Influence of surrounding vegetation on insect herbivory: A matter of spatial scale and herbivore specialisation

The diversity of surrounding vegetation is thought to modify the interactions between a focal plant and its herbivores, disrupting (associational resistance) or enhancing (associational susceptibility) host plant location and colonisation. We compared the effects of host plant concentration on herbivory by generalist and specialist insects feeding on oak seedlings by increasing local concentration of seedlings. We also assessed the effects of the composition and structure of surrounding vegetation, both at stand and local levels. The damage caused by generalist leaf-feeding insects depended on the structure of plant communities at stand level, and increased with tree cover. By contrast, infestation by specialist leaf miners was affected by local understorey vegetation surrounding oak seedlings, and decreased with increasing shrub cover and stratification diversity. Leaf mine abundance was higher at higher oak seedling density, giving support to the host concentration hypothesis. However, the abundance of these specialist herbivores was also negatively correlated with damage caused by the generalist external leaf-feeders, suggesting competitive interactions.     

Temporal dynamics in non-additive responses of arthropods to host-plant genotypic diversity

Genotypic diversity within host‐plant populations has been linked to the diversity of associated arthropod communities, but the temporal dynamics of this relationship, along with the underlying mechanisms, are not well understood. In this study, we employed a common garden experiment that manipulated the number of genotypes within patches of Solidago altissima, tall goldenrod, to contain 1, 3, 6 or 12 genotypes m^?2 and measured both host‐plant and arthropod responses to genotypic diversity throughout an entire growing season. Despite substantial phenological changes in host plants and in the composition of the arthropod community, we detected consistent positive responses of arthropod diversity to host‐plant genotypic diversity throughout all but the end of the growing season. Arthropod richness and abundance increased with genotypic diversity by up to～65%. Furthermore, arthropod responses were non‐additive for most of the growing season, with up to 52% more species occurring in mixtures than the number predicted by summing the number of arthropods associated with component genotypes in monoculture. Non‐additive arthropod responses were likely driven by concurrent non‐additive increases in host‐plant aboveground biomass. Qualitative differences among host‐plant genotypes were also important early in the season, when specialist herbivores dominated the arthropod community. Neither arthropod diversity nor flower number was associated with genotypic diversity at the end of the growing season, when generalist floral‐associated herbivores dominated. Taken together, these results show that focusing on the temporal dynamics in the quantity and quality of co‐occurring host‐plant genotypes and associated community composition can help uncover the mechanisms that link intraspecific host‐plant diversity to the structure of arthropod communities. Furthermore, consistent non‐additive effects in genotypically diverse plots may limit the predictability of the arthropod community based solely on the genetic make‐up of a host‐plant patch.     

Additive and non‐additive effects of birch genotypic diversity on arthropod herbivory in a long‐term field experiment

Herbivores are important drivers of plant population dynamics and community composition in natural and managed systems. Intraspecific genetic diversity of long‐lived plants like trees might shape patterns of herbivory by different guilds of herbivores that trees experience through time. However, previous studies on plant genetic diversity effects on herbivores have been largely short‐term. We investigated how tree genotypic variation and diversity influence herbivory of silver birch Betula pendula in a long‐term field experiment. Using clones of eight genotypes, we constructed experimental plots consisting of one, two, four or eight genotypes, and measured damage by five guilds of arthropod herbivores twice a year over three different years (four, six and nine years after the experiment was established). Genotypes varied significantly for most types of herbivore damage, but genotype resistance rankings often shifted over time, and none of the clones was more resistant than all others to all types of herbivores. At the plot level, birch genotypic diversity had significant positive additive effect on leaf rollers and negative non‐additive effects on chewing herbivores and gall makers. In contrast, leaf‐mining and leaf‐tying damage was not influenced by birch genotypic diversity. Within diverse plots, the direction of genotypic diversity effects varied depending on birch genotype, some having lower and some having higher herbivory in mixed stands. This research highlights the importance of long‐term studies including different feeding guilds of herbivores to understand the effects of plant genetic diversity on arthropod communities. Different responses of various feeding guilds to genotypic diversity and shifts in resistance of individual genotypes over time indicate that genotypic mixtures are unlikely to result in overall reduction in herbivory over time.     

Native plant diversity increases herbivory to non-natives

There is often an inverse relationship between the diversity of a plant community and the invasibility of that community by non-native plants. Native herbivores that colonize novel plants may contribute to diversity–invasibility relationships by limiting the relative success of non-native plants. Here, we show that, in large collections of non-native oak trees at sites across the USA, non-native oaks introduced to regions with greater oak species richness accumulated greater leaf damage than in regions with low oak richness. Underlying this trend was the ability of herbivores to exploit non-native plants that were close relatives to their native host. In diverse oak communities, non-native trees were on average more closely related to native trees and received greater leaf damage than those in depauperate oak communities. Because insect herbivores colonize non-native plants that are similar to their native hosts, in communities with greater native plant diversity, non-natives experience greater herbivory.     

Conditional outcomes in plant–herbivore interactions: neighbours matter

Spatial distribution of palatable and unpalatable plants can influence the foraging behaviour of herbivores, thereby changing plant‐damage probabilities. Moreover, the immediate proximity to certain plants can benefit other plants that grow below them, where toxicity or spines act as a physical barrier or concealment against herbivores. This paper presents the results of a multi‐scale experiment performed to test the effect of shrubs as protectors of tree saplings against herbivores and the mechanism involved in Mediterranean ecosystems. We performed a factorial design in two mountain ranges, similar in physiognomy and vegetation, planting saplings of a palatable tree, the maple (Acer opalus subsp. granatense), and an unpalatable tree, the black pine (Pinus nigra), under three different types of shrubs. We considered four experimental microhabitats: highly palatable shrub (Amelanchier ovalis), palatable but spiny shrub (Crataegus monogyna or Prunus ramburii), unpalatable spiny shrub (Berberis vulgaris subsp. australis) and control (gaps of bare soil without shrubs). Three main factors were found to determine the probability of sapling attack: sapling palatability, experimental microhabitat and plot. Palatable saplings (maples) were browsed much more than unpalatable ones (pines). The degree of protection provided by the shrub proved greater as its palatability decreased with respect to sapling palatability, the unpalatable spiny shrub being the safest microhabitat for palatable saplings and bare soil for unpalatable ones. The differences found in number of attacked saplings between plots may be attributable to differences in herbivore pressure. The community context in which interaction takes place, namely the characteristics of the neighbours and the intensity of herbivore pressure, are determining factors for understanding and predicting the damage undergone by a target plant species. The mechanism that best explains these results is associational avoidance of saplings that grow near to unpalatable shrubs. It is necessary to introduce this neighbour effect in theoretical models and food‐web approaches that analyse the plant–herbivore relationships, since it can strongly determine not only the intensity of the interaction, but also the spatial distribution and diversity of the plant community.     

Partitioning of herbivore hosts across time and food plants promotes diversification in the Megastigmus dorsalis oak gall parasitoid complex

Communities of insect herbivores and their natural enemies are rich and ecologically crucial components of terrestrial biodiversity. Understanding the processes that promote their origin and maintenance is thus of considerable interest. One major proposed mechanism is ecological speciation through host‐associated differentiation (HAD), the divergence of a polyphagous species first into ecological host races and eventually into more specialized daughter species. The rich chalcid parasitoid communities attacking cynipid oak gall wasp hosts are structured by multiple host traits, including food plant taxon, host gall phenology, and gall structure. Here, we ask whether the same traits structure genetic diversity within supposedly generalist parasitoid morphospecies. We use mitochondrial DNA sequences and microsatellite genotypes to quantify HAD for Megastigmus (Bootanomyia) dorsalis, a complex of two apparently generalist cryptic parasitoid species attacking oak galls. Ancient Balkan refugial populations showed phenological separation between the cryptic species, one primarily attacking spring galls, and the other mainly attacking autumn galls. The spring species also contained host races specializing on galls developing on different host‐plant lineages (sections Cerris vs. Quercus) within the oak genus Quercus. These results indicate more significant host‐associated structuring within oak gall parasitoid communities than previously thought and support ecological theory predicting the evolution of specialist lineages within generalist parasitoids. In contrast, UK populations of the autumn cryptic species associated with both native and recently invading oak gall wasps showed no evidence of population differentiation, implying rapid recruitment of native parasitoid populations onto invading hosts, and hence potential for natural biological control. This is of significance given recent rapid range expansion of the economically damaging chestnut gall wasp, Dryocosmus kuriphilus, in Europe.     

Comparison of tree genotypic diversity and species diversity effects on different guilds of insect herbivores

Although the effects of plant diversity on herbivores are contingent upon herbivore traits and the source of plant diversity (e.g. intra‐ and interspecific), most studies have analyzed these effects separately. We compared the effects of genotypic diversity of big‐leaf mahogany Swietenia macrophylla with that of tree species diversity on two specialist caterpillars (Hypsipyla grandella stem borers and Phyllocnistis meliacella leaf miners) and three generalist leafhoppers (Cicadellidae) feeding on mahogany in a large‐scale (7.2 ha) forest diversity experiment in southern Mexico. The experiment consisted of fifty‐nine 21 × 21‐m plots, with 64 tree saplings each (3‐m spacing between plants). Plots were either mahogany monocultures or species polycultures of four species (including mahogany) and – within each of these two plot types – mahogany was represented by either one or four genotypes. Throughout a five‐month period, beginning six months after planting, we measured mahogany growth and monitored herbivore and predator (spider) abundance. We found no effect of mahogany genotypic diversity on either specialist caterpillars or generalist leafhoppers, and this result was consistent across levels of tree species diversity. In contrast, species diversity had significant effects on both specialists but neither of the generalist herbivores. Specifically, species diversity lowered H. grandella attack at the middle of the sampling season, but increased attack at the end of the season, whereas P. meliacella abundance was consistently reduced. Such effects were not mediated by effects of species diversity on plant growth (of which there were none), but rather through resource heterogeneity. Diversity did not influence spider abundance. This study is one of few to directly compare sources of plant diversity, and uniquely compares such effects among herbivores with contrasting life histories (e.g. diet breadths). Overall, we demonstrate that plant species diversity effects outweigh those of genotypes, and our results suggest that such effects are stronger on specialist than generalist herbivores.     

Chemotype of <Emphasis Type="Italic">Senecio jacobaea</Emphasis> affects damage by pathogens and insect herbivores in the field

The evolution of the diversity of plant secondary metabolites is still poorly understood. To determine whether natural enemies could exert selection on plant secondary chemistry, pathogen infestation and invertebrate herbivory were measured on 10 genotypes of Senecio jacobaea (Tansy Ragwort) at two experimental field sites during a 2-year period. The genotypes represented two chemotypes based on the presence of the pyrrolizidine alkaloids (PA) jacobine and erucifoline. At one site, Heteren, mainly generalist herbivores were present. Here, damage was limited and did not differ among genotypes or chemotypes. At the other site, Meijendel, several specialists attacked the plants. Damage increased during the year, with a peak in July when most damage was caused by the specialist moth Tyria jacobaeae. At this peak there was no difference in damage among chemotypes. In the months prior to T. jacobaeae damage, chemotypes with jacobine were more severely attacked by specialists than the chemotypes without jacobine. Total damage during that period was positively correlated with both total PA concentration and jacobine concentration. Probably plant vigor also played a role in host preference since damage per individual plant was positively correlated with plant size. Our results suggest that total PA concentration and specifically jacobine had a positive effect on specialist feeding, indicating ecological costs involved in the production of PAs. Ecological costs related to plant secondary compounds could explain why not all individuals produce high levels of these compounds. In addition, differences in specialist herbivore pressures among sites may contribute to the variation in secondary metabolites among populations.     

EXTREME HOST PLANT CONSERVATISM DURING AT LEAST 20 MILLION YEARS OF HOST PLANT PURSUIT BY OAK GALLWASPS

Diversification of insect herbivores is often associated with coevolution between plant toxins and insect countermeasures, resulting in a specificity that restricts host plant shifts. Gall inducers, however, bypass plant toxins and the factors influencing host plant associations in these specialized herbivores remain unclear. We reconstructed the evolution of host plant associations in Western Palaearctic oak gallwasps (Cynipidae: Cynipini), a species-rich lineage of specialist herbivores on oak ( Quercus ). (1) Bayesian analyses of sequence data for three genes revealed extreme host plant conservatism, with inferred shifts between major oak lineages (sections Cerris and Quercus ) closely matching the minimum required to explain observed diversity. It thus appears that the coevolutionary demands of gall induction constrain host plant shifts, both in cases of mutualism (e.g., fig wasps, yucca moths) and parasitism (oak gallwasps). (2) Shifts between oak sections occurred independently in sexual and asexual generations of the gallwasp lifecycle, implying that these can evolve independently. (3) Western Palaearctic gallwasps associated with sections Cerris and Quercus diverged at least 20 million years ago (mya), prior to the arrival of oaks in the Western Palaearctic from Asia 5–7 mya. This implies an Asian origin for Western Palaearctic gallwasps, with independent westwards range expansion by multiple lineages.     

Genotype × Herbivore Effect on Leaf Litter Decomposition in Betula Pendula Saplings: Ecological and Evolutionary Consequences and the Role of Secondary Metabolites

Plant genetic variation and herbivores can both influence ecosystem functioning by affecting the quantity and quality of leaf litter. Few studies have, however, investigated the effects of herbivore load on litter decomposition at plant genotype level. We reduced insect herbivory using an insecticide on one half of field-grown Betula Pendula saplings of 17 genotypes, representing random intrapopulation genetic variation, and allowed insects to naturally colonize the other half. We hypothesized that due to induced herbivore defence, saplings under natural herbivory produce litter of higher concentrations of secondary metabolites (terpenes and soluble phenolics) and have slower litter decomposition rate than saplings under reduced herbivory. We found that leaf damage was 89 and 53% lower in the insecticide treated saplings in the summer and autumn surveys, respectively, which led to 73% higher litter production. Litter decomposition rate was also affected by herbivore load, but the effect varied from positive to negative among genotypes and added up to an insignificant net effect at the population level. In contrast to our hypothesis, concentrations of terpenes and soluble phenolics were higher under reduced than natural herbivory. Those genotypes, whose leaves were most injured by herbivores, produced litter of lowest mass loss, but unlike we expected, the concentrations of terpenes and soluble phenolics were not linked to either leaf damage or litter decomposition. Our results show that (1) the genetic and herbivore effects on B. pendula litter decomposition are not mediated through variation in terpene or soluble phenolic concentrations and suggest that (2) the presumably higher insect herbivore pressure in the future warmer climate will not, at the ecological time scale, affect the mean decomposition rate in genetically diverse B. pendula populations. However, (3) due to the significant genetic variation in the response of decomposition to herbivory, evolutionary changes in mean decomposition rate are possible.     

Diversity and abundance patterns of phytophagous insect communities on alien and native host plants in the Brassicaceae

The herbivore load (abundance and species richness of herbivores) on alien plants is supposed to be one of the keys to understand the invasiveness of species. We investigate the phytophagous insect communities on cabbage plants (Brassicaceae) in Europe. We compare the communities of endophagous and ectophagous insects as well as of Coleoptera and Lepidoptera on native and alien cabbage plant species. Contrary to many other reports, we found no differences in the herbivore load between native and alien hosts. The majority of insect species attacked alien as well as native hosts. Across insect species, there was no difference in the patterns of host range on native and on alien hosts. Likewise the similarity of insect communities across pairs of host species was not different between natives and aliens. We conclude that the general similarity in the community patterns between native and alien cabbage plant species are due to the chemical characteristics of this plant family. All cabbage plants share glucosinolates. This may facilitate host switches from natives to aliens. Hence the presence of native congeners may influence invasiveness of alien plants.     

Intraspecific chemical diversity among neighbouring plants correlates positively with plant size and herbivore load but negatively with herbivore damage

Intraspecific plant diversity can modify the properties of associated arthropod communities and plant fitness. However, it is not well understood which plant traits determine these ecological effects. We explored the effect of intraspecific chemical diversity among neighbouring plants on the associated invertebrate community and plant traits. In a common garden experiment, intraspecific diversity among neighbouring plants was manipulated using three plant populations of wild cabbage that differ in foliar glucosinolates. Plants were larger, harboured more herbivores, but were less damaged when plant diversity was increased. Glucosinolate concentration differentially correlated with generalist and specialist herbivore abundance. Glucosinolate composition correlated with plant damage, while in polycultures, variation in glucosinolate concentrations among neighbouring plants correlated positively with herbivore diversity and negatively with plant damage levels. The results suggest that intraspecific variation in secondary chemistry among neighbouring plants is important in determining the structure of the associated insect community and positively affects plant performance.     

北京山区辽东栎林的径级结构、种群起源及生态学意义

 北京山区的辽东栎(Quercus wutaishanica)林径级结构的调查分析结果表明,辽东栎林是稳定性的群落类型,其更新主要通过萌生来完成。能够与辽东栎长期共存的伴生种有色木(Acer mono)、大果榆(Ulmus macrocarpa)、大叶白蜡(Fraxinus rhynchophylla)等。萌生在辽东栎种群中十分普遍,除辽东栎伐桩、枯木和大树根际有较多的萌生茎干外,多年生实生苗也有萌生现象发生。辽东栎林的更新在辽东栎总体种群的幼树阶段(径级2)存在着明显的瓶颈现象,但萌生特性使辽东栎林在更新时或受到干扰后形成较大的萌生种群,使辽东栎的大幼树能够得到迅速的补充,顺利地通过瓶颈进行更新,保持群落的相对稳定。一定数量的辽东栎实生种群的存在对于维持辽东栎的遗传多样性、种群整体活力和群落的稳定,提高辽东栎种群对环境变化的反应能力等方面有着十分重要的生态学意义。     

The genetics of chutes and ladders: a community genetics approach to tritrophic interactions

Community genetics research has firmly established that intraspecific genetic variation in single populations can have large extended ecological consequences for populations and entire communities of organisms. Here, we sought to understand the bottom‐up effects of plant genetic variation on herbivore preference and performance, and the top–down control of predators on herbivores and their joint effects on plant fitness and evolution. Following three ecological genetics field experiments we detected heritable variation in plant traits that influenced both the preference and performance of a specialist weevil on Oenothera biennis. However, the weevil's preference and performance were not genetically correlated among O. biennis plant genotypes. Although predators and parasitoids were abundant, predators had no detectable effect on weevil performance because high egg and larval mortality was caused by non‐predatory factors such as intraspecific competition. Finally, neither the specialist weevil nor predators influenced plant fitness. Our results suggest that the focal tritrophic community studied here is primarily shaped by the bottom–up effects of plant genetic variation on herbivores, while top–down effects have no clear impacts on O. biennis fitness or evolution. We suggest that future studies should incorporate plant intraspecific genetic variation as a fundamental part of tritrophic interactions including their eco‐evolutionary dynamics.     

Tests for Parasite-mediated Frequency-dependent Selection in Natural Populations of an Asexual Plant Species

Genetic variation in plant populations for resistance to pathogens and herbivores might be maintained by parasite-mediated negative frequency-dependent selection (FDS). But it is difficult to observe the time-lagged oscillations between host and parasite genotypes that should result from FDS. To evaluate the potential for FDS, we tested for local adaptation of parasites to common clones, the role of host genetic diversity in resistance to parasites, and genetic correlations among fitness, parasitism, and the frequency of host clones. We studied three populations of Arabis holboellii, a short-lived apomictic (asexual by seed) plant attacked by rust fungi and insect herbivores. To estimate clone frequency, we used polymorphic allozyme markers on 200 individuals in each population in 1990 and in 2000. We also recorded levels of parasitism and host fitness (fruit production). Only the rust fungi showed evidence for local host adaptation; they usually increased in incidence as a function of clone frequency, and they tracked temporal change in clone frequency. In further support of FDS, parasitism was lower in populations with higher genetic diversity. However, total parasitism (herbivory and disease combined) decreased as host clone frequency and fitness increased. Thus, although the highly virulent rust pathogen showed potential for driving the cycles that result from FDS, this apparently does not occur in the populations studied because the host clones were also attacked by herbivores.Co-ordinating editor: J.F. Stuefer     

Individual tree traits shape insect and disease damage on oak in a climate‐matching tree diversity experiment

Diversifying planted forests by increasing genetic and species diversity is often promoted as a method to improve forest resilience to climate change and reduce pest and pathogen damage. In this study, we used a young tree diversity experiment replicated at two sites in the UK to study the impacts of tree diversity and tree provenance (geographic origin) on the oak (Quercus robur) insect herbivore community and a specialist biotrophic pathogen, oak powdery mildew. Local UK, French, and Italian provenances were planted in monocultures, provenance mixtures, and species mixes, allowing us to test whether: (a) local and nonlocal provenances differ in their insect herbivore and pathogen communities, and (b) admixing trees leads to associational effects on insect herbivore and pathogen damage. Tree diversity had variable impacts on foliar organisms across sites and years, suggesting that diversity effects can be highly dependent on environmental context. Provenance identity impacted upon both herbivores and powdery mildew, but we did not find consistent support for the local adaptation hypothesis for any group of organisms studied. Independent of provenance, we found tree vigor traits (shoot length, tree height) and tree apparency (the height of focal trees relative to their surroundings) were consistent positive predictors of powdery mildew and insect herbivory. Synthesis. Our results have implications for understanding the complex interplay between tree identity and diversity in determining pest damage, and show that tree traits, partially influenced by tree genotype, can be important drivers of tree pest and pathogen loads.     

Tree species diversity influences herbivore abundance and damage: meta-analysis of long-term forest experiments

Plant monocultures are commonly believed to be more susceptible to herbivore attacks than stands composed of several plant species. However, few studies have experimentally tested the effects of tree species diversity on herbivory. In this paper, we present a meta-analysis of uniformly collected data on insect herbivore abundance and damage on three tree species (silver birch, black alder and sessile oak) from seven long-term forest diversity experiments in boreal and temperate forest zones. Our aim was to compare the effects of forest diversity on herbivores belonging to different feeding guilds and inhabiting different tree species. At the same time we also examined the variation in herbivore responses due to tree age and sampling period within the season, the effects of experimental design (plot size and planting density) and the stability of herbivore responses over time. Herbivore responses varied significantly both among insect feeding guilds and among host tree species. Among insect feeding guilds, only leaf miner densities were consistently lower and less variable in mixed stands as compared to tree monocultures regardless of the host tree species. The responses of other herbivores to forest diversity depended largely on host tree species. Insect herbivory on birch was significantly lower in mixtures than in birch monocultures, whereas insect herbivory on oak and alder was higher in mixtures than in oak and alder monocultures. The effects of tree species diversity were also more pronounced in older trees, in the earlier part of the season, at larger plots and at lower planting density. Overall our results demonstrate that forest diversity does not generally and uniformly reduce insect herbivory and suggest instead that insect herbivore responses to forest diversity are highly variable and strongly dependent on the host tree species and other stand characteristics as well as on the type of the herbivore.     

Plant Size as Determinant of Species Richness of Herbivores,Natural Enemies and Pollinators across 21 Brassicaceae Species

Large plants are often more conspicuous and more attractive for associated animals than small plants, e.g. due to their wider range of resources. Therefore, plant size can positively affect species richness of associated animals, as shown for single groups of herbivores, but studies usually consider intraspecific size differences of plants in unstandardised environments. As comprehensive tests of interspecific plant size differences under standardised conditions are missing so far, we investigated effects of plant size on species richness of all associated arthropods using a common garden experiment with 21 Brassicaceae species covering a broad interspecific plant size gradient from 10 to 130 cm height. We recorded plant associated ecto- and endophagous herbivores, their natural enemies and pollinators on and in each aboveground plant organ, i.e. flowers, fruits, leaves and stems. Plant size (measured as height from the ground), the number of different plant organ entities and their biomass were assessed. Increasing plant size led to increased species richness of associated herbivores, natural enemies and pollinating insects. This pattern was found for ectophagous and endophagous herbivores, their natural enemies, as well as for herbivores associated with leaves and fruits and their natural enemies, independently of the additional positive effects of resource availability (i.e. organ biomass or number of entities and, regarding natural enemies, herbivore species richness). We found a lower R² for pollinators compared to herbivores and natural enemies, probably caused by the high importance of flower characteristics for pollinator species richness besides plant size. Overall, the increase in plant height from 10 to 130 cm led to a 2.7-fold increase in predicted total arthropod species richness. In conclusion, plant size is a comprehensive driver of species richness of the plant associated arthropods, including pollinators, herbivores and their natural enemies, whether they are endophagous or ectophagous or associated with leaves or fruits.     

Geographic association and temporal variation of chemical and physical defense and leaf damage in Datura stramonium

The evolution of plant defense traits has traditionally been explained trough the “coevolutionary arms race” between plants and herbivores. According to this, specialist herbivores have evolved to cope effectively with the defensive traits of their host plants and may even use them as a cue for host location. We analyzed the geographic association between leaf trichomes, two tropane alkaloids (putative resistance traits), and leaf damage by herbivores in 28 populations of Datura stramonium in central Mexico. Since the specialist leaf beetles Epitrix parvula and Lema trilineata are the main herbivores of D. stramonium in central Mexico, we predicted a positive association between plant defense and leaf damage across populations. Also, if physical environmental conditions (temperature or precipitation) constrain the expression of plant defense, then the geographic variation in leaf damage should be explained partially by the interaction between defensive traits and environmental factors. Furthermore, we studied the temporal and spatial variation in leaf trichome density and leaf damage in five selected populations of D. stramonium sampled in two periods (1997 vs. 2007). We found a positive association between leaf trichomes density and atropine concentration with leaf damage across populations. The interaction between defensive traits and water availability in each locality had a significant effect on the geographic variation in leaf damage. Differences among populations in leaf trichome density are maintained over time. Our results indicate that local plant–herbivore interaction plays an important role in shaping the geographic and temporal variation in plant defense in D. stramonium.     

Shoot development and extension of Quercus serrata saplings in response to insect damage and nutrient conditions

BACKGROUND AND AIMS: Plants have the ability to compensate for damage caused by herbivores. This is important to plant growth, because a plant cannot always avoid damage, even if it has developed defence mechanisms against herbivores. In previous work, we elucidated the herbivory-induced compensatory response of Quercus (at both the individual shoot and whole sapling levels) in both low- and high-nutrient conditions throughout one growing season. In this study, we determine how the compensatory growth of Quercus serrata saplings is achieved at different nutrient levels. METHODS: Quercus serrata saplings were grown under controlled conditions. Length, number of leaves and percentage of leaf area lost on all extension units (EUs) were measured. KEY RESULTS: Both the probability of flushing and the length of subsequent EUs significantly increased with an increase in the length of the parent EU. The probability of flushing increased with an increase in leaf damage of the parent EU, but the length of subsequent EUs decreased. This indicates that EU growth is fundamentally regulated at the individual EU level. The probabilities of a second and third flush were significantly higher in plants in high-nutrient soil than those in low-nutrient soil. The subsequent EUs of damaged saplings were also significantly longer at high-nutrient conditions. CONCLUSIONS: An increase in the probability of flushes in response to herbivore damage is important for damaged saplings to produce new EUs; further, shortening the length of EUs helps to effectively reproduce foliage lost by herbivory. The probability of flushing also varied according to soil nutrient levels, suggesting that the compensatory growth of individual EUs in response to local damage levels is affected by the nutrients available to the whole sapling.