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.     

Above- and belowground insect herbivores differentially affect soil nematode communities in species-rich plant communities

Interactions between above‐ and belowground invertebrate herbivores alter plant diversity, however, little is known on how these effects may influence higher trophic level organisms belowground. Here we explore whether above‐ and belowground invertebrate herbivores which alter plant community diversity and biomass, in turn affect soil nematode communities. We test the hypotheses that insect herbivores 1) alter soil nematode diversity, 2) stimulate bacterial‐feeding and 3) reduce plant‐feeding nematode abundances. In a full factorial outdoor mesocosm experiment we introduced grasshoppers (aboveground herbivores), wireworms (belowground herbivores) and a diverse soil nematode community to species‐rich model plant communities. After two years, insect herbivore effects on nematode diversity and on abundance of herbivorous, bacterivorous, fungivorous and omni‐carnivorous nematodes were evaluated in relation to plant community composition. Wireworms did not affect nematode diversity despite enhanced plant diversity, while grasshoppers, which did not affect plant diversity, reduced nematode diversity. Although grasshoppers and wireworms caused contrasting shifts in plant species dominance, they did not affect abundances of decomposer nematodes at any trophic level. Primary consumer nematodes were, however, strongly promoted by wireworms, while community root biomass was not altered by the insect herbivores. Overall, interaction effects of wireworms and grasshoppers on the soil nematodes were not observed, and we found no support for bottom‐up control of the nematodes. However, our results show that above‐ and belowground insect herbivores may facilitate root‐feeding rather than decomposer nematodes and that this facilitation appears to be driven by shifts in plant species composition. Moreover, the addition of nematodes strongly suppressed shoot biomass of several forb species and reduced grasshopper abundance. Thus, our results suggest that nematode feedback effects on plant community composition, due to plant and herbivore parasitism, may strongly depend on the presence of insect herbivores.     

Plant genotypic diversity reduces the rate of consumer resource utilization

While plant species diversity can reduce herbivore densities and herbivory, little is known regarding how plant genotypic diversity alters resource utilization by herbivores. Here, we show that an invasive folivore—the Japanese beetle (Popillia japonica)—increases 28 per cent in abundance, but consumes 24 per cent less foliage in genotypic polycultures compared with monocultures of the common evening primrose (Oenothera biennis). We found strong complementarity for reduced herbivore damage among plant genotypes growing in polycultures and a weak dominance effect of particularly resistant genotypes. Sequential feeding by P. japonica on different genotypes from polycultures resulted in reduced consumption compared with feeding on different plants of the same genotype from monocultures. Thus, diet mixing among plant genotypes reduced herbivore consumption efficiency. Despite positive complementarity driving an increase in fruit production in polycultures, we observed a trade-off between complementarity for increased plant productivity and resistance to herbivory, suggesting costs in the complementary use of resources by plant genotypes may manifest across trophic levels. These results elucidate mechanisms for how plant genotypic diversity simultaneously alters resource utilization by both producers and consumers, and show that population genotypic diversity can increase the resistance of a native plant to an invasive herbivore.     

The effects of long-term nitrogen loading on grassland insect communities

Just as long-term nitrogen loading of grasslands decreases plant species richness and increases plant biomass, we have found that nitrogen loading decreases insect species richness and increases insect abundances. We sampled 54 plots that had been maintained at various rates of nitrogen addition for 14 years. Total insect species richness and effective insect diversity, as well as herbivore and predator species richness, were significantly, negatively related to the rate of nitrogen addition. However, there was variation in trophic responses to nitrogen. Detritivore species richness increased as nitrogen addition increased, and parasitoids showed no response. Insect abundances, measured as the number of insects and insect biovolume (an estimate of biomass), were significantly, positively related to the rate of nitrogen addition, as were the abundances of herbivores and detritivores. Parasitoid abundance was negatively related to the rate of nitrogen addition. Changes in the insect community were correlated with changes in the plant community. As rates of nitrogen addition increased, plant species richness decreased, plant productivity and plant tissue nitrogen increased, and plant composition shifted from C₄ to C₃ grass species. Along this gradient, total insect species richness and effective insect diversity were most strongly, positively correlated with plant species richness. Insect biovolume was negatively correlated with plant species richness. Responses of individual herbivores varied along the nitrogen gradient, but numbers of 13 of the 18 most abundant herbivores were positively correlated with their host plant biomass. Although insect communities did not respond as strongly as plant communities, insect species richness, abundance, and composition were impacted by nitrogen addition. This study demonstrates that long-term nitrogen loading affects the entire food chain, simplifying both plant and insect communities. Received: 18 May 1999 / Accepted: 5 January 2000     

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.     

N availability does not modify plant-mediated responses of Trichoplusia ni to elevated CO2

Aims Elevated CO₂ and increased N availability can alter a variety of plant physiological processes leading to changes in the nutritional quality of leaf tissue for herbivores. Numerous experiments have examined the responses of herbivores to environmental change; however the potential effects of simultaneous change in multiple factors on leaf-chewing insect herbivores are less well understood. The plant-mediated effects of elevated CO₂ and high N on the performance of a generalist leaf-chewing insect herbivore, Trichoplusia ni, were investigated.Methods Newly hatched T. ni larvae were introduced to Amaranthus viridis and Polygonum persicaria plants grown under ambient and elevated CO₂ and low and high N conditions. Insect performance was assessed by measuring larvae weight after ten days of feeding. Plant photosynthesis, biomass, leaf area and specific leaf weight were measured to determine the effects of elevated CO₂, N and insect feeding on plant performance.Important findings Elevated CO₂ did not have strong effects on plant or insect performance, only affecting a few responses under low or high N conditions, but not both. Growth under high nitrogen improved almost all measures of plant performance. Trichoplusia ni performed significantly better on Amaranthus viridis (C 4) compared to Polygonum persicaria (C 3), despite similar leaf C:N ratios in both species. The performance of T. ni caterpillars was only improved by the high nitrogen treatment when they were feeding on P. persicaria, the host they performed poorly on. The only interactions between N and CO₂ affecting plant performance were seen for leaf photosynthesis of P. persicaria and leaf area of A. viridis. Contrary to the predictions, there were no significant CO₂ by N interactions affecting T. ni performance.     

Community heterogeneity and the evolution of interactions between plants and insect herbivores

Plant communities vary tremendously in terms of productivity, species diversity, and genetic diversity within species. This vegetation heterogeneity can impact both the likelihood and strength of interactions between plants and insect herbivores. Because altering plant-herbivore interactions will likely impact the fitness of both partners, these ecological effects also have evolutionary consequences. We review several hypothesized and well-documented mechanisms whereby variation in the plant community alters the plant-herbivore interaction, discuss potential evolutionary outcomes of each of these ecological effects, and conclude by highlighting several avenues for future research. The underlying theme of this review is that the neighborhood of plants is an important determinant of insect attack, and this results in feedback effects on the plant community. Because plants exert selection on herbivore traits and, reciprocally, herbivores exert selection on plant-defense traits, variation in the plant community likely contributes to spatial and temporal variation in both plant and insect traits, which could influence macroevolutionary patterns.     

The deleterious effects of salinity stress on leafminers and their freshwater host

Abstract.  1. Salinity is an important cause of abiotic stress in wetland communities yet little is known about its consequences for freshwater plants and their insect herbivores. The goal of this study was to test the effect of salinity on a leafmining insect, Cerodontha iridiphora , and its herbaceous host plant, Iris hexagona .
2. Leafminer performance was evaluated on irises grown in control and saline treatments, and the effects of salinity and herbivory on leaf quality and mortality was measured.
3. Leafminer density and size were significantly lower on irises grown in saline water compared with freshwater.
4. Both salinity and herbivory accelerated leaf senescence and mortality, and their combined effects increased tissue loss by an order of magnitude compared with controls.
5. Leafminers acted as nutrient sinks. The undamaged regions of mined leaves contained 40% less nitrogen than unmined leaves, providing a mechanism for the premature leaf mortality.
6. Salinity was detrimental to the performance and survival of both the iris leafminer and its host plant. We propose that glycophytic host plants and their insect herbivores will suffer more than halophytic communities from environmental salinity because they lack the adaptive mechanisms to tolerate this potent physiological stress.     

Positive Effects of Plant Genotypic and Species Diversity on Anti-Herbivore Defenses in a Tropical Tree Species

Despite increasing evidence that plant intra- and inter-specific diversity increases primary productivity, and that such effect may in turn cascade up to influence herbivores, there is little information about plant diversity effects on plant anti-herbivore defenses, the relative importance of different sources of plant diversity, and the mechanisms for such effects. For example, increased plant growth at high diversity may lead to reduced investment in defenses via growth-defense trade-offs. Alternatively, positive effects of plant diversity on plant growth may lead to increased herbivore abundance which in turn leads to a greater investment in plant defenses. The magnitude of trait variation underlying diversity effects is usually greater among species than among genotypes within a given species, so plant species diversity effects on resource use by producers as well as on higher trophic levels should be stronger than genotypic diversity effects. Here we compared the relative importance of plant genotypic and species diversity on anti-herbivore defenses and whether such effects are mediated indirectly via diversity effects on plant growth and/or herbivore damage. To this end, we performed a large-scale field experiment where we manipulated genotypic diversity of big-leaf mahogany (Swietenia macrophylla) and tree species diversity, and measured effects on mahogany growth, damage by the stem-boring specialist caterpillar Hypsipyla grandella, and defensive traits (polyphenolics and condensed tannins in stem and leaves). We found that both forms of plant diversity had positive effects on stem (but not leaf) defenses. However, neither source of diversity influenced mahogany growth, and diversity effects on defenses were not mediated by either growth-defense trade-offs or changes in stem-borer damage. Although the mechanism(s) of diversity effects on plant defenses are yet to be determined, our study is one of the few to test for and show producer diversity effects on plant chemical defenses.     

Woody plant phylogenetic diversity mediates bottom–up control of arthropod biomass in species-rich forests

Global change is predicted to cause non-random species loss in plant communities, with consequences for ecosystem functioning. However, beyond the simple effects of plant species richness, little is known about how plant diversity and its loss influence higher trophic levels, which are crucial to the functioning of many species-rich ecosystems. We analyzed to what extent woody plant phylogenetic diversity and species richness contribute to explaining the biomass and abundance of herbivorous and predatory arthropods in a species-rich forest in subtropical China. The biomass and abundance of leaf-chewing herbivores, and the biomass dispersion of herbivores within plots, increased with woody plant phylogenetic diversity. Woody plant species richness had much weaker effects on arthropods, but interacted with plant phylogenetic diversity to negatively affect the ratio of predator to herbivore biomass. Overall, our results point to a strong bottom–up control of functionally important herbivores mediated particularly by plant phylogenetic diversity, but do not support the general expectation that top–down predator effects increase with plant diversity. The observed effects appear to be driven primarily by increasing resource diversity rather than diversity-dependent primary productivity, as the latter did not affect arthropods. The strong effects of plant phylogenetic diversity and the overall weaker effects of plant species richness show that the diversity-dependence of ecosystem processes and interactions across trophic levels can depend fundamentally on non-random species associations. This has important implications for the regulation of ecosystem functions via trophic interaction pathways and for the way species loss may impact these pathways in species-rich forests.     

Small mammal herbivores mediate the effects of soil nitrogen and invertebrate herbivores on grassland diversity

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Genotypic selection shapes patterns of within-species diversity in experimental plant populations

1.  An increasing body of evidence suggests that within-species diversity plays an important role for community and ecosystem functioning, alters complex trophic interactions and affects patterns of species diversity and coexistence. Nonetheless, we lack a good understanding of how genotypic trait variation translates into shifts in the relative abundance of genotypes within populations.
2.  In this study, we show that genotypic selection strongly alters dominance relationships among genotypes over a period of 5 years. This resulted in remarkably consistent changes in the proportional representation of genotypes, and in a concomitant decline of diversity and evenness in our experimental populations.
3.  High growth rates and the production of large offspring were positively associated with genotypic performance. Vegetative abundances of genotypes translated monotonically into flowering frequencies.
4.   Synthesis . We conclude that genotypic selection markedly affects patterns of diversity and consistently alters genotypic abundance and mean trait distributions in plant populations over a relatively short period of time.     

Impact of avian and arthropod predation on lepidopteran caterpillar densities and plant productivity in an ephemeral agroecosystem

Abstract.  1. Most studies evaluating the combined impact of spiders and other predators on herbivore densities in agroecosystems have focused primarily on their trophic connections with invertebrate predators (e.g. carabids, chrysopids); however linkages among spiders and vertebrate predators may also help structure the population dynamics of insect herbivores. A field experiment was conducted to examine the impact of avian and spider predation on lepidopteran caterpillar densities and plant productivity within a Brassica agroecosystem.
2. Arthropod abundance, leaf-chewing damage, and final plant productivity associated with broccoli, Brassica oleracea L. (var. italica ), were recorded for four treatments: (1) bird present but spiders removed; (2) both birds and spiders present; (3) birds excluded, spiders present; and (4) birds and spiders both excluded.
3. Densities of Artogeia rapae L. (Lepidoptera: Pieridae) and Trichoplusia ni Hübner (Lepidoptera: Noctuidae) large caterpillars and post feeding stages were reduced significantly by bird predation. The abundance of large caterpillars was also reduced on spider-inhabited plants during early plant growth; however the assemblage of birds and spiders did not suppress caterpillar densities more significantly than either predator alone.
4. Plants protected by birds, spiders, and birds plus spiders sustained less folivory attributable to leaf chewing caterpillars than check plants. Plant productivity was also greater for predator-protected plants than check plants.
5. Although spiders and parasitoids were responsible for some of the mortality inflicted upon lepidopteran caterpillars, it was concluded that in this study system, birds are the most important natural enemies of folivores.     

Resource-Mediated Indirect Effects of Grassland Management on Arthropod Diversity

Intensive land use is a driving force for biodiversity decline in many ecosystems. In semi-natural grasslands, land-use activities such as mowing, grazing and fertilization affect the diversity of plants and arthropods, but the combined effects of different drivers and the chain of effects are largely unknown. In this study we used structural equation modelling to analyse how the arthropod communities in managed grasslands respond to land use and whether these responses are mediated through changes in resource diversity or resource quantity (biomass). Plants were considered resources for herbivores which themselves were considered resources for predators. Plant and arthropod (herbivores and predators) communities were sampled on 141 meadows, pastures and mown pastures within three regions in Germany in 2008 and 2009. Increasing land-use intensity generally increased plant biomass and decreased plant diversity, mainly through increasing fertilization. Herbivore diversity decreased together with plant diversity but showed no response to changes in plant biomass. Hence, land-use effects on herbivore diversity were mediated through resource diversity rather than quantity. Land-use effects on predator diversity were mediated by both herbivore diversity (resource diversity) and herbivore quantity (herbivore biomass), but indirect effects through resource quantity were stronger. Our findings highlight the importance of assessing both direct and indirect effects of land-use intensity and mode on different trophic levels. In addition to the overall effects, there were subtle differences between the different regions, pointing to the importance of regional land-use specificities. Our study underlines the commonly observed strong effect of grassland land use on biodiversity. It also highlights that mechanistic approaches help us to understand how different land-use modes affect biodiversity.     

Dietary shift and lowered biomass gain of a generalist herbivore in species-poor experimental plant communities

Species loss of primary producers is likely to affect processes on other trophic levels. We studied consumption and individual performance of the generalist herbivore Parapleurus alliaceus (Orthoptera) in relation to the species richness of primary producers. Adult grasshoppers were caged and left to feed for 2 weeks on experimental grassland communities ranging in plant species richness from one, two, four, eight to 32 species. Low plant diversity had a negative effect on both plant community biomass and on biomass gain of female grasshoppers, feeding to produce eggs (male grasshoppers did not gain biomass during the feeding period). This was surprising because plots with high plant diversity had a low proportion of grass biomass and grasshoppers preferentially selected grasses, leading to a greater exploitation of grasses in experimental communities of higher diversity. Thus, the concurrent increase in non-grass species in the diet from these high-diversity communities must have been beneficial to the generalist herbivore. In addition to the positive effects of plant diversity, the presence of legumes in a mixture with grasses further enhanced the biomass gain of grasshoppers at a given level of diversity. These findings suggest that plant species loss may lead to shifts in herbivore population sizes, reducing those of generalists and benefiting specialists of the remaining plant species. Our results further suggest that generalist herbivores, by having feeding preferences, can also change the relative abundances of plant species with different functional characteristics. This may feedback on both composition and diversity of plant communities.     

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.     

Plant Diversity Impacts Decomposition and Herbivory via Changes in Aboveground Arthropods

Loss of plant diversity influences essential ecosystem processes as aboveground productivity, and can have cascading effects on the arthropod communities in adjacent trophic levels. However, few studies have examined how those changes in arthropod communities can have additional impacts on ecosystem processes caused by them (e.g. pollination, bioturbation, predation, decomposition, herbivory). Therefore, including arthropod effects in predictions of the impact of plant diversity loss on such ecosystem processes is an important but little studied piece of information. In a grassland biodiversity experiment, we addressed this gap by assessing aboveground decomposer and herbivore communities and linking their abundance and diversity to rates of decomposition and herbivory. Path analyses showed that increasing plant diversity led to higher abundance and diversity of decomposing arthropods through higher plant biomass. Higher species richness of decomposers, in turn, enhanced decomposition. Similarly, species-rich plant communities hosted a higher abundance and diversity of herbivores through elevated plant biomass and C:N ratio, leading to higher herbivory rates. Integrating trophic interactions into the study of biodiversity effects is required to understand the multiple pathways by which biodiversity affects ecosystem functioning.     

Insects affect relationships between plant species richness and ecosystem processes

This study examined whether insects can alter relationships between plant species diversity and ecosystem function in grassland communities, by (i) altering biomass across a plant diversity gradient, (ii) altering relative abundances of plant species, or (iii) altering ecosystem function directly. We measured herbivore damage on seminatural grassland plots planted with 1, 2, 4, 8, or 12 plant species, and compared plant biomass in a subset of these plots with replicates in which insect levels were reduced. Plant biomass and herbivore damage increased with species richness. Reducing insect populations resulted in greater evenness of relative plant species abundances and revealed a strong positive relationship between plant species richness and above-ground biomass. Reducing insects also changed the relationship between plant species richness and decomposition. Plant species mixtures and their relative abundances partially explained plant biomass results, but not decomposition results. These results suggest that insects can alter relationships between plant diversity and ecosystem processes through all three mechanisms.     

Does insect herbivory on oak depend on the diversity of tree stands?

Studies on the effects of plant diversity on insect herbivory have produced conflicting results. Plant diversity has been reported to cause positive and negative responses of herbivores. Explanations for these conflicting responses include not only various population-level processes but also changes in plant quality that lead to changes in herbivore performance. In a tree diversity experiment, we investigated the effects of tree diversity on insect herbivory on oak in general and whether the effects of tree diversity on herbivore damage are reflected by the performance (leaf consumption, growth) of the generalist herbivore Lymantria dispar. Our study showed that the feeding damage caused by naturally occurring herbivores on oak trees decreased with increasing diversity of tree stands. The performance of L. dispar on oak leaves was not affected by tree diversity, neither in field nor laboratory experiments. Our results can be explained by the various processes behind the hypothesis of associational resistance.     

Community- Weighted Mean Plant Traits Predict Small Scale Distribution of Insect Root Herbivore Abundance

Small scale distribution of insect root herbivores may promote plant species diversity by creating patches of different herbivore pressure. However, determinants of small scale distribution of insect root herbivores, and impact of land use intensity on their small scale distribution are largely unknown. We sampled insect root herbivores and measured vegetation parameters and soil water content along transects in grasslands of different management intensity in three regions in Germany. We calculated community-weighted mean plant traits to test whether the functional plant community composition determines the small scale distribution of insect root herbivores. To analyze spatial patterns in plant species and trait composition and insect root herbivore abundance we computed Mantel correlograms. Insect root herbivores mainly comprised click beetle (Coleoptera, Elateridae) larvae (43%) in the investigated grasslands. Total insect root herbivore numbers were positively related to community-weighted mean traits indicating high plant growth rates and biomass (specific leaf area, reproductive- and vegetative plant height), and negatively related to plant traits indicating poor tissue quality (leaf C/N ratio). Generalist Elaterid larvae, when analyzed independently, were also positively related to high plant growth rates and furthermore to root dry mass, but were not related to tissue quality. Insect root herbivore numbers were not related to plant cover, plant species richness and soil water content. Plant species composition and to a lesser extent plant trait composition displayed spatial autocorrelation, which was not influenced by land use intensity. Insect root herbivore abundance was not spatially autocorrelated. We conclude that in semi-natural grasslands with a high share of generalist insect root herbivores, insect root herbivores affiliate with large, fast growing plants, presumably because of availability of high quantities of food. Affiliation of insect root herbivores with large, fast growing plants may counteract dominance of those species, thus promoting plant diversity.