Aphid-Induced Volatiles and Subsequent Attraction of Natural Enemies Varies among Sorghum Cultivars.

The production of herbivore-induced plant volatiles (HIPVs) is a type of indirect defense used by plants to attract natural enemies and reduce herbivory by insect pests. In many crops little is known about genotypic variation in HIPV production or how this may affect natural enemy attraction. In this study, we identified and quantified HIPVs produced by 10 sorghum (Sorghum bicolor) cultivars infested with a prominent aphid pest, the sorghum aphid (Melanaphis sorghi Theobald). Volatiles were collected using dynamic headspace sampling techniques and identified and quantified using GC-MS. The total amounts of volatiles induced by the aphids did not differ among the 10 cultivars, but overall blends of volatiles differed significantly in composition. Most notably, aphid herbivory induced higher levels of methyl salicylate (MeSA) emission in two cultivars, whereas in four cultivars, the volatile emissions did not change in response to aphid infestation. Dual-choice olfactometer assays were used to determine preference of the aphid parasitoid, Aphelinus nigritus, and predator, Chrysoperla rufilabris, between plants of the same cultivar that were un-infested or infested with aphids. Two aphid-infested cultivars were preferred by natural enemies, while four other cultivars were more attractive to natural enemies when they were free of aphids. The remaining four cultivars elicited no response from parasitoids. Our work suggests that genetic variation in HIPV emissions greatly affects parasitoid and predator attraction to aphid-infested sorghum and that screening crop cultivars for specific predator and parasitoid attractants has the potential to improve the efficacy of biological control.

Macroevolution of protective coloration across caterpillars reflects relationships with host plants.

A critical function of animal coloration is avoiding attack, either by warning predators or reducing detectability. Evolution of these divergent strategies may depend on prey palatability and apparency to predators: conspicuous coloration may be favoured if species are distasteful, or habitats make hiding difficult; by contrast, camouflage may be effective if prey lack defences or environments are visually complex. For insect herbivores, host plants provide both chemical defence and the background against which they are detected or obscured; thus, plant traits may be key to coloration in these foundational terrestrial organisms. We use 1808 species of larval Lepidoptera to explore macroevolution of protective coloration strategy. We find that colour and pattern evolve jointly in caterpillars, similar to an array of species across the animal kingdom, while individual elements of coloration evolve closely with diet ecology. Consistent with key tenets of plant defence and plant-herbivore coevolutionary theory, conspicuous colours are associated with herbaceous host plants-thought to be defended by toxins-while camouflage colours and patterns are associated with woody plants and grasses. Contrary to theory, dietary specialization is not associated with conspicuous coloration. Our results add valuable insights into the evolutionary forces shaping colour and pattern in nature.

The consequence of leaf life span to virus infection of herbivorous insects.

Many herbivorous insects die of pathogen infections, though the role of plant traits in promoting the persistence of these pathogens as an indirect interaction is poorly understood. We tested whether winter leaf retention of bush lupines (Lupinus arboreus) promotes the persistence of a nucleopolyhedroviruses, thereby increasing the infection risk of caterpillars (Arctia virginalis) feeding on the foliage during spring. We also investigated whether winter leaf retention reduces viral exposure of younger caterpillars that live on the ground, as leaf retention prevents contaminated leaves from reaching the ground. We surveyed winter leaf retention of 248 lupine bush canopies across twelve sites and examined how it related to caterpillar infection risk, herbivory, and inflorescence density. We also manipulated the amount of lupine litter available to young caterpillars in a feeding experiment to emulate litterfall exposure in the field. Greater retention of contaminated leaves from the previous season increased infection rates of caterpillars in early spring. Higher infection rates reduced herbivory and increased plant inflorescence density by summer. Young caterpillars exposed to less litterfall were more likely to starve to death but less likely to die from infection, further suggesting foliage mediated exposure to viruses. We speculate that longer leaf life span may be an unrecognized trait that indirectly mediates top-down control of herbivores by facilitating epizootics.

Oviposition Preference and Performance of a Specialist Herbivore Is Modulated by Natural Enemies, Larval Odors, and Immune Status.

Insect herbivores frequently must balance host plant quality and the risk of attack by their natural enemies when making oviposition decisions. Yet, which factor is more important remains unresolved in plant-insect ecology. Here, we report the oviposition preference and larval performance of the brassicaceous specialist Plutella xylostella, in the context of plant quality (cabbage Brassica oleracea vs. mustard B. juncea) and associated natural enemies. Despite the greater larval weight and adult lifespan on cabbage, ovipositing females strongly preferred mustard. Both the egg parasitoid Trichogrammatoidea bactrae and the larval ectoparasitoid Bracon brevicornis are more likely to attack P. xylostella that feed on cabbage; thus, mustard represents enemy-reduced space from these two parasitoids. However, larval diet had no impact on the parasitism rate of specialist Cotesia vestalis. Feeding on mustard improved larval immune responses. The total hemocyte number, diversity, and phenoloxidase activity were higher in mustard-fed larvae which increased their survival against the entomopathogen, Bacillus thuringiensis. Interestingly, host plants altered the larval body odor profile. Mustard-fed larvae emitted allyl isothiocyanate (AITC) and butyl isothiocyanate (BITC) while cabbage-fed larvae emitted dimethyl disulphide (DMDS) and dimethyl trisulphide (DMTS) that served as short-range cues for larval parasitoids. For B. brevicornis, host body odor guided oviposition choice was crucial as their fitness was affected by the host larval diet. Although C. vestalis showed a clear preference towards volatiles emitted by mustard fed larvae, their fitness was unaltered. Taken together, our results illustrate that P. xylostella prefers to lay eggs on mustard plants providing enemy-reduced space from some, but not all, natural enemies.

An unbiased approach elucidates variation in (S)-(+)-linalool, a context-specific mediator of a tri-trophic interaction in wild tobacco.

Plant volatile organic compounds (VOCs) mediate many interactions, and the function of common VOCs is especially likely to depend on ecological context. We used a genetic mapping population of wild tobacco, Nicotiana attenuata, originating from a cross of 2 natural accessions from Arizona and Utah, separated by the Grand Canyon, to dissect genetic variation controlling VOCs. Herbivory-induced leaf terpenoid emissions varied substantially, while green leaf volatile emissions were similar. In a field experiment, only emissions of linalool, a common VOC, correlated significantly with predation of the herbivore Manduca sexta by native predators. Using quantitative trait locus mapping and genome mining, we identified an (S)-(+)-linalool synthase (NaLIS). Genome resequencing, gene cloning, and activity assays revealed that the presence/absence of a 766-bp sequence in NaLIS underlies the variation of linalool emissions in 26 natural accessions. We manipulated linalool emissions and composition by ectopically expressing linalool synthases for both enantiomers, (S)-(+)- and (R)-(-)-linalool, reported to oppositely affect M. sexta oviposition, in the Arizona and Utah accessions. We used these lines to test ovipositing moths in increasingly complex environments. The enantiomers had opposite effects on oviposition preference, but the magnitude of the effect depended strongly both on plant genetic background, and complexity of the bioassay environment. Our study reveals that the emission of linalool, a common VOC, differs by orders-of-magnitude among geographically interspersed conspecific plants due to allelic variation in a linalool synthase, and that the response of a specialist herbivore to linalool depends on enantiomer, plant genotype, and environmental complexity.

Vector-borne plant pathogens modify top-down and bottom-up effects on insect herbivores.

Ecological theory predicts that host-plant traits affect herbivore population growth rates, which in turn modulates predator-prey interactions. However, while vector-borne plant pathogens often alter traits of both host plants and vectors, a few studies have assessed how pathogens may act as interaction modifiers within tri-trophic food webs. By applying a food web motif framework, we assessed how a vector-borne plant pathogen (Pea-enation mosaic virus, PEMV) modified both bottom-up (plant-herbivore) and top-down (predator-prey) interactions. Specifically, we assessed trophic interactions with PEMV-infectious Acyrthosiphon pisum (pea aphid) vectors compared to non-infectious aphids in a factorial experiment that manipulated predator and plant communities. We show that PEMV altered bi-trophic relationships, whereby on certain plant species, PEMV reduced vector performance but also increased their susceptibility to predators. However, on other plant species, PEMV weakened top-down control or increased vector performance. Our results suggest that vector-borne plant pathogens are important interaction modifiers for plant-herbivore-predator dynamics: host-plant response to viruses can decrease herbivore abundance by reducing herbivore performance, but also increase herbivore abundance by weakening top-down control. Broadly speaking, trophic interactions that regulate herbivore outbreaks appear to be modified for herbivores actively transmitting viruses to host plants. Consequently, management and monitoring of outbreaking herbivores should consider the infection status of focal populations.

Plant silicon application alters leaf alkaloid concentrations and impacts parasitoids more adversely than their aphid hosts.

Grasses accumulate large amounts of silicon (Si) which acts as a highly effective physical defence against insect herbivory, however recent evidence shows that Si supplementation also modifies plant secondary metabolite concetrations. Changes in plant secondary metabolites concentrations can have cascading effects on higher trophic levels, such as parasitoids, as they are dependent on the host herbivore for growth and development. However, relatively little is known about how Si application affects higher trophic levels. We examined the effects of Si addition on alkaloid content in leaves of Phalaris aquatica (Poaceae) and the effect on interactions between an aphid (Rhopalosiphum padi) and its parasitoid (Aphidius colemani). Si supplementation had no effect on aphid abundance or parasitism rate. Adult aphids, aphid mummies (parasitised aphids) and the emergent parasitoids were, however, significantly smaller on Si+ plants. Parasitoid traits (size and emergence) were correlated with aphid mummy size. Si addition reduced parasitoid emergence rate and size due to reduced host mummy size, in addition, significantly fewer females emerged from mummies on Si+ plants. Aphid infestation significantly altered alkaloids concentrations, reducing gramine by 80% while increasing tryptamine by 91% in Si- plants. Si addition reduced aphid-induced tryptamine concentrations by 64% and increased 5-MeO-tryptamine by over 800% in control and 142% in aphid infested plants. Our results show that while Si addition has modest impacts on the herbivore, it significantly alters secondary metabolites and has stronger effects on the higher trophic level through changes in the quality of the parasitised host.

Intraspecific difference among herbivore lineages and their host-plant specialization drive the strength of trophic cascades.

Trophic cascades - the indirect effect of predators on non-adjacent lower trophic levels - are important drivers of the structure and dynamics of ecological communities. However, the influence of intraspecific trait variation on the strength of trophic cascade remains largely unexplored, which limits our understanding of the mechanisms underlying ecological networks. Here we experimentally investigated how intraspecific difference among herbivore lineages specialized on different host plants influences trophic cascade strength in a terrestrial tri-trophic system. We found that the occurrence and strength of the trophic cascade are strongly influenced by herbivores' lineage and host-plant specialization but are not associated with density-dependent effects mediated by the growth rate of herbivore populations. Our findings stress the importance of intraspecific heterogeneities and evolutionary specialization as drivers of trophic cascade strength and underline that intraspecific variation should not be overlooked to decipher the joint influence of evolutionary and ecological factors on the functioning of multi-trophic interactions.

When do herbivorous insects compete? A phylogenetic meta-analysis.

When herbivorous insects interact, they can increase or decrease each other's fitness. As it stands, we know little of what causes this variation. Classic competition theory predicts that competition will increase with niche overlap and population density. And classic hypotheses of herbivorous insect diversification predict that diet specialists will be superior competitors to generalists. Here, we test these predictions using phylogenetic meta-analysis. We estimate the effects of diet breadth, population density and proxies of niche overlap: phylogenetic relatedness, physical proximity and feeding-guild membership. As predicted, we find that competition between herbivorous insects increases with population density as well as phylogenetic and physical proximity. Contrary to predictions, competition tends to be stronger between than within feeding guilds and affects specialists as much as generalists. This is the first statistical evidence that niche overlap increases competition between herbivorous insects. However, niche overlap is not everything; complex feeding guild effects indicate important indirect interactions.

Environmental gradients and the evolution of tri-trophic interactions.

Long-standing theory predicts herbivores and predators should drive selection for increased plant defences, such as the specific production of volatile organic compounds for attracting predators near the site of damage. Along elevation gradients, a general pattern is that herbivores and predators are abundant at low elevation and progressively diminish at higher elevations. To determine whether plant adaptation along such a gradient influences top-down control of herbivores, we manipulated soil predatory nematodes, root herbivore pressure and plant ecotypes in a reciprocal transplant experiment. Plant survival was significantly higher for low-elevation plants, but only when in the presence of predatory nematodes. Using olfactometer bioassays, we showed correlated differential nematode attraction and plant ecotype-specific variation in volatile production. This study not only provides an assessment of how elevation gradients modulate the strength of trophic cascades, but also demonstrates how habitat specialisation drives variation in the expression of indirect plant defences.

Seasonal timing in honey bee colonies: phenology shifts affect honey stores and varroa infestation levels.

Increasingly frequent warm periods during winter, which are associated with climate change, may cause mismatches between the colony phenology of the western honey bee, Apis mellifera L., and their floral resources. Warmer winter periods can also affect colony brood rearing activity and consequently the reproduction of the invasive brood parasite Varroa destructor Anderson and Trueman. Until now little is known about the effects of climate change on biotic interactions in such a multitrophic system comprising flowering plants, a pollinator, and its parasite. We performed a reciprocal translocation experiment with honey bee colonies to simulate climate change-induced phenology shifts. Honey bee brood phenology was highly sensitive to environmental conditions in late winter. Colonies in which phenology was experimentally delayed had smaller worker populations in early spring and reduced amounts of stored honey during the following months. During summer, the varroa load in colonies with non-shifted phenology was three times higher than in colonies with delayed phenology. High varroa loads during summer were negatively correlated with worker population growth. Despite a remarkable resilience of colony development to phenology shifts, our results show that mismatches between the phenology of honey bee colonies and flowering plants can affect the build-up of resource stores. Further, an advanced onset of brood rearing activity after hibernation can reinforce the negative impact of the brood parasite V. destructor. We conclude that trade-offs between synchronisation with earlier flower phenology and prolonged brood phases with build-up of varroa populations might constrain the honey bees' capability to adapt to climate warming.

Quantitative measure of fitness in tri-trophic interactions and its influence on diet breadth of insect herbivores.

Herbivore-plant interactions should be studied using a tri-trophic approach, but we lack a quantitative measure of the combined effect of top-down and bottom-up forces on herbivore fitness. We propose the combination of the bi-trophic fitness slopes as a tri-trophic fitness measure. We use the relationship between fitness associated with top-down and bottom-up forces and the frequency of host plant use to calculate the top-down and bottom-up fitness slopes, which we then combine to obtain three possible directions of tri-trophic slopes. A positive tri-trophic slope indicates that herbivores have overall greater tri-trophic fitness on the more frequently used hosts. A null tri-trophic fitness slope indicates that herbivores have similar fitness on all host plants. A negative tri-trophic slope indicates that herbivores have generally lower fitness on the more frequently used hosts. We tested the explanation power of our method using data from the literature that tested herbivore host shifts and experimentally using a generalist herbivore with variable diet breadth across populations. We found that in host shifts, herbivores have higher tri-trophic fitness on the novel host, while in generalist populations, herbivores use most frequently the best host available. We present applications in other research areas and consider the limitations of our approach. Our approach is a first step towards a comprehensive model of multiple selective forces acting on the evolution of interactions.

Dual-guild herbivory disrupts predator-prey interactions in the field.

Plant defenses often mediate whether competing chewing and sucking herbivores indirectly benefit or harm one another. Dual-guild herbivory also can muddle plant signals used by specialist natural enemies to locate prey, further complicating the net impact of herbivore-herbivore interactions in naturally diverse settings. While dual-guild herbivore communities are common in nature, consequences for top-down processes are unclear, as chemically mediated tri-trophic interactions are rarely evaluated in field environments. Combining observational and experimental approaches in the open field, we test a prediction that chewing herbivores interfere with top-down suppression of phloem feeders on Brassica oleracea across broad landscapes. In a two-year survey of 52 working farm sites, we found that parasitoid and aphid densities on broccoli plants positively correlated at farms where aphids and caterpillars rarely co-occurred, but this relationship disappeared at farms where caterpillars commonly co-occurred. In a follow-up experiment, we compared single and dual-guild herbivore communities at four local farm sites and found that caterpillars (P. rapae) caused a 30% reduction in aphid parasitism (primarily by Diaeretiella rapae), and increased aphid colony (Brevicoryne brassicae) growth at some sites. Notably, in the absence of predators, caterpillars indirectly suppressed, rather than enhanced, aphid growth. Amid considerable ecological noise, our study reveals a pattern of apparent commensalism: herbivore-herbivore facilitation via relaxed top-down suppression. This work suggests that enemy-mediated apparent commensalism may override constraints to growth induced by competing herbivores in field environments, and emphasizes the value of placing chemically mediated interactions within their broader environmental and community contexts.

The effect of predator presence on the behavioral sequence from host selection to reproduction in an invulnerable stage of insect prey.

Predator-prey interactions primarily focus on prey life-stages that are consumed. However, animals in less vulnerable life-stages might also be influenced by the presence of a predator, making our understanding of predation-related impacts across all life-stages of prey essential. It has been previously demonstrated that Podisus maculiventris is a voracious predator of eggs and larvae of Leptinotarsa decemlineata, and that larvae will alter their behavior to avoid predation. However, the adult beetles are not readily consumed by P. maculiventris, raising the question of whether they will respond to predators to protect themselves or their offspring. Here, we examine the effect of predation risk by P. maculiventris, on three adult behaviors of L. decemlineata; colonization, oviposition, and feeding, and the resulting impact on host plant damage. In an open-field test, there was no difference in natural beetle colonization between plots with predation risk and control treatments. However, subsequent host plant damage by adult beetles was 63.9% less in predation risk treatments. Over the lifetime of adult beetles in field mesocosms, per capita feeding was 23% less in the predation risk treatment. Beetle oviposition was 37% less in the presence of predators in a short-term, greenhouse assay, and marginally reduced in longer term field mesocosms. Our results indicate that predation risk can drive relatively invulnerable adult herbivores to adjust behaviors that affect themselves (feeding) and their offspring (oviposition). Thus, the full impact of predator presence must be considered across the prey life cycle.

Costs and benefits of omnivore-mediated plant protection: effects of plant-feeding on Salix growth more detrimental than expected.

Predators can decrease herbivore damage to plants, and this is often assumed to be beneficial to plant growth/reproduction without actual quantification. Moreover, previous studies have been biased towards strict carnivores and neglected the role of omnivorous predators in prey-suppression. Here, we examined the costs (reduction in growth) and benefits (increase in growth) of enemy-mediated plant protection via the omnivorous (prey and plant-feeding) Orthotylus marginalis, relative to herbivory by a detrimental insect pest of Salix spp. plantations, the beetle Phratora vulgatissima. In a first experiment, we compared the cost of adult beetle versus omnivore nymph plant-feeding, and assessed the (non-) additive effects of the two types of damage. In a second experiment, we quantified the reduction in plant damage resulting from beetle-egg feeding by omnivorous nymphs and subsequent benefits to plants. We found that plant-feeding by omnivores negatively affected plant growth and this effect was similar to the cost imposed by beetle herbivory. Furthermore, simultaneous damage effects were additive and more detrimental than individual effects. While egg-predation by omnivore nymphs completely prevented beetle damage to plants, there was no difference in plant growth relative to only herbivore-damaged plants and growth was still reduced compared to control plants. Thus, despite herbivore suppression, there was no benefit to plant growth of omnivore-mediated plant protection and the negative effects of omnivore plant-feeding remained. These results are a first for an omnivorous enemy, and provide novel and timely insights on the underlying assumptions of tri-trophic associations and their use for biocontrol of insect pests.

Effects of plant intraspecific diversity across three trophic levels: Underlying mechanisms and plant traits.

PREMISE OF STUDY: Although there is increasing recognition of the effects of plant intraspecific diversity on consumers, the mechanisms by which such effects cascade-up to higher trophic levels remain elusive. METHODS: We evaluated the effects of plant (lima bean, Phaseolus lunatus) intraspecific diversity on a suite of insect herbivores (leaf-chewers, aphids, and seed-eating beetles) and their third trophic-level associates (parasitoids and aphid-tending ants). We established plots of three plants, classified as monocultures of one population source or polycultures with mixtures of three of the four population sources (N = 16 plots per level of diversity). Within each plot, plants were individually placed in pots and canopy contact was prevented, therefore eliminating diversity effects on consumers arising from changes in plant traits due to plant physical interactions. KEY RESULTS: Plant diversity reduced damage by leaf-chewers as well as aphid abundance, and the latter effect in turn reduced ant abundance. In contrast, plant diversity increased the abundance of seed-eating beetles, but did not influence their associated parasitoids. There were no effects of diversity on seed traits potentially associated with seed predation, suggesting that differences in early season herbivory between monocultures and polycultures (a likely mechanism of diversity effects on plants since plant interactions were prevented) did not drive concomitant changes in plant traits. CONCLUSIONS: This study emphasizes that effects of plant intraspecific diversity on consumers are contingent upon differences in associate responses within and among higher trophic levels and suggests possible mechanisms by which such effects propagate up this food web.

Species interactions and plant polyploidy.

Polyploidy is a common mode of speciation that can have far-reaching consequences for plant ecology and evolution. Because polyploidy can induce an array of phenotypic changes, there can be cascading effects on interactions with other species. These interactions, in turn, can have reciprocal effects on polyploid plants, potentially impacting their establishment and persistence. Although there is a wealth of information on the genetic and phenotypic effects of polyploidy, the study of species interactions in polyploid plants remains a comparatively young field. Here we reviewed the available evidence for how polyploidy may impact many types of species interactions that range from mutualism to antagonism. Specifically, we focused on three main questions: (1) Does polyploidy directly cause the formation of novel interactions not experienced by diploids, or does it create an opportunity for natural selection to then form novel interactions? (2) Does polyploidy cause consistent, predictable changes in species interactions vs. the evolution of idiosyncratic differences? (3) Does polyploidy lead to greater evolvability in species interactions? From the scarce evidence available, we found that novel interactions are rare but that polyploidy can induce changes in pollinator, herbivore, and pathogen interactions. Although further tests are needed, it is likely that selection following whole-genome duplication is important in all types of species interaction and that there are circumstances in which polyploidy can enhance the evolvability of interactions with other species.

Do birds see the forest for the trees? Scale-dependent effects of tree diversity on avian predation of artificial larvae.

The enemies hypothesis states that reduced insect herbivory in mixed-species stands can be attributed to more effective top-down control by predators with increasing plant diversity. Although evidence for this mechanism exists for invertebrate predators, studies on avian predation are comparatively rare and have not explicitly tested the effects of diversity at different spatial scales, even though heterogeneity at macro- and micro-scales can influence bird foraging selection. We studied bird predation in an established forest diversity experiment in SW Finland, using artificial larvae installed on birch, alder and pine trees. Effects of tree species diversity and densities on bird predation were tested at two different scales: between plots and within the neighbourhood around focal trees. At the neighbourhood scale, birds preferentially foraged on focal trees surrounded by a higher diversity of neighbours. However, predation rates did not increase with tree species richness at the plot level and were instead negatively affected by tree height variation within the plot. The highest probability of predation was observed on pine, and rates of predation increased with the density of pine regardless of scale. Strong tree species preferences observed may be due to a combination of innate bird species preferences and opportunistic foraging on profitable-looking artificial prey. This study therefore finds partial support for the enemies hypothesis and highlights the importance of spatial scale and focal tree species in modifying trophic interactions between avian predators and insect herbivores in forest ecosystems.

Conifer Monoterpene Chemistry during an Outbreak Enhances Consumption and Immune Response of an Eruptive Folivore.

Changes in the chemical composition of plant defense compounds during herbivory can impact herbivore resource allocation patterns and thereby herbivore survival, growth, and immune response against endoparasitoid infection. Few studies have investigated folivore responses to changes in plant chemistry that occur under outbreak conditions in mature conifer systems. Using data from an earlier observational field study, we carried out laboratory bioassays to test how variation in monoterpenes in piñon pine trees (Pinus edulis, Pinaceae) during an outbreak affects growth, consumption, and immune response of a specialist herbivore, the Southwestern tiger moth (Lophocampa ingens, Arctiidae). Larvae were fed on artificial diets containing four monoterpenes at concentrations that mimicked those observed in undamaged and herbivore-damaged trees in situ during an outbreak. Damaged trees contained 30% lower total monoterpene concentrations, likely reflecting volatile losses as observed in a previous field study Trowbridge et al. (Ecology 95:1591-1603, Trowbridge et al. 2014). Herbivores reared on diets mimicking terpene concentrations in the needles of damaged trees exhibited an approximately 60% increase in consumption relative to larvae reared on diets characteristic of trees without herbivore damage. Higher consumption was accompanied by a 40% increase in immune response with no change in growth rate. These observations suggest preferential resource allocation towards immunity and/or a strong genetic component that determines growth under these conditions. These outcomes, which favor the herbivore, point to: (i) a potential positive feedback mechanism that may increase L. ingens's chance of escaping parasitism during the early phases of an outbreak; and (ii) the important role of monoterpenes in mediating conifer-folivore interactions specifically for P. edulis, which has suffered large-scale drought-induced mortality events exacerbated by the presence of insects.

Plant protection and biotremology: fundamental and applied aspects.

There is overwhelming evidence that synthetic pesticides have a negative impact on the environment and human health, emphasizing the need for novel and sustainable methods for plant protection. A growing body of literature reports that plants interact through substrate-borne vibrations with arthropod pests and mutualistic arthropods that provide biological control and pollination services. Here, we propose a new theoretical framework that integrates insights from biological control, the ecology of fear, and plant-borne vibrations, to address plant-insect interactions and explore new, sustainable opportunities to improve plant health and productivity.