent of many plant species and a key defence priming signal in maize. We used indole-deficient maize mutants and synthetic indole at physiologically relevant doses to document the impact of the volatile on the generalist herbivore Spodoptera littoralis. Our experiments demonstrate that indole directly decreases food consumption, plant damage and survival of S. littoralis caterpillars. Surprisingly, exposure to volatile indole increased caterpillar growth. Furthermore, we show that S. littoralis caterpillars and adults consistently avoid indole-producing plants in olfactometer experiments, feeding assays and oviposition trials. Synthesis. Together, these results provide a potential evolutionary trajectory by which the release of a HIPV as a direct defence precedes its use as a cue by herbivores and an alert signal by plants. Furthermore, our experiments show that the effects of a plant secondary metabolite on weight gain and food consumption can diverge in a counterintuitive manner, which implies that larval growth can be a poor proxy for herbivore fitness and plant resistance.

Associative learning is known to modify foraging behavior in numerous parasitic wasps. This is in agreement with optimal foraging theory, which predicts that the wasps will adapt their responses to specific cues in accordance with the rewards they receive while perceiving these cues. Indeed, the generalist parasitoid Cotesia marginiventris shows increased attraction to a specific plant odor after perceiving this odor during contact with hosts. This positive associative learning is common among many parasitoids, but little is known about the effects of unrewarding host searching events on the attractiveness of odors. To study this, preferences of female C. marginiventris for herbivore-induced odors of three plant species were tested in a six-arm olfactometer after the wasps perceived one of these odors either i) without contacting any caterpillars, ii) while contacting the host caterpillar Spodoptera littoralis, or iii) while contacting the non-host caterpillar Pieris rapae. The results confirm the effects of positive associative learning, but showed no changes in innate responses to the host-induced odors after "negative" experiences. Hence, a positive association is made during an encounter with hosts, but unsuccessful host-foraging experiences do not necessarily lead to avoidance learning in this generalist parasitoid.

In response to insect herbivory, plants mobilize various defenses. Defense responses include the release of herbivore-induced plant volatiles (HIPVs) that can serve as signals to alert undamaged tissues and to attract natural enemies of the herbivores. Some HIPVs can have a direct negative impact on herbivore survival, but it is not well understood by what mechanisms. Here, we tested the hypothesis that exposure to HIPVs renders insects more susceptible to natural pathogens. Exposure of the caterpillars of the noctuid Spodoptera exigua to indole and linalool, but not exposure to (Z)-3-hexenyl acetate, increased the susceptibility to Spodoptera exigua multiple nucleopolyhedrovirus (SeMNPV). We also found that exposure to indole, but not exposure to linalool or (Z)-3-hexenyl acetate, increased the pathogenicity of Bacillus thuringiensis. Additional experiments revealed significant changes in microbiota composition after forty-eight hours of larval exposure to indole. Overall, these results provide evidence that certain HIPVs can strongly enhance the susceptibility of caterpillars to pathogens, possibly through effects on the insect gut microbiota. These findings suggest a novel mechanism by which HIPVs can protect plants from herbivorous insects. IMPORTANCE Multitrophic interactions involving insect pests, their natural enemies, microorganisms, and plant hosts are increasingly being recognized as relevant factors in pest management. In response to herbivory attacks, plants activate a wide range of defenses that aim to mitigate the damage. Attacked plants release herbivore-induced plant volatiles (HIPVs), which can act as priming signals for other plants and attract natural enemies of herbivores, and which may have a direct negative impact on herbivore survival. In the present work, we show that exposure of the insects to the induced volatiles could increase the insects’ susceptibility to the entomopathogens naturally occurring in the plant environment. These findings suggest a novel role for plant volatiles by influencing insect interactions with natural pathogens, probably mediated by alterations in the insect microbiota composition. In addition, this work provides evidence for selectable plant traits (production of secondary metabolites) that can have an influence on the ecology of the pests and could be relevant in the improvement of pest management strategies using natural entomopathogens.

Entomopathogenic nematodes (EPNs) occur in natural and agricultural soils and contribute to the control of soil dwelling insect pests. Ecosystems differ in their soil parameters (pH, moisture, texture, etc.) and microhabitat conditions (presence of roots, plant cover, belowground fauna, etc.), which can affect the EPN soil food web assemblage in an unpredictable manner. By comparing soils that represent different ecological scenarios, we can link the natural distribution of the EPN to abiotic and biotic factors that shape the assemblages of soil food web to unravel the factors that determine EPN abundance and activity. We hypothesized that the EPN soil food web assemblages are affected by agricultural management practices and that this should be reflected when compared to natural ecosystems, such as forests and grasslands. By combining traditional and state-of-the-art molecular tools, we targeted 25 soil organisms to explore EPN food web in 40 Swiss georeferenced sites that can be categorized as “agricultural area”, “forest”, and “grassland”. EPNs abundance, richness and diversity did not vary among the different ecosystems. However, EPN activity (capability to kill insects) was higher in grasslands. Similarly, the free-living nematodes (FLNs) that compete with EPN for the cadavers were much more abundant in grassland, while reduced in the forest soils, suggesting contrasting conditions between these two natural areas. Nematophagous fungi (NF) were more diverse and present in agricultural areas, suggesting that the abiotic conditions in the agricultural areas may favor NF presence. Six soil properties and environmental factors shaped the EPN soil food web in Swiss soils (pH, elevation, clay content, soil water content, temperature, and rainfall), which explained 54% of the community variation in multivariate analysis. These observations were consistent with the key abiotic variables described for subtropical and Mediterranean regions. Identifying the links between the abiotic and biotic factors in very distinct areas can be the basis for predicting EPN soil food web assemblages. Such information can be of value for developing strategies to favour EPN resilience in a changing environment and enhance their capacity as biocontrol agents., This work was supported by an economic stimulus grant (51NF40-144621) and by the National Research Program 68 (NRP68) grant (406840_143065) from the Swiss National Science Foundation. GJ was supported by an assistantship from the University of Neuchâtel and XC was supported by a PhD fellowship from the Commission Fédérale des Bourses pour Etudiants Etrangères CFBE (Confédération Suisse). RCH was supported by The Government of Portugal (Investigator Program award, IF/00552/2014).

Summary When exotic herbivores invade new environments, they have the potential to interfere with native predator-prey relationships. This interference can be indirect, via changes induced in native host plants, and may have cascading consequences at the community level. Here, we investigate the impact of the presence of an exotic generalist insect herbivore, the African cotton leafworm Spodoptera littoralis, on the outcome of native tritrophic interactions between the plant Brassica rapa, the herbivore Pieris brassicae and its parasitoid Cotesia glomerata. In olfactometer choice tests, plants damaged by S. littoralis and plants damaged by both S. littoralis and P. brassicae were consistently less attractive to the parasitoid than plants damaged by P. brassicae alone. Chemical analyses revealed that B. rapa volatiles typically induced by S. littoralis dominated the headspace in case of dual herbivore infestation. In behavioural observations in petri dishes, C. glomerata wasps mistakenly attacked S. littoralis larvae significantly more often when P. brassicae was present, and attacks on both herbivores were comparable in terms of time (attack duration) and energy investment (number of eggs injected). Successful parasitism of S. littoralis was never observed, but larvae exposed to C. glomerata for 24 h exhibited reduced survivorship, possibly as a result of failed parasitism attempts. In tents with herbivore-infested B. rapa plants, the presence of S. littoralis did not have an effect on the parasitism rates of P. brassicae by C. glomerata, regardless of whether the two species were on the same or on different plants. Field experiments in newly invaded environments are necessary to evaluate the realized impact of S. littoralis. Synthesis. Our study illustrates that exotic herbivores can impact native tritrophic interactions associated with Brassica rapa, even if they cannot be used as prey by native natural enemies. The mechanisms behind such effects, in particular chemical interference with foraging cues via changes in herbivore-induced plant volatiles, have the potential to be quite general, and their long-term effects on native communities should not be underestimated.

We believe that replicating studies in ecology and evolution is extremely valuable, but replication within species and systems is troublingly rare, and even ‘quasi-replications’ in different systems are often insufficient. We make a case for supporting multiple types of replications and point out that the current incentive structure needs to change if ecologists and evolutionary biologist are to value scientific replication sufficiently.

In a previous study, we observed no spatial genetic structure in Mexican populations of the parasitoids Chelonus insularis Cresson (Hymenoptera: Braconidae) and Campoletis sonorensis Cameron (Hymenoptera: Ichneumonidae) by using microsatellite markers. In the current study, we investigated whether for these important parasitoids of the fall armyworm (Lepidoptera: Noctuidae) there is any genetic structure at a larger scale. Insects of both species were collected across the American continent and their phylogeography was investigated using both nuclear and mitochondrial markers. Our results suggest an ancient north-south migration of C. insularis, whereas no clear pattern could be determined for C. sonorensis. Nonetheless, the resulting topology indicated the existence of a cryptic taxon within this later species: a few Canadian specimens determined as C. sonorensis branch outside a clade composed of the Argentinean Chelonus grioti Blanchard, the Brazilian Chelonus flavicincta Ashmead, and the rest of the C. sonorensis individuals. The individuals revealing the cryptic taxon were collected from Trichoplusia ni (Hübner) (Lepidoptera: Noctuidae) on tomato (Lycopersicon spp.) and may represent a biotype that has adapted to the early season phenology of its host. Overall, the loosely defined spatial genetic structure previously shown at a local fine scale also was found at the larger scale, for both species. Dispersal of these insects may be partly driven by wind as suggested by genetic similarities between individuals coming from very distant locations. Fil: Violaine, Jourdie. Universite de Neuchatel; Suiza Fil: Virla, Eduardo Gabriel. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Tucumán. Planta Piloto de Procesos Industriales Microbiológicos; Argentina Fil: Murillo, Henry. University of Windsor. Department of Biological Sciences; Canadá Fil: Bento, José Mauricio. Universidade de Sao Paulo; Brasil Fil: Turlings, Ted C. J.. Universite de Neuchatel; Suiza Fil: Alvarez, Nadir. Universite de Neuchatel; Suiza

Wood ants fight pathogens by incorporating tree resin with antimicrobial properties into their nests. They also produce large quantities of formic acid in their venom gland, which they readily spray to defend or disinfect their nest. Mixing chemicals to produce powerful antibiotics is common practice in human medicine, yet evidence for the use of such “defensive cocktails” by animals remains scant. Here, we test the hypothesis that wood ants enhance the antifungal activity of tree resin by treating it with formic acid. In a series of experiments, we document that (i) tree resin had much higher inhibitory activity against the common entomopathogenic fungus Metarhizium brunneum after having been in contact with ants, while no such effect was detected for other nest materials; (ii) wood ants applied significant amounts of endogenous formic and succinic acid on resin and other nest materials; and (iii) the application of synthetic formic acid greatly increased the antifungal activity of resin, but had no such effect when applied to inert glass material. Together, these results demonstrate that wood ants obtain an effective protection against a detrimental microorganism by mixing endogenous and plant‐acquired chemical defenses. In conclusion, the ability to synergistically combine antimicrobial substances of diverse origins is not restricted to humans and may play an important role in insect societies.

Conservation agriculture is arising as an alternative to conventional agriculture with the aim to have a reduced impact on the environment. This includes the use of cover crops to conserve soil quality by limiting soil erosion, adding organic matter, and enhancing soil nutrient content, as well as water availability, which are all factors that can greatly influence the presence of soil organisms. In the current study, we investigated whether winter cover crops can enhance the persistence of entomopathogenic nematodes (EPN) over the winter season. In a first trial we augmented EPN populations in plots without (bared) and with the cover crops, pea (Pisum sativum) or mustard (Brassica juncea). In late autumn, individual mini-plots in each of the three treatments were supplemented with infective juveniles (IJs) of either Heterorhabditis bacteriophora or Steinernema feltiae. In a second trial we studied naturally occurring EPN in plots without (bared) and with the cover crop pea (Pisum sativum) followed by planting of winter wheat. To identify and quantify EPN, we analyzed soil samples using quantitative real time PCR (qPCR) at three time points over the winter season. We also measured the activity of augmented EPN by baiting the soil with wax moth larvae, Galleria mellonella. In addition, we used the qPCR method to investigate the presence of nematophagous fungi (NF), free-living nematodes (FLN) and ectoparasitic bacteria, all of which can interfere with EPN performance. Numbers of naturally occurring EPN in the investigated fields were very low (< 1 EPN per 100 g of soil). The cover crops only had a significant positive effect on the numbers of augmented S. feltiae found in early winter. No striking effect was found for H. bacteriophora. Yet, augmentation was found to enhance the insect-suppressiveness of the soil, as the recorded EPN infectivity after the winter was slightly higher than what was observed in autumn, one month after application. The numbers of FLN, which compete for insect cadavers, was higher in spring than in early winter. These FLN and other antagonists may be important in reducing EPN numbers. In conclusion, the effect of cover crops on EPN persistence was only evident during early-winter and was only significant in the plots augmented with S. feltiae. Moreover, we found that higher numbers of EPN in agricultural soils do not necessarily translate into high infectivity, which is the key factor determining their effectiveness in controlling soil pests. Economic stimulus grant(51NF40-144621) from the Swiss National Science Foundation; National Research Program 68 (NRP68) grant (406840_143065) from the Swiss National Science Foundation info:eu-repo/semantics/publishedVersion

In agricultural ecosystems, pest insects, pathogens, and reduced soil fertility pose major challenges to crop productivity and are responsible for significant yield losses worldwide. Management of belowground pests and diseases remains particularly challenging due to the complex nature of the soil and the limited reach of conventional agrochemicals. Boosting the presence of beneficial rhizosphere organisms is a potentially sustainable alternative and may help to optimize crop health and productivity. Field application of single beneficial soil organisms has shown satisfactory results under optimal conditions. This might be further enhanced by combining multiple beneficial soil organisms, but this remains poorly investigated. Here, we inoculated wheat plots with combinations of three beneficial soil organisms that have different rhizosphere functions and studied their effects on crop performance. Plant beneficial Pseudomonas bacteria, arbuscular mycorrhizal fungi (AMF), and entomopathogenic nematodes (EPN), were inoculated individually or in combinations at seeding, and their effects on plant performance were evaluated throughout the season. We used traditional and molecular identification tools to monitor their persistence over the cropping season in augmented and control treatments, and to estimate the possible displacement of native populations. In three separate trials, beneficial soil organisms were successfully introduced into the native populations and readily survived the field conditions. Various Pseudomonas, mycorrhiza, and nematode treatments improved plant health and productivity, while their combinations provided no significant additive or synergistic benefits compared to when applied alone. EPN application temporarily displaced some of the native EPN, but had no significant long-term effect on the associated food web. The strongest positive effect on wheat survival was observed for Pseudomonas and AMF during a season with heavy natural infestation by the frit fly, Oscinella frit, a major pest of cereals. Hence, beneficial impacts differed between the beneficial soil organisms and were most evident for plants under biotic stress. Overall, our findings indicate that in wheat production under the test conditions the three beneficial soil organisms can establish nicely and are compatible, but their combined application provides no additional benefits. Further studies are required, also in other cropping systems, to fine-tune the functional interactions among beneficial soil organisms, crops, and the environment., The project was funded by grants of the Swiss National Research Program 68 on the “Sustainable Use of Soil as a Resource” from the Swiss National Science Foundation (406840_143065 awarded to TT and FM, 406840_143141 awarded to CK and MM, 406840_143097 to MvdH, OF, and WC). GJ was supported by an assistantship from the University of Neuchâtel, and a Swiss Government Excellence Scholarships supported XC for Foreign Scholars. The Government of Portugal supports RCH with an Investigator Program award (IF/00552/2014).

Volatile organic compounds emitted by plants represent the largest part of biogenic volatile organic compounds (BVOCs) released into our atmosphere. Plant volatiles are formed through many biochemical pathways, constitutively and after stress induction. In recent years, our understanding of the functions of these molecules has made constant and rapid progress. From being considered in the past as a mere waste of carbon, BVOCs have now emerged as an essential element of an invisible language that is perceived and exploited by the plants' enemies, the enemies of plant enemies, and neighbouring plants. In addition, BVOCs have important functions in protecting plants from abiotic stresses. Recent advances in our understanding of the role of BVOC in direct and indirect defences are driving further attention to these emissions. This special issue gathers some of the latest and most original research that further expands our knowledge of BVOC. BVOC emissions and functions in (1) unexplored terrestrial (including the soil) and marine environments, (2) in changing climate conditions, and (3) under anthropic pressures, or (4) in complex trophic communities are comprehensively reviewed. Stepping up from scientific awareness, the presented information shows that the manipulation and exploitation of BVOC is a realistic and promising strategy for agricultural applications and biotechnological exploitations.

Sex pheromones have rarely been studied in parasitoids, and it remains largely unknown how male and female parasitoids locate each other. We investigated possible attraction (and repellency) between the sexes of two braconid wasps belonging to the same genus, the gregarious parasitoid, Cotesia glomerata (L.), and the solitary parasitoid, Cotesia marginiventris (Cresson). Males of both species were strongly attracted to conspecific virgin females. Interestingly, in C. glomerata, the males were repelled by mated females, as well as by males of their own species. This repellency of mated females was only evident hours after mating, implying a change in pheromone composition. Males of C. marginiventris were also no longer attracted, but not repelled, by mated females. Females of both species showed no attraction to the odors of conspecific individuals, male or female, and C. glomerata females even appeared to be repelled by mated males. Moreover, the pheromones were found to be highly specific, as males were not attracted by females of the other species. Males of Cotesia glomerata even avoided the pheromones of female Cotesia marginiventris, indicating the recognition of non-conspecific pheromones. We discuss these unique responses in the context of optimal mate finding strategies in parasitoids.

The great spotted cuckoo (Clamator glandarius) is an important brood parasite of carrion crows (Corvus corone corone) in northern Spain. We recently found that, unlike what is commonly known for cuckoo-host interactions, the great spotted cuckoo has no negative impact on average crow fitness in this region. The explanation for this surprising effect is a repulsive secretion that the cuckoo chicks produce when they are harassed and that may protect the brood against predation. Here, we provide details on the chemical composition of the cuckoo secretion, as well as conclusive evidence that the dominating volatile chemicals in the secretion are highly repellent to model species representative of common predators of the crows. These results support the notion that, in this particular system, the production of a repulsive secretion by the cuckoo chicks has turned a normally parasitic interaction into a mutualistic one.

Agricultural management practices can modify soil properties in ways that may disrupt the abundance and activity of beneficial organisms in the soil. We assessed the impact of different soil management practices on entomopathogenic nematodes (EPN), which have great potential as biological control agents against root-feeding insects. Soils were sampled during spring and autumn 2013 in all 96 plots of a long-term Swiss field trial (DOK experiment). By combining a traditional insect-baiting technique and real-time qPCR analyses, we identified and quantified over 20 soil-dwelling species (or genera). This allowed us to investigate how communities of natural EPN populations and their associated natural enemies and competitors are affected by (i) three crop types (wheat, maize and grass-clover ley) and (ii) farming systems, i.e. conventional, organic and biodynamic, which differed in fertilization, and pesticide use. We also determined the effects on soils' microbial biomass in terms of carbon (C-mic) and nitrogen (N-mic) and applied spatial distribution analysis (SADIE) to uncover patterns of aggregations and associations of the study organisms. Although manure based farming systems increased microbial biomass, the systems did not influence the presence of EPN or their antagonists. EPN was more abundant in winter-wheat plots than in maize and grass-clover ley plots. Overall, very low numbers of EPN were recorded, implying that their natural presence would not be sufficient to have a satisfactory suppressive effect on root-feeding pests and the application of EPN would therefore be an appropriate measure to protect yields in case of root pest outbreaks. (C) 2016 Elsevier B.V. All rights reserved. Grant (51NF40-144621) and by the National Research Program 68 (NRP68) grant (406840_143065)

In response to insect attack, plants release complex blends of volatile compounds. These volatiles serve as foraging cues for herbivores, predators and parasitoids, leading to plant-mediated interactions within and between trophic levels. Hence, plant volatiles may be important determinants of insect community composition. To test this, we created rice lines that are impaired in the emission of two major signals, S-linalool and (E)-β-caryophyllene. We found that inducible S-linalool attracted predators and parasitoids as well as chewing herbivores, but repelled the rice brown planthopper Nilaparvata lugens, a major pest. The constitutively produced (E)-β-caryophyllene on the other hand attracted both parasitoids and planthoppers, resulting in an increased herbivore load. Thus, silencing either signal resulted in specific insect assemblages in the field, highlighting the importance of plant volatiles in determining insect community structures. Moreover, the results imply that the manipulation of volatile emissions in crops has great potential for the control of pest populations.

In most agro-ecosystems the organisms that feed on plant roots have an important impact on crop yield and can impose tremendous costs to farmers. Similar to aboveground pests, they rely on a broad range of chemical cues to locate their host plant. In their turn, plants have co-evolved a large arsenal of direct and indirect defense to face these attacks. For instance, insect herbivory induces the synthesis and release of specific volatile compounds in plants. These volatiles have been shown to be highly attractive to natural enemies of the herbivores, such as parasitoids, predators, or entomopathogenic nematodes. So far few of the key compounds mediating these so-called tritrophic interactions have been identified and only few genes and biochemical pathways responsible for the production of the emitted volatiles have been elucidated and described. Roots also exude chemicals that directly impact belowground herbivores by altering their behavior or development. Many of these compounds remain unknown, but the identification of, for instance, a key compound that triggers nematode egg hatching to some plant parasitic nematodes has great potential for application in crop protection. These advances in understanding the chemical emissions and their role in ecological signaling open novel ways to manipulate plant exudates in order to enhance their natural defense properties. The potential of this approach is discussed, and we identify several gaps in our knowledge and steps that need to be taken to arrive at ecologically sound strategies for belowground pest management.

Background: Entomopathogenic nematodes (EPNs) are tiny parasitic worms that parasitize insects, in which they reproduce. Their foraging behavior has been subject to numerous studies, most of which have proposed that, at short distances, EPNs use chemicals that are emitted directly from the host as host location cues. Carbon dioxide (CO2) in particular has been implicated as an important cue. Recent evidence shows that at longer distances several EPNs take advantage of volatiles that are specifically emitted by roots in response to insect attack. Studies that have revealed these plant-mediated interactions among three trophic levels have been met with some disbelief. Scope: This review aims to take away this skepticism by summarizing the evidence for a role of root volatiles as foraging cues for EPNs. To reinforce our argument, we conducted olfactometer assays in which we directly compared the attraction of an EPN species to CO2 and two typical inducible root volatiles. Conclusions: The combination of the ubiquitous gas and a more specific root volatile was found to be considerably more attractive than one of the two alone. Hence, future studies on EPN foraging behavior should take into account that CO2 and plant volatiles may work in synergy as attractants for EPNs. Recent research efforts also reveal prospects of exploiting plant-produced signals to improve the biological control of insect pests in the rhizosphere

Apparent competition, through the action of shared natural enemies, is frequently suggested as a possible mechanism underlying the impact of invasive alien species on native species, but examples are rare, particularly in insects. A previous study showed that the beech leaf mining weevil, Orchestes fagi, was significantly less abundant close to horse-chestnut trees infested by the invasive horse-chestnut leaf mining moth, Cameraria ohridella, compared to control sites. Apparent competition through the sharing of natural enemies was proposed as a potential mechanism underlying this effect. To test the occurrence of apparent competition between the two leaf miner species, three observational studies and one experimental manipulation were carried out in Switzerland during 3 years. The total mortality, parasitism, predation and parasitoid diversity of larvae and pupae of O. fagi were compared between sites with and without horse-chestnut trees severely attacked by C. ohridella. Total mortality and predation rates of O. fagi were not significantly different between sites with and sites without C. ohridella. Despite a large overlap between the parasitoid complexes of the two leaf miners, parasitism of O. fagi was found to be positively influenced by the presence of horse-chestnuts infested by C. ohridella in only one of the four studies and only for 1 year. Similarly, parasitoid diversity was not higher near infested horse-chestnut trees compared to control sites. Thus, little evidence for apparent competition was found. Possible reasons, including possible insufficiencies in the experimental circumstances and design, are discussed.

Determining links between plant defence strategies is important to understand plant evolution and to optimize crop breeding strategies. Although several examples of synergies and trade-offs between defence traits are known for plants that are under attack by multiple organisms, few studies have attempted to measure correlations of defensive strategies using specific single attackers. Such links are hard to detect in natural populations because they are inherently confounded by the evolutionary history of different ecotypes. We therefore used a range of 20 maize inbred lines with considerable differences in resistance traits to determine if correlations exist between leaf and root resistance against pathogens and insects. Aboveground resistance against insects was positively correlated with the plant's capacity to produce volatiles in response to insect attack. Resistance to herbivores and resistance to a pathogen, on the other hand, were negatively correlated. Our results also give first insights into the intraspecific variability of root volatiles release in maize and its positive correlation with leaf volatile production. We show that the breeding history of the different genotypes (dent versus flint) has influenced several defensive parameters. Taken together, our study demonstrates the importance of genetically determined synergies and trade-offs for plant resistance against insects and pathogens.

Predation Favors Parasitism Parasitism in birds often results in ejection or starvation of the host's nestlings. Consequently, many host bird species have evolved protective behavior such as mobbing and parasite egg rejection. Curiously, some host species show no parasite avoidance behaviors; for example, the crow Corvus corone corone tolerates cuckoo chicks among its own brood. In a long-term study, Canestrari et al. (p. 1350 ) found that crow nests containing a cuckoo chick had lower rates of predation because the parasite's chicks secrete a noxious repellent substance. Overall, in years of high predation pressure, the presence of cuckoos improves the crow's breeding success, but when there are fewer predators around, parasitism reduces crow fitness.

Induced changes in plant quality can mediate indirect interactions between herbivores. Although the sequence of attack by different herbivores has been shown to influence plant responses, little is known about how this affects the herbivores themselves. We therefore investigated how induction by the leaf herbivore Spodoptera frugiperda influences resistance of teosinte (Zea mays mexicana) and cultivated maize (Zea mays mays) against root-feeding larvae of Diabrotica virgifera virgifera. The importance of the sequence of arrival was tested in the field and laboratory. Spodoptera frugiperda infestation had a significant negative effect on colonization by D. virgifera larvae in the field and weight gain in the laboratory, but only when S. frugiperda arrived on the plant before the root herbivore. When S. frugiperda arrived after the root herbivore had established, no negative effects on larval performance were detected. Yet, adult emergence of D. virgifera was reduced even when the root feeder had established first, indicating that the negative effects were not entirely absent in this treatment. The defoliation of the plants was not a decisive factor for the negative effects on root herbivore development, as both minor and major leaf damage resulted in an increase in root resistance and the extent of biomass removal was not correlated with root-herbivore growth. We propose that leaf-herbivore-induced increases in feeding-deterrent and/or toxic secondary metabolites may account for the sequence-specific reduction in root-herbivore performance. Synthesis. Our results demonstrate that the sequence of arrival can be an important determinant of plant-mediated interactions between insect herbivores in both wild and cultivated plants. Arriving early on a plant may be an important strategy of insects to avoid competition with other herbivores. To fully understand plant-mediated interactions between insect herbivores, the sequence of arrival should be taken into account. © 2011 The Authors. Journal of Ecology © 2011 British Ecological Society.

Plant chemistry can strongly influence interactions between herbivores and their natural enemies, either by providing volatile compounds that serve as foraging cues for parasitoids or predators, or by affecting the quality of herbivores as hosts or prey. Through these effects plants may influence parasitoid population genetic structure. We tested for a possible specialization on specific crop plants in Chelonus insularis and Campoletis sonorensis, two primary parasitoids of the fall armyworm, Spodoptera frugiperda. Throughout Mexico, S. frugiperda larvae were collected from their main host plants, maize and sorghum and parasitoids that emerged from the larvae were used for subsequent comparison by molecular analysis. Genetic variation at eight and 11 microsatellites were respectively assayed for C. insularis and C. sonorensis to examine isolation by distance, host plant and regional effects. Kinship analyses were also performed to assess female migration among host-plants. The analyses showed considerable within population variation and revealed a significant regional effect. No effect of host plant on population structure of either of the two parasitoid species was found. Isolation by distance was observed at the individual level, but not at the population level. Kinship analyses revealed significantly more genetically related—or kin—individuals on the same plant species than on different plant species, suggesting that locally, mothers preferentially stay on the same plant species. Although the standard population genetics parameters showed no effect of plant species on population structure, the kinship analyses revealed that mothers exhibit plant species fidelity, which may speed up divergence if adaptation were to occur.

Herbivore-damaged plants emit volatile organic compounds that attract natural enemies of the herbivores. This form of indirect plant defence occurs aboveground as well as belowground, but it remains unclear how simultaneous feeding by different herbivores attacking leaves and roots may affect the production of the respective defence signals. We employed a setup that combines trapping of volatile organic signals and simultaneous measurements of the attractiveness of these signals to above and belowground natural enemies. Young maize plants were infested with either the foliar herbivore Spodoptera littoralis, the root herbivore Diabrotica virgifera virgifera, or with both these important pest insects. The parasitic wasp Cotesia marginiventris and the entomopathogenic nematode Heterorhabditis megidis were strongly attracted if their respective host was feeding on a plant, but this attraction was significantly reduced if both herbivores were on a plant. The emission of the principal root attractant was indeed reduced due to double infestation, but this was not evident for the leaf volatiles. The parasitoid showed an ability to learn the differences in odour emissions and increased its response to the odour of a doubly infested plant after experiencing this odour during an encounter with hosts. This first study to measure effects of belowground herbivory on aboveground tritrophic signalling and vice-versa reemphasizes the important role of plants in bridging interactions between spatially distinct components of the ecosystem.

1 Laboratory of Entomology, Wageningen University, Wageningen, Netherlands, Department of Terrestrial Ecology, Netherlands Institute of Ecology (NIOO-KNAW), Wageningen, Netherlands, 3 R&D Microbiology, Koppert Biological Systems, Berkel en Rodenrijs, Netherlands, Department of Soil Microbiology and Symbiotic Systems, Estacion Experimental del Zaidin, CSIC, Granada, Spain, 5 Laboratory of Functional Ecology, Institute of Biology, University of Neuchâtel, Neuchâtel, Switzerland, 6 Laboratory of Fundamental and Applied Research in Chemical Ecology, Institute of Biology, University of

The selection of oviposition sites by syrphids and other aphidophagous insects is influenced by the presence of con- and heterospecific competitors. Chemical cues play a role in this selection process, some of them being volatile semiochemicals. Yet, little is known about the identity and specificity of chemical signals that are involved in the searching behavior of these predators. In this study, we used olfactometer bioassays to explore the olfactory responses of gravid females and larvae of the syrphid Sphaerophoria rueppellii, focussing on volatiles from conspecific immature stages, as well as odors from immature stages of the competing coccinellid Adalia bipunctata. In addition, a multiple-choice oviposition experiment was conducted to study if females respond differently when they can also sense their competitors through visual or tactile cues. Results showed that volatiles from plants and aphids did not affect the behavior of second-instars, whereas adult females strongly preferred odors from aphid colonies without competitors. Odors from conspecific immature stages had a repellent effect on S. rueppellii adult females, whereas their choices were not affected by volatiles coming from immature heterospecific A. bipunctata. The results imply that the syrphid uses odors to avoid sites that are already occupied by conspecifics. They did not avoid the odor of the heterospecific competitor, although in close vicinity they were found to avoid laying eggs on leaves that had traces of the coccinellid. Apparently adult syrphids do not rely greatly on volatile semiochemicals to detect the coccinellid, but rather use other stimuli at close range (e. g., visual or non-volatile compounds) to avoid this competitor. RAJ was supported by JAE Predoc CSIC scholarship (JAEPre_08_00457). The Authors acknowledge the financial support from the Spanish Ministry of Science and Education (AGL2005-01449/AGR project) and the Swiss National Science Foundation (SNF 31003A-122132).

In response to feeding by arthropods, plants actively and systemically emit various volatile substances. It has been proposed that these herbivore-induced volatiles (HIPVs) can be exploited in agricultural pest control because they might repel herbivores and because they serve as attractants for the enemies of the herbivores. Indeed, recent studies with transgenic plants confirm that odour emissions can be manipulated in order to enhance the plants' attractiveness to beneficial arthropods. An additional advantage of manipulating HIPV emissions could be their effects on neighbouring plants, as a rapidly increasing number of studies show that exposure to HIPVs primes plants for augmented defence expression. Targeting the right volatiles for enhanced emission should lead to ecologically and economically sound ways of combating important pests.

1. In studies on optimal foraging strategies, long-range decisions in the pursuit of resource are rarely considered. This is also the case for sympatric parasitoids, which may be confronted with the decision to accept or reject host larvae that are already parasitized by a competing species. They can be expected to reject already parasitized hosts if it is likely that they will lose the resulting intrinsic competition. However, examples of such interspecific host discrimination are rare. 2. We propose that parasitoids that are not egg-limited should reject inferior hosts only if it saves them time, and that this will be achieved mainly when the parasitoids are able to detect competitors from a distance. We tested this hypothesis using the sympatric parasitoids Cotesia marginiventris (Cresson) and Campoletis sonorensis (Cameron). 3. C. sonorensis was found to be the superior intrinsic competitor but, upon contact with a host larva, both wasps readily accepted hosts that had already been parasitized by the other species. However, in an olfactometer experiment, C. marginiventris females were found to strongly avoid the odour of their superior competitor. 4. These results are in accordance with a time optimization scenario, whereby the inferior competitor accepts competition if it costs only an egg, but avoids competition if it may save time that can be allocated to the search for more profitable hosts. 5. Models on host discrimination strategies in parasitoids had not yet considered discrimination from a distance. Long-range foraging decisions can also be expected for other organisms that have to choose between resources of varying suitability and profitability.

Oreina cacaliae (Schrank) (Coleoptera: Chrysomelidae) has a 2-year life cycle that it has to complete within the short warm seasons of the harsh alpine environment. Three years of field observations and experiments revealed that not all beetles overwintered in the soil next to their principal host Adenostyles alliariae (Asteraceae), as was previously assumed, but that many O. cacaliae left their host in autumn and flew to overwintering sites that were extensively sun-exposed. In spring, these individuals became active 2 months earlier than their conspecifics that had remained in the soil close to the host plant. These early beetles flew from their hibernation sites against the direction of the prevailing wind. After a random landing in snow, they walked to the spring host Petasites paradoxus (Asteraceae) and fed on its floral stalks, the only plant parts present at that time. A few weeks later, they took flight again to locate newly emerging A. alliariae on which they would feed and deposit larvae as did individuals that had overwintered close to A. alliariae. Leaves of A. alliariae contain pyrrolizidine alkaloids (PAs), which the beetles sequester for their own defence. The dominating PA (seneciphylline) was also found to be present in the floral stalks of P. paradoxus. With additional behavioural assays in the field and laboratory, we demonstrated the importance of plant odours in the short-range host location process. This study reveals a unique hibernation behaviour in which part of the beetle population uses exceptionally warm locations from which they emerge in spring, long before all the snow has melted. This early, but risky emergence allows them to exploit a second, highly suitable host plant, which they locate first by wind-guided flight and then by odour-guided walking. The well-fed beetles then use odour again to move to their principal host plant, on which they reproduce.

“He is lost for science.” Those were the words from a highly respected evolutionary ecologist when he learned, almost 30 years ago, that I would travel to the US to conduct a PhD project in chemical ecology at the USDA in Gainesville, Florida. Agro-ecosystems were commonly avoided like the plague by respected or respectable ecologists. An obsession with the F-word can be blamed for this. The general perception has long been that fundamental research is far more worthy than applied research, and that this can only be conducted in “natural” systems. This seems a bit silly if we consider that more than 50 % of the terrestrial surface of our planet is being exploited and modified in some way or another by humans (http://data.worldbank.org), and it is time to accept that most of the relevant contemporary evolution is taking place in these exploited areas. We also can wonder if it really is so bad to devote our research efforts to “unnatural” systems in order to address questions of importance to our society. Even the respected ecologist that criticized my career move eventually sold out and switched to applied research, initially most likely because of funding opportunities, but also perhaps because fundamental research was more and more considered to be compatible with trendy societal issues like climate change, invasive organisms, and biodiversity. What does this have to do with the Journal of Chemical Ecology and its 40th anniversary? It is my impression that the Journal and the scientists that contribute to it havemade an important trendsetting contribution to what is considered to be important research. When I left Holland for my adventure in Florida, I entered the world of chemical ecology and applied entomology. I joined the group of JimTumlinson, one of about a dozen scientists that have paved the way for the field of chemical ecology and its journal. Their work on insect pheromones was groundbreaking and led to the very successful application of pheromones in insect pest control. Much of their work was published in the Journal of Chemical Ecology (JCE), but it also led to papers in journals with a slightly higher impact, like Nature and Science. Chemical ecology also was one of the first disciplines that was highly interdisciplinary, combining behavioral ecology, insect physiology, as well as analytical and synthetic chemistry. This extent of interdisciplinarity was a rarity at the time. In those early days, this very young discipline received considerable respect and attracted an important base of enthusiastic young scientists, also those that had a background in ecology and evolutionary biology. I argue that this new infusion caused a shift in the general focus of research in chemical ecology. It became more and more evident that not only pheromones, but also plant-derived chemicals are of utmost importance for the foraging behavior and performance of insects. Plants and the evolutionary processes that determined their interactions with insects started to dominate the field. This notion finds support in the types of papers that appeared in JCE. Consulting the Web-of-Science teaches us that indeed pheromone research largely dominated the earlier days of the field. The number of papers on pheromones has not decreased over the years, but other topics have been introduced (Fig. 1). One of those is evolutionary ecology. The term evolution rarely occurred in earlier work, but, as Fig. 2 shows us, was an important aspect of much of the work that was published in JCE in the nineties, and it is still going strong. It seems roughly correct to say that most of this evolutionary work was inspired by a general interest in the evolution of plantinsect interactions. With the involvement of molecular plant physiologists, the work has revealed a tremendously sophisticated network of plant defense responses to herbivory and pathogen infections. This knowledge has nowopened theway to... application, againwith a great emphasis on controlling insect pests. If you have read this article until here, it probably not only means that many of your friends and family regularly tell you to get a life, but also that you are probably genuinely interested in the field of chemical ecology. In short, chemical ecologists read the Journal of Chemical Ecology and this is the final point that I would like to make. These days we are playing the numbers games with our publications in order to impress our peers, superiors, and funding agencies. Impact is what counts, and this is not always to the benefit of our Journal, but it should be. There is a false perception that we are better off publishing in journals that are a few numbers higher in average impact. If your intention is to maximize your h-index, this is the wrong notion. I would argue that your colleagues are more likely to read and cite your papers if they are published in JCE than in journals with a considerably higher impact factor. For myself this is certainly true. I searched for the average number of citations of the twenty JCE papers that I have been author on and found that it is 50 % higher than for the average of my papers published in journals that have an impact factor between 3 and 9. I like to think that I was not entirely lost for science when I decided for a career in chemical ecology. Youwon’t be either, especially if you publish in the Journal of Chemical Ecology. Fig. 1 Pheromones 1977 to 2013

Plant volatiles induced by insect feeding are known to attract natural enemies of the herbivores. Six maize inbred lines that showed distinctly different patterns of volatile emission in laboratory assays were planted in randomized plots in the Central Mexican Highlands to test their ability to recruit parasitic wasps under field conditions. The plants were artificially infested with neonate larvae of the fall armyworm Spodoptera frugiperda, and two of its main endoparasitoids, Campoletis sonorensis and Cotesia marginiventris, were released in the plots. Volatiles were collected from equally treated reference plants in the neighbourhood of the experimental field. The cumulative amount of 36 quantified volatile compounds determined for each line was in good accordance with findings from the laboratory; there was an almost 15-fold difference in total emission between the two extreme lines. We found significant differences among the lines with respect to the numbers of armyworms recovered from the plants, their average weight gain and parasitism rates. Average weight of the caterpillars was negatively correlated with the average total amount of volatiles released by the six inbred lines. However, neither total volatile emission nor any specific single compound within the blend could explain the differential parasitism rates among the lines, with the possible exception of (E)-2-hexenal for Campoletis sonorensis and methyl salicylate for Cotesia marginiventris. Herbivore-induced plant volatiles and/or correlates thereof contribute to reducing insect damage of maize plants through direct plant defence and enhanced attraction of parasitoids, alleged indirect defence. The potential to exploit these volatiles for pest control deserves to be further evaluated.

The effect of the alien horse-chestnut leaf miner, Cameraria ohridella, on native fauna was studied by comparing the species richness of native leaf miner communities and the abundance of selected native leaf miner species in the presence and absence of horse-chestnut trees infested by C. ohridella, in various environments in Europe. The species richness of native leaf miner communities in Switzerland was lower at sites where C. ohridella was present than at control sites. In Switzerland, France and Bulgaria, several native leaf miner species were significantly less abundant in the vicinity of infested horse-chestnuts. The native species most affected by the presence of the invasive alien species were those occurring early in the year and sharing their parasitoid complex with C. ohridella. These results suggest apparent competition mediated by shared natural enemies because these are the only link between C. ohridella and native leaf miners using other food resources.

Chemical cues enable female parasitic wasps to locate the eggs, larvae, or other life stages of the insects in or on which they place their eggs. These chemical signals, or semiochemicals, may be produced by the hosts and/or by the plants on which the hosts feed. The composition of the chemical signal often differs with different species of hosts or with different plants. New evidence suggests that the wasps exploit semiochemicals emitted by plants in response to insect herbivore feeding. The wasps learn to respond to the different blends of chemicals that indicate the location of their hosts and they can be trained to respond to a specific odor blend. Thus, it may be possible to increase their effectiveness for biological control by conditioning them, prior to their release, to search for a target pest in

1 Native natural enemies have the potential to control fall armyworm Spodoptera frugiperda (Smith) in tropical maize grown in Mexico, where this insect pest causes severe economic losses to farmers. It has been proposed that enhancing herbivore-induced volatile emissions in maize plants may help to increase the effectiveness of natural enemies, which use these volatiles to locate their prey. This will only be of immediate benefit to farmers if the activity of the natural enemies results in a direct reduction in herbivory. Here we report on field surveys for the most common natural enemies in a tropical maize-growing region in Mexico and the potential effects of these enemies on herbivory by fall armyworm. 2 Caterpillars were collected in maize fields near Poza Rica in the state of Veracruz during January and February 1999, 2000 and 2001. Plants were either naturally infested by S. frugiperda, or artificially infested with laboratory-reared larvae. Ten species of parasitoids emerged from the collected larvae and eight species of predators that are known to feed on larvae and eggs were observed on the plants. Campoletis sonorensis (Cameron) (Hymenoptera: Ichneumonidae) was the dominant parasitoid species, in 1999 and 2001. 3 Of the nine larval parasitoids collected, six (all solitary) are known to reduce herbivory, whereas one causes the host to eat more (for two species this is not known). This implies that enhancing the effectiveness of solitary endoparasitoids may benefit subsistence farmers in developing countries by immediately reducing herbivory. The overall benefit for the plant resulting from parasitoid activity also has important implications for the evolutionary role of parasitoids in contributing to selection pressures that shape indirect defences in plants.

Plants have evolved intricate strategies to withstand attacks by herbivores and pathogens. Although it is known that plants change their primary and secondary metabolism in leaves to resist and tolerate aboveground attack, there is little awareness of the role of roots in these processes. This is surprising given that plant roots are responsible for the synthesis of plant toxins, play an active role in environmental sensing and defense signaling, and serve as dynamic storage organs to allow regrowth. Hence, studying roots is essential for a solid understanding of resistance and tolerance to leaf-feeding insects and pathogens. Here, we highlight this function of roots in plant resistance to aboveground attackers, with a special focus on systemic signaling and insect herbivores.

Responses of individual females of the parasitoidCotesia marginiventris to the odors of four different complexes of host larvae feeding on leaves were observed in a four-arm olfactometer. The plant-host complexes were composed of fall armyworm (FAW) larvae or cabbage looper (CL) larvae feeding on either corn or cotton seedlings. Prior to testing, each female was given a brief foraging experience on a plant-host complex and was then exposed to the odors of the same complex in the olfactometer. The experienced females responded to familiar odors in a dose-related manner, and these responses were virtually identical to all four complexes. Preferences for the odors of one of two plant-host complexes were tested in dual choice situations. Generally, FAW odors were preferred over CL odors and corn odors over cotton odors. A short foraging experience significantly affected the females' odor preferences in favor of the odors released by the experienced complex. Additional experiments revealed that neither longer bouts of experience nor bouts that included ovipositions resulted in a stronger change in preference. Experience affected preference in combinations where only the host species was varied as well as in combinations where only the plant species was varied. The results, therefore, strongly indicate that both the plants and the hosts somehow are involved in the production and/or release of the semiochemicals that attractC. marginiventris.