Types Of Herbivores Research Articles

Disentangling herbivore impacts on Populus tremuloides: a comparison of native ungulates and cattle in Canada’s Aspen Parkland

Ungulates impact woody species' growth and abundance but little is understood about the comparative impacts of different ungulate species on forest expansion in savanna environments. Replacement of native herbivore guilds with livestock [i.e., beef cattle (Bos taurus)] has been hypothesized as a factor facilitating trembling aspen (Populus tremuloides Michx.) encroachment into grasslands of the Northern Great Plains. We used a controlled herbivory study in the Parklands of western Canada to compare the impact of native ungulates and cattle on aspen saplings. Native ungulate treatments included a mixed species guild and sequences of herbivory by different ungulates [bison (Bison bison subsp. bison), elk (Cervus elaphus) then deer (Odocoileus hemionus); or deer, elk, then bison]. Herbivory treatments were replicated in three pastures, within which sets of 40 marked aspen saplings (<1.8 m) were tracked along permanent transects at 2-week intervals, and compared to a non-grazed aspen stand. Stems were assessed for mortality and incremental damage (herbivory, leader breakage, stem abrasion and trampling). Final mortality was greater with exposure to any type of herbivore, but remained similar between ungulate treatments. However, among all treatments, the growth of aspen was highest with exposure only to cattle. Herbivory of aspen was attributed primarily to elk within the native ungulate treatments, with other forms of physical damage, and ultimately sapling mortality, associated with exposure to bison. Overall, these results indicate that native ungulates, specifically elk and bison, have more negative impacts on aspen saplings and provide evidence that native and domestic ungulates can have different functional effects on woody plant dynamics in savanna ecosystems.

Herbivory and habitat association of tree seedlings in lowland evergreen rainforest on white-sand and terra-firme in the upper Rio Negro

Background: It has been proposed that the interaction between herbivory and soil nutrient availability drives habitat association of tree species in Peruvian Amazonia. Nevertheless, there is no empirical evidence that this interaction holds across other Amazonian regions. Aims: We address this knowledge gap by testing whether the interaction between herbivory and soil nutrient contributes to habitat association of tree species in white-sand and terra-firme forests in the upper Rio Negro, Brazil. Methods: We conducted a reciprocal transplanting field experiment in which we controlled for the presence of herbivores. We tested for differences in tree-seedling growth and herbivory among seven white-sand and seven terra-firme habitat-specialist species. Additionally, we assessed whether tree seedlings differed in their functional traits. Results: We found no empirical evidence that an interaction between herbivory and soil nutrients shapes habitat association in white-sand and terra-firme forests of the upper Rio Negro. Tree seedlings showed higher mortality when growing in their non-typical habitat. Growth and herbivory were similar regardless of the presence or absence of herbivore protection and type of soil. Conclusions: We suggest that the overall differences in soil nutrient status between white-sand and terra-firme forests in the upper Rio Negro are insufficient to trigger an interaction between herbivory and soil nutrient availability.

Omnipresence of leaf herbivory by invertebrates and leaf infection by fungal pathogens in agriculturally used grasslands of the Swiss Alps, but low plant damage

Agriculturally used grasslands in the Alps are characterised by a trade-off between high fodder production in some and high plant species richness in others. In contrast to plant species richness and production, however, little is known on the relevance of biological interactions between plants, invertebrate herbivores, and fungal pathogens for grasslands in the Alps. At the time when the vegetation was fully developed, but prior to agricultural use, we examined whether leaf damage by herbivory and fungal pathogen infection, and their diversity, are affected by plant functional group, land use, and altitude. Moreover, we studied whether extent and diversity of leaf damage are related to each other, to plant species richness, and to standing crop. We recorded the leaf area damaged by ten types of herbivory and five types of fungal pathogen infection on 12,054 plant leaves of legumes, other forbs, and graminoids collected in 215 grassland parcels in 12 valleys in the Swiss Alps. With 83 % of all leaves infested, herbivory and fungal pathogen infection were omnipresent. However, only 2.7 % leaf area was damaged by herbivory and 1.2 % by fungal pathogens. Damage by herbivory was highest on legumes, and damage by fungal pathogens was highest on graminoids. More leaf damage by herbivory occurred in traditionally mown sites and at lower altitudes, while damage by fungal pathogen infection was independent of land use and altitude. Most types of herbivory were found on legumes and on leaves from fertilised sites, whereas the number of fungal pathogen types was highest on graminoids and in unfertilised sites. Larger standing crop was associated with higher leaf damage and diversity of herbivory types per leaf. Neither damage by herbivory nor by fungal pathogens was correlated with plant species diversity. In more plant species rich parcels, the number of herbivory types was lower at the leaf level, but tended to be higher at the parcel level. Our results highlight the omnipresence of plant–herbivore and plant–pathogen interactions. They suggest that current land use changes from mowing to grazing or to abandonment decrease the diversity of herbivory, and that fertilisation decreases pathogen diversity. As our results did not reveal conservation conflicts between diversities of plants, herbivores, and fungal pathogens, and as the damage from herbivory and pathogens is generally low, we conclude that for protecting the high diversity of plant–herbivore and plant–pathogen interactions a diverse low-intensity land use should be maintained.

Aboveground–belowground herbivore interactions: a meta‐analysis

Research investigating interactions between aboveground (AG) and below-ground (BG) herbivores has been central to characterizing AG-BG linkages in terrestrial ecosystems, with many of these interactions forming the basis of complex food webs spanning the two subsystems. Despite the growing literature on the effects of AG and BG herbivores on each other, underlying patterns have been difficult to identify due to a high degree of context dependency. In this study, we present the first quantitative meta-analysis of AG and BG herbivore interactions. Previous global predictions, specifically that BG herbivores normally promoted AG herbivore performance and AG herbivores normally reduced BG herbivore performance, were not supported. Instead, the meta-analysis identified four factors that determined the outcome of AG-BG interactions. (1) Sequence of herbivore arrival on host plants was important, with BG herbivores promoting AG herbivore performance only when introduced to the plant simultaneously, whereas AG herbivores had negative effects on BG herbivores only when introduced first. (2) AG herbivores negatively affected BG herbivore survival but tended to increase population growth rates. (3) AG herbivores negatively affected BG herbivore performance on annual plants, but not on perennials, and these effects were observed more consistently in laboratory than field studies. (4) The type of herbivore was also important, with BG insect herbivores belonging to the order Diptera (i.e., true flies) having the strongest negative effects on AG herbivores. Coleoptera (i.e., beetles) species were the most widely investigated BG herbivores and had positive impacts on AG Homoptera (e.g., aphids), but negative effects on AG Hymenoptera (e.g., sawflies). The strongest negative outcomes for BG herbivores were seen when the AG herbivore was a Coleoptera species. We found no evidence for publication bias in AG-BG herbivore interaction literature and conclude that several biological and experimental factors are important for predicting the outcome of AG-BG herbivore interactions. The sequence of herbivore arrival on the host plant was among the most influential.

Induction of extrafloral nectar depends on herbivore type in invasive and native Chinese tallow seedlings

Although induced defenses are widespread in nature, and a potentially important strategy used by invasive plants, it is unclear how induced defenses vary among populations and whether the intensity and duration of induced defenses depends on herbivore type. For invasive plants, low herbivore loads in their introduced ranges can lead to differences in herbivore defense compared to their native ranges, but we currently know little about how induced defenses vary among native and invasive populations. We conducted a greenhouse experiment to examine variation in one type of induced defense, extrafloral nectar (EFN) production, among native and invasive populations of Chinese tallow tree, Triadica sebifera. We experimentally manipulated herbivory from an exotic generalist scale insect, a native generalist caterpillar, both herbivores, or neither and then examined EFN production by Triadica. Damage from leaf-chewing caterpillars resulted in strongly induced EFN in both native and invasive populations while damage from phloem-feeding scales did not. Extrafloral nectar production and dissolved solute content peaked 4 days after caterpillar herbivory for both native and invasive populations. Number and proportion of leaves producing EFN, EFN volume and concentration of dissolved solutes were similar among native and invasive populations. These results suggest that selection for indirect defenses may be different than selection for other defenses in the introduced ranges of invasive plants, as constitutive and induced EFN production is retained in invasive populations.

In situ estimates of waterhyacinth leaf tissue nitrogen using a SPAD-502 chlorophyll meter

Previous investigations with Eichhornia crassipes and its biological control agents Neochetina bruchi and N. eichhorniae identified leaf tissue nitrogen (N) as a driver in their interactions. However, traditional methods for determining plant tissue N content are cumbersome, time-consuming, and destructive–and thus unsuited for rapid in situ evaluations. We therefore tested the utility of a hand-held chlorophyll meter as a means of producing in situ estimates of N in the leaves of this floating aquatic weed. The Minolta SPAD-502 chlorophyll meter provided excellent estimates (F=385.96, P<0.0001) of leaf tissue N levels. SPAD readings varied within leaves (F=78.66, P<0.0001), so average readings per leaf were used. The relationship between SPAD readings and tissue N levels was affected by the phenological stage of the leaf (F=102.79, P<0.0001), but not leaf size (F=0.75, P=0.3867). The estimates were also unaffected by fertilizer level (F=0.95, P=0.4354), but were marginally affected by herbivory (F=3.86, P=0.0505). Thus, with suitable calibration (e.g., different field sites, presence and type of herbivory) output from the SPAD-502 could be used to provide consistent estimates for the nitrogen content of E. crassipes leaves.

Herbivore species and grazing intensity regulate community composition and an encroaching woody plant in semi-arid rangeland

Grazing by livestock can influence ecosystems in various ways, including altering plant communities, influencing woody plant encroachment, and determining livestock productivity. Evaluating long term effects of grazing on plant composition is valuable not only to understand herbivory on rangelands but to be able to address the primary factors that can threaten long term livestock productivity. We examined plant species composition and woody plant encroachment 45 years after the initiation of differing grazing treatments within a semiarid savanna of the southern Great Plains, USA. Grazing treatments varied in herbivore type (domestic cattle, sheep, and goats vs. goats only) and grazing intensity (heavy, moderate, and no-herbivory). All individual trees of Juniperus ashei Buchholz, the encroaching woody plant of the area, were removed prior to treatment initiation. Moderate and heavy grazing by a combination of species resulted in similar plant communities, while a history of heavy browsing by goats only and no-herbivory resulted in more distinct communities. Cover of J. ashei did not differ between mixed grazing and no-herbivory treatments, indicating that grazing was not responsible for woody plant encroachment. J. ashei cover within the browsed treatment was a third less compared to other treatments; compositional differences within this treatment are possibly due to reduced cover of woody vegetation. Declines in livestock productivity of the area are likely related to compositional changes resulting from increased woody plants. Livestock production within this semi-arid rangeland is likely unsustainable without management of woody plant encroachment, as communities tend to a closed canopy woodland.

Insect herbivory along environmental gradients

There is a general assumption in the literature that insect herbivory increases towards the tropics, but decreases with increasing altitude. Similar generalities have been identified along other environmental gradients, such as resource, temperature, climatic and biotic gradients. However there is growing evidence in the scientific literature that such generalities are not consistent. This could be due to a number of reasons including the lack of consistency in the way herbivory is assessed such as different methodologies used by researchers, or fundamental differences in leaf damage caused by different types of insect herbivores. Here we assess 61 publications researching insect herbivory along a range of environmental gradients (both biotic and abiotic) and review the methods that researchers have used to collected their data. We found leaf chewing from samples collected in North America dominated the field and most studies assessed herbivory on a single host plant species. Thirty three percent of the studies assessed latitudinal gradients, while 10% assessed altitudinal gradients. Insect herbivory was most commonly expressed as percentage leaf damage using point herbivory. Fewer studies measured a range of different types of herbivory (such as sap sucking, leaf mining, galling, and root feeding) as leaves aged. From our synthesis, we hope that future research into insect herbivory along environmental gradients will take into account herbivory other than just leaf chewing, such as sap sucking, which may cause more damage to plants. Future research should also assess herbivory as a rate, rather than just a single point in time as damage to a young leaf may be more costly to a plant than damage to a mature or senescing leaf. Measurements of plant traits will also assist in comparing herbivory across habitats, plant species, and within species physiological variation. The true impacts that insects have on plants via herbivory along environmental gradients are still poorly understood.

Herbivore-Induced Changes in Tomato (Solanum lycopersicum) Primary Metabolism: A Whole Plant Perspective

Induced changes in primary metabolism are important plant responses to herbivory, providing energy and metabolic precursors for defense compounds. Metabolic shifts also can lead to reallocation of leaf resources to storage tissues, thus increasing a plant's tolerance. We characterized whole-plant metabolic responses of tomato (Solanum lycopersicum) 24h after leaf herbivory by two caterpillars (the generalist Helicoverpa zea and the specialist Manduca sexta) by using GC-MS. We measured 56 primary metabolites across the leaves, stems, roots, and apex, comparing herbivore-attacked plants to undamaged plants and mechanically damaged plants. Induced metabolic change, in terms of magnitude and number of individual concentration changes, was stronger in the apex and root tissues than in undamaged leaflets of damaged leaves, indicating rapid and significant whole-plant responses to damage. Helicoverpa zea altered many more metabolites than M. sexta across most tissues, suggesting an enhanced plant response to H. zea herbivory. Helicoverpa zea herbivory strongly affected concentrations of defense-related metabolites (simple phenolics and precursor amino acids), while M. sexta altered metabolites associated with carbon and nitrogen transport. We conclude that herbivory induces many systemic primary metabolic changes in tomato, and that changes often are specific to a single tissue or type of herbivore. The potential implications of primary metabolic changes are discussed in relation to resistance and tolerance.

Development of the Pasture Simulation Model for assessing livestock production under climate change

To simulate climate change impacts on pastures and domestic herbivores as well as feedbacks to the atmosphere in terms of greenhouse gas emissions, we have improved a process-based biogeochemical pasture model, PaSim. The overall aim was to simulate the meat and milk production of cattle (suckler cows with their calves, dairy cows and heifers) in response to climate and management, as well as feedbacks to the atmosphere through enteric methane emissions. Herbage intake at grazing was calculated from animal characteristics, herbage availability, diet digestibility and air temperature. With suckler cows, milk production and changes in daily liveweight and body condition were calculated from net energy balance. The net energy intake of dairy cows and their body reserves at turnout to pasture were used to simulate milk production at pasture, daily liveweight and body condition changes, taking into account cow energy requirements and intake capacity. Heifer growth was determined from heifer net energy intake and liveweight. Net energy intake was used to assess enteric methane production through a conversion factor, which depends both on the energy level of the diet and on the herbivore type. The model was assessed against experimental data for animal performance and methane emissions at grazing. Predictions show good agreement with observations. On average, the root mean square error was 6.5, 4 and 2.5% for the liveweights of suckler cows, suckler calves and heifers, respectively, 18% for dairy milk production and 12% for enteric methane emissions. By comparing this new version of the PaSim model to the previous version, we show that a greater accuracy in animal performance modelling improves the sensitivity of the model to interannual climate variability. However, long term (30 years) projections of climate change impacts on grasslands and of radiative feedbacks to the atmosphere are not significantly modified. The originality and the validity domain of the model are discussed.

Induced responses in the neotropical shrub Bauhinia brevipes Vogel: does early season herbivory function as cue to plant resistance?

Models of plant phenotypic plasticity suggest that past or current environment provide information about future environments, and plants respond to past or current herbivory by increasing their resistance to herbivores. In this study, we determined whether damage to Bauhinia brevipes leaves caused by free-feeding and galling herbivores early in the season predicts the frequency of attack throughout the season, examining the occurrence of induced responses and resistance during 2 years in 50 individual plants. We found that damage early in the season was a reasonable predictor of the number of galls and scars that would be initiated throughout the season. Galling attack rates were significantly higher in 1997 than in 1998, but the opposite pattern was observed for attack rates by free-feeding herbivores between years. Also, leaf nutritional quality differed significantly among three sampling dates, and a trend for an increase in tannin concentration was observed from the beginning to the end of the season. A negative correlation was found between the number of scars caused by chewing herbivores at the end of the season and tannin concentration, but there was no relationship between tannin concentration and percentage of galled leaves or number of Contarinia sp. galls. Furthermore, likelihood of gall survivorship was inversely related to the number of galls recorded early in the season in both years—an indicator of induced plant resistance. This study suggests that early season damage was a reasonable predictor of damage caused by two types of herbivores throughout the season, and that in systems where herbivory is fairly unpredictable within a short temporal scale; the expression of induced responses and resistance is favored.

Differential responses of vertebrate and invertebrate herbivores to traits of New Zealand subalpine shrubs

Plant traits are influenced by herbivore diet selection, but little is known about how traits are affected by different types of herbivores. We related eight traits of 27 subalpine shrub species in South Island, New Zealand, to damage of these shrubs by introduced red deer (Cervus elaphus) and native invertebrate herbivores using phylogenetically explicit modeling. Deer preferentially consumed species that grew quickly, were low in foliar tannins, or had high leaf area per unit mass. However, these traits did not trade off against each other; rather, they could be related to different multivariate defense strategies. Although the proportion of leaves damaged by leaf-chewing invertebrates also increased with stem growth, invertebrates did not damage the same fast growing species as those preferred by deer. Other traits may also be important in determining herbivore preferences, as suggested by the high proportion of variation in herbivory explained by phylogeny. Last, we found that the composition of invertebrate herbivore communities was more similar among closely related shrubs, and consequently, the range of invertebrate-plant associations may change if introduced deer shift plant composition toward slow-growing species. Overall, our results demonstrate the importance of herbivore type and coevolved interactions for the adaptive significance of plant traits.

Herbivore-induced resource sequestration in plants: why bother?

Herbivores can cause numerous changes in primary plant metabolism. Recent studies using radioisotopes, for example, have found that insect herbivores and related cues can induce faster export from leaves and roots and greater partitioning into tissues inaccessible to foraging herbivores. This process, termed induced resource sequestration, is being proposed as an important response of plants to cope with herbivory. Here, we review the evidence for resource sequestration and suggest that associated allocation and ecological costs may limit the benefit of this response because resources allocated to storage are not immediately available to other plant functions or may be consumed by other enemies. We then present a conceptual model that describes the conditions under which benefits might outweigh costs of induced resource sequestration. Benefits and costs are discussed in the context of differences in plant life-history traits and biotic and abiotic conditions, and new testable hypotheses are presented to guide future research. We predict that intrinsic factors related to life history, ontogeny and phenology will alter patterns of induced sequestration. We also predict that induced sequestration will depend on certain external factors: abiotic conditions, types of herbivores, and trophic interactions. We hope the concepts presented here will stimulate more focused research on the ecological and evolutionary costs and benefits of herbivore-induced resource sequestration.

Increased reproductive capacity and physical defense but decreased tannin content in an invasive plant

Abstract Plant invasions create novel plant–insect interactions. The EICA (evolution of increased competitive ability) hypothesis proposes that invasive plants will reallocate resources from defense to growth and/or reproduction because they have escaped from their co‐evolved insect natural enemies. Testing multiple herbivory by monophagous and oligophagous herbivores and simultaneous measurement of various plant traits will provide new insights into the evolutionary change of invasive plants. In this context, we conducted a common garden experiment to compare plant growth and reproduction, chemical and physical defense, and plant responses to herbivory by different types of herbivores between invasive North American populations and native East Asian populations of mile‐a‐minute weed, Persicaria perfoliata. We found that invasive mile‐a‐minute exhibited lower biomass, flowered earlier and had greater reproductive output than plants from the native range. Compared with native populations, plants from invasive populations had lower tannin content, but exhibited higher prickle density on nodes and leaves. Thus our results partially support the EICA hypothesis. When exposed to the monophagous insect, Rhinoncomimus latipes and the oligophagous insects, Gallerucida grisescens and Smaragdina nigrifrons, more damage by herbivory was found on invasive plants than on natives. R. latipes, G. grisescens and S. nigrifrons had strong, moderate and weak impacts on the growth and reproduction of mile‐a‐minute, respectively. The results indicate that mile‐a‐minute may have evolved a higher reproductive capacity in the introduced range, and this along with a lack of oligophagous and monophagous herbivores in the new range may have contributed to its invasiveness in North America.

Determinants of spatial variation in fire return period in a semiarid African savanna

Fire pattern is predicted to vary across an African savanna in accordance with spatial variation in rainfall through its effects on fuel production, vegetation type (on account of differences in fuel load and in flammability), and distribution of herbivores (because of their effects on fuel load). These predictions were examined for the 23 651-ha Mkuzi Game Reserve, KwaZulu-Natal, based on a 37-year data set. Fire return period varied from no occurrence to a fire every 1.76 years. Approximately 75% of the reserve experienced a fire approximately every 5 years, 25% every 4.1–2.2 years and less than 1% every 2 years on average. Fire return period decreased in relation to an increase in mean annual rainfall. For terrestrial vegetation types, median fire return periods decreased with increasing herbaceous biomass, from forest that did not burn to grasslands that burnt every 2.64 years. Fire was absent from some permanent wetlands but seasonal wetlands burnt every 5.29 years. Grazer biomass above 0.5 animal units ha–1 had a limiting influence on the maximum fire frequency of fire-prone vegetation types. The primary determinant of long-term spatial fire patterns is thus fuel load as determined by mean rainfall, vegetation type, and the effects of grazing herbivores.

Ecophysiological Responses of Giant Reed (Arundo donax) to Herbivory

AbstractThe effect of invasive species might be lessened if herbivores reduced transpiration and growth rates; however, simply removing photosynthetic material might not ensure that the transpiration rate of active leaf tissue decreases. We assessed whether biological control has an injurious effect on the target plant species, giant reed (Arundo donax), by quantifying leaf photosynthetic and transpiration responses to two herbivores: an armored scale, Rhizaspidiotus donacis, and a stem-galling wasp, Tetramesa romana. Herbivory by a sap-feeding scale and a stem-galling wasp both separately and together, reduces the rates of leaf level physiological processes in A. donax. The effect of the wasp increases with density and reduces photosynthesis by reducing the carboxylation rate of ribulose-1,5-bisphosphate carboxylase oxygenase, which controls CO2 fixation in photosynthesis. The scale insect reduces photosynthesis by decreasing the maximum rate of electron transport, which determines how much light energy can be captured in photosynthesis. The effect of the armored scale takes approximately 5 mo after infestation, which coincides with generation time. When both insects are present at the same time, the effect of their herbivory appears additive after time for the scale to reproduce. We conclude that a combination of two herbivores can have a stronger physiological effect than one type of herbivore, likely because of their different effects on leaf function.

A tritrophic signal that attracts parasitoids to host-damaged plants withstands disruption by non-host herbivores

BackgroundVolatiles emitted by herbivore-infested plants are highly attractive to parasitoids and therefore have been proposed to be part of an indirect plant defense strategy. However, this proposed function of the plant-provided signals remains controversial, and it is unclear how specific and reliable the signals are under natural conditions with simultaneous feeding by multiple herbivores. Phloem feeders in particular are assumed to interfere with plant defense responses. Therefore, we investigated how attack by the piercing-sucking cicadellid Euscelidius variegatus influences signaling by maize plants in response to the chewing herbivore Spodoptera littoralis.ResultsThe parasitoid Cotesia marginiventris strongly preferred volatiles of plants infested with its host S. littoralis. Overall, the volatile emissions induced by S. littoralis and E. variegatus were similar, but higher levels of certain wound-released compounds may have allowed the wasps to specifically recognize plants infested by hosts. Expression levels of defense marker genes and further behavioral bioassays with the parasitoid showed that neither the physiological defense responses nor the attractiveness of S. littoralis infested plants were altered by simultaneous E. variegatus attack.ConclusionsOur findings imply that plant defense responses to herbivory can be more robust than generally assumed and that ensuing volatiles convey specific information about the type of herbivore that is attacking a plant, even in complex situations with multiple herbivores. Hence, the results of this study support the notion that herbivore-induced plant volatiles may be part of a plant's indirect defense stratagem.

Resource allocation to defence and growth are driven by different responses to generalist and specialist herbivory in an invasive plant

Summary1. Invasive plants often have novel biotic interactions in their introduced ranges. These interactions, including less frequent herbivore attacks, may convey a competitive advantage over native plants. Invasive plants may vary in defence strategies (resistance vs. tolerance) or in response to the type of herbivore (generalists vs. specialists), but no study to date has examined this broad set of traits simultaneously.2. Here, we examined resistance and tolerance of Chinese tallow (Triadica sebifera) populations from the introduced and native ranges to generalist (Cnidocampa flavescens) and specialist herbivores (Gadirtha inexacta) in the native range.3. In a field common‐garden test of resistance, caterpillars of each species were raised on plants from native and invasive populations. We found the specialist grew larger on and consumed more mass of invasive plant populations than native populations, while the generalist showed the same performance between them. The results were consistent with our laboratory bioassay using excised leaves. Chemical analyses showed that the invasive plants had lower tannin content and higher ratio of carbohydrate to protein than those of their native counterparts, suggesting that plants from invasive populations have altered chemistry that has a larger impact on specialist than on generalist resistance.4. To test for differences in herbivore tolerance, plants were first defoliated by specialist or generalist herbivory and then allowed to regrow for 100 days in a field common garden. We found that plants from invasive populations had greater herbivore tolerance than native populations, especially for tolerance to generalists. They also grew more rapidly than native counterparts in the absence of herbivory.5. Synthesis.The results of these experiments indicate that differences in selective pressures between ranges have caused dramatic reductions in resistance to specialist herbivores and those changes in plant secondary chemistry likely underlie these differences. The greater tolerance of invasive populations to herbivory appears to at least partly reflect an increase in growth rate in the introduced range. The greater tolerance to generalist herbivores suggests the intriguing possibility of selection for traits that allow plants to tolerate generalist herbivores more than specialist herbivores.

Variation and fitness costs for tolerance to different types of herbivore damage in Boechera stricta genotypes with contrasting glucosinolate structures

• Analyses of plant tolerance in response to different modes of herbivory are essential to an understanding of plant defense evolution, yet are still scarce. Allocation costs and trade-offs between tolerance and plant chemical defenses may influence genetic variation for tolerance. However, variation in defenses also occurs for the presence or absence of discrete chemical structures; yet, the effects of intraspecific polymorphisms on tolerance to multiple herbivores have not been evaluated. • Here, in a glasshouse experiment, we investigated the variation for tolerance to different types of herbivore damage, and direct allocation costs, in 10 genotypes of Boechera stricta (Brassicaceae), a wild relative of Arabidopsis, with contrasting foliar glucosinolate chemical structures (methionine-derived glucosinolates vs glucosinolates derived from branched-chain amino acids). • We found significant genetic variation for tolerance to different types of herbivore. Structural variations in the glucosinolate profile did not influence tolerance to damage, but predicted plant fitness. Levels of constitutive and induced glucosinolates varied between genotypes with different structural profiles, but we did not detect any cost of tolerance explaining the genetic variation in tolerance among genotypes. • Trade-offs between plant tolerance to multiple herbivores may not explain the existence of intermediate levels of tolerance to damage in plants with contrasting chemical defensive profiles.

The Ontogeny of Plant Defense and Herbivory: Characterizing General Patterns Using Meta‐Analysis

Defense against herbivores often changes dramatically as plants develop. Hypotheses based on allocation theory and herbivore selection patterns predict that defense should increase or decrease, respectively, across ontogeny, and previous research partly supports both predictions. Thus, it remains unclear which pattern is more common and what factors contribute to variability among studies. We conducted a meta-analysis of 116 published studies reporting ontogenetic patterns in plant defense traits and herbivory. Patterns varied depending on plant life form (woody, herbaceous, grass), type of herbivore (insect, mollusk, mammal), and type of defense trait (secondary chemistry, physical defense, tolerance). In woody plants, chemical defense increased during the seedling stage, followed by an increase in physical defenses during the vegetative juvenile stage. Mammalian herbivores showed a strong preference for mature compared to juvenile tissues in woody plants. Herbs experienced a significant increase in secondary chemistry across the entire ontogenetic trajectory, although the magnitude of increase was greatest during the seedling stage. Correspondingly, mollusks preferred young compared to older herbs. Future research investigating growth/defense trade-offs, allometry, herbivore selection patterns, and ecological costs would shed light on the mechanisms driving the ontogenetic patterns observed.