Associations of plant fitness, leaf chemistry, and damage suggest selection mosaic in plant-herbivore interactions

The geographic mosaic theory of coevolution states that variation in species interactions forms the raw material for coevolutionary processes, which take place over large geographic scales. One key assumption underlying the process of coevolution in plant-herbivore interactions is that herbivores exert selection on their host plants and that this selection varies among plant populations. We examined spatial variation in the existence and strength of phenotypic selection on host plant resistance exerted by specialist herbivores in 17 archipelago populations of the perennial herb Vincetoxicum hirundinaria (Asclepiadaceae). In these highly fragmented populations, V. hirundinaria is consumed by the larvae of two specialist herbivores: the folivorous moth Abrostola asclepiadis and the seed predator Euphranta connexa. Selection imposed on host plants by these herbivores was examined by analyzing the associations between levels of herbivory, plant fitness, and contents of a number of leaf chemicals reflecting plant resistance to and quality for the herbivores. We found extensive spatial variation in the levels of herbivory and in plant fitness. More importantly, the impact of both leaf herbivory and seed predation on plant fitness varied among plant populations, indicating spatial variation in phenotypic selection. In addition, leaf chemistry varied widely among plant populations, reflecting spatial variation in plant quality as food for the herbivores. However, leaf compounds influenced folivory similarly in all the studied plant populations, and interestingly, some of the compounds were associated with the intensity of seed predation. Finally, some of the leaf compounds were associated with plant fitness, and the strength and direction of these associations varied among plant populations. The observed spatial variation in the strength of the interactions between V. hirundinaria and its specialist herbivores suggests a geographic selection mosaic. Because the occurrence and strength of spatial variation varied between the two specialist herbivores, our results highlight the importance of considering multiple enemies when trying to understand evolution of interactions between plants and their herbivores.     

Sensitivity of plant-pollinator-herbivore communities to changes in phenology

For many species in seasonal environments, warmer springs associated with anthropogenic climate change are causing phenological changes. Within ecological communities, the timing of interactions among species may be altered if the species experience asymmetrical phenological shifts. We present a model that examines the consequences of changes in the relative timing of herbivory and pollination in a community of herbivores and pollinators that share a host plant. Our model suggests that phenological shifts can alter the abundances of these species and, in some cases, their population dynamics. If historical patterns of interactions in a community change and herbivores become active before pollinators, the community could see a reduction in pollinators and an increase in herbivores, while if pollinators become active before herbivores, there could be a loss of stable coexistence. Previous studies have warned of the potential for climate change to cause large phenological mismatches whereby species that depend on one another become so separated in time that they can no longer interact. Our results suggest that climate change-induced phenological shifts can have a major impact on communities even in cases where complete phenological mismatches do not occur.     

Combining optimal defense theory and the evolutionary dilemma model to refine predictions regarding plant invasion

Optimal defense theory posits that plants with limited resources deploy chemical defenses based on the fitness value of different tissues and their probability of attack. However, what constitutes optimal defense depends on the identity of the herbivores involved in the interaction. Generalists, which are not tightly coevolved with their many host plants, are typically deterred by chemical defenses, while coevolved specialists are often attracted to these same chemicals. This imposes an "evolutionary dilemma" in which generalists and specialists exert opposing selection on plant investment in defense, thereby stabilizing defenses at intermediate levels. We used the natural shift in herbivore community composition that typifies many plant invasions to test a novel, combined prediction of optimal defense theory and the evolutionary dilemma model: that the within-plant distribution of defenses reflects both the value of different tissues (i.e., young vs. old leaves) and the relative importance of specialist and generalist herbivores in the community. Using populations of Verbascum thapsus exposed to ambient herbivory in its native range (where specialist and generalist chewing herbivores are prevalent) and its introduced range (where only generalist chewing herbivores are prevalent), we illustrate significant differences in the way iridoid glycosides are distributed among young and old leaves. Importantly, high-quality young leaves are 6.5x more highly defended than old leaves in the introduced range, but only 2x more highly defended in the native range. Additionally, defense levels are tracked by patterns of chewing damage, with damage restricted mostly to low-quality old leaves in the introduced range, but not the native range. Given that whole-plant investment in defense does not differ between ranges, introduced mullein may achieve increased fitness simply by optimizing its within-plant distribution of defense in the absence of certain specialist herbivores.     

Additive effects of aboveground polyphagous herbivores and soil feedback in native and range-expanding exotic plants

Plant biomass and plant abundance can be controlled by aboveground and belowground natural enemies. However, little is known about how the aboveground and belowground enemy effects may add up. We exposed 15 plant species to aboveground polyphagous insect herbivores and feedback effects from the soil community alone, as well as in combination. We envisaged three possibilities: additive, synergistic, or antagonistic effects of the aboveground and belowground enemies on plant biomass. In our analysis, we included native and phylogenetically related range-expanding exotic plant species, because exotic plants on average are less sensitive to aboveground herbivores and soil feedback than related natives. Thus, we examined if lower sensitivity of exotic plant species to enemies also alters aboveground-belowground interactions. In a greenhouse experiment, we exposed six exotic and nine native plant species to feedback from their own soil communities, aboveground herbivory by polyphagous insects, or a combination of soil feedback and aboveground insects and compared shoot and root biomass to control plants without aboveground and belowground enemies. We observed that for both native and range-expanding exotic plant species effects of insect herbivory aboveground and soil feedback added up linearly, instead of enforcing or counteracting each other. However, there was no correlation between the strength of aboveground herbivory and soil feedback. We conclude that effects of polyphagous aboveground herbivorous insects and soil feedback add up both in the case of native and related range-expanding exotic plant species, but that aboveground herbivory effects may not necessarily predict the strengths of soil feedback effects.     

Dealing with double trouble: consequences of single and double herbivory in Brassica juncea

In their natural environment, plants are often attacked simultaneously by many insect species. The specificity of induced plant responses that is reported after single herbivore attacks may be compromised under double herbivory and this may influence later arriving herbivores. The present study focuses on the dynamics of induced plant responses induced by single and double herbivory, and their effects on successive herbivores. Morphological (leaf length, area and trichome density) and chemical changes (leaf alkenyl and indole glucosinolates) in Brassica juncea were evaluated 4, 10, 14 and 20 days after damage by the specialist Plutella xylostella alone, or together with the generalist Spodoptera litura. To assess the biological effect of the plant’s responses, the preference and performance of both herbivores on previously induced plants were measured. We found that alkenyl glucosinolates were induced 20 days after damage by P. xylostella alone, whereas their levels were elevated as early as 4 days after double herbivory. Trichome density was increased in both treatments, but was higher after double herbivory. Interestingly, there was an overall decrease in indole glucosinolates and an increase in leaf size due to damage by P. xylostella, which was not observed during double damage. S. litura preferred and performed better on undamaged plants, whereas P. xylostella preferred damaged plants and performed better on plants damaged 14 and 10 days after single and double herbivory, respectively. Our results suggest that temporal studies involving single versus multiple attacker situations are necessary to comprehend the role of induced plant responses in plant–herbivore interactions.     

Direct and indirect effects of CO2, nitrogen, and community diversity on plant-enemy interactions

Resource abundance and plant diversity are two predominant factors hypothesized to influence the amount of damage plants receive from natural enemies. Many impacts of these environmental variables on plant damage are likely indirect and result because both resource availability and diversity can influence plant traits associated with attractiveness to herbivores or susceptibility to pathogens. We used a long-term, manipulative field experiment to investigate how carbon dioxide (CO2) enrichment, nitrogen (N) fertilization, and plant community diversity affect plant traits and the amount of herbivore and pathogen damage experienced by the common prairie legume Lespedeza capitata. We detected little evidence that CO2 or N affected plant traits; however, plants growing in high-diversity treatments (polycultures) were taller, were less pubescent, and produced thinner leaves (higher specific leaf area). Interestingly, we also detected little evidence that CO2 or N affect damage. Plants growing in polycultures compared to monocultures, however, experienced a fivefold increase in damage from generalist herbivores, 64% less damage from specialist herbivores, and 91% less damage from pathogens. Moreover, within diversity treatments, damage by generalist herbivores was negatively correlated with pubescence and often was positively correlated with plant height, while damage by specialist herbivores typically was positively correlated with pubescence and negatively associated with height. These patterns are consistent with changes in plant traits driving differences in herbivory between diversity treatments. In contrast, changes in measured plant traits did not explain the difference in disease incidence between monocultures and polycultures. In summary, our data provide little evidence that CO2 or N supply alter damage from natural enemies. By contrast, plants grown in monocultures experienced greater specialist herbivore and pathogen damage but less generalist herbivore damage than plants grown in diverse communities. Part of this diversity effect was mediated by changes in plant traits, many of which likely are plastic responses to diversity treatments, but some of which may be the result of evolutionary changes in response to these long-term experimental manipulations.     

Damage-induced resistance in sagebrush: volatiles are key to intra- and interplant communication

Airborne communication between individuals, called "eavesdropping" in this paper, can cause plants to become more resistant to herbivores when a neighbor has been experimentally clipped. The ecological relevance of this result has been in question, since individuals may be too far apart for this interaction to affect many plants in natural populations. We investigated induced resistance to herbivory in sagebrush, Artemisia tridentata, caused by experimental clipping of the focal plant and its neighbors. We found no evidence for systemic induced resistance when one branch was clipped and another branch on the same plant was assayed for naturally occurring damage. In this experiment, air contact and plant age were not controlled. Previous work indicated that sagebrush received less damage when a neighboring upwind plant within 15 cm had been experimentally clipped. Here we found that pairs of sagebrush plants that were up to 60 cm apart were influenced by experimental clipping of a neighbor. Furthermore, we observed that most individuals had conspecific neighbors that were much closer than 60 cm. Air contact was essential for communication; treatments that reduced airflow between neighboring individuals, either because of wind direction or bagging, prevented induced resistance. Airflow was also necessary for systemic induced resistance among branches within an individual. Reports from the literature indicated that sagebrush is highly sectorial, as are many desert shrubs. Branches within a sagebrush plant do not freely exchange material via vascular connections and apparently cannot rely on an internal signaling pathway for coordinating induction of resistance to herbivores. Instead, they may use external, volatile cues. This hypothesis provides a proximal explanation for why sagebrush does not demonstrate systemic induced resistance without directed airflow, and why airborne communication between branches induces resistance.     

Herbivory limits recruitment in an old-field seed addition experiment

Environmental variability can promote coexistence by creating establishment sites for rare plants, but low diversity in anthropogenic grasslands suggests that this variability may be eliminated (homogenization hypothesis) or inaccessible (barrier hypothesis). We explore these alternatives on the northern Great Plains, where 11 million hectares have been transformed by multiple environmental changes, but the causes of species loss are unclear. In a degraded grassland, we increased environmental variability by manipulating competition and herbivory along gradients of fertility and disturbance, and we circumvented dispersal barriers by adding 1.2 million seeds of five functionally distinct species at varying densities. The experiment ended after 12 weeks due to the direct and indirect effects of unapparent small native herbivores, which were barriers to population establishment by the added species. The direct cause of recruitment failure was browsing. The indirect cause was associated with competition from invasive plants that appeared to be more tolerant or resistant to herbivory. Variability in fertility, disturbance, propagule pressure, and competition had relatively minor impacts on colonization by the added species because herbivores controlled recruitment in most environments. Recruitment outside the herbivore exclosures was mostly by unpalatable exotics, suggesting a possible link between invasion success and herbivore resistance for some introduced plants.     

Population and leaf-level variation of iridoid glycosides in the invasive weed Verbascum thapsus L. (common mullein): implications for herbivory by generalist insects

Plant–insect interactions, which are strongly mediated by chemical defenses, have the potential to shape invasion dynamics. Despite this, few studies have quantified natural variation in key defensive compounds of invasive plant populations, or how those defenses relate to levels of herbivory. Here, we evaluated variation in the iridoid glycosides aucubin and catalpol in rosette plants of naturally occurring, introduced populations of the North American invader, Verbascum thapsus L. (common mullein; Scrophulariaceae). We examined two scales that are likely to structure interactions with insect herbivores—among populations and within plant tissues (i.e., between young and old leaves). We additionally estimated the severity of damage incurred at these scales due to insect chewing herbivores (predominantly grasshoppers and caterpillars), and evaluated the relationship between iridoid glycoside content and leaf damage. We found significant variation in iridoid glycoside concentrations among populations and between young and old leaves, with levels of herbivory strongly tracking leaf-level investment in defense. Specifically, across populations, young leaves were highly defended by iridoids (averaging 6.5× the concentration present in old leaves, and containing higher proportions of the potentially more toxic iridoid, catalpol) and suffered only minimal damage from generalist herbivores. In contrast, old leaves were significantly less defended and accordingly more substantially utilized. These findings reveal that quantitative variation in iridoid glycosides is a key feature explaining patterns of herbivory in an introduced plant. In particular, these data support the hypothesis that defenses limit the ability of generalists to feed on mullein’s well-defended young leaves, resulting in minimal losses of high-quality tissue, and increasing performance of this introduced species.     

Herbivores and edaphic factors constrain the realized niche of a native plant

Biotic interactions, such as competition and herbivory, can limit plant species ranges to a subset of edaphically suitable habitats, termed the realized niche. Here we explored the role that herbivores play in restricting the niche of serpentine ecotypes of the native California annual Collinsia sparsiflora. We planted seeds from four populations into a range of natural field environments that varied in the presence/absence of naturally occurring C. sparsiflora and in predicted suitability for growth and survival of the serpentine ecotype of C. sparsiflora. Path analysis was then used to model the direct and herbivore-mediated indirect effects of environmental variables on the survival of C. sparsiflora serpentine ecotypes. We found that C. sparsiflora received more herbivory when planted into areas where serpentine ecotypes of C. sparsiflora were not predicted to persist, and that increased herbivory was associated with decreased survival, suggesting that herbivores may limit the distribution of C. sparsiflora serpentine ecotypes. Additionally, we demonstrated that edaphic environmental variables impacted the survival of C. sparsiflora serpentine ecotypes both directly and indirectly, by altering interactions with herbivores. These indirect effects were probably trait-mediated and probably occurred because edaphic factors may influence plant traits that, in turn, alter attractiveness to herbivores. Although the magnitude of direct effects exceeded the magnitude of indirect effects, many strong herbivore-mediated indirect effects were detected. Thus, interactions between the abiotic environment and insect herbivory contributed to restricting the niche of C. sparsiflora serpentine ecotypes to a subset of available habitat.     

Herbivore species, infestation time, and herbivore density affect induced volatiles in tea plants

In response to insect herbivory, plants emit volatiles that are used by the herbivores’ natural enemies to locate their host or prey. Herbivore attack also enhances tea aroma. Herbivore-induced plant volatiles (HIPVs) vary both quantitatively and qualitatively with infestation duration and herbivore density. Thus, whether HIPVs can reliably communicate the identities of herbivores is of interest. Here, we studied the tea plant volatiles induced by the tea leafhopper (Empoasca vitis, a piercing–sucking insect), the tea geometrid (Ectropis oblique, a chewing insect), and methyl jasmonate (MeJA, a plant hormone). Geometrid feeding induced more complex volatile blends than did leafhopper infestation. The volatiles induced by both herbivores significantly increased in quality and quantity with time during the first 16 h of infestation, after which the profiles of induced volatile blends and the emission of induced compounds varied diurnally. (E)-β-Ocimene displayed a unique rhythm in which emission peaked at night. The amount of HIPVs significantly increased, while their profiles changed little, with herbivore density. Overall, the leafhoppers and geometrids induced significantly different volatiles from tea plants, while the HIPV profiles varied with a circadian rhythm and were similar at different herbivore densities. Our findings also suggest a new method of enhancing tea flavor using exogenously applied plant hormones, because the volatiles induced by leafhoppers and MeJA were similar in general composition.     

Herbivory reduces plant interactions with above- and belowground antagonists and mutualists

Herbivores affect plants through direct effects, such as tissue damage, and through indirect effects that alter species interactions. Interactions may be positive or negative, so indirect effects have the potential to enhance or lessen the net impacts of herbivores. Despite the ubiquity of these interactions, the indirect pathways are considerably less understood than the direct effects of herbivores, and multiple indirect pathways are rarely studied simultaneously. We placed herbivore effects in a comprehensive community context by studying how herbivory influences plant interactions with antagonists and mutualists both aboveground and belowground. We manipulated early-season aboveground herbivore damage to Cucumis sativus (cucumber, Cucurbitaceae) and measured interactions with subsequent aboveground herbivores, root-feeding herbivores, pollinators, and arbuscular mycorrhizal fungi (AMF). We quantified plant growth and reproduction and used an enhanced pollination treatment to determine if plants were pollen limited. Increased herbivory reduced interactions with both antagonists and mutualists. Plants with high levels of early herbivory were significantly less likely to suffer leaf damage later in the summer and tended to be less attacked by root herbivores. Herbivory also reduced pollinator visitation, likely due to fewer and smaller flowers, and reduced AMF colonization. The net effect of herbivory on plant growth and reproduction was strongly negative, but lower fruit and seed production were not due to reduced pollinator visits, because reproduction was not pollen limited. Although herbivores influenced interactions between plants and other organisms, these effects appear to be weaker than the direct negative effects of early-season tissue loss.     

Antagonistic interactions between plant competition and insect herbivory

Interspecific competition between plants and herbivory by specialized insects can have synergistic effects on the growth and performance of the attacked host plant. We tested the hypothesis that competition between plants may also negatively affect the performance of herbivores as well as their top-down effect on the host plant. In such a case, the combined effects of competition and herbivory may be less than expected from a simple multiplicative response. In other words, competition and herbivory may interact antagonistically. In a greenhouse experiment, Poa annua was grown in the presence or absence of a competitor (either Plantago lanceolata or Trifolium repens), as well as with or without a Poa-specialist aphid herbivore. Both competition and herbivory negatively affected Poa growth. Competition also reduced aphid density on Poa. This effect could in part be explained by changes in the biomass and the nitrogen content of Poa shoots. In treatments with competitors, reduced aphid densities alleviated the negative effect of herbivory on above- and belowground Poa biomass. Hence, we were able to demonstrate an antagonistic interaction between plant-plant interspecific competition and herbivory. However, response indices suggested that antagonistic interactions between competition and herbivory were contingent on the identity of the competitor. We found the antagonistic effect only in treatments with T. repens as the competitor. We conclude that both competitor identity and the herbivore's ability to respond with changes in its density or activity to plant competition affect the magnitude and direction (synergistic vs. antagonistic) of the interaction between competition and herbivory on plant growth.     

Protection from Natural Enemies in Managing Rare Plant Species

Abstract: Natural enemies such as pathogens, herbivores, and seed predators can substantially limit the abundance of plants, including rare species. Vulnerability to particular enemies is likely to differ between life-history stages. We hypothesized that short-term protection of juvenile plants from herbivores can be used to increase population growth of rare species and thus improve the probability of long-term persistence. Using the federally listed (threatened) Pitcher's thistle ( Cirsium pitcheri ) as a model, we experimentally excluded insect herbivores from juvenile rosettes to evaluate the potential benefits of deliberate insect control as a tool for management of rare species. Herbivore effects varied spatially across the local environment. Excluding insects in portions of the habitat where herbivory was high had direct benefits, including a 53% decrease in juvenile plant mortality (60% to 7%) and a 10-fold increase in seed production of juveniles that matured and flowered. In other areas, where herbivore-induced juvenile mortality was relatively low, excluding insects either increased seed production of plants that flowered or had no major effect. Our data also suggest indirect benefits to the metapopulation via potential improvement in dispersal among patches. Temporal variation in growing conditions occurred between years, suggesting that multiple-year exclusions would be most effective. Our study suggests that small–scale manipulation of often inconspicuous interactions between rare plants and their natural enemies can be an effective, relatively low-cost tool for the management and restoration of rare plant species.     

Shifts in positive and negative plant interactions along a grazing intensity gradient

Isolating the single effects and net balance of negative and positive species effects in complex interaction networks is a necessary step for understanding community dynamics. Facilitation and competition have both been found to operate in harsh environments, but their relative strength may be predicted to change along gradients of herbivory. Moreover, facilitation effects through habitat amelioration and protection from herbivory may act together determining the outcome of neighborhood plant-plant interactions. We tested the hypothesis that grazing pressure alters the balance of positive and negative interactions between palatable and unpalatable species by increasing the strength of positive indirect effects mediated by associational resistance to herbivory. We conducted a two-year factorial experiment in which distance (i.e., spatial association) from the nearest unpalatable neighbor (Stipa speciosa) and root competition were manipulated for two palatable grasses (Poa ligularis and Bromus pictus), at three levels of sheep grazing (none, moderate, and high) in a Patagonian steppe community. We found that grazing shifted the effect of Stipa on both palatable grasses, from negative (competition) in the absence of grazing to positive (facilitation) under increasing herbivore pressure. In ungrazed sites, belowground competition was the dominant interaction, as shown by a significant reduction in performance of palatable grasses transplanted near to Stipa tussocks. In grazed sites, biomass of palatable plants was greater near than far from Stipa regardless of competition treatment. Proximity to Stipa reduced the amount of herbivory suffered by palatable grasses, an indirect effect that was stronger under moderate than under intense grazing. Our results demonstrate that facilitation, resulting mainly from protection against herbivory, is the overriding effect produced by unpalatable neighbors on palatable grasses in this rangeland community. This finding challenges the common view that abiotic stress amelioration should be the predominant type of facilitation in arid environments and highlights the role of herbivory in modulating complex neighborhood plant interactions in grazing systems.     

Habitat patch shape, not corridors, determines herbivory and fruit production of an annual plant

Habitat corridors confer many conservation benefits by increasing movement of organisms between habitat patches, but the benefits for some species may exact costs for others. For example, corridors may increase the abundance of consumers in a habitat to the detriment of the species they consume. In this study we assessed the impact of corridors on insect herbivory of a native plant, Solanum americanum, in large-scale, experimentally fragmented landscapes. We quantified leaf herbivory and assessed fruit production as a proxy for plant fitness. We also conducted field surveys of grasshoppers (Orthoptera), a group of abundant, generalist herbivores that feed on S. americanum, and we used exclosure cages to explicitly link grasshopper herbivory to fruit production of individual S. americanum. The presence of corridors did not increase herbivory or decrease plant fruit production. Likewise, corridors did not increase grasshopper abundance. Instead, patches in our landscapes with the least amount of edge habitat and the greatest amount of warmer "core" area had the highest levels of herbivory, the largest cost to plant fruit production as a result of herbivory, and the most grasshoppers. Thus habitat quality, governed by patch shape, can be more important than connectivity for determining levels of herbivory and the impact of herbivory on plant fitness in fragmented landscapes.     

Selecting for Tolerance against Pathogens and Herbivores to Enhance Success of Reintroduction and Translocation

Abstract: Some species have insufficient defenses against climate change, emerging infectious diseases, and non‐native species because they have not been exposed to these factors over their evolutionary history, and this can decrease their likelihood of persistence. Captive breeding programs are sometimes used to reintroduce individuals back into the wild; however, successful captive breeding and reintroduction can be difficult because species or populations often cannot coexist with non‐native pathogens and herbivores without artificial selection. In captive breeding programs, breeders can select for host defenses that prevent or reduce pathogen or herbivore burden (i.e., resistance) or traits that limit the effects of parasitism or herbivory on host fitness (i.e., tolerance). We propose that selection for host tolerance may enhance the success of reintroduction or translocation because tolerant hosts generally have neutral effects on introduced pathogens and herbivores. The release of resistant hosts would have detrimental effects on their natural enemies, promoting rapid evolution to circumvent the host resistance that may reduce the long‐term probability of persistence of the reintroduced or translocated species. We examined 2 case studies, one on the pathogenic amphibian chytrid fungus ( Batrachochytrium dendrobatidis [Bd]) and the other on the herbivorous cactus moth ( Cactoblastis cactorum) in the United States, where it is not native. In each case study, we provide recommendations for how captive breeders and managers could go about selecting for host tolerance. Selecting for tolerance may offer a promising tool to rescue hosts species from invasive natural enemies as well as new natural enemies associated with climate change‐induced range shifts.     

Fitness related diet-mixing by intraspecific host-plant-switching of specialist insect herbivores

Generalist insect herbivores may profit by feeding on a mixture of plant species that differ in nutritional quality. Herbivore performance can also be affected by intraspecific host plant variation. However, it is unknown whether conspecific plant individuals differ sufficiently to promote diet-mixing behavior in specialist herbivores. We experimentally tested this "specialist diet-mixing hypothesis" for specialist caterpillars (Chrysopsyche imparilis, Lasiocampidae) in a West African savanna. The caterpillars switched regularly between host tree individuals (Combretum fragrans, Combretaceae). To examine whether switching benefited caterpillar performance via diet-mixing, the caterpillars were reared either on leaves from several plant individuals (mixed diet) or on leaves from a single plant. The strongest effect of diet-mixing was found for fecundity, with females reared on a mixed diet laying significantly more eggs than sisters receiving a single-plant diet. In addition, a mixed diet decreased variability in egg size and increased the growth of second-instar caterpillars. Supplementary food choice experiments were conducted to assess a potential influence of lowered host quality (induced by herbivory) on caterpillar behavior; no such effect was found. By linking intraspecific host-switching behavior and herbivore performance, this study provides new information on the relevance of intraspecific plant variation for herbivorous insects.     

Plant chemistry and insect sequestration

Most plant families are distinguished by characteristic secondary metabolites, which can function as putative defence against herbivores. However, many herbivorous insects of different orders can make use of these plant-synthesised compounds by ingesting and storing them in their body tissue or integument. Such sequestration of putatively unpalatable or toxic metabolites can enhance the insects’ own defence against enemies and may also be involved in reproductive behaviour. This review gives a comprehensive overview of all groups of secondary plant metabolites for which sequestration by insect herbivores belonging to different orders has been demonstrated. Sequestered compounds include various aromatic compounds, nitrogen-containing metabolites such as alkaloids, cyanogenic glycosides, glucosinolates and other sulphur-containing metabolites, and isoprenoids such as cardiac glycosides, cucurbitacins, iridoid glycosides and others. Sequestration of plant compounds has been investigated most in insects feeding or gathering on Apocynaceae s.l. (Apocynoideae, Asclepiaoideae), Aristolochiaceae, Asteraceae, Boraginaceae, Fabaceae and Plantaginaceae, but it also occurs for some gymnosperms and even lichens. In total, more than 250 insect species have been shown to sequester plant metabolites from at least 40 plant families. Sequestration predominates in the Coleoptera and Lepidoptera, but also occurs frequently in the orders Heteroptera, Hymenoptera, Orthoptera and Sternorrhyncha. Patterns of sequestration mechanisms for various compound classes and common or individual features occurring in different insect orders are highlighted. More research is needed to elucidate the specific transport mechanisms and the physiological processes of sequestration in various insect species.     

Insect herbivores, density dependence, and the performance of the perennial herb Solanum carolinense

How insect herbivores affect plant performance is of central importance to basic and applied ecology. A full understanding of herbivore effects on plant performance requires understanding interactions (if any) of herbivore effects with plant density and size because these interactions will be critical for determining how herbivores influence plant population size. However, few studies have considered these interactions, particularly over a wide enough range of densities to detect nonlinear effects. Here we ask whether plant density and herbivores influence plant performance linearly or nonlinearly, how plant density affects herbivore damage, and how herbivores alter density dependence in transitions between plant size classes. In a large field experiment, we manipulated the density of the herbaceous perennial plant Solanum carolinense and herbivore presence in a fully crossed design. We measured plant size, sexual reproduction, and damage to plants in two consecutive years, and asexual reproduction of new stems in the second year, allowing us to characterize both plant performance and rates of transition between plant size classes across years. We found nonlinear effects of plant density on damage. Damage by herbivores and plant density both influenced sexual and asexual reproduction of S. carolinense; these effects were mostly mediated via effects on plant size. Importantly, we found that herbivores altered the pattern of linear density dependence in some transition rates (including survival and asexual reproduction) between plant size classes. These results suggest that understanding the ecological or evolutionary effects of herbivores on plant populations requires consideration of plant density and plant size, because feedbacks between density, herbivores, and plant size may complicate longer-term dynamics.