Herbivore‐induced plant responses in Brassica oleracea prevail over effects of constitutive resistance and result in enhanced herbivore attack

1. Plant responses to herbivore attack may have community‐wide effects on the composition of the plant‐associated insect community. Thereby, plant responses to an early‐season herbivore may have profound consequences for the amount and type of future attack. 2. Here we studied the effect of early‐season herbivory by caterpillars of Pieris rapae on the composition of the insect herbivore community on domesticated Brassica oleracea plants. We compared the effect of herbivory on two cultivars that differ in the degree of susceptibility to herbivores to analyse whether induced plant responses supersede differences caused by constitutive resistance. 3. Early‐season herbivory affected the herbivore community, having contrasting effects on different herbivore species, while these effects were similar on the two cultivars. Generalist insect herbivores avoided plants that had been induced, whereas these plants were colonised preferentially by specialist herbivores belonging to both leaf‐chewing and sap‐sucking guilds. 4. Our results show that community‐wide effects of early‐season herbivory may prevail over effects of constitutive plant resistance. Induced responses triggered by prior herbivory may lead to an increase in susceptibility to the dominant specialists in the herbivorous insect community. The outcome of the balance between contrasting responses of herbivorous community members to induced plants therefore determines whether induced plant responses result in enhanced plant resistance.     

The ecology and evolution of host-plant resistance to insects

Genetic techniques have yielded new insights into plant-herbivore coevolution. Quantitative genetic tests of herbivory theory reveal that in some cases insect herbivores impose selection on resistance traits. Also, some resistance traits are costly while others appear not to be, and genetic models can explain these results. Genetic variation in plant resistance influences insect community structure by modifying interactions of herbivores with competitors and natural enemies. Therefore, models of multispecies coevolution are more realistic than pairwise coevolutionary models. Ecological genetics will facilitate further theoretical and empirical exploration of multispecies coevolution of plants and herbivores.     

PAIRWISE VERSUS DIFFUSE NATURAL SELECTION AND THE MULTIPLE HERBIVORES OF SCARLET GILIA,IPOMOPSIS AGGREGATA

Recent theoretical studies have argued that plant-herbivore coevolution proceeds in a diffuse rather than a pairwise manner in multispecies interactions when at least one of two conditions are met: (1) genetic correlations exist between plant resistances to different herbivore species; and (2) ecological interactions between herbivores sharing a host plant cause nonadditive impacts of herbivory on plant fitness. We present results from manipulative field experiments investigating the single and interactive fitness effects of three types of herbivory on scarlet gilia (Ipomopsis aggregata) over two years of study. We utilize these data to test whether selection imposed by herbivore attack on date of first flowering is pairwise (independent) or diffuse (dependent) in nature. Our results reveal complex patterns of the fitness effect of herbivores. Simulated early season browsing had a strong negative fitness effect on plants and also reduced subsequent insect attack. Surprisingly, this ecological interaction did not translate into significant interactions between clipping and insect manipulations on plant fitness. However, we detected a significant interaction between seed fly and caterpillar herbivory on plant fitness, with the negative effect of either insect being greatest when occurring alone. These results suggest that herbivore-imposed selection may have pairwise and diffuse components. In our selection analysis of flowering phenology, we discovered significant pairwise linear selection imposed by clipping, diffuse linear selection imposed by insects, and diffuse nonlinear selection imposed by clipping and insect attack acting simultaneously. Our results reveal that the evolution of flowering phenology in scarlet gilia may be in response to diffuse and pairwise natural selection imposed by multiple herbivores. We discuss the evolution of resistance characters in light of diffuse versus pairwise forms of linear and nonlinear selection and stress the complexity of selection imposed by suites of interacting species.     

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.     

Combined effects of patch size and plant nutritional quality on local densities of insect herbivores

Plant–insect interactions occur in spatially heterogeneous habitats. Understanding how such interactions shape density distributions of herbivores requires knowledge on how variation in plant traits (e.g. nutritional quality) affects herbivore abundance through, for example, affecting movement rates and aggregation behaviour. We studied the effects of plant patch size and herbivore-induced differences in plant nutritional quality on local densities of insect herbivores for two Brassica oleracea cultivars, i.e. white cabbage and Brussels sprouts. Early season herbivory as a treatment resulted in measurable differences in glucosinolate concentrations in both cultivars throughout the season. Herbivore induction and patch size both influenced community composition of herbivores in both cultivars, but the effects differed between species. Flea beetles (Phyllotreta spp.) were more abundant in large than in small patches, and this patch response was more pronounced on white cabbage than on Brussels sprouts. Herbivore-induction increased densities in all patches. Thrips tabaci was also more abundant in large patches and densities of this species were higher on Brussels sprouts than on white cabbage. Thrips densities were lower on induced than on control plants of both cultivars and this negative effect of induction tended to be more pronounced in large than in small patches. Densities of the cabbage moth (Mamestra brassicae) were lower on Brussels sprouts than on white cabbage and lower on herbivore-induced than on uninduced plants, with no effect of patch size. No clear effects of patch size and induction were found for aphids. This study shows that constitutive and herbivore-induced differences in plant traits interact with patch responses of insect herbivores.     

Similar responses of insect herbivores to leaf fluctuating asymmetry

Fluctuating asymmetry (FA) represents small, random variation from symmetry and it has been used as an indicator of plant quality and susceptibility to herbivory. In this study, the effects of FA on the responses of distinct herbivore species belonging to several guilds were examined along an environmental gradient in south Florida. This approach was chosen because it relies on a multi-species approach to the study of fluctuating asymmetry and patterns of herbivory between and within plants along an environmental gradient of salinity and plant stress. To examine differences in FA between and within plant communities, seven plant species were investigated. Four of these plants were coastal species and three species occurred in upland communities. Levels of FA were assessed before herbivory and plants were followed for the whole herbivory season in 2006. Coastal plants exhibited significantly higher salt concentration, higher percentage of asymmetric leaves and higher asymmetry levels than upland plants. Herbivore abundance varied widely amongst the seven species studied, but quantitative syntheses of our results indicated significant and positive responses of insect herbivores to leaf asymmetry: insects were 25.11% more abundant on more asymmetric plants and stronger effects of asymmetry were observed for leaf miners compared to gall-formers. As demonstrated by other recent studies, FA might be used as a reliable stress indicator, leading to similar responses of insect herbivores to variation in leaf symmetry.     

Effects of clipping and N fertilization on insect herbivory and infestation by pathogenic fungi on bilberry

Interactions among herbivores or between herbivores and other plant natural enemies, such as fungal pathogens, range from competition to facilitation. Moreover, the outcome of these interactions depends on the ecological context where they occur. In this study we examined the effects of clipping, as a surrogate of herbivory by ungulates, on the damage caused by two types of natural enemies (herbivorous insects and foliar fungal pathogens) on bilberry, Vaccinium myrtillus, in combination with nitrogen (N) fertilization representing current N atmospheric deposition. To examine whether the responses of both these natural enemies were mediated by changes in the plant, we estimated the effects of the treatments on bilberry growth and branching and on chlorophyll content as proxy of N content in leaves. Clipping increased the proportion of leaves damaged by herbivorous insects regardless of whether it was combined with N fertilization or not in 2008. In 2007 and 2009 repeated damage to the shrub also facilitated insect herbivory but only under N applications. Regarding fungal infestation incidence, clipping decreased the proportion of infected leaves in all the years considered but only in fertilized plots. Our results suggest that vertebrate herbivores facilitate insect herbivory and reduce fungal infestation but that these effects are dependent on nutritional conditions. Moreover, we found a negative residual correlation between insect herbivory and fungal infestation on bilberry leaves. Therefore, interactions between insect herbivores and fungal pathogens could be implicated in the final outcome of interactions between browsing ungulates and both bilberry natural enemies.     

Induced defences in the neotropical tree Bauhinia brevipes (Vog.) to herbivory: effects of damage-induced changes on leaf quality and insect attack

Induced defences to herbivory are physical, nutritional, and allelochemical traits that change in plants following damage or stress, and that reduce the performance and/or preference of herbivores. The aim of this study was to verify the occurrence and effect of induced responses in Bauhinia brevipes (Vog.) (Leguminosae) which defend it against herbivores, through the manipulation of its leaves, and their effects on herbivore foraging behaviour. We selected 15 plants in the field, and three shoots per plant were subjected to one of three treatments: (1) damaged shoots (simulation of the main types of foliar herbivory and insect exclusion); (2) damaged control shoots (insect exclusion); and (3) control shoots (not manipulated). Water and nitrogen content, tannin concentration, levels of herbivory, and shoot growth rates were compared among treatments. Leaf quality varied among treatments. Damaged leaves showed higher tannin concentration, and lower water and nitrogen content compared to undamaged leaves. On the other hand, they experienced higher rates of herbivory than leaves on control shoots. Moreover, shoots that were experimentally induced showed a higher increase in final shoot length. These results suggest that simulated herbivory on B. brevipes reduced the nutritional quality of its leaves and increased the amount of secondary compounds, therefore altering insect herbivore attack and increasing shoot performance.     

Indirect effects of deer on insect pests and soybean plants

1. White-tailed deer (Odocoileus virginianus Zimmermann) and insect pests negatively affect soybean production; however, little is known about how these herbivores potentially interact to affect soybean yield. Previous studies have shown deer browse on non-crop plants affects insect density and insect-mediated leaf damage, which together reduce plant reproductive output. In soybeans, reproductive output is influenced by direct and indirect interactions of different herbivores.
2. Here, we quantified indirect interactions between two groups of herbivores (mammals and insects) and their effects on soybean growth and yield. We examined responses of insect pest communities along a gradient of deer herbivory (29% to 49% browsed stems) in soybean monocultures.
3. Structural equation models showed that deer browse had direct negative effects on soybean plant height and yield. Deer browse indirectly decreased insect-mediated leaf damage by reducing plant height. Deer browse also indirectly increased pest insect abundance through reductions in plant height. Similarly, deer herbivory had an indirect positive effect on leaf carbon: nitrogen ratios through changes in plant height, thereby decreasing leaf nutrition.
4. These results suggest that pest insect abundance may be greater on soybean plants in areas of higher deer browse, but deer browse may reduce insect herbivory through reduced leaf nutrition.

     

Effects of Nitrogen Deposition on Insect Herbivory: Implications for Community and Ecosystem Processes

The deposition of anthropogenically fixed nitrogen (N) from the atmosphere onto land and plant surfaces has strong influences on terrestrial ecosystem processes. Although recent research has expanded our understanding of how N deposition affects ecosystems directly, less attention has been directed toward the investigation of how N deposition may affect ecosystems indirectly by modifying interactions among organisms. Empirical evidence suggests that there are several mechanisms by which N deposition may affect interactions between plants and insect herbivores. The most likely mechanisms are deposition-induced shifts in the quality and availability of host plant tissues. We discuss the effects of N deposition on host plant chemistry, production, and phenology, and we review the evidence for the effects of N deposition on insect herbivores at the individual, population, and community levels. In general, N deposition has positive effects on individual insect performance, probably due to deposition-induced improvements in host plant chemistry. These improvements include increased N and decreased carbon-based defensive compound concentrations. The evidence to date suggests that N deposition may also have a positive effect on insect populations. These effects may have considerable ecological, as well as economic consequences if the rates of herbivory on economically important timber species continue to increase. Deposition-induced changes in plant–herbivore relationships may affect community and ecosystem processes. However, we predict that the larger-scale consequences of interactions between N deposition and herbivory will vary based on site-specific factors. In addition, interactions between N deposition and other global-scale changes may lead to nonadditive effects on patterns of herbivory.     

Asymmetric effects of a leaf‐chewing herbivore on aphid population growth

1. Plant responses to herbivory are often specific to the feeding guild of the attacking herbivore. These phytochemical responses to herbivore damage can affect herbivore performance and activity. Comprehensive studies on the ecological consequences of multi‐herbivore plant interactions are key to understanding plant–herbivore community dynamics. 2. This study examined how feeding damage by co‐occurring herbivores from separate feeding guilds, Myzus persicae (Sulzer), a sucking herbivore, and Leptinotarsa decemlineata (Say), a chewing herbivore, alter plant chemistry and indirectly affect herbivore performance. Performance was measured when each insect fed on plants individually, sequentially, or simultaneously in laboratory and field experiments. Phytohormone and glycoalkaloid content were measured for each feeding sequence to evaluate plant responses to herbivory by each guild. Mid‐season and end‐of‐season tuber yield were evaluated in the field study. 3. Damage by L. decemlineata negatively impacted M. persicae performance in both laboratory and field settings. Damage by M. persicae did not affect L. decemlineata performance in laboratory assays. However, L. decemlineata performance was positively affected by M. persicae herbivory in the field, but this effect was temporary. Although phytohormones and plant defences varied across treatments, they provide little resolution on interaction outcomes. 4. These results confirm that the presence of multiple feeding guilds on a single plant can affect these chewing and sucking herbivores differentially, but given the variability in our phytochemical analyses compared with other studies, the mechanism remains unclear. The study's findings show that aphids are negatively affected by chewing herbivores across systems, while aphids temporarily affected beetles positively.     

Effects of elevated CO2 and O3 on leaf damage and insect abundance in a soybean agroecosystem

By altering myriad aspects of leaf chemistry, increasing concentrations of CO₂ and O₃ in the atmosphere derived from human activities may fundamentally alter the relationships between insect herbivores and plants. Because exposure to elevated CO₂ can alter the nutritional value of leaves, some herbivores may increase consumption rates to compensate. The effects of O₃ on leaf nutritional quality are less clear; however, increased senescence may also reduce leaf quality for insect herbivores. Additionally, changes in secondary chemistry and the microclimate of leaves may render plants more susceptible to herbivory in elevated CO₂ and O₃. Damage to soybean (Glycine max L.) leaves and the size and composition of the insect community in the plant canopy were examined in large intact plots exposed to elevated CO₂ (~550 μmol mol⁻¹) and elevated O₃ (1.2*ambient) in a fully factorial design with a Soybean Free Air Concentration Enrichment system (SoyFACE). Leaf area removed by folivorous insects was estimated by digital photography and insect surveys were conducted during two consecutive growing seasons, 2003 and 2004. Elevated CO₂ alone and in combination with O₃ increased the number of insects and the amount of leaf area removed by insect herbivores across feeding guilds. Exposure to elevated CO₂ significantly increased the number of western corn rootworm (Diabrotica virgifera) adults (foliage chewer) and soybean aphids (Aphis glycines; phloem feeder). No consistent effect of elevated O₃ on herbivory or insect population size was detected. Increased loss of leaf area to herbivores was associated with increased carbon-to-nitrogen ratio and leaf surface temperature. Soybean aphids are invasive pests in North America and new to this ecosystem. Higher concentrations of CO₂ in the atmosphere may increase herbivory in the soybean agroecosystem, particularly by recently introduced insect herbivores. Handling editor: Gary Felton.     

Impacts of urbanization on insect herbivory and plant defences in oak trees

Systematic comparisons of species interactions in urban versus rural environments can improve our understanding of shifts in ecological processes due to urbanization. However, such studies are relatively uncommon and the mechanisms driving urbanization effects on species interactions (e.g. between plants and insect herbivores) remain elusive. Here we investigated the effects of urbanization on leaf herbivory by insect chewers and miners associated with the English oak Quercus robur by sampling trees in rural and urban areas throughout most of the latitudinal distribution of this species. In performing these comparisons, we also controlled for the size of the urban areas (18 cities) and gathered data on CO₂ emissions. In addition, we assessed whether urbanization affected leaf chemical defences (phenolic compounds) and nutritional traits (phosphorus and nitrogen), and whether such changes correlated with herbivory levels. Urbanization significantly reduced leaf chewer damage but did not affect leaf miners. In addition, we found that leaves from urban locations had lower levels of chemical defences (condensed and hydrolysable tannins) and higher levels of nutrients (nitrogen and phosphorus) compared to leaves in rural locations. The magnitude of urbanization effects on herbivory and leaf defences was not contingent upon city size. Importantly, while the effects of urbanization on chemical defences were associated with CO₂ emissions, changes in leaf chewer damage were not associated with either leaf traits or CO₂ levels. These results suggest that effects of urbanization on herbivory occur through mechanisms other than changes in the plant traits measured here. Overall, our simultaneous assessment of insect herbivory, plant traits and abiotic correlates advances our understanding of the main drivers of urbanization effects on plant–herbivore interactions.     

Präadaptation, Wirtskreiserweiterung and Parallel-Cladogenese in der Evolution von phytophagen Insekten1, 2

Preadaptation, host shifts and parallel cladogenesis in the evolution of phytophagous insects In this contribution we investigate the possibilities to apply concepts developed for the evolution of animal parasites to insect-plant systems. We compare host parasite systems in animals with plant-herbivore systems and list similarities and differences. The terms preadaptation, predisposition, expansion and contraction of host ranges, and parallel cladogenesis are discussed. We enumerate general preadaptations for the evolution of herbivory in insects and preadaptations for shifts between herbivory and entomophagy. Examples are given for expansions of host ranges based on phytochemical or structural characters of host plants. Cases of parallel cladogenesis in herbivoreparasitoid systems and plant-herbivore systems are compiled from the literature. An analysis of the insect fauna of the “thistles” (Cynaroideae) in the Palearctic and Nearctic demonstrates the importance of the evolutionary history of the plant taxa and of the existence of preadapted pools of herbivores for the evolution of guilds of specialized herbivores. The members of the Curculionid taxon Cleoninae provide examples for multiple colonizations and radiations of herbivores on the Cynaroideae. The taxonomic and biological relationships of the weevil genera Rhinocyllus, Bangasternus and Larinus which exploit the flower heads of Cynaroideae, can be interpreted as result of a basic parallel cladogenesis between herbivore and host. A gelelectrophoretic analysis of Larinus spp. supports this hypothesis.     

Latitudinal variation in resistance and tolerance to herbivory in the perennial herb Lythrum salicaria is related to intensity of herbivory and plant phenology

Both the length of the growing season and the intensity of herbivory often vary along climatic gradients, which may result in divergent selection on plant phenology, and on resistance and tolerance to herbivory. In Sweden, the length of the growing season and the number of insect herbivore species feeding on the perennial herb Lythrum salicaria decrease from south to north. Previous common‐garden experiments have shown that northern L. salicaria populations develop aboveground shoots earlier in the summer and finish growth before southern populations do. We tested the hypotheses that resistance and tolerance to damage vary with latitude in L. salicaria and are positively related to the intensity of herbivory in natural populations. We quantified resistance and tolerance of populations sampled along a latitudinal gradient by scoring damage from natural herbivores and fitness in a common‐garden experiment in the field and by documenting oviposition and feeding preference by specialist leaf beetles in a glasshouse experiment. Plant resistance decreased with latitude of origin, whereas plant tolerance increased. Oviposition and feeding preference in the glasshouse and leaf damage in the common‐garden experiment were negatively related to damage in the source populations. The latitudinal variation in resistance was thus consistent with reduced selection from herbivores towards the northern range margin of L. salicaria. Variation in tolerance may be related to differences in the timing of damage in relation to the seasonal pattern of plant growth, as northern genotypes have developed further than southern have when herbivores emerge in early summer.     

Mechanisms and ecological implications of plant-mediated interactions between belowground and aboveground insect herbivores

Plant-mediated interactions between belowground (BG) and aboveground (AG) herbivores have received increasing interest recently. However, the molecular mechanisms underlying ecological consequences of BG–AG interactions are not fully clear yet. Herbivore-induced plant defenses are complex and comprise phytohormonal signaling, gene expression and production of defensive compounds (defined here as response levels), each with their own temporal dynamics. Jointly they shape the response that will be expressed. However, because different induction methods are used in different plant-herbivore systems, and only one or two response levels are measured in each study, our ability to construct a general framework for BG–AG interactions remains limited. Here we aim to link the mechanisms to the ecological consequences of plant-mediated interactions between BG and AG insect herbivores. We first outline the molecular mechanisms of herbivore-induced responses involved in BG–AG interactions. Then we synthesize the literature on BG–AG interactions in two well-studied plant-herbivore systems, Brassica spp. and Zea mays, to identify general patterns and specific differences. Based on this comprehensive review, we conclude that phytohormones can only partially mimic induction by real herbivores. BG herbivory induces resistance to AG herbivores in both systems, but only in maize this involves drought stress responses. This may be due to morphological and physiological differences between monocotyledonous (maize) and dicotyledonous (Brassica) species, and differences in the feeding strategies of the herbivores used. Therefore, we strongly recommend that future studies explicitly account for these basic differences in plant morphology and include additional herbivores while investigating all response levels involved in BG–AG interactions.     

Interactions among fungal endophytes,grasses and herbivores

The interaction between two species often depends on the presence or absence of a third species. One widespread three-species interaction involves fungal endophytes infecting grasses and the herbivores that feed upon them. The endophytes are allied with the fungal family Clavicipitaceae and grow systemically in intercellular spaces in above-ground plant tissues including seeds. Like relatedClaviceps species, the endophytes produce a variety of alkaloids that make the host plants toxic or distasteful to herbivores. A large number of grass species are infected, especially cool-season grasses in temperate areas. Field and laboratory studies have shown that herbivores avoid infected plants in choice trials and suffer increased mortality and decreased growth on infected grasses in feeding experiments. Resistance to herbivores may provide a selective advantage to infected plants in competitive interactions with noninfected plants. Recent studies have shown that differential herbivory can reverse competitive hierarchies among plant species. Both endophyte-infected and noninfected tall fescue grass (Festuca arundinacea) are outcompeted by orchardgrass (Dactylis glomerata) in the absence of insect herbivory. However, when herbivores are present infected tall fescue outcompetes orchardgrass. These results suggest that the frequency of infection in grass species and grassland communities will increase over time. Several studies are reviewed illustrating increases in infection frequency within grass populations subject to herbivore pressure. Endophytic fungi may be important regulators of plant-herbivore interactions and so indirectly affect the structure and dynamics of plant communities.     

Interactive effects of elevation,species richness and extreme climatic events on plant–pollinator networks

Plant–pollinator interactions are essential for the functioning of terrestrial ecosystems, but are increasingly affected by global change. The risks to such mutualistic interactions from increasing temperature and more frequent extreme climatic events such as drought or advanced snow melt are assumed to depend on network specialization, species richness, local climate and associated parameters such as the amplitude of extreme events. Even though elevational gradients provide valuable model systems for climate change and are accompanied by changes in species richness, responses of plant–pollinator networks to climatic extreme events under different environmental and biotic conditions are currently unknown. Here, we show that elevational climatic gradients, species richness and experimentally simulated extreme events interactively change the structure of mutualistic networks in alpine grasslands. We found that the degree of specialization in plant–pollinator networks (H2′) decreased with elevation. Nonetheless, network specialization increased after advanced snow melt at high elevations, whereas changes in network specialization after drought were most pronounced at sites with low species richness. Thus, changes in network specialization after extreme climatic events depended on climatic context and were buffered by high species richness. In our experiment, only generalized plant–pollinator networks changed in their degree of specialization after climatic extreme events. This indicates that contrary to our assumptions, network generalization may not always foster stability of mutualistic interaction networks.     

Spatial Coupling of Plant and Herbivore Dynamics: The Contribution of Herbivore Dispersal to Transient and Persistent "Waves" of Damage

Spatial variations in the abundance of insect herbivores and in herbivore damage are both striking an commonplace. The standard explanations for heterogeneity in herbivore attack emphasize spatial variations in plant genetype, soils, or physical environment. Here I examine an alternative hypothesis-that heterogeneity arises in plant-herbivore systems, even in homogeneous environments, as a result of the direct coupling of herbivore movement to herbivore density and plant quality. Using a mathematical model for plant quality and herbivore growth and dispersal, I demonstrate how spatial instabilities about homogeneous steady state values result in both transient and stationary waves of damage to the plant. Key herbivore movement behaviors include the tendendy for herbivores to aggregate over a range of spatial scales for increased feeding efficiency and the tendency for herbivores to move up gradients in plant quality (herbivory-taxis). My approach translates the biased "random walk" behavior of individual herbivores into a continuum partial differential equation model. Analytical and numerical methods are used to demonstrate the nature of the spatio-temporal variations in plant quality and herbivore density.     

Environmental and genetic control of insect abundance and herbivory along a forest elevational gradient

Environmental conditions and plant genotype may influence insect herbivory along elevational gradients. Plant damage would decrease with elevation as temperature declines to suboptimal levels for insects. However, host plants at higher elevations may exhibit traits that either reduce or enhance leaf quality to insects, with uncertain net effects on herbivory. We examined folivory, insect abundance and leaf traits along six replicated elevational ranges in Nothofagus pumilio forests of the northern Patagonian Andes, Argentina. We also conducted a reciprocal transplant experiment between low- and high-elevation sites to test the extent of environmental and plant genetic control on insect abundance and folivory. We found that insect abundance, leaf size and specific leaf area decreased, whereas foliar phosphorous content increased, from low-, through mid- to high-elevation sites. Path analysis indicated that changes in both insect abundance and leaf traits were important in reducing folivory with increasing elevation and decreasing mean temperature. At both planting sites, plants from a low-elevation origin experienced higher damage and supported greater insect loads than plants from a high-elevation origin. The differences in leaf damage between sites were twofold larger than those between plant origins, suggesting that local environment was more important than host genotype in explaining folivory patterns. Different folivore guilds exhibited qualitatively similar responses to elevation. Our results suggest an increase in insect folivory on high-elevation N. pumilio forests under future climate warming scenarios. However, in the short-term, folivory increases might be smaller than expected from insect abundance only because at high elevations herbivores would encounter more resistant tree genotypes.