Response of mirid predators to synthetic herbivore‐induced plant volatiles

Zoophytophagous plant bugs feed on plant tissue as a source of water and nutrients, besides feeding on prey. By phytophagy, mirid predators activate plant defense responses through different pathways, resulting, among others, in the release of herbivore‐induced plant volatiles (HIPVs). These compounds could repel herbivores and attract parasitoids and predators, and synthetic versions could potentially be used in biological control. Nevertheless, little is known about the influence of synthetic volatiles on mirid attraction. Using Y‐tube olfactometer trials, we evaluated the responses of Nesidiocoris tenuis (Reuter), Macrolophus pygmaeus (Rambur), and Dicyphus bolivari Lindberg (Hemiptera: Miridae), important natural enemies used to control various greenhouse pests, to 10 synthetic versions of HIPVs released from tomato (Solanum lycopersicum L., Solanaceae) plants induced by N. tenuis and M. pygmaeus. Nesidiocoris tenuis responded to five of the 10 HIPVs, whereas M. pygmaeus and D. bolivari responded to four of the 10 HIPVs. Two green leaf volatiles, (Z)‐3‐hexenyl propanoate and (Z)‐3‐hexenyl acetate, and the ester methyl salicylate (MeSA) were attractive to all three mirid predator species. Our results demonstrate that the volatiles released by tomato plants activated by N. tenuis and M. pygmaeus phytophagy are attractive to their conspecifics and also to D. bolivari. Further studies should evaluate the potential of these compounds to attract predatory mirids in the field.     

Heritability of life‐history traits in the spruce budworm

The heritability of life‐history traits is of particular importance for insects that are very dependent on host conditions. Severe defoliation caused by the spruce budworm negatively impacts its food source, which in turn imposes environmental constraints on the insect. The heritability of those traits can help elucidate this species' evolutionary process. Heritability also helps identify which traits exhibit significant additive variance and can be key to understanding natural selection effects. Individuals were reared under laboratory conditions over three generations on an artificial diet. Heritability was estimated by parent–offspring regression. Fertility and fecundity demonstrated significant heritability followed by larval development, while pupal mass showed minimal heritable variation. These results suggest an important percent of additive variance in life‐history traits. This study contributes to our understanding of the relationship of this forest pest to its environmental conditions. This study also reveals an important genetic architectural structure of life‐history traits in the spruce budworm.     

An ecogenomic analysis of herbivore‐induced plant volatiles in Brassica juncea

Upon herbivore feeding, plants emit complex bouquets of induced volatiles that may repel insect herbivores as well as attract parasitoids or predators. Due to differences in the temporal dynamics of individual components, the composition of the herbivore‐induced plant volatile (HIPV) blend changes with time. Consequently, the response of insects associated with plants is not constant either. Using Brassica juncea as the model plant and generalist Spodoptera spp. larvae as the inducing herbivore, we investigated herbivore and parasitoid preference as well as the molecular mechanisms behind the temporal dynamics in HIPV emissions at 24, 48 and 72 h after damage. In choice tests, Spodoptera litura moth preferred undamaged plants, whereas its parasitoid Cotesia marginiventris favoured plants induced for 48 h. In contrast, the specialist Plutella xylostella and its parasitoid C. vestalis preferred plants induced for 72 h. These preferences matched the dynamic changes in HIPV blends over time. Gene expression analysis suggested that the induced response after Spodoptera feeding is mainly controlled by the jasmonic acid pathway in both damaged and systemic leaves. Several genes involved in sulphide and green leaf volatile synthesis were clearly up‐regulated. This study thus shows that HIPV blends vary considerably over a short period of time, and these changes are actively regulated at the gene expression level. Moreover, temporal changes in HIPVs elicit differential preferences of herbivores and their natural enemies. We argue that the temporal dynamics of HIPVs may play a key role in shaping the response of insects associated with plants.     

Shifting and extension of phenological periods with increasing temperature along elevational transects in southern Bavaria

The impact of global warming on phenology has been widely studied, and almost consistently advancing spring events have been reported. Especially in alpine regions, an extraordinary rapid warming has been observed in the last decades. However, little is known about phenological phases over the whole vegetation period at high elevations. We observed 12 phenological phases of seven tree species and measured air temperature at 42 sites along four transects of about 1000 m elevational range in the years 2010 and 2011 near Garmisch‐Partenkirchen, Germany. Site‐ and species‐specific onset dates for the phenological phases were determined and related to elevation, temperature lapse rates and site‐specific temperature sums. Increasing temperatures induced advanced spring and delayed autumn phases, in which both yielded similar magnitudes. Delayed leaf senescence could therefore have been underestimated until now in extending the vegetation period. Not only the vegetation period, but also phenological periods extended with increasing temperature. Moreover, sensitivity to elevation and temperature strongly depends on the specific phenological phase. Differences between species and groups of species (deciduous, evergreen, high elevation) were found in onset dates, phenological response rates and also in the effect of chilling and forcing temperatures. Increased chilling days highly reduced forcing temperature requirements for deciduous trees, but less for evergreen trees. The problem of shifted species associations and phenological mismatches due to species‐specific responses to increasing temperature is a recent topic in ecological research. Therefore, we consider our findings from this novel, dense observation network in an alpine area of particular importance to deepen knowledge on phenological responses to climate change.     

Warming delays the phenological sequences of an autumn‐flowering invader

Phenology can play an important role in driving plant invasions; however, little is known about how climate warming, nitrogen (N) deposition, and invasion stages influence the phenological sequences of autumn‐flowering invaders in a subtropical climate. Accordingly, we conducted an experiment to address the effects of experimental warming, N‐addition, and community types on the first inflorescence buds, flowering, seed‐setting, and dieback of invasive Solidago canadensis. Warming delayed the onset of first inflorescence buds, flowering, seed‐setting, and dieback; N‐addition did not influence these four phenophases; community types influenced the onset of first seed‐setting but not the other phenological phases. Seed‐setting was more sensitive to experimental manipulations than the other phenophases. The onset of first inflorescence buds, flowering, and seed‐setting was marginally or significantly correlated with ramet height but not ramet numbers. Our results suggest that future climate warming might delay the phenological sequences of autumn‐flowering invaders and some phenophases can shift with invasion stages.     

Caterpillar‐induced plant volatiles attract conspecific herbivores and a generalist predator

Plants release volatiles in response to caterpillar feeding that attracts natural enemies of the herbivores, a tritrophic interaction which has been considered to be an indirect plant defence against herbivores. On the other hand, the caterpillar‐induced plant volatiles have been reported to either repel or attract conspecific adult herbivores. This work was undertaken to investigate the response of both herbivores and natural enemies to caterpillar‐induced plant volatiles in apple orchards. We sampled volatile compounds emitted from uninfested apple trees, and apple trees infested with generalist herbivore the pandemis leafroller moth, Pandemis pyrusana (Lepidoptera, Tortricidae) larvae using headspace collection and analysed by gas chromatography/mass spectrometry. Infested apple trees uniquely release six compounds (benzyl alcohol, phenylacetonitrile, phenylacetaldehyde, 2‐phenylethanol, indole and (E)‐nerolidol). These compounds were tested on two species of herbivores and one predator in apple orchards. Binary blends of phenylacetonitrile + acetic acid or 2‐phenylethanol + acetic acid attracted a large number of conspecific male and female adult herbivores. The response of pandemis leafroller to herbivore‐induced plant volatiles (HIPVs) was so pronounced that over one thousand and seven hundred conspecific male and female adult herbivores were caught in traps baited with HIPVs in three‐day trapping period. In addition, significantly higher number of male and female obliquebanded leafroller, Choristoneura rosaceana (Lepidoptera, Tortricidae), was caught in traps baited a binary blend of 2‐phenylethanol + acetic acid, or a ternary blend contains 2‐phenylethanol and phenylacetonitrile + acetic acid. This result challenges the current paradigm hypothesized that HIPVs repel herbivores and question the indirect defensive function proposed for these compounds. On the other hand, a ternary blend of phenylacetonitrile and 2‐phenylethanol + acetic acid attracted the largest numbers of the general predator, the common green lacewing, Chrysoperla plorabunda. To our knowledge, this is the first record of the direct attraction of conspecific adult herbivores as well as a predator to the caterpillar‐induced plant volatiles in the field.     

Responses of Thrips tabaci to odours of herbivore‐induced cotton seedlings

Herbivore‐induced changes in plants have been widely viewed as defensive responses against further insect attack. However, changes in plants as a consequence of herbivore feeding can elicit various responses in herbivores; these are variable, context dependent, and often unpredictable. In this laboratory study, the responses of Thrips tabaci Lindeman (Thysanoptera: Thripidae) to volatiles emitted by intact and herbivore‐damaged or mechanically damaged cotton seedlings [Gossypium hirsutum L. (Malvaceae)] were investigated in dual‐choice olfactometer assays. Thrips tabaci showed increased attraction to seedlings subject to foliar mechanical damage and those with foliar damage inflicted by conspecifics or Tetranychus urticae Koch (Acari: Tetranychidae), upon which it preys. However, T. tabaci did not discriminate between intact seedlings and those with foliar damage inflicted by Helicoverpa armigera Hübner (Lepidoptera: Noctuidae), two other species of thrips, Frankliniella schultzei Trybom and Frankliniella occidentalis Pergrande (Thysanoptera: Thripidae), or those with root damage inflicted by Tenebrio molitor L. (Coleoptera: Tenebrionidae). Attraction of T. tabaci was also affected by herbivore density on damaged plants. That is, seedlings damaged by higher densities of T. urticae or T. tabaci were more attractive than seedlings damaged by lower densities of the corresponding arthropod. Although attracted to plants damaged by conspecifics or T. urticae, T. tabaci showed greater attraction to seedlings damaged by T. urticae than to seedlings damaged by conspecifics. Results are discussed in the context of the responses of F. schultzei and F. occidentalis to herbivore‐induced cotton seedlings, highlighting the complexity, variability, and unpredictability of the responses of even closely related species of insects to plants under herbivore attack.     

Warming, plant phenology and the spatial dimension of trophic mismatch for large herbivores

Temporal advancement of resource availability by warming in seasonal environments can reduce reproductive success of vertebrates if their own reproductive phenology does not also advance with warming. Indirect evidence from large-scale analyses suggests, however, that migratory vertebrates might compensate for this by tracking phenological variation across landscapes. Results from our two-year warming experiment combined with seven years of observations of plant phenology and offspring production by caribou (Rangifer tarandus) in Greenland, however, contradict evidence from large-scale analyses. At spatial scales relevant to the foraging horizon of individual herbivores, spatial variability in plant phenology was reduced--not increased--by both experimental and observed warming. Concurrently, offspring production by female caribou declined with reductions in spatial variability in plant phenology. By highlighting the spatial dimension of trophic mismatch, these results reveal heretofore unexpected adverse consequences of climatic warming for herbivore population ecology.     

Predicting bird phenology from space: satellite‐derived vegetation green‐up signal uncovers spatial variation in phenological synchrony between birds and their environment

Population‐level studies of how tit species (Parus spp.) track the changing phenology of their caterpillar food source have provided a model system allowing inference into how populations can adjust to changing climates, but are often limited because they implicitly assume all individuals experience similar environments. Ecologists are increasingly using satellite‐derived data to quantify aspects of animals' environments, but so far studies examining phenology have generally done so at large spatial scales. Considering the scale at which individuals experience their environment is likely to be key if we are to understand the ecological and evolutionary processes acting on reproductive phenology within populations. Here, we use time series of satellite images, with a resolution of 240 m, to quantify spatial variation in vegetation green‐up for a 385‐ha mixed‐deciduous woodland. Using data spanning 13 years, we demonstrate that annual population‐level measures of the timing of peak abundance of winter moth larvae (Operophtera brumata) and the timing of egg laying in great tits (Parus major) and blue tits (Cyanistes caeruleus) is related to satellite‐derived spring vegetation phenology. We go on to show that timing of local vegetation green‐up significantly explained individual differences in tit reproductive phenology within the population, and that the degree of synchrony between bird and vegetation phenology showed marked spatial variation across the woodland. Areas of high oak tree (Quercus robur) and hazel (Corylus avellana) density showed the strongest match between remote‐sensed vegetation phenology and reproductive phenology in both species. Marked within‐population variation in the extent to which phenology of different trophic levels match suggests that more attention should be given to small‐scale processes when exploring the causes and consequences of phenological matching. We discuss how use of remotely sensed data to study within‐population variation could broaden the scale and scope of studies exploring phenological synchrony between organisms and their environment.     

Use of herbivore‐induced plant volatiles as search cues by Tiphia vernalis and Tiphia popilliavora to locate their below‐ground scarabaeid hosts

Japanese beetle, Popillia japonica Newman, and oriental beetle, Anomala orientalis (Waterhouse) (both Coleoptera: Scarabaeidae) are considered invasive species and have been reported as key pests of urban landscapes in the Northeastern USA. Tiphia vernalis Rohwer and Tiphia popilliavora Rohwer (Hymenoptera: Tiphiidae) were introduced as biocontrol agents against these beetles. These parasitic wasps burrow into the soil and search for grubs. When a host is found, the wasp attaches an egg in a location that is specific for the wasp species. It is unknown if these wasps can detect patches of concealed hosts from a distance above ground and what role, if any, herbivore‐induced plant volatiles play in their host location. This study evaluated the responses of female T. vernalis and T. popilliavora to grub‐infested and healthy plants in Y‐tube olfactometer bioassays. Also the effect of root herbivory on the composition of turfgrass (Poaceae) volatile profiles was investigated by collecting volatiles from healthy and grub‐infested grasses. Tiphia wasps were highly attracted to volatiles emitted by grub‐infested tall fescue (Festuca arundinacea Schreb.) and Kentucky bluegrass (Poa pratensis L.) over healthy grasses. In contrast, wasps did not exhibit a significant preference for grub‐infested perennial ryegrass (Lolium perenne L.) as compared with the control plants. The terpene levels emitted by grub‐infested Kentucky bluegrass and tall fescue were greater than that of control plants. Low levels of terpenes were observed for both test and control perennial ryegrass. The elevated levels of terpenes emitted by grub‐infested Kentucky bluegrass and tall fescue coincided with the attractiveness to the tiphiid wasps. Here, we provide evidence that plant exposure to root‐feeding insects P. japonica and A. orientalis resulted in an increase in terpenoid levels in turfgrasses, which strongly attracts their above‐ground parasitoids.     

Birds exploit herbivore‐induced plant volatiles to locate herbivorous prey

Arthropod herbivory induces plant volatiles that can be used by natural enemies of the herbivores to find their prey. This has been studied mainly for arthropods that prey upon or parasitise herbivorous arthropods but rarely for insectivorous birds, one of the main groups of predators of herbivorous insects such as lepidopteran larvae. Here, we show that great tits (Parus major) discriminate between caterpillar‐infested and uninfested trees. Birds were attracted to infested trees, even when they could not see the larvae or their feeding damage. We furthermore show that infested and uninfested trees differ in volatile emissions and visual characteristics. Finally, we show, for the first time, that birds smell which tree is infested with their prey based on differences in volatile profiles emitted by infested and uninfested trees. Volatiles emitted by plants in response to herbivory by lepidopteran larvae thus not only attract predatory insects but also vertebrate predators.     

Temporal changes in olfactory and oviposition responses of the diamondback moth to herbivore‐induced host plants

Temporal changes in the pre‐ and post‐alighting responses of mated female diamondback moth, Plutella xylostella L. (Lepidoptera: Plutellidae), to two species of Brassica (Brassicaceae) host plants induced by larval feeding were studied using olfactometer and oviposition assays. Females displayed strong olfactory and oviposition preferences for herbivore‐induced common cabbage (Brassica oleracea var. capitata L. cv. sugarloaf) plants over intact plants; these preferences decreased with time and disappeared by the 7th day after induction. In herbivore‐induced common cabbage plants, eggs were clustered near feeding damage on the younger leaves (leaves 5–7), whereas in intact plants, eggs were clustered on the stem and lower leaves (leaves 1–4) . However, as the time interval between larval feeding and oviposition increased, more eggs were laid on the lower leaves of induced plants. This demonstrates a change in egg distribution from the pattern associated with induced plants to that associated with intact plants. In contrast, females displayed strong olfactory and oviposition preferences for intact Chinese cabbage [Brassica rapa ssp. pekinensis (Lour.) Hanelt cv. Wombok] plants over induced plants; these preferences decreased with time and disappeared by the 5th day after induction. More eggs were laid on the upper leaves (leaves 4–6) than on the lower leaves (leaves 1–3) of intact Chinese cabbage plants at first, but the distribution changed over time until there were no significant differences in the egg count between upper and lower leaves by the 4th day post induction. For both host plant species, pre‐alighting responses of moths were reliable indicators of post‐alighting responses on the first 2 days post induction. The results suggest that temporal changes in a plant's profile (chemical or otherwise) following herbivory may influence attractiveness to an insect herbivore and be accompanied by changes in olfactory and oviposition preferences.     

Warming off southwestern Japan linked to distributional shifts of subtidal canopy‐forming seaweeds

To assess distributional shifts of species in response to recent warming, historical distribution records are the most requisite information. The surface seawater temperature (SST) of Kochi Prefecture, southwestern Japan on the western North Pacific, has significantly risen, being warmed by the Kuroshio Current. Past distributional records of subtidal canopy‐forming seaweeds (Laminariales and Fucales) exist at about 10‐year intervals from the 1970s, along with detailed SST datasets at several sites along Kochi's >700 km coastline. In order to provide a clear picture of distributional shifts of coastal marine organisms in response to warming SST, we observed the present distribution of seaweeds and analyzed the SST datasets to estimate spatiotemporal SST trends in this coastal region. We present a large increase of 0.3°C/decade in the annual mean SST of this area over the past 40 years. Furthermore, a comparison of the previous and present distributions clearly showed the contraction of temperate species' distributional ranges and expansion of tropical species' distributional ranges in the seaweeds. Although the main temperate kelp Ecklonia (Laminariales) had expanded their distribution during periods of cooler SST, they subsequently declined as the SST warmed. Notably, the warmest SST of the 1997–98 El Niño Southern Oscillation event was the most likely cause of a widespread destruction of the kelp populations; no recovery was found even in the present survey at the formerly habitable sites where warm SSTs have been maintained. Temperate Sargassum spp. (Fucales) that dominated widely in the 1970s also declined in accordance with recent warming SSTs. In contrast, the tropical species, S. ilicifolium, has gradually expanded its distribution to become the most conspicuously dominant among the present observations. Thermal gradients, mainly driven by the warming Kuroshio Current, are presented as an explanation for the successive changes in both temperate and tropical species' distributions.     

Climate variation alters the synchrony of host–parasitoid interactions

Observed changes in mean temperature and increased frequency of extreme climate events have already impacted the distributions and phenologies of various organisms, including insects. Although some research has examined how parasitoids will respond to colder temperatures or experimental warming, we know relatively little about how increased variation in temperature and humidity could affect interactions between parasitoids and their hosts. Using a study system consisting of emerald ash borer (EAB), Agrilus planipennis, and its egg parasitoid Oobius agrili, we conducted environmentally controlled laboratory experiments to investigate how increased seasonal climate variation affected the synchrony of host–parasitoid interactions. We hypothesized that increased climate variation would lead to decreases in host and parasitoid survival, host fecundity, and percent parasitism (independent of host density), while also influencing percent diapause in parasitoids. EAB was reared in environmental chambers under four climate variation treatments (standard deviations in temperature of 1.24, 3.00, 3.60, and 4.79°C), while O. agrili experiments were conducted in the same environmental chambers using a 4 × 3 design (four climate variation treatments × 3 EAB egg densities). We found that EAB fecundity was negatively associated with temperature variation and that temperature variation altered the temporal egg laying distribution of EAB. Additionally, even moderate increases in temperature variation affected parasitoid emergence times, while decreasing percent parasitism and survival. Furthermore, percent diapause in parasitoids was positively associated with humidity variation. Our findings indicate that relatively small changes in the frequency and severity of extreme climate events have the potential to phenologically isolate emerging parasitoids from host eggs, which in the absence of alternative hosts could lead to localized extinctions. More broadly, these results indicate how climate change could affect various life history parameters in insects, and have implications for consumer–resource stability and biological control.     

Lengthening of the duration of xylogenesis engenders disproportionate increases in xylem production

In cold climates, the expected global warming will lead to earlier cambial resumptions in spring, with a resultant lengthening of the growing season but unknown consequences on forest productivity. The phenological traits of cambium activity and xylem formation were analyzed at a short time scale along a thermal gradient represented by an alti‐latitudinal range from the 48th to 53rd parallels and covering the whole closed black‐spruce [Picea mariana (Mill.) BSP] forest in Quebec, Canada. A hypothesis was tested that warmer temperatures influence cambium phenology, allowing longer duration and higher intensity of growth, and resulting in proportionally increased xylem production. From April to October 2012, cell division in cambium and post‐cambial differentiation of xylem were observed on anatomical sections obtained from microcores collected weekly from the stem of fifty trees. The southern and warmer site was characterized by the highest radial growth, which corresponded to both the highest rates and longest durations of cell production. The differences in terms of xylem phenology and growth were marginal between the other sites. Xylem growth was positively correlated with rate and duration of cell production, with the latter explaining most variability in growth. Within the range analyzed, the relationship between temperature and most phenological phases of xylogenesis was linear. On the contrary, temperature was related with cell production according to an exponential pattern. Periods of xylogenesis of 14 days longer (+13.1%) corresponded to a massive increase in cell production (33 cells, +109%). This disproportionate change occurred at a May–September average temperature of ca. 14 °C and a snow‐free period of 210–235 days. At the lower boundary of the distribution of black spruce, small environmental changes allowing marginal lengthening of the period of cell division could potentially lead to disproportionate increases in xylem cell production, with substantial consequences for the productivity of this boreal species.     

AMP‐activated protein kinase counteracts brain‐derived neurotrophic factor‐induced mammalian target of rapamycin complex 1 signaling in neurons

Growth factors and nutrients, such as amino acids and glucose, regulate mammalian target of rapamycin complex 1 (mTORC1) signaling and subsequent translational control in a coordinated manner. Brain‐derived neurotrophic factor (BDNF), the most prominent neurotrophic factor in the brain, activates mTORC1 and induces phosphorylation of its target, p70S6 kinase (p70S6K), at Thr389 in neurons. BDNF also increases mammalian target of rapamycin‐dependent novel protein synthesis in neurons. Here, we report that BDNF‐induced p70S6K activation is dependent on glucose, but not amino acids, sufficiency in cultured cortical neurons. AMP‐activated protein kinase (AMPK) is the molecular background to this specific nutrient dependency. Activation of AMPK, which is induced by glucose deprivation, treatment with pharmacological agents such as 2‐Deoxy‐d ‐glucose, metformin, and 5‐aminoimidazole‐4‐carboxamide ribonucleoside or forced expression of a constitutively active AMPKα subunit, counteracts BDNF‐induced phosphorylation of p70S6K and enhanced protein synthesis in cortical neurons. These results indicate that AMPK inhibits the effects of BDNF on mTORC1‐mediated translation in neurons.

     

Climate change affecting oil palm agronomy,and oil palm cultivation increasing climate change,require amelioration

Palm oil is used in various valued commodities and is a large global industry worth over US$ 50 billion annually. Oil palms (OP) are grown commercially in Indonesia and Malaysia and other countries within Latin America and Africa. The large‐scale land‐use change has high ecological, economic, and social impacts. Tropical countries in particular are affected negatively by climate change (CC) which also has a detrimental impact on OP agronomy, whereas the cultivation of OP increases CC. Amelioration of both is required. The reduced ability to grow OP will reduce CC, which may allow more cultivation tending to increase CC, in a decreasing cycle. OP could be increasingly grown in more suitable regions occurring under CC. Enhancing the soil fauna may compensate for the effect of CC on OP agriculture to some extent. The effect of OP cultivation on CC may be reduced by employing reduced emissions from deforestation and forest degradation plans, for example, by avoiding illegal fire land clearing. Other ameliorating methods are reported herein. More research is required involving good management practices that can offset the increases in CC by OP plantations. Overall, OP‐growing countries should support the Paris convention on reducing CC as the most feasible scheme for reducing CC.     

Plant chemistry underlies herbivore‐mediated inbreeding depression in nature

The cost of inbreeding (inbreeding depression, ID) is an important variable in the maintenance of reproductive variation. Ecological interactions such as herbivory could modulate this cost, provided that defence traits harbour deleterious mutations and herbivores are responsible for differences in fitness. In the field, we manipulated the presence of herbivores on experimentally inbred and outcrossed plants of Solanum carolinense (horsenettle) for three years. Damage was greater on inbred plants, and ID for growth and fitness was significantly greater under herbivory. Inbreeding reduced phenolic expression both qualitatively (phytochemical diversity) and quantitatively, indicating deleterious load at loci related to the biosynthesis of defence compounds. Our results indicate that inbreeding effects on plant–herbivore interactions are mediated by changes to functional plant metabolites, suggesting that variation in inbreeding could be a predictor of defence trait variation. The magnitude of herbivore‐mediated, ecological ID indicates that herbivores could maintain outcrossing mating systems in nature.