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Upon parasitism, many parasitoids inject symbiotic viruses and venom into their host. Thereby, they modify the hosts physiology including its saliva composition and, in turn, plant perception of herbivory. It has been hypothesised that parasitoids manipulate plant responses to increase their host performance and maximise their own fitness. However, it is still unclear whether parasitoids are under selective pressure to increase plant quality or whether indirect changes in plants due to parasitism are a by-product of the physiological changes induced in their host. We tested whether the parasitoids Hyposoter ebeninus and Cotesia glomerata manipulate induced plant responses through the host Pieris brassicae caterpillars to increase their own performance. During their entire lifespan, parasitised herbivores were fed with leaf material of Brassica oleracea plants that were left untreated or continuously exposed to feeding by either unparasitised, conspecific or heterospecific parasitised caterpillars. We measured the development time, weight, clutch size and mortality of parasitoids that emerged from caterpillars as proxy for their performance. Both parasitoid species did not perform significantly better when their host was fed with leaves from plants continuously induced by a conspecific parasitoid relative to unparasitised caterpillars. However, parasitoid species asymmetrically affected each other's performance through plant-mediated interactions. Our results do not support the hypothesis of parasitoids manipulating plant responses for their own benefit, suggesting that indirect plant-mediated interactions among parasitoids may be a by-product of host manipulation. However, our work confirms the significance of parasitoid-induced plant-mediated interactions in insect communities that to date are still understudied.

The phenotypic plasticity of flowering plants in response to herbivore damage to vegetative tissues can affect plant interactions with flower-feeding organisms. Such induced systemic responses are probably regulated by defence-related phytohormones that signal flowers to alter secondary chemistry that affects resistance to florivores. Current knowledge on the effects of damage to vegetative tissues on plant interactions with florivores and the underlying mechanisms is limited. We compared the preference and performance of two florivores on flowering Brassica nigra plants damaged by one of three herbivores feeding from roots or leaves. To investigate the underlying mechanisms, we quantified expression patterns of marker genes for defence-related phytohormonal pathways, and concentrations of phytohormones and glucosinolates in buds and flowers. Florivores displayed contrasting preferences for plants damaged by herbivores feeding on roots and leaves. Chewing florivores performed better on plants damaged by folivores, but worse on plants damaged by the root herbivore. Chewing root and foliar herbivory led to specific induced changes in the phytohormone profile of buds and flowers. This resulted in increased glucosinolate concentrations for leaf-damaged plants, and decreased glucosinolate concentrations for root-damaged plants. The outcome of herbivore–herbivore interactions spanning from vegetative tissues to floral tissues is unique for the inducing root/leaf herbivore and receiving florivore combination.

Species-specific responses to the environment can moderate the strength of interactions between plants, herbivores and parasitoids. However, the ways in which characteristics of plants, such as genotypic variation in herbivore induced volatiles (HIPVs) that attract parasitoids, affect trophic interactions in different contexts of plant patch size and plant neighbourhood is not well understood. We conducted a factorial field experiment with white cabbage Brassica oleracea accessions that differ in the attractiveness of their HIPVs for parasitoids, in the context of different patch sizes and presence or absence of surrounding Brassica nigra plants. Parasitism rates of experimentally introduced Pieris brassicae caterpillars and the presence of naturally occurring Pieris spp. caterpillars in the plots were assessed throughout the growing season. The abundance of Pieris caterpillars was neither affected by cabbage accession nor plot size. Later in the season, when B. nigra plants had senesced, fewer caterpillars were found on cabbage plants in plots with a B. nigra border. Parasitism rates fluctuated over the season, and were not affected by plot size. However, the B. nigra border negatively affected parasitism rates on the accession that is less attractive to the parasitoid Cotesia glomerata, but not on the more attractive accession. Our results show that plant variation in HIPVs can differentially influence herbivores and parasitoids depending on characteristics of the surrounding vegetation context. These findings underscore the importance of considering the interaction between focal plant traits and neighbourhood context to reliably predict trophic cascades.

Plants balance the allocation of resources between growth and defence to optimize fitness in a competitive environment. Perception of neighbour‐detection cues, such as a low ratio of red to far‐red (R:FR) radiation, activates a suite of shade‐avoidance responses that include stem elongation and upward leaf movement, whilst simultaneously downregulating defence. This downregulation is hypothesized to benefit the plant either by mediating the growth‐defence balance in favour of growth in high plant densities or, alternatively, by mediating defence of individual leaves such that those most photosynthetically productive are best protected. To test these hypotheses, we used a 3D functional–structural plant model of Brassica nigra that mechanistically simulates the interactions between plant architecture, herbivory, and the light environment. Our results show that plant‐level defence expression is a strong determinant of plant fitness and that leaf‐level defence mediation by R:FR can provide a fitness benefit in high densities. However, optimal plant‐level defence expression does not decrease monotonically with plant density, indicating that R:FR mediation of defence alone is not enough to optimize defence between densities. Therefore, assessing the ecological significance of R:FR‐mediated defence is paramount to better understand the evolution of this physiological linkage and its implications for crop breeding.

Plants need to deal with antagonists, such as herbivores, while maintaining interactions with mutualists, such as pollinators that help plants to maximize their reproductive output. Although many plant species have inducible defences to save metabolic costs of defence in the absence of herbivores, plant responses induced by herbivore attack can have ecological costs. For example, herbivore-induced responses can affect flower traits and alter interactions with flower visitors. Such plant-mediated interactions between herbivores and flower visitors can affect plant reproductive output. Current knowledge on the generality and specificity of plant-mediated herbivore-flower-visitor interactions and its consequences for plant fitness is limited. In this study, we investigated whether a broad range of herbivores feeding on the annual plant Brassica nigra affect interactions with flower visitors, whether the direction of interactions is predicted by the feeding modes (chewing and sap-feeding) and sites (above- and belowground) of the herbivores, and whether it results in fitness consequences for the plant. Our results show that attack of B. nigra by a range of different herbivores influenced plant interactions with mutualist pollinators and an antagonist florivore, the pollen beetle Meligethes aeneus. Pollinator community composition was affected by herbivory, whereas overall pollinator attraction was maintained. Pollinator community composition of uninfested plants differed from that of chewing and root herbivore-infested plants. Main responders in the pollinator communities to changes induced by herbivory were syrphid flies, bumblebees, and solitary bees. Although the preference of pollen beetle adults was not affected by herbivory, beetle larvae performed best on plants infested with the nematode Heterodera schachtii. The changes in pollinator community composition and syrphid fly visitation can explain the observed increase in seed set of root herbivore-infested plants. Interactions of flowering B. nigra plants with mutualist and antagonist insects are well integrated and conflicting interactions do not reduce reproductive output. Our results suggest some degree of specificity in herbivore-flower-visitor interactions with consequences for plant fitness. Specificity of plant responses were determined at the species level as well as the herbivore functional group level, and differed depending on the flower visitor. Because plant reproduction was affected by indirect plant-mediated interactions, these can potentially result in selection on plant strategies to optimize growth, defence and reproduction. A plain language summary is available for this article.

1. Parasitoids are known to utilise learning of herbivore-induced plant volatiles (HIPVs) when foraging for their herbivorous host. In natural situations these hosts share food plants with other, non-suitable herbivores (non-hosts). Simultaneous infestation of plants by hosts and non-hosts has been found to result in induction of HIPVs that differ from host-infested plants. Each non-host herbivore may have different effects on HIPVs when sharing the food plant with hosts, and thus parasitoids may learn that plants with a specific non-host herbivore also contain the host. 2. This study investigated the adaptive nature of learning by a foraging parasitoid that had acquired oviposition experience on a plant infested with both hosts and different non-hosts in the laboratory and in semi-field experiments. 3. In two-choice preference tests, the parasitoid Cotesia glomerata shifted its preference towards HIPVs of a plant-host-non-host complex previously associated with an oviposition experience. It could, indeed, learn that the presence of its host is associated with HIPVs induced by simultaneous feeding of its host Pieris brassicae and either the non-host caterpillar Mamestra brassicae or the non-host aphid Myzus persicae. However, the learned preference found in the laboratory did not translate into parasitisation preferences for hosts accompanying non-host caterpillars or aphids in a semi-field situation. 4. This paper discusses the importance of learning in parasitoid foraging, and debates why observed learned preferences for HIPVs in the laboratory may cancel out under some field experimental conditions.

1. Plants are frequently under attack by multiple insect herbivores, which may interact indirectly through herbivore-induced changes in the plant's phenotype. The identity, order, and timing of herbivore arrivals may influence the outcome of interactions between two herbivores. How these aspects affect, in turn, subsequently arriving herbivores that feed on double herbivore-induced plants has not been widely investigated. 2. This study tested whether the order and timing of arrival of two inducing herbivores from different feeding guilds affected the preference and performance of a subsequently arriving third herbivore, caterpillars of Mamestra brassicae L. (Lepidoptera: Noctuidae). Aphids [Brevicoryne brassicae L. (Hemiptera: Aphididae)] and caterpillars [Plutella xylostella L. (Lepidoptera: Yponomeutidae)] were introduced onto wild Brassica oleracea L. (Brassicaceae) plants in different sequences and with different arrival times. The effects of these plant treatments on M. brassicae caterpillars were assessed in pairwise preference tests and no-choice performance tests. 3. The caterpillars of M. brassicae preferred to feed from undamaged plants rather than double herbivore-induced plants. Compared with undamaged plants, they preferred plant material on which aphids had arrived first followed by caterpillars, whereas they avoided plant material with the reverse order of herbivore arrival. Performance of the caterpillars increased with increasing arrival time between herbivore infestations in double herbivore-induced plants. Although M. brassicae grew faster on plants induced by aphids than on those induced by caterpillars alone, its performance was not affected by the order of previous herbivore arrival. 4. These results imply that the timing of colonisation by multiple herbivores determines the outcome of plant-mediated herbivore-herbivore interactions.

Chemical communication is ubiquitous. The identification of conserved structural elements in visual and acoustic communication is well established, but comparable information on chemical communication displays (CCDs) is lacking. We assessed the phenotypic integration of CCDs in a meta‐analysis to characterize patterns of covariation in CCDs and identified functional or biosynthetically constrained modules. Poorly integrated plant CCDs (i.e. low covariation between scent compounds) support the notion that plants often utilize one or few key compounds to repel antagonists or to attract pollinators and enemies of herbivores. Animal CCDs (mostly insect pheromones) were usually more integrated than those of plants (i.e. stronger covariation), suggesting that animals communicate via fixed proportions among compounds. Both plant and animal CCDs were composed of modules, which are groups of strongly covarying compounds. Biosynthetic similarity of compounds revealed biosynthetic constraints in the covariation patterns of plant CCDs. We provide a novel perspective on chemical communication and a basis for future investigations on structural properties of CCDs. This will facilitate identifying modules and biosynthetic constraints that may affect the outcome of selection and thus provide a predictive framework for evolutionary trajectories of CCDs in plants and animals., The project was supported by the Graduate School ‘Evolutionary Networks: Organisms, Reactions, Molecules’ (E‐Norm) of the Heinrich‐Heine‐University, Düsseldorf, Germany, the Max Planck Society and the Deutsche Forschungsgemeinschaft (DFG JU 2856/1‐1 and DFG JU 2856/2‐2).

Extensive research has been conducted to reveal how species diversity affects ecosystem functions and services. Yet, consequences of diversity loss for ecosystems as a whole as well as for single community members are still difficult to predict. Arthropod communities typically are species-rich, and their species interactions, such as those between herbivores and their predators or parasitoids, may be particularly sensitive to changes in community composition. Parasitoids forage for herbivorous hosts by using herbivore-induced plant volatiles (indirect cues) and cues produced by their host (direct cues). However, in addition to hosts, non-suitable herbivores are present in a parasitoid's environment which may complicate the foraging process for the parasitoid. Therefore, ecosystem changes in the diversity of herbivores may affect the foraging efficiency of parasitoids. The effect of herbivore diversity may be mediated by either species numbers per se, by specific species traits, or by both. To investigate how diversity and identity of non-host herbivores influence the behaviour of parasitoids, we created environments with different levels of non-host diversity. On individual plants in these environments, we complemented host herbivores with 1–4 non-host herbivore species. We subsequently studied the behaviour of the gregarious endoparasitoid Cotesia glomerata L. (Hymenoptera: Braconidae) while foraging for its gregarious host Pieris brassicae L. (Lepidoptera: Pieridae). Neither non-host species diversity nor non-host identity influenced the preference of the parasitoid for herbivore-infested plants. However, after landing on the plant, non-host species identity did affect parasitoid behaviour, whereas non-host diversity did not. One of the non-host species, Trichoplusia ni Hübner (Lepidoptera: Noctuidae), reduced the time the parasitoid spent on the plant as well as the number of hosts it parasitized. We conclude that non-host herbivore species identity has a larger influence on C. glomerata foraging behaviour than non-host species diversity. Our study shows the importance of species identity over species diversity in a multitrophic interaction of plants, herbivores, and parasitoids.

Changes in the arthropod community structure can be attributed to differences in constitutively expressed plant traits or those that change depending on environmental conditions such as herbivory. Early-season herbivory may have community-wide effects on successive insect colonisation of host plants and the identity of the initially inducing insect may determine the direction and strength of the effects on the dynamics and composition of the associated insect community. 2. Previous studies have addressed the effect of early infestation with a chewing herbivore. In the present study, the effect of early infestation was investigated with a phloem-feeding aphid [Brevicoryne brassicae L. (Hemiptera, Aphididae)] on the insect community associated with three wild cabbage (Brassica oleracea L.) populations, which are known to differ in defence chemistry, throughout the season in field experiments. 3. Aphid infestation had asymmetric effects on the associated insect community and only influenced the abundance of the natural enemies of aphids, but not that of chewing herbivores and their natural enemies. The effect size of aphid infestation further depended on the cabbage population. 4. Aphid feeding has been previously reported to promote host-plant quality for chewing herbivores, which has been attributed to antagonism between the two major defence signalling pathways controlled by the hormones salicylic acid (SA) and jasmonic acid (JA), respectively. Our results show no effects of early infestation by aphids on chewing herbivores, suggesting the absence of long-term JA–SA antagonism. 5. Investigating the effects of the identity of an early-season coloniser and genotypic variation among plant populations on insect community dynamics are important in understanding insect–plant community ecology.

1. In nature, competitive interactions occur when different species exploit similar niches. Parasitic wasps (parasitoids) often have narrow host ranges and need to cope with competitors that use the same host species for development of their offspring. When larvae of different parasitoid species develop in the same host, this leads to intrinsic and often contest com.petition. Thus far, m.ost studies on intrinsic competition have focused on primary parasitoids. However, competition among primary.hyperparasitoids, parasitic wasps that use primary parasitoids as a host, has been little studied. 2. This study investigated intrinsic competition between two primary hyperparasitoids, the gregarious Baryscapus galactopus and the solitary Mesochoru,s gemellu,s, which lay their eggs in primary parasitoid larvae of tesla rubecula, while those in turn are developing inside their herbivore host, Pieris rapae. The aims were to identify: (i) which hyperparasitoid is the superior competitor; and (ii) whether oviposition sequence affects the outcome of intrinsic competition. 3. The results show that B. galactopus won 70% of contests when the two hyperparasitoids parasitised the host at the same time, and 90% when 13. galactopus oviposited first. When M. gemellus had a 48 h head start, the two hyperparasitoids had an equal chance to win the competition. This suggests that M. galactopus is an intrinsically superior competitor to M. gemellu,s, Moreover, the outcome of competition is affected by time lags in oviposition events. 4. In contrast to what has been reported for primary parasitoids, we found that a. gregarious hyperparasitoid species had a competitive advantage over a solitary species.

Plant–insect interactions typically take place in complex settings of interactions among multiple trophic levels as well as multiple species in each trophic level. The complex interaction network may strongly impact on extrapolations of resistance traits to have a defensive function. For example, the induced response plants express to their current attacker often enhances resistance to that attacker, but may make a plant more susceptible to attack by another herbivore. Hence, the defensive function or plant fitness benefit of the response to a single attacker may be misinterpreted from pairwise interactions. Moreover, plant physiological responses to a first stress by herbivory may hamper the response to a second stress and lead to conclusions of maladaptation in plant defence responses. In light of the entire community of attackers and beneficial organisms the plant interacts with, the susceptibility to some attackers may be a consequence of adaptations that reduce fitness costs of herbivory when considering the full sweep of species that affect plant fitness. A similar argumentation may apply for indirect resistance in which predators or parasitoids dampen the effect of herbivores on plants. Plant volatiles that attract third trophic level organisms such as parasitoids may at the same time attract enemies of the parasitoids in the fourth trophic level, hyperparasitoids, which again dampen the effect of parasitoids on herbivores. In addition, the effectiveness of predators and parasitoids may be dependent on habitat complexity. Here, I plea for studies on the full plant-associated community to understand the fitness outcome of an (induced) plant trait and hence coin it induced direct or indirect plant defence.

1. In nature, several parasitoid species often exploit the same stages of a common herbivore host species and are able to coexist despite competitive interactions amongst them. Less is known about the direct effects of resource quality on intrinsic interactions between immature parasitoid stages. The present study is based on the hypothesis that variation in the quality or type of plant resources on which the parasitoids indirectly develop may be complementary and thus facilitate niche segregation favouring different parasitoids in intrinsic competition under different dietary regimes. 2. The present study investigated whether two herbivore species, the cabbage butterflies Pieris brassicae and Pieris rapae (Pieridae), and the quality of two important food plants, Brassica oleracea and Brassica nigra (Brassicaceae), affect the outcome of intrinsic competition between their primary larval endoparasitoids, the gregarious Cotesia glomerata (Braconidae) and the solitary Hyposoter ebeninus (Ichneumonidae). 3. Hyposoter ebeninus is generally an intrinsically superior competitor over C.glomerata. However, C.glomerata survived more antagonistic encounters with H.ebeninus when both developed in P.brassicae rather than in P.rapae caterpillars, and while its host was feeding on B.nigra rather than B.oleracea. Moreover, H.ebeninus benefitted from competition by its higher survival in multiparasitised hosts. 4. These results show that both plant and herbivore species mediate the battleground on which competitive interactions between parasitoids are played out and may affect the outcomes of these interactions in ways that enable parasitoids to segregate their niches. This in turn may promote coexistence among parasitoid species that are associated with the same herbivore host.

1. Arthropod communities are structured by complex interactions between bottom-up (resource-based) and top-down (natural enemy-based) forces. Their relative importance in shaping arthropod communities, however, continues to be under debate. Bottom-up and top-down forces can be affected by intraspecific plant variation, for example by differences in concentrations of secondary metabolites that affect herbivore abundance through plant quality (bottom-up) or attract natural enemies of these herbivores (top-down). 2. Our objective was to investigate whether herbivore abundance is more strongly affected by plant-mediated bottom-up or top-down forces. 3. We used a model system of four cultivars of Brassica oleracea that show a high degree of variation in several plant traits, resistance to herbivores and attraction of natural enemies. During two field seasons, we recorded the abundance of several herbivorous and carnivorous insect species. To assess the relative importance of bottom-up and top-down forces, we quantified chemical and morphological traits of the cultivars (bottom-up) and assessed parasitization of herbivores and predator oviposition on plants inoculated with a controlled number of herbivores (top-down). 4. We show that intraspecific variation in plant chemistry and morphology consistently affects the abundance of insect herbivores and their natural enemies, resulting in cascading effects on tritrophic interactions in the associated insect community. Foliar profiles of glucosinolates and leaf toughness appeared most important for these effects. Brassica oleracea cultivars that harboured the largest numbers of herbivores also harboured the largest numbers of natural enemies. Differences in the fraction of herbivores parasitized and in predator oviposition on plants inoculated with a controlled number of herbivores could not explain the differences in natural abundance of herbivores. 5. Although abundance of herbivores is most likely influenced by a combination of bottom-up and top-down forces, it appears that in the tritrophic system investigated, bottom-up forces (plant chemistry and morphology) were more important for herbivore abundance than plant-mediated top-down forces (attraction and arrestment of natural enemies).

Communities are riddled with indirect species interactions and these interactions can be modified by organisms that are parasitic or symbiotic with one of the indirectly interacting species. By inducing plant responses, herbivores are well known to alter the plant quality for subsequent feeders. The reduced performance of herbivores on induced plants cascades into effects on the performance of higher trophic level organisms such as parasitoids that develop inside herbivores. Parasitoids themselves may also, indirectly, interact with the host plant by affecting the behaviour and physiology of their herbivorous host. Here, we show that, through their herbivorous host, larvae of two parasitoid species differentially affect plant phenotypes leading to asymmetric interactions among parasitoid larvae developing in different hosts that feed on the same plant. Our results show that temporally separated parasitoid larvae are involved in indirect plant-mediated interactions by a network of trophic and non-trophic relationships.

Plants display a wide range of chemical defences that may differ in effectiveness against generalist and specialist insect herbivores. Host plant-specific secondary chemicals such as glucosinolates (GS) in Brassicaceae typically reduce the performance of generalist herbivores, whereas specialists have adaptations to detoxify these compounds. The concentration of glucosinolates may also alter upon herbivory, allowing the plant to tailor its response to specifically affect the performance of the attacking herbivore. We studied the performance of three Lepidoptera species, two specialists [ Pieris rapae L. (Pieridae), Plutella xylostella L. (Yponomeutidae)] and one generalist [ Mamestra brassicae L. (Noctuidae)], when feeding on eight cultivars of Brassica oleracea L. and a native congener ( Brassica nigra L.) and related this to the GS content. We tested the hypotheses (i) that a generalist herbivore is more affected by high GS concentrations, and (ii) that generalist feeding has a stronger effect on GS levels. Although performance of the three herbivores was different on the B. oleracea cultivars, M. brassicae and P. xylostella had a similar ranking order of performance on the eight cultivars. In most of the cultivars, the concentration of indole GS was significantly higher after feeding by P. rapae or M. brassicae than after P. xylostella feeding. As a consequence, the total concentration of GS in the cultivars showed a different ranking order for each herbivore species. The generalist M. brassicae performed equally well as the specialist P. xylostella on cultivars with high concentrations of GS. Our findings suggest that secondary metabolites other than GSs or differences in nutrient levels affect performance of the species studied.

Plants may be visited by many species of insects during their life-time. These insects include harmful herbivores above and belowground as well as beneficial natural enemies of herbivores and pollinators. Moreover, these interactions may take place sequentially or simultaneously. Responses of plants in interactions with one species result in phenotypic changes that subsequently affect the performance and behaviour of other species. Plants, thus, mediate interactions among their associated community members and such indirect trait-mediated interactions affect the structure of insect communities. As a consequence, plant-mediated species interactions may also affect plant fitness.