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Native to South Africa, fireweed (Senecio madagascariensis Poiret; Asteraceae) is an annual or short‐lived perennial herb that is highly invasive in Australia, where it is a target for biological control. Preliminary research indicates that fireweed may be undergoing adaptive changes along its invasion gradient in Australia. Changes, such as a shift in plant chemical defence, may influence fireweed's interaction with specialist insect herbivores, and hence the efficacy of a biocontrol program. As the testing of biocontrol candidates is a rigorous and lengthy process, confirmation of fireweed's susceptibility to insect attack may assist in fast tracking candidate suitability, where changes in plant defensive chemistry are found. In a large experimental field study in fireweed's native range, we compared the susceptibility of Australian and South African populations to insect natural enemy attack (in terms of plant biomass, insect richness and composition). Simultaneously, we assessed the variation in plant pyrrolizidine alkaloid concentrations and composition to rationalise any differences in natural enemy attack between populations. We found no significant differences in plant biomass across fireweed populations, despite significantly higher alkaloid concentrations in Australian populations. The composition of the endophagous (specialist) insect herbivore assemblage was related to alkaloid composition, but there were only minor differences in the assemblage between Australian and South African fireweed populations. Moreover, we found no significant differences in the abundance, richness or Shannon diversity indices of endophagous or ectophagous (generalist) insect herbivores between Australian and South African fireweed populations. Our results suggest that despite variations in chemical defence, Australian fireweed populations displayed neither reduced nor increased susceptibility to specialist insect herbivore attack, supporting the notion that if host‐specific biocontrol agents are released, their impact in Australia is unlikely to be influenced by any changes in plant chemical defence. [ABSTRACT FROM AUTHOR]

Host plant specialisation can promote evolutionary divergence between herbivore populations associated with different plant species. While the mechanisms by which specialist species exploit their hosts have been studied widely across taxa, less is known about the mechanisms that allow intraspecific variants to arise and to be maintained across spatial and temporal scales. To understand whether adaptations to plant defences against herbivory contribute to the co-existence of genetically distinct populations of an herbivore, we investigate the interaction between honeysuckle (Lonicera periclymenum) and sympatric specialist and generalist populations of the spider mite Tetranychus urticae. We found that mite folivory induces the production of sticky droplets on honeysuckle, which have a defensive role: they increase mite mortality directly, and potentially indirectly by increasing the arrestment of a predator. We show that droplet induction and the preference to feed on honeysuckle depend on mite genotype, where the generalist avoids this host and the specialist suppresses droplet production. These traits are heritable and dominant in F1 hybrids between generalists and specialists. Selection pressure from honeysuckle and differences in host preference likely reduce the opportunity of mating encounters on this host. We propose that the interplay between selection from host plant defences and ecological barriers to hybridisation contribute to the persistence of genetically distinct populations of a single species in sympatry.

My goal is to investigate global patterns in the structure of interaction networks of insect herbivores and their host plants. Specifically, I seek to determine whether intensification of land use and the dominance of exotic host plant species influence the structure and robustness (i.e., resistance to co-extinctions) of interaction networks of insect herbivores and host plants. I also ask whether latitude has an influence on the structure and robustness of these interactions. I compiled 90 local plant-herbivore networks distributed worldwide, spanning different taxonomic groups of plants and insects and several herbivore guilds. My results showed that intensification of land use was associated with dominance of exotic plant species and can impoverish the species richness and taxonomic diversity of insect herbivores in the networks. Moreover, land use intensification surprisingly increases network specialization by decreasing connectance and nestedness, and increases modularity; while the increase in the proportion of exotic hosts had opposite effects. These changes in the network structure may be due to the proportionately greater loss of generalist herbivores relative to specialists. Land use intensification also decreases the robustness of plant-herbivore networks, while the proportion of exotic host plant species increases, which is an intriguing result that contradicts previous studies. Controlling for anthropic effects that can act on the networks, my results show that plant–herbivore networks are structured independently of latitude, suggesting that the factors that influence the interactions between host plants and insect herbivores are latitudinally invariant.

Novel interactions between introduced oaks and their natural enemies across different continents provide an opportunity to test the enemy release hypothesis (ERH) at local and global scales. Based on the ERH, we assessed the impacts of native seed‐feeding insects on introduced and native oaks within and among continents. We combined a common‐garden experiment in China and biogeographic literature surveys to measure seed predation by insects and the proportion of acorn embryos surviving after insect infestation among 4 oak species with different geographical origins: Quercus mongolica origin from China, Q. robur and Q. petraea from Europe, and Q. rubra from North America. Mostly supporting the ERH, oaks in introduced continents escaped seed predation compared to those in native continents and compared to other native oaks in introduced continents. Common‐garden comparisons showed that total acorn infestation rate of introduced Q. rubra (section Lobatae) was considerably lower than that of native oaks (section Quercus) in China and Europe, likely because of the differences in seed traits associated with different oak sections. Literature surveys showed that seed predation of introduced oaks was lower in the introduced continent than in the native continent. Embryo survival was higher in introduced Q. rubra than native oaks in China and Poland. However, insect seed predation of recently introduced Q. rubra in China was similar to that in Europe, which is not consistent with the ERH. Our results suggest that reduced acorn attack by native insects and higher embryo survival after acorn damage could increase the establishment success or invasion risk of introduced oaks in non‐native continents. [ABSTRACT FROM AUTHOR]

Structural diversity is an emerging dimension of biodiversity that accounts for size variations in organs among individuals in a community. Previous studies show significant effects of structural diversity on forest growth, but its effects on forest mortality are not known, particularly at a large scale.To address this knowledge gap, we quantified structural diversity using stem structural diversity (SSD) based on both tree diameter and height. We obtained U.S. Forest Service Forest Inventory and Analysis (FIA) data from over 2400 plots across southcentral U.S. forests that have suffered a recent drought. Using data from multiple sampling times, we calculated SSD and compared the relative importance of SSD, species diversity, functional diversity and other stand attributes in determining tree mortalities caused by fire, insects and diseases. We also used FIRETEC, a physics‐based fire model, to test the effect of SSD on canopy consumption by fire.Our results showed that (1) SSD was positively associated with tree mortalities caused by all three disturbances; (2) species richness was negatively associated with insect‐ and disease‐caused mortalities; (3) functional diversity was negatively associated with fire‐ and disease‐caused mortalities and (4) more phylogenetically related species had more similar mortality rates by insect and disease but not fire. Moreover, the FIRETEC model showed increasing canopy consumption by fire in stands with greater SSD.Together, the different tree mortalities during drought associated with SSD more consistently than the other biodiversity metrics were evaluated.Synthesis. Our results suggest that SSD could be considered in modelling forest dynamics and planning management to sustain forest health under disturbances. [ABSTRACT FROM AUTHOR]

Recent studies suggest that changes in leaf traits due to interactions between plants affect resource utilisation by herbivores, as well as herbivore distribution. However, this has not yet been confirmed experimentally. Here, we investigated the effects of phenotypic plasticity in leaf traits of Rumex obtusifolius (host plant) in response to intraspecific and interspecific interactions on the distribution of two leaf beetles, Gastrophysa atrocyanea (specialist herbivore) and Galerucella grisescens (generalist herbivore).We investigated the local population density of R. obtusifolius plants and the presence of leaf beetles on the plants at five study sites. Leaf chemicals (condensed tannins and total phenolics) were compared between aggregated and solitary R. obtusifolius plants. To clarify the effects of the interaction environment of R. obtusifolius plants on their leaf traits and on resource utilisation by the leaf beetles, we compared leaf chemicals and preferences of adult leaf beetles among treatments where R. obtusifolius experienced intraspecific interaction, interspecific interaction or no interaction in cultivation experiments. Finally, we evaluated the independent and combined effects of patch size and intraspecific interaction on leaf beetle distribution in mesocosm experiments.In the field, the presence of the specialist leaf beetle G. atrocyanea was positively correlated with the local population density (rosette overlap ratio) of R. obtusifolius plants; however, there was no correlation in the case of the generalist leaf beetle G. grisescens. In the cultivation experiments, plants in the intraspecific interaction treatment increased their leaf concentrations of condensed tannins and total phenolics, and G. atrocyanea consumed more of these leaves than leaves in other treatments. Similar results were observed in the field. In the mesocosm experiments, larger numbers of G. atrocyanea were distributed on R. obtusifolius plants exposed to below‐ground intraspecific interaction than on plants not exposed to intraspecific interaction.Our results provide experimental evidence that leaf‐trait changes in response to intraspecific interaction between host plants influence specialist herbivore distribution. This highlights the need to integrate plant–plant interactions into our understanding of plant–animal interactions. A free Plain Language Summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

Current theory (e.g., consumer‐controlled theory) predicts that nutrient enrichment typically amplifies herbivory and thereby suppresses the growth and expansion of invasive plants. Herbivores can facilitate plant regrowth in the native community by stimulating complementary growth or ameliorating habitat conditions (e.g., by increasing soil oxygen and nutrient availability), but whether they have similar positive effects on invasive plants, especially under nutrient enrichment, remains unknown. Using a field nitrogen (N)‐enrichment × crab exclusion experiment, we evaluated and compared the effects of both N enrichment and crab herbivory on the growth performance of a global invasive cordgrass, Spartina alterniflora, and a co‐occurring native plant, Phragmites australis. We found that crabs consistently suppressed P. australis by density and aboveground biomass regardless of N enrichment. In contrast, for S. alterniflora, the negative effects of crabs under ambient N were replaced by positive effects under N enrichment, with crabs stimulating complementary increases in density and aboveground biomass. The differing effects between the N treatments were driven by crab burrowing activity, which increased soil N availability, and the nutrient‐use efficiency of S. alterniflora. Our findings revealed that native herbivores can have opposing effects on native and invasive plants, which broadens our understanding of how exotic plants can achieve dominance in a changing world. [ABSTRACT FROM AUTHOR]

Do hotspots of plant biodiversity translate into hotspots in the abundance and diversity of large mammalian herbivores? A common expectation in community ecology is that the diversity of plants and animals should be positively correlated in space, as with the latitudinal diversity gradient and the geographic mosaic of biodiversity. Whether this pattern 'scales down' to landscape-level linkages between the diversity of plants or the activities of highly mobile megafauna has received less attention. We investigated spatial associations between plants and large herbivores by integrating data from a plant-DNA-barcode phylogeny, camera traps, and a comprehensive map of woody plants across the 1.2-km2 Mpala Forest Global Earth Observatory (ForestGEO) plot, Kenya. Plant and large herbivore communities were strongly associated with an underlying soil gradient, but the richness of large herbivore species was negatively correlated with the richness of woody plants. Results suggest thickets and steep terrain create associational refuges for plants by deterring megaherbivores from browsing on otherwise palatable species. Recent work using dietary DNA metabarcoding has demonstrated that large herbivores often directly control populations of the plant species they prefer to eat, and our results reinforce the important role of megaherbivores in shaping vegetation across landscapes. [ABSTRACT FROM AUTHOR]

Predicting how plants allocate to different anti-herbivore defences in response to elevated carbon dioxide (CO2) concentrations is important for understanding future patterns of crop susceptibility to herbivory. Theories of defence allocation, especially in the context of environmental change, largely overlook the role of silicon (Si), despite it being the major anti-herbivore defence in the Poaceae. We demonstrated that elevated levels of atmospheric CO2 (e[CO2]) promoted plant growth by 33% and caused wheat (Triticum aestivum) to switch from Si (-19%) to phenolic (+44%) defences. Despite the lower levels of Si under e[CO2], resistance to the global pest Helicoverpa armigera persisted; relative growth rates (RGRs) were reduced by at least 33% on Si-supplied plants, irrespective of CO2 levels. RGR was negatively correlated with leaf Si concentrations. Mandible wear was c. 30% higher when feeding on Si-supplemented plants compared to those feeding on plants with no Si supply. We conclude that higher carbon availability under e[CO2] reduces silicification and causes wheat to increase concentrations of phenolics. However, Si supply, at all levels, suppressed the growth of H. armigera under both CO2 regimes, suggesting that shifts in defence allocation under future climate change may not compromise herbivore resistance in wheat. [ABSTRACT FROM AUTHOR]

Invasive species are major drivers of ecosystem degradation globally. How invasive herbivore impacts differ from native herbivore impacts remains understudied. We examined the relationships between herbivore sign and vegetation height, foliage density, cover of forbs, weeds, bare ground, and soil compaction across environmental and herbivore activity gradients in the mainland Australian Alps. We detected native and exotic herbivore sign at 32.8% and 94.0% of sites, respectively. Total herbivore activity was primarily attributed to exotic herbivores and was associated with elevation and grassland type. Greater horse (exotic) activity was associated with lower vegetation height, lower foliage density, higher forb cover, and higher soil compaction. Greater rabbit and hare (exotic) activity was associated with lower vegetation height, lower foliage density, and a higher cover of bare ground. Greater total herbivore activity was associated with greater weed cover. Neither deer (exotic) nor kangaroo and wallaby (native) activity was related to response variables. We demonstrate that exotic herbivores dominate mammalian herbivory in these grasslands, which evolved without analogous hooved species. Given the restricted distribution and high endemism of these ecosystems, and associations between exotic herbivores and characteristics of degraded grasslands, we recommend landscape‐scale exotic herbivore management, focusing on maintaining ground cover and vegetation structure. [ABSTRACT FROM AUTHOR]

Plant–herbivore interactions account for much of the biodiversity observed in an ecosystem. Plants may alter defense expression dependent on the specific herbivore present. Many studies focus on the optimization of these defenses against a single herbivore, but in an ecological context, there may be a myriad of herbivores at any given time. We designed a study to examine defenses employed by swamp white oak (Quercus bicolor) against insects and large mammals. We established six deer exclosures with trees located inside and outside each exclosure. Outside the exclosures, physical constraints only allow deer to feed up to a browseline on trees, whereas insects can potentially access all portions of the tree. Trees inside exclosures were only accessible to insect herbivores. Furthermore, insecticides were applied to half of the exclosures to create insect‐free (i.e., insecticide applied), deer‐free (i.e., inside exclosures), deer‐only (i.e., outside exclosures with insecticide), and herbivore‐free (i.e., inside exclosures with insecticide) environments. This design allowed us to analyze simultaneous defensive responses of individual trees to mammal and insect herbivores. We predicted that because insects represent a continuous threat, plants should invest in fixed concentrations of defenses (e.g., polyphenols, tannins, and condensed tannins), even when deer are absent. When deer are present, plants should increase investment in defenses in tissues that deer have access to. We found that plant defense strategies varied spatially on individual trees. When only insects were present, nutritional constituents (e.g., nitrogen and non‐structural carbohydrates) did not change but antinutritional constituents increased. Contrastingly, when deer were present, both nutritional and antinutritional constituents decreased. We found a decrease in insect functional group diversity when deer were present. Our findings suggest that plants may partition defense strategies depending on the most prominent herbivore pressure. However, changes in insect functional group diversity may result from either changes in plant defense strategy or intraspecific competition among multiple herbivores. Future studies should investigate patterns of chemical heterogeneity displayed by plants in response to diverse herbivore pressures. [ABSTRACT FROM AUTHOR]

Summary: The ability to predict future risks is essential for many organisms, including plants. Plants can gather information about potential future herbivory by detecting volatiles that are emitted by herbivore‐attacked neighbors. Several individual volatiles have been identified as active danger cues. Recent work has also shown that plants may integrate multiple volatiles into their defense responses. Here, I discuss how the integration of multiple volatiles can increase the capacity of plants to predict future herbivore attack. I propose that integration of multiple volatile cues does not occur at the perception stage, but may through downstream early defense signaling and then be further consolidated by hormonal crosstalk. Exploring plant volatile cue integration can facilitate our understanding and utilization of chemical information transfer. [ABSTRACT FROM AUTHOR]

Questions: Ungulate herbivory and its impact on one plant species may be driven by other plants in the neighbourhood. Although such neighbourhood effects have long been recognized, there is limited understanding of how spatial variation in these effects relates to local plant density and distances between plants. Our goal was to examine if the density of or distance to conspecific and heterospecific neighbouring woody individuals produces neighbourhood effects on bark stripping by deer, and how these effects differ in spatial scale and type (reduce/increase herbivory). Location: Lowland old-growth forest, Czech Republic. Methods: Woody individuals were assessed for evidence of bark stripping and the level of damage done by deer in a square 4-ha plot, mapping all woody stems ≥1 cm diameter at breast height. The spatial distribution of stripping was used to examine neighbourhood effects using spatial point pattern methods. Results: The density and distance to neighbours and species identity of focal and neighbouring individuals contributed to a large spatial variation in neighbourhood effects (up to distances of 2-30 m from neighbours). Neighbourhood effects arose mainly from the interactions between neighbouring shrub species and all tree species, and interactions between individual species pairs. Neighbourhood effects more often increased than reduced the stripping of focal individuals. The stripping of Ulmus laevis, Acer campestre and Crataegus laevigata increased near conspecifics, while the stripping of all tree species was more likely near Ulmus laevis, Fraxinus angustifolia and shrubs. Heterospecific tree species were less susceptible to stripping near Tilia cordata and especially Carpinus betulus. Conclusion: Our study provides new insight into neighbourhood effects by showing how and at which spatial scales woody plant interactions modify deer herbivory. We advocate considering the life stage and relevant plant traits with multivariate spatial methods in order to elucidate the role of neighbourhood effects in complex plant communities. [ABSTRACT FROM AUTHOR]

Herbivore‐induced changes in both leaf silicon‐based defence and nutrient levels are potential mechanisms through which grazers alter the quality of their own grass supply. In tundra grasslands, herbivores have been shown to increase nutrient contents of grasses; yet, it is an open question whether they also increase grass silicon‐based defence levels. Here, we asked if, and to what extent, herbivores affect silicon content and silicon:nutrient ratios of grasses found in tundra grasslands.We performed an herbivore‐interaction field‐experiment spanning four tundra‐grassland sites. At each site, we established reindeer‐open and reindeer‐exclusion plots in tundra‐patches that had been disturbed or not by small rodents during the previous winter, for a total of 96 plots. We randomly collected over 1,150 leaf samples of inherently silicon‐rich and silicon‐poor grass species throughout a growing season and analysed silicon, nitrogen and phosphorus contents of each leaf.Small‐rodent winter disturbance did not affect grass silicon content, but increased grass quality (i.e. lowered silicon:nutrient ratios) by enhancing nutrient levels of both silicon‐rich (+20%–22%) and silicon‐poor (+26%–34%) grasses. Reindeer summer herbivory increased the quality of silicon‐rich grasses by decreasing their silicon content (−7%). However, the two herbivores together offset both these quality increments in silicon‐rich grasses, thus reducing their quality towards the level of those found in the absence of herbivores and further enhancing their silicon:nutrient ratios (+13%–22%) relative to silicon‐poor grasses.Synthesis. We provide the first community‐level, field‐based assessment of how herbivory‐driven changes in both leaf silicon‐based defence and nutrient levels alter grass‐forage quality in tundra grasslands. Herbivores did not promote a net silicon accumulation in grasses, but rather enhanced their overall quality. Yet, the magnitude of these quality increments varied depending on the herbivore(s) involved and differed between silicon‐rich and silicon‐poor grasses. Such differential herbivory‐induced changes in grass quality between and within tundra‐patches may mediate plant–herbivore interactions by altering herbivore forage patterns and food choices. In tundra‐patches utilised by both herbivores, the quality of silicon‐rich grasses was further decreased relative to that of silicon‐poor grasses. This could provide an advantage against herbivory, potentially being one of the pathways through which tundra‐grassland vegetation states dominated by silicon‐rich grasses are maintained by herbivores. [ABSTRACT FROM AUTHOR]

Theory predicts that a large fraction of phytochemical diversity—the richness of individual chemical compounds produced by plants—governs the complexity of interactions between plants and their herbivores. While the effect of specific classes of chemical compounds on plant resistance against herbivores has been largely documented, the effect of community‐level variation in phytochemical diversity on plant–herbivore interactions has so far received minimal consideration.We hypothesized that plant communities bearing on average higher levels of phytochemical diversity should sustain lower herbivory rates, overall. Yet, the magnitude of this effect could vary across different environmental conditions, potentially because of climate‐mediated effects on phytochemical production and changes in herbivore community richness and composition.To address these hypotheses, we used previous knowledge of species‐level phytochemical make‐up for more than 400 plant species of the Swiss Alps. Using common garden experiments, we estimated season‐wide herbivore damage on low (average 3,500 unique molecules) and high (average 4,500 unique molecules) phytochemical diversity plant communities that were planted in the colline, mountain and alpine vegetation sites along two elevation transects in the Alps.We found that high phytochemical diversity plant communities showed reduced levels of herbivore damage in the colline (low elevation) sites, but this pattern reversed in the alpine (high elevation) sites. Our results suggest that the outcome of phytochemical diversity on plant–herbivore interactions depends on the characteristics of the local herbivore communities, together with trade‐offs between chemical defences and other plant traits (i.e. physical defences and plant palatability).Synthesis. Phytochemical diversity is a key component of functional diversity, influencing community composition and dynamics. We show that the effect of phytochemical diversity on herbivory is environmental‐dependent, generating ecological switches when moving from low to high elevation. Through upward movement of plants under climate change, phytochemical community structure will be likely modified, ultimately disrupting local community assembly processes. [ABSTRACT FROM AUTHOR]

Summary: Overabundant deer populations can cause severe ecological impacts. To inform management decisions and assess the effectiveness of mitigation strategies, land managers require a rapid and cost‐effective method for collecting data on both deer density and impacts. Here, we describe a field method for rapidly and concurrently assessing indices of deer density and impacts in forested habitats, and using field data collected in 2019–2020, we demonstrate its performance. The method combines commonly used protocols for surveying deer faecal pellets, vegetation impact and floristic attributes, without relying on indicator species. The survey involves assessment of 150‐m transects in forested habitats along which 30 plots are surveyed for faecal pellets of deer and sympatric herbivores (here, Swamp Wallaby, Wallabia bicolor), and impacts on woody trees and shrubs at different heights. In total, we surveyed 154 transects comprising 15,941 plant impact assessments and assessed the methods performance using basic and comparative statistics, and power analyses. Almost all plant species were similarly impacted at the local (transect) level. Indices of deer and Swamp Wallaby density explained similar amounts of variance in impacts <1 m, while the deer index explained much more of the variance in impacts >1 m. We detected differences in vegetation impact and other floristic attributes between four locations surveyed in 2020 from a modest sample size (23–24 transects). The method is simple to implement and can be applied in a wide range of forest types. In comparison to alternatives (e.g. fenced exclosures), this method presents a rapid and cost‐effective means of collecting information on deer density and impacts, and detecting large differences in impact. We suggest that the proposed method enables rapid landscape‐scale assessments of deer impacts suitable for evaluating management interventions. [ABSTRACT FROM AUTHOR]

Introduction: The study of herbivory is fundamental in ecology and includes how plants invest in strategies and mechanisms to reduce herbivore damage. However, there is still a lack of information about how the environment, plant density, and functional traits influence herbivory in aquatic ecosystems. Objective: To assess if there is a relationship between herbivory, environmental variables, and plant traits two species of Montrichardia, a neotropical aquatic plant. Methods: In September 2018, we studied 78 specimens of Montrichardia arborescens and 18 of Montrichardia linifera, in 18 sites in Melgaço, Pará, Brazil. On each site, we measured water depth, distance to the margin, and plant density. From plants, we measured plant height and leaf thickness, and photographed the leaves to calculate the specific leaf area and percentage herbivory. To identify anatomical structures, we collected fully expanded leaves from three individuals per quadrat. Results: For M. arborescens, plants with thicker leaves and higher specific leaf area have less herbivore damage. For M. linifera, plants from deeper sites and with thicker leaves had more herbivore damage, while plants that grew farther from the margin had less damage. We found anatomical structures related to defense, such as idioblast cells with phenolic compounds, and cells with solid inclusions that can contribute to avoiding severe damage. Conclusions: Herbivory in these Montrichardia species can be explained by a combination of plant and environmental traits (patch isolation and water depth). The main plant traits are leaf thickness and area, but chemical compounds and solid inclusions also help Montrichardia to sustain less damage than other macrophytes. [ABSTRACT FROM AUTHOR]

Abstract Many insects use plant‐borne vibrations to obtain important information about their environment, such as where to find a mate or a prey, or when to avoid a predator. Plant species can differ in the way they vibrate, possibly affecting the reliability of information, and ultimately the decisions that are made by animals based on this information. We examined whether the production, transmission, and possible perception of plant‐borne vibrational cues is affected by variation in leaf traits. We recorded vibrations of 69 Spodoptera exigua caterpillars foraging on four plant species that differed widely in their leaf traits (cabbage, beetroot, sunflower, and corn). We carried out a transmission and an airborne noise absorption experiment to assess whether leaf traits influence amplitude and frequency characteristics, and background noise levels of vibrational chewing cues. Our results reveal that species‐specific leaf traits can influence transmission and potentially perception of herbivore‐induced chewing vibrations. Experimentally‐induced vibrations attenuated stronger on plants with thicker leaves. Amplitude and frequency characteristics of chewing vibrations measured near a chewing caterpillar were, however, not affected by leaf traits. Furthermore, we found a significant effect of leaf area, water content and leaf thickness—important plant traits against herbivory, on the vibrations induced by airborne noise. On larger leaves higher amplitude vibrations were induced, whereas on thicker leaves containing more water airborne noise induced higher peak frequencies. Our findings indicate that variation in leaf traits can be important for the transmission and possibly detection of vibrational cues.

Abstract The enemy release hypothesis (ERH) attributes the success of some exotic plant species to reduced top‐down effects of natural enemies in the non‐native range relative to the native range. Many studies have tested this idea, but very few have considered the simultaneous effects of multiple kinds of enemies on more than one invasive species in both the native and non‐native ranges. Here, we examined the effects of two important groups of natural enemies–insect herbivores and soil biota–on the performance of Tanacetum vulgare (native to Europe but invasive in the USA) and Solidago canadensis (native to the USA but invasive in Europe) in their native and non‐native ranges, and in the presence and absence of competition. In the field, we replicated full‐factorial experiments that crossed insecticide, T. vulgare–S. canadensis competition, and biogeographic range (Europe vs. USA) treatments. In greenhouses, we replicated full‐factorial experiments that crossed soil sterilization, plant–soil feedback, and biogeographic range treatments. We evaluated the effects of experimental treatments on T. vulgare and S. canadensis biomass. The effects of natural enemies were idiosyncratic. In the non‐native range and relative to populations in the native range, T. vulgare escaped the negative effects of insect herbivores but not soil biota, depending upon the presence of S. canadensis; and S. canadensis escaped the negative effects of soil biota but not insect herbivores, regardless of competition. Thus, biogeographic escape from natural enemies depended upon the enemies, the invader, and competition. Synthesis: By explicitly testing the ERH in terms of more than one kind of enemy, more than one invader, and more than one continent, this study enhances our nuanced perspective of how natural enemies can influence the performance of invasive species in their native and non‐native ranges.

Abstract To understand how comprehensive plant defense phenotypes will respond to global change, we investigated the legacy effects of elevated CO2 on the relationships between chemical resistance (constitutive and induced via mechanical damage) and regrowth tolerance in four milkweed species (Asclepias). We quantified potential resistance and tolerance trade‐offs at the physiological level following simulated mowing, which are relevant to milkweed ecology and conservation. We examined the legacy effects of elevated CO2 on four hypothesized trade‐offs between the following: (a) plant growth rate and constitutive chemical resistance (foliar cardenolide concentrations), (b) plant growth rate and mechanically induced chemical resistance, (c) constitutive resistance and regrowth tolerance, and (d) regrowth tolerance and mechanically induced resistance. We observed support for one trade‐off between plant regrowth tolerance and mechanically induced resistance traits that was, surprisingly, independent of CO2 exposure. Across milkweed species, mechanically induced resistance increased by 28% in those plants previously exposed to elevated CO2. In contrast, constitutive resistance and the diversity of mechanically induced chemical resistance traits declined in response to elevated CO2 in two out of four milkweed species. Finally, previous exposure to elevated CO2 uncoupled the positive relationship between plant growth rate and regrowth tolerance following damage. Our data highlight the complex and dynamic nature of plant defense phenotypes under environmental change and question the generality of physiologically based defense trade‐offs.

Abstract Plant defense suppression is an offensive strategy of herbivores, in which they manipulate plant physiological processes to increase their performance. Paradoxically, defense suppression does not always benefit the defense‐suppressing herbivores, because lowered plant defenses can also enhance the performance of competing herbivores and can expose herbivores to increased predation. Suppression of plant defense may therefore entail considerable ecological costs depending on the presence of competitors and natural enemies in a community. Hence, we hypothesize that the optimal magnitude of suppression differs among locations. To investigate this, we studied defense suppression across populations of Tetranychus evansi spider mites, a herbivore from South America that is an invasive pest of solanaceous plants including cultivated tomato, Solanum lycopersicum, in other parts of the world. We measured the level of expression of defense marker genes in tomato plants after infestation with mites from eleven different T. evansi populations. These populations were chosen across a range of native (South American) and non‐native (other continents) environments and from different host plant species. We found significant variation at three out of four defense marker genes, demonstrating that T. evansi populations suppress jasmonic acid‐ and salicylic acid‐dependent plant signaling pathways to varying degrees. While we found no indication that this variation in defense suppression was explained by differences in host plant species, invasive populations tended to suppress plant defense to a smaller extent than native populations. This may reflect either the genetic lineage of T. evansi—as all invasive populations we studied belong to one linage and both native populations to another—or the absence of specialized natural enemies in invasive T. evansi populations.

Abstract Plants have the capacity to alter their phenotype in response to environmental factors, such as herbivory, a phenomenon called phenotypic plasticity. However, little is known on how plant responses to herbivory are modulated by environmental variation along ecological gradients. To investigate this question, we used bilberry (Vaccinium myrtillus L.) plants and an experimental treatment to induce plant defenses (i.e., application of methyl jasmonate; MeJA), to observe ecological responses and gene expression changes along an elevational gradient in a boreal system in western Norway. The gradient included optimal growing conditions for bilberry in this region (ca. 500 m a.s.l.), and the plant's range limits at high (ca. 900 m a.s.l.) and low (100 m a.s.l.) elevations. Across all altitudinal sites, MeJA‐treated plants allocated more resources to herbivory resistance while reducing growth and reproduction than control plants, but this response was more pronounced at the lowest elevation. High‐elevation plants growing under less herbivory pressure but more resource‐limiting conditions exhibited consistently high expression levels of defense genes in both MeJA‐treated and untreated plants at all times, suggesting a constant state of “alert.” These results suggest that plant defense responses at both the molecular and ecological levels are modulated by the combination of climate and herbivory pressure, such that plants under different environmental conditions differentially direct the resources available to specific antiherbivore strategies. Our findings are important for understanding the complex impact of future climate changes on plant–herbivore interactions, as this is a major driver of ecosystem functioning and biodiversity.

Abstract Negative frequency‐dependent selection (NFDS) has been shown to maintain polymorphism in a diverse array of traits. The action of NFDS has been confirmed through modeling, experimental approaches, and genetic analyses. In this study, we investigated NFDS in the wild using morph‐frequency changes spanning a 20‐year period from over 30 dimorphic populations of Datura wrightii. In these populations, plants either possess glandular (sticky) or non‐glandular (velvety) trichomes, and the ratio of these morphs varies substantially among populations. Our method provided evidence that NFDS, rather than drift or migration, is the primary force maintaining this dimorphism. Most populations that were initially dimorphic remained dimorphic, and the overall mean and variance in morph frequency did not change over time. Furthermore, morph‐frequency differences were not related to geographic distances. Together, these results indicate that neither directional selection, drift, or migration played a substantial role in determining morph frequencies. However, as predicted by negative frequency‐dependent selection, we found that the rare morph tended to increase in frequency, leading to a negative relationship between the change in the frequency of the sticky morph and its initial frequency. In addition, we found that morph‐frequency change over time was significantly correlated with the damage inflicted by two herbivores: Lema daturaphila and Tupiochoris notatus. The latter is a specialist on the sticky morph and damage by this herbivore was greatest when the sticky morph was common. The reverse was true for L. daturaphila, such that damage increased with the frequency of the velvety morph. These findings suggest that these herbivores contribute to balancing selection on the observed trichome dimorphism.

Abstract Plant–herbivore interactions account for much of the biodiversity observed in an ecosystem. Plants may alter defense expression dependent on the specific herbivore present. Many studies focus on the optimization of these defenses against a single herbivore, but in an ecological context, there may be a myriad of herbivores at any given time. We designed a study to examine defenses employed by swamp white oak (Quercus bicolor) against insects and large mammals. We established six deer exclosures with trees located inside and outside each exclosure. Outside the exclosures, physical constraints only allow deer to feed up to a browseline on trees, whereas insects can potentially access all portions of the tree. Trees inside exclosures were only accessible to insect herbivores. Furthermore, insecticides were applied to half of the exclosures to create insect‐free (i.e., insecticide applied), deer‐free (i.e., inside exclosures), deer‐only (i.e., outside exclosures with insecticide), and herbivore‐free (i.e., inside exclosures with insecticide) environments. This design allowed us to analyze simultaneous defensive responses of individual trees to mammal and insect herbivores. We predicted that because insects represent a continuous threat, plants should invest in fixed concentrations of defenses (e.g., polyphenols, tannins, and condensed tannins), even when deer are absent. When deer are present, plants should increase investment in defenses in tissues that deer have access to. We found that plant defense strategies varied spatially on individual trees. When only insects were present, nutritional constituents (e.g., nitrogen and non‐structural carbohydrates) did not change but antinutritional constituents increased. Contrastingly, when deer were present, both nutritional and antinutritional constituents decreased. We found a decrease in insect functional group diversity when deer were present. Our findings suggest that plants may partition defense strategies depending on the most prominent herbivore pressure. However, changes in insect functional group diversity may result from either changes in plant defense strategy or intraspecific competition among multiple herbivores. Future studies should investigate patterns of chemical heterogeneity displayed by plants in response to diverse herbivore pressures.

Ecological theory and empirical research show that both direct lethal effects and indirect non-lethal effects can structure the composition of communities. While the direct effects of grazers on marine phytoplankton communities are well studied, their indirect effects are still poorly understood. Direct and indirect effects are inherently difficult to disentangle in plankton food webs. In this study we evaluate the indirect effects of copepod grazers on community function and structure using isolated chemical alarm signals, copepodamides. We expose intact summer and spring communities to direct grazing from copepods, or to chemical alarm cues without the presence of grazers in controlled experiments. The effects of direct grazing on ecosystem function were moderate in both experiments as indicated by levels of chlorophyll and primary production. Indirect and direct effects resulted in changes in the composition of both the eukaryote and prokaryote communities as shown by metabarcoding of 18S and 16S rRNA. Size structure analysis suggests that direct grazing and copepodamide exposure both favoured smaller organisms (< 10-15 μm) corroborating the size-structuring effect of copepod grazers. We conclude that the well-established effect of copepods on phytoplankton communities results from a combination of direct and indirect effects. This is a first attempt to isolate indirect effects of copepods on community structure and the results suggest that a full mechanistic understanding of the structuring effect of copepods will require insights to both direct and indirect effects of consumers as demonstrated for other ecosystems components., Competing Interests: None., (© The Author(s) 2024. Published by Oxford University Press.)

Bioassay with an insect herbivore is a common approach to studying plant defense. While measuring insect growth rate as a negative indicator of plant defense levels is simple and straightforward, analysing more detailed feeding behavior parameters of insects, such as feeding rates, leaf area consumed per feeding event, intervals between feeding events, and spatio-temporal patterns of feeding sites on leaves, is more informative. However, such observations are generally time consuming and labor-intensive. Here, we provide a semi-automated system for quantifying feeding behavior parameters of insects feeding on plant leaves. Automated photo scanners record the time-course development of feeding marks on leaves. An image analysis pipeline processes the scanned images and extracts leaf area. By analysing changes in leaf area over time, it detects insect feeding events and calculates the leaf area consumed during each feeding event, providing quantitative parameters of the feeding behavior of insects. In addition, it visualizes spatio-temporal changes in feeding sites, providing a measure of the complex behavior of insects on leaves. Using this analysis pipeline, we demonstrate that Arabidopsis trichomes reduce insect feeding rate, but not feeding duration or intervals between feeding events. Our image acquisition system requires only a photo scanner and a laptop computer and does not require any specialized equipment. The analysis software is provided as an ImageJ macro and R package and is available at no cost. Taken together, our work provides a scalable method for quantitative assessment of the feeding behavior of insects on leaves, facilitating understanding of plant defense mechanisms., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

Plant chemodiversity, the diversity of plant-specialized metabolites, is an important dimension of biodiversity. However, there are so far few mathematical models to test verbal hypotheses on how chemodiversity evolved. Here, we develop such a model to test predictions of five hypotheses: the 'fluctuating selection hypothesis', the 'dominance reversal hypothesis', the interaction diversity hypothesis, the synergy hypothesis, and the screening hypothesis. We build a population genetic model of a plant population attacked by herbivore species whose occurrence fluctuates over time. We study the model using mathematical analysis and individual-based simulations. As predicted by the 'dominance reversal hypothesis', chemodiversity can be maintained if alleles conferring a defense metabolite are dominant with respect to the benefits, but recessive with respect to costs. However, even smaller changes in dominance can maintain polymorphism. Moreover, our results underpin and elaborate predictions of the synergy and interaction diversity hypotheses, and, to the extent that our model can address it, the screening hypotheses. By contrast, we found only partial support for the 'fluctuating selection hypothesis'. In summary, we have developed a flexible model and tested various verbal models for the evolution of chemodiversity. Next, more mechanistic models are needed that explicitly consider the organization of metabolic pathways., (© 2024 The Author(s). New Phytologist © 2024 New Phytologist Foundation.)

Plant-soil interactions have bottom-up and top-down effects within a plant community. Heavy metal pollution can change plant-soil interactions, directly influence bottom-up effects and indirectly affect herbivores within the community. In turn, herbivores can affect plant-soil interactions through top-down effects. However, the combined effects of heavy metals and herbivores on soil enzymes, plants and herbivores have rarely been reported. Therefore, the effects of lead (Pb), Spodoptera litura and their combined effects on soil enzyme activities, pakchoi nutrition, defence compounds and S. litura fitness were examined here. Results showed that Pb, S. litura and their combined effects significantly affected soil enzymes, pakchoi and S. litura . Specifically, exposure to double stress (Pb and S. litura ) decreased soil urease, phosphatase and sucrase activities compared with controls. Furthermore, the soluble protein and sugar contents of pakchoi decreased, and the trypsin inhibitor content and antioxidant enzyme activity increased. Finally, the S. litura development period was extended, and survival, emergence rates and body weight decreased after exposure to double stress. The combined stress of Pb and S. litura significantly decreased soil enzyme activities. Heavy metal accumulation in plants may create a superposition or synergistic effect with heavy metal-mediated plant chemical defence, further suppressing herbivore development. Pb, S. litura and their combined effects inhibited soil enzyme activities, improved pakchoi resistance and reduced S. litura development. The results reveal details of soil-plant-herbivore interactions and provide a reference for crop pest control management in the presence of heavy metal pollution.

Premise: Polyploidy is a widespread mutational process in angiosperms that may alter population performance of not only plants but also their interacting species. Yet, knowledge of whether polyploidy affects plant-herbivore dynamics is scarce. Here, we tested whether aphid herbivores exhibit preference for diploid or neopolyploid plants, whether polyploidy impacts plant and herbivore performance, and whether these interactions depend on the plant genetic background., Methods: Using independently synthesized neotetraploid strains paired with their diploid progenitors of greater duckweed (Spirodela polyrhiza), we evaluated the effect of neopolyploidy on duckweed's interaction with the water-lily aphid (Rhopalosiphum nymphaeae). Using paired-choice experiments, we evaluated feeding preference of the herbivore. We then evaluated the consequences of polyploidy on aphid and plant performance by measuring population growth over multiple generations., Results: Aphids preferred neopolyploids when plants were provided at equal abundances but not at equal surface areas, suggesting the role of plant population surface area in driving this preference. Additionally, neopolyploidy increased aphid population performance, but this result was dependent on the plant's genetic lineage. Lastly, the impact of herbivory on neopolyploid vs. diploid duckweed varied greatly with genetic lineage, where neopolyploids appeared to be variably tolerant compared to diploids, sometimes mirroring the effect on herbivore performance., Conclusions: By experimentally testing the impacts of polyploidy on trophic species interactions, we showed that polyploidization can impact the preference and performance of herbivores on their plant hosts. These results have significant implications for the establishment and persistence of plants and herbivores in the face of plant polyploidy., (© 2024 The Authors. American Journal of Botany published by Wiley Periodicals LLC on behalf of Botanical Society of America.)

Animals interact with nutrient cycles by consuming and depositing nutrients, interactions studied separately in nutritional ecology and zoogeochemistry. Recent theoretical work bridges these disciplines, highlighting that animal-driven nutrient recycling could be crucial in helping animals meet their nutritional needs. When animals exhibit site fidelity, they consistently deposit nutrients, potentially improving vegetation quality. We investigated this potential feedback by analysing changes in forage nitrogen stocks following simulated caribou calving. We found that forage nitrogen stocks increased after 2 weeks and remained elevated after 1 year, a change due to increased forage quality, not quantity. We also developed a nutrient budget within calving grounds, demonstrating that natal fluid and calf carcasses contribute substantial nitrogen subsidies. We, thus, highlight a positive zoogeochemical feedback whereby nutrients deposited during calving become bioavailable during lactation and provide evidence that site fidelity creates a biogeochemical boomerang in which animals deposit nutrients that can be reused later., (© 2024 John Wiley & Sons Ltd.)

Plants employ diverse anti-herbivore defences that can covary to form syndromes consisting of multiple traits. Such syndromes are hypothesized to impact herbivores more than individual defences. We studied 16 species of lowland willows occurring in central Europe and explored if their chemical and physical traits form detectable syndromes. We tested for phylogenetic trends in the syndromes and explored whether three herbivore guilds (i.e., generalist leaf-chewers, specialist leaf-chewers, and gallers) are affected more by the detected syndromes or individual traits. The recovered syndromes showed low phylogenetic signal and were mainly defined by investment in concentration, richness, or uniqueness of structurally related phenolic metabolites. Resource acquisition traits or inducible volatile organic compounds exhibited a limited correlation with the syndromes. Individual traits composing the syndromes showed various correlations to the assemblages of herbivores from the three studied guilds. In turn, we found some support for the hypothesis that defence syndromes are composed of traits that provide defence against various herbivores. However, individual traits rather than trait syndromes explained more variation for all studied herbivore assemblages. The detected negative correlations between various phenolics suggest that investment trade-offs may occur primarily among plant metabolites with shared metabolic pathways that may compete for their precursors. Moreover, several traits characterizing the recovered syndromes play additional roles in willows other than defence from herbivory. Taken together, our findings suggest that the detected syndromes did not solely evolve as an anti-herbivore defence., (© 2024. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

The relationship between herbivory and latitude may differ between native and introduced populations of invasive plants, which can generate latitudinal heterogeneity in the strength of enemy release. However, still little is known about how latitudinal heterogeneity in herbivore pressure influences latitudinal variation in defense phenotypes of invasive plants. We tested how latitudinal patterns in multi‐variate defense syndromes differed between native (Argentinian) and introduced (Chinese) populations of the invasive herb Alternanthera philoxeroides. In addition, to better understand the drivers underlying latitudinal patterns, we also tested whether associations of defense syndromes with climate and herbivory differed between native and introduced ranges. We found that native plant populations clustered into three main defense syndromes associated with latitude. In contrast, we only found two defense syndromes in the introduced range. One matched the high‐latitude syndrome from the native range, but was distributed at both the northern and southern range limits in the introduced range. The other was unique to the introduced range and occurred at mid‐latitudes. Climatic conditions were associated with variation in syndromes in the native range, and climatic conditions and herbivory were associated with variation in syndromes in the introduced range. Together, our results demonstrate that plants may under the new environmental conditions in the introduced range show latitudinal patterns of defense syndromes that are different from those in their native range. This emphasizes that geographical dependence of population differentiation should be explicitly considered in studies on the evolution of defense in invasive plants. [ABSTRACT FROM AUTHOR]

SUMMARY: Crop damage by herbivorous insects remains a significant contributor to annual yield reductions. Following attack, maize (Zea mays) responds to herbivore‐associated molecular patterns (HAMPs) and damage‐associated molecular patterns (DAMPs), activating dynamic direct and indirect antiherbivore defense responses. To define underlying signaling processes, comparative analyses between plant elicitor peptide (Pep) DAMPs and fatty acid–amino acid conjugate (FAC) HAMPs were conducted. RNA sequencing analysis of early transcriptional changes following Pep and FAC treatments revealed quantitative differences in the strength of response yet a high degree of qualitative similarity, providing evidence for shared signaling pathways. In further comparisons of FAC and Pep responses across diverse maize inbred lines, we identified Mo17 as part of a small subset of lines displaying selective FAC insensitivity. Genetic mapping for FAC sensitivity using the intermated B73 × Mo17 population identified a single locus on chromosome 4 associated with FAC sensitivity. Pursuit of multiple fine‐mapping approaches further narrowed the locus to 19 candidate genes. The top candidate gene identified, termed FAC SENSITIVITY ASSOCIATED (ZmFACS), encodes a leucine‐rich repeat receptor‐like kinase (LRR‐RLK) that belongs to the same family as a rice (Oryza sativa) receptor gene previously associated with the activation of induced responses to diverse Lepidoptera. Consistent with reduced sensitivity, ZmFACS expression was significantly lower in Mo17 as compared to B73. Transient heterologous expression of ZmFACS in Nicotiana benthamiana resulted in a significantly increased FAC‐elicited response. Together, our results provide useful resources for studying early elicitor‐induced antiherbivore responses in maize and approaches to discover gene candidates underlying HAMP sensitivity in grain crops. Significance Statement: Expression analyses and forward genetics approaches were used to identify a maize locus and receptor kinase gene candidate associated with response sensitivity to fatty acid–amino acid conjugate (FAC) elicitors widely present in insect oral secretions. Despite broad overlap, herbivore‐associated molecular pattern (HAMP) responses can be decoupled from damage‐associated molecular pattern (DAMP) responses, providing defined resources to investigate early signaling events underlying plant responses to diverse lepidoptera herbivores. [ABSTRACT FROM AUTHOR]

Arbuscular mycorrhizal fungi (AMF) develop strong mutualistic associations with roots of about 70% of all vascular plants. By modifying host plant nutrient uptake, growth and defense, AMF also indirectly influence the aboveground herbivorous insect community. Since plant response to AMF depends on the extent of mycorrhization, outcomes of the bottom-up effects of AMF on herbivores are extremely variable and not well understood. We thus tested whether the generalist phloem feeder Bemisia tabaci (tobacco whitefly) was affected by the colonization of habanero pepper seedlings (Capsicum chinense) by different densities of the AMF Rhizophagus irregularis. After AMF inoculation (with control, low and high doses of liquid inoculum), seedlings were grown in a greenhouse without (control) and with whiteflies (10 adults per seedling). We measured plant traits and growth and biomass allocations at 2, 16 and 30 days after transplantation of emerged seedlings. We estimated whitefly adult, egg and nymph densities 28 days after transplantation. B. tabaci abundance significantly increased after C. chinense inoculation with low AMF density (120% increase for adults, 97% for eggs) through an augmentation of seedling nutritional status. By enhancing plant tolerance and primary metabolism, the higher density of AMF did not affect B. tabaci fitness on seedlings. We highlight here that whitefly abundance on mycorrhizal C. chinense varies widely depending on AMF inoculum concentration. [ABSTRACT FROM AUTHOR]

Summary: Understanding the complexities of pasture responses (the overall seasonal gains or losses in herbage mass) under different environmental conditions is critical to the effective management of total grazing pressure, whether to achieve agricultural, cultural or biodiversity conservation outcomes. Whilst large‐scale management programmes for kangaroos (Macropus spp. and Osphranter spp.) occur across much of south‐eastern Australia, there is limited understanding of how such programmes influence the overall pasture response across different environmental contexts in the temperate environment. This study investigates the effects of potential grass productivity (measured as the growth of pasture protected from large herbivore grazing), initial herbage mass (measured as the attached, above‐ground component of grass tussocks), kangaroo density (measured at the site scale) and season on the overall pasture response in plots dominated by six common grass types from south‐eastern Australia. Using generalised additive mixed modelling, we demonstrate potential grass productivity (related to but not synonymous with grass palatability) to be the major predictor variable for pasture responses across all six grass types. Kangaroo density, either directly or via an interaction with initial herbage mass, was related to overall pasture responses for most grasses in spring and remained an important variable in favoured areas dominated by preferred Kangaroo Grass (Themeda triandra) and Redleg Grass (Bothriochloa macra) throughout the autumn survey period. Plots dominated by grazing‐tolerant species, including wallaby grasses (Rytidosperma spp.) and Phalaris (Phalaris aquatica; an introduced pasture grass), showed marginally positive pasture responses under higher kangaroo densities, whilst plots with Wild Oats (Avena fatua; an introduced annual grass) and less preferred native spear grasses (Austrostipa spp.) showed no significant effect of kangaroo density on pasture response. Results support a context‐specific mix of 'bottom‐up' (resource‐driven) and 'top‐down' (consumer‐driven) processes in temperate grassy ecosystems, highlighting the importance of considering landscape productivity, existing herbage mass and grass species composition when informing kangaroo management decisions. [ABSTRACT FROM AUTHOR]

Host plant selection by herbivores is driven by a complex array of cues, including leaf traits and previous leaf damage. Herbivore‐associated cues to host selection at the plant and leaf scale aid understanding of mechanisms responsible for host preference that might translate into increased performance, as well as processes structuring herbivore populations mediated by interactions. We investigated how changes induced by a galling insect in the tropical fern Cyathea phalerata act as repellent or attractant cues for sawfly feeding and the effects of leaf size on herbivory levels.We recorded gall abundance, damage by chewers, leaf size, plant nutritional quality, phenolic concentration and leaf anatomical traits between galled and non‐galled leaf samples.Galled samples contained less N, higher levels of phenolics and higher C/N ratio. However, leaf‐chewing damage did not differ between galled and non‐galled leaves. The gall structure was avoided by chewers, as it had high concentrations of phenolics, lignification and suberization. Larger leaves sustained higher gall abundance, but leaf size did not have a significant effect on chewer damage. A co‐occurrence index calculated for both guilds indicated that galls and chewers exhibited a distribution that did not differ from random, reinforcing that the two guilds on C. phalerata do not show patterns of repulsion such as those maintained by interspecific competition.Sawflies dismissing chemical cues indicate that the increase in phenolics caused by galling insects does not generate increased protection of the galled pinnules. Our results highlight ferns as key resources for herbivores and as a potential plant group to study new research avenues on plant–insect interactions. [ABSTRACT FROM AUTHOR]

Many theories regarding the evolution of inducible resistance in plants have an implicit spatial component, but most relevant population dynamic studies ignore spatial dynamics. We examined a spatially explicit model of plant inducible resistance and herbivore population dynamics to explore how realistic features of resistance and herbivore responses influence spatial patterning. Both transient and persistent spatial patterns developed in all models examined, where patterns manifested as wave-like aggregations of herbivores and variation in induction levels. Patterns arose when herbivores moved away from highly induced plants, there was a lag between damage and deployment of induced resistance, and the relationship between herbivore density and strength of the induction response had a sigmoid shape. These mechanisms influenced pattern formation regardless of the assumed functional relationship between resistance and herbivore recruitment and mortality. However, in models where induction affected herbivore mortality, large-scale herbivore population cycles driven by the mortality response often co-occurred with smaller scale spatial patterns driven by herbivore movement. When the mortality effect dominated, however, spatial pattern formation was completely replaced by spatially synchronized herbivore population cycles. Our results present a new type of ecological pattern formation driven by induced trait variation, consumer behavior, and time delays that has broad implications for the community and evolutionary ecology of plant defenses.

Herbivory is a key innovation in insects, yet has only evolved in one-third of living orders. The evolution of herbivory likely involves major behavioral changes mediated by remodeling of canonical chemosensory modules. Herbivorous flies in the genus Scaptomyza (Drosophilidae) are compelling species in which to study the genomic architecture linked to the transition to herbivory because they recently evolved from microbe-feeding ancestors and are closely related to Drosophila melanogaster. We found that Scaptomyza flava, a leaf-mining specialist on plants in the family (Brassicaceae), was not attracted to yeast volatiles in a four-field olfactometer assay, whereas D. melanogaster was strongly attracted to these volatiles. Yeast-associated volatiles, especially short-chain aliphatic esters, elicited strong antennal responses in D. melanogaster, but weak antennal responses in electroantennographic recordings from S. flava. We sequenced the genome of S. flava and characterized this species' odorant receptor repertoire. Orthologs of odorant receptors, which detect yeast volatiles in D. melanogaster and mediate critical host-choice behavior, were deleted or pseudogenized in the genome of S. flava. These genes were lost step-wise during the evolution of Scaptomyza. Additionally, Scaptomyza has experienced gene duplication and likely positive selection in paralogs of Or67b in D. melanogaster. Olfactory sensory neurons expressing Or67b are sensitive to green-leaf volatiles. Major trophic shifts in insects are associated with chemoreceptor gene loss as recently evolved ecologies shape sensory repertoires.

Increased ungulate browsing alters the composition of plant communities and modifies forest ecosystems worldwide. Ungulates alter their diet following changes in availability of plant species; however, we know little about how browse selection and plant community composition change with different stages of deer establishment. Here, we provide insight into this area of study by combining multiple approaches: comparison of the understory plant community, analysis of records of browsing damage, and DNA barcoding of sika deer feces at 22 sites in forests in northern Japan varying in when deer were first established. The coverage of vegetation and number of plant species were only lower at sites where deer were present for more than 20 years, while the difference in plant coverage among deer establishment years varied among plant species. Deer diet differed across establishment years, but was more affected by the site, thereby indicating that food selection by deer could change over several years after deer establishment. Plant life form and plant architecture explained the difference in plant coverage across establishment years, but large variability was observed in deer diet within the two categories. Integrating these results, we categorized 98 plant taxa into six groups that differed in vulnerability to deer browsing (degree of damage and coverage). The different responses to browsing among plant species inferred from this study could be a first step in predicting the short- and long-term responses of forest plant communities to deer browsing.

Aims: Plants with a history of overgrazing show trait-mediated legacy effects. These legacy effects strongly influence the growth dynamics and stress tolerance of plants, thus affecting ecosystem functioning. Long-term overgrazing has dramatic effects on plant growth and carbon assimilation via asexual propagation. However, the link between nitrogen (N) absorbability and assimilation with grazing-induced plant legacy effects remains largely unknown. Methods: We investigated the strength of legacy effects induced by long-term overgrazing on N uptake and metabolism in the clonal plant Leymus chinensis, and its associated changes at the physiological and molecular levels. These tests were conducted in both field and greenhouse experiments. Results: The clonal offspring of overgrazed L. chinensis were significantly smaller than the control offspring, with lower individual N uptake and utilisation efficiency, indicating that the N dynamics were affected by plant legacy effects. The response ratios of root length and biomass to N patches in the clonal offspring of overgrazed L. chinensis were significantly higher than those of the control, indicating that root nutrient foraging plasticity increased in response to grazing-induced N heterogeneity. Moreover, the observed plant legacy effects decreased N acquirement but significantly increased N assimilation by increasing N resorption efficiency, with biotic stress memory activated at the enzymatic and transcriptional levels. Conclusions: We propose that multigenerational exposure of perennial plants to herbivore foraging can produce a legacy effect on N uptake, which offers insights into the potential resilience of grasslands to overgrazing. [ABSTRACT FROM AUTHOR]

Recovery of green turtles (Chelonia mydas), mega‐herbivores that consume seagrasses, is resulting in dramatic ecosystem‐wide changes as meadows are returned to a natural grazed state. The green turtle grazing strategy, with long‐term cultivation of meadows and high foraging site fidelity, is distinct from other terrestrial and aquatic mega‐herbivores and may affect seagrass compensatory growth responses. Identifying the mechanisms of compensatory growth responses to grazing is essential to understand the functioning of plant systems under natural grazing regimes.In a naturally grazed Caribbean seagrass ecosystem, we identify a mechanism for compensatory growth responses to grazing by evaluating relationships between Thalassia testudinum morphology and growth, grazing intensity, and canopy light dynamics in grazed and ungrazed areas.The morphological characteristics that explain variability in T. testudinum growth differed between grazed and ungrazed areas. In grazed areas, T. testudinum leaf linear growth, leaf area growth, and productivity:biomass (P:B) significantly increased as above‐ground biomass decreased; P:B also increased with shoot density. Mass growth in grazed areas exhibited an increasing trend with shoot density and was maintained above a threshold of 2.5 g dry mass m−2 above‐ground biomass. In ungrazed areas, trends for mass growth and P:B with above‐ground biomass and shoot density were opposite to those in grazed areas. In grazed areas, shoot density significantly increased with grazing intensity while above‐ground biomass decreased and leaf area index (LAI) was not affected. Light availability at canopy height was greater in grazed areas than in ungrazed areas, and canopy light attenuation increased with shoot density in grazed areas.Synthesis. Grazing removes above‐ground biomass, which increases light availability and stimulates leaf growth and turnover (i.e. compensatory growth). Shoot density increases with grazing intensity, maintaining LAI and canopy light harvesting potential. This maximizes the potential for leaf photosynthetic activity and provides the plant with the capacity to sustain mass growth and support a compensatory growth response to grazing. This study presents novel insight for assessing the underlying mechanisms of plant compensatory growth responses to cultivation grazing and proposes potential thresholds that may be used to evaluate the sustainability of in situ grazing pressure by a recovering mega‐herbivore. [ABSTRACT FROM AUTHOR]

Herbivory is a key process structuring vegetation in savannas, especially in Africa where large mammal herbivore communities remain intact. Exclusion experiments consistently show that herbivores impact savanna vegetation, but effect size variation has resisted explanation, limiting our understanding of the past, present and future roles of herbivory in savanna ecosystems.Synthesis of vegetation responses to herbivore exclusion shows that herbivory decreased grass abundance by 57.0% and tree abundance by 30.6% across African savannas.The magnitude of herbivore exclusion effects scaled with herbivore abundance: more grazing herbivores resulted in larger grass responses and more browsing herbivores in larger tree responses. However, existing experiments are concentrated in semi‐arid savannas (400–800‐mm rainfall) and soils data are mostly lacking, which makes disentangling environmental constraints a challenge and priority for future research.Observed herbivore impacts were ~2.1× larger than existing estimates modelled based on consumption. Wildlife metabolic rates may be higher than are usually used for estimating consumption, which offers one clear avenue for reconciling estimated herbivore consumption with observed herbivore impacts. Plant‐soil feedbacks, plant community composition, and the phenological or demographic timing of herbivory may also influence vegetation productivity, thereby magnifying herbivore impacts.Because herbivore abundance so closely predicts vegetation impact, changes in herbivore abundance through time are likely predictive of the past and future of their impacts. Grazer diversity in Africa has declined from its peak 1 million years ago and wild grazer abundance has declined historically, suggesting that grazing likely had larger impacts in the past than it does today.Current wildlife impacts are dominated by small‐bodied mixed feeders, which will likely continue into the future, but the magnitude of top‐down control may also depend on changing climate, fire and atmospheric CO2.Synthesis. Herbivore biomass determines the magnitude of their impacts on savanna vegetation, with effect sizes based on direct observation that outstrip existing modelled estimates across African savannas. Findings suggest substantial ecosystem impacts of herbivory and allow us to generate evidence‐based hypotheses of the past and future impacts of herbivores on savanna vegetation. [ABSTRACT FROM AUTHOR]

Restoration of dryland ecosystems is often limited by low seedling establishment and survival. Defoliation caused by insects and small mammals could be an overlooked cause of seedling mortality. In the sagebrush steppe, we examined the effect of seedling defoliation on the survival of perennial grasses commonly used as restoration materials. Under field conditions, seedlings of three perennial bunchgrass species (non-native Agropyron cristatum , and native grasses Poa secunda and Pseudoroegneria spicata) were defoliated at two intensities (30 % and 70 % leaf length removal) and frequencies (one or two clippings) and compared to a non-defoliated control. Following emergence the first year, clippings occurred at the two-leaf stage; a second clipping occurred 1 month later for repeated defoliation treatments. We monitored seedling survival and tillering for 2 years. We expected higher defoliation intensity and frequency to reduce survival for all species, but only a few treatments reduced Po. secunda survival. Conversely, larger-statured Triticeae (wheatgrasses) benefited from some defoliation treatments. In both years, A. cristatum survival increased with repeated defoliation at both intensities. Defoliation did not affect Ps. spicata survival in the first year, but a single defoliation in the second year resulted in increased survival. In both A. cristatum and Ps. spicata , higher-intensity defoliation reduced the boost to survival resulting from defoliation frequency. Seedlings with more tillers had greater survival probabilities, but tiller number was unaffected by defoliation. Further research may elucidate mechanisms seedlings use to compensate for or benefit from defoliation. In the meantime, managers should aim to select defoliation-tolerant species if they anticipate herbivory will be problematic for restoration sites. [ABSTRACT FROM AUTHOR]

The consequences of the introduction of invasive plants for the diet of herbivorous insects have been little explored in nature where, potentially, abiotic and biotic factors operate. In this study, we examined the larval performance of two Neotropical Danaini butterflies when using either a native or an exotic Apocynaceae species as host plant in both field and laboratory experiments. Hosts greatly differ in their amount of latex exudation and other physicochemical traits, as well as in the amount of evolutionary time they have interacted with herbivores. First, herbivore performance on the hosts was investigated under laboratory conditions. Larvae of both Danaini species took more time to develop on the exotic host; larval survivorship did not vary between hosts. Second, first instar survivorship on both hosts was evaluated in two field sites, one site per host. To do so, in both sites half of the larvae were bagged (protected against both abiotic and biotic factors) while the remainder were nonbagged (exposed). The interaction between larval exposure with the use of the exotic host reduced larval survival. We concluded that the combined effects of host plant traits and abiotic factors reduced survival of herbivores in field conditions. Therefore, the performance of herbivores when using hosts of different origins should be considered together with the multiple ecological factors found in natural environments, as these factors can modify the result of plant-herbivore interactions. [ABSTRACT FROM AUTHOR]

A central challenge in ecology and evolutionary biology is to understand how variation in abiotic and biotic factors combine to shape the distribution, abundance, and diversity of focal species. Environmental gradients, whether natural (e.g., latitude, elevation, ocean proximity) or anthropogenic (e.g., land-use intensity, urbanization), provide compelling settings for addressing this challenge. However, not all organisms are amenable to the observational and experimental approaches required for untangling the factors that structure species along gradients. Here we highlight herbaceous plants in the genus Plantago as models for studying the ecology and evolution of species interactions along abiotic gradients. Plantago lanceolata and P. major are native to Europe and Asia but distributed globally, and they are established models for studying population ecology and interactions with herbivores, pathogens, and soil microbes. Studying restricted range congeners in comparison with those cosmopolitan species can provide insight into abiotic and biotic determinants of range size and population structure. We highlight one such species, P. rugelii , which is endemic to eastern North America. We give an overview of the literature on these focal Plantago species and explain why they are logical candidates for studies of species interactions across environmental gradients. Finally, we emphasize collaborative and community science approaches that can facilitate such research and note the amenability of Plantago for authentic research projects in science education. [ABSTRACT FROM AUTHOR]

Despite increasing evidence that herbivore‐mediated indirect effects play a major role in plant competition, it is unclear how and when they contribute to plant invasiveness. The outcomes of herbivore‐mediated indirect effects are primarily dependent on the environment and have complex interactions with the direct interactions between plants.We evaluated the herbivore‐mediated indirect effects of Solidago altissima (Asteraceae) on other co‐occurring native Asteraceae species at multiple sites in a native range [the United States (US)] and in an introduced range (Japan) using common garden experiments. We examined the effect of S. altissima on herbivore damage, above‐ground vegetative and reproductive traits of the co‐occurring Asteraceae species at sites with different densities of a herbivore, the lace bug, Corythucha marmorata, which is native to the US and introduced in Japan.We observed increased lace bug herbivory in plants grown with S. altissima in Japan but decreased lace bug herbivory in the US. We did not find consistent effect of S. altissima on aboveground vegetative production of co‐occurring plants in either ranges. Flower production decreased in plants grown with S. altissima in Japan because of both direct competition and increased herbivory. Direct competition and apparent competition via lace bug herbivory interacted to produce a strong negative effect of S. altissima on co‐occurring plants only in Japan.Synthesis. The findings imply that evolutionary history and the local environment jointly affect herbivore‐mediated indirect effects, which play a large role in shaping the pattern of herbivory and plant competition. Another implication of our results is that exotic herbivores may reinforce the negative effects of exotic plants on native plants under high herbivore density environments. [ABSTRACT FROM AUTHOR]

Identifying food web linkages between biocontrol agents of invasive plants and native species is crucial for predicting indirect non‐target effects. Biocontrol insects can integrate into food webs within recipient habitats and influence native insects through apparent competition (altering shared natural enemies) or density‐mediated exploitation competition (changing density of native plants). However, whether and how trait‐mediated exploitation competition (modifying native plant chemicals and volatiles profiles) can produce indirect non‐target effects remains largely overlooked, despite plant phenotypic responses to insect herbivory being common and widely documented.The flea beetle Agasicles hygrophila was introduced into China for management of alligator weed Alternanthera philoxeroides, but it also attacks the native congener sessile joyweed Alternanthera sessilis, which may cause indirect non‐target effects on the native tortoise beetle Cassida piperata that also feeds on sessile joyweed. Here, we examined the relationships among abundances of the flea beetle and tortoise beetle, and coverage of sessile joyweed in the field. Then, we investigated the impact of flea beetle herbivory on tortoise beetle development and oviposition, as well as on sessile joyweed primary metabolites and leaf volatiles.Tortoise beetle abundance was not related to sessile joyweed coverage, but they were less abundant on plants with more flea beetles in the field survey, and produced fewer offspring on plants with more prior flea beetle damage in the field cage experiment. Tortoise beetle development was inhibited by prior flea beetle herbivory in bioassays in enemy‐free conditions and they preferred to oviposit on sessile joyweed that had experienced little or no flea beetle damage. Flea beetle herbivory decreased sessile joyweed foliar glucose and protein, and substantially changed its leaf volatile blend.Synthesis. Our results show that the flea beetle has major indirect non‐target effects on the tortoise beetle through trait‐mediated exploitation competition, rather than apparent competition or density‐mediated exploitation competition. Our results demonstrate a new example for indirect non‐target effects of biocontrol agents. Furthermore, our results indicate that minor brief negative impacts of biocontrol agents on non‐target plants might propagate to higher trophic levels and such negative impacts can strengthen with increasing intensity of the direct non‐target effect. [ABSTRACT FROM AUTHOR]