Host functional and phylogenetic composition rather than host diversity structure plant–herbivore networks

Declining plant diversity alters ecological networks, such as plant–herbivore interactions. However, our knowledge of the potential mechanisms underlying effects of plant species loss on plant–herbivore network structure is still limited. We used DNA barcoding to identify herbivore–host plant associations along declining levels of tree diversity in a large‐scale, subtropical biodiversity experiment. We tested for effects of tree species richness, host functional and phylogenetic diversity, and host functional (leaf trait) and phylogenetic composition on species, phylogenetic and network composition of herbivore communities. We found that phylogenetic host composition and related palatability/defence traits but not tree species richness significantly affected herbivore communities and interaction network complexity at both the species and community levels. Our study indicates that evolutionary dependencies and functional traits of host plants determine the composition of higher trophic levels and corresponding interaction networks in species‐rich ecosystems. Our findings highlight that characteristics of the species lost have effects on ecosystem structure and functioning across trophic levels that cannot be predicted from mere reductions in species richness.     

Host‐plant genotypic diversity and community genetic interactions mediate aphid spatial distribution

Genetic variation in plants can influence the community structure of associated species, through both direct and indirect interactions. Herbivorous insects are known to feed on a restricted range of plants, and herbivore preference and performance can vary among host plants within a species due to genetically based traits of the plant (e.g., defensive compounds). In a natural system, we expect to find genetic variation within both plant and herbivore communities and we expect this variation to influence species interactions. Using a three‐species plant‐aphid model system, we investigated the effect of genetic diversity on genetic interactions among the community members. Our system involved a host plant (Hordeum vulgare) that was shared by an aphid (Sitobion avenae) and a hemi‐parasitic plant (Rhinanthus minor). We showed that aphids cluster more tightly in a genetically diverse host‐plant community than in a genetic monoculture, with host‐plant genetic diversity explaining up to 24% of the variation in aphid distribution. This is driven by differing preferences of the aphids to the different plant genotypes and their resulting performance on these plants. Within the two host‐plant diversity levels, aphid spatial distribution was influenced by an interaction among the aphid's own genotype, the genotype of a competing aphid, the origin of the parasitic plant population, and the host‐plant genotype. Thus, the overall outcome involves both direct (i.e., host plant to aphid) and indirect (i.e., parasitic plant to aphid) interactions across all these species. These results show that a complex genetic environment influences the distribution of herbivores among host plants. Thus, in genetically diverse systems, interspecific genetic interactions between the host plant and herbivore can influence the population dynamics of the system and could also structure local communities. We suggest that direct and indirect genotypic interactions among species can influence community structure and processes.     

Herbivore community promotes trait evolution in a leaf beetle via induced plant response

Several recent studies have emphasised that community composition alters species trait evolution. Here, we demonstrate that differences in composition of local herbivore communities lead to divergent trait evolution of the leaf beetle Plagiodera versicolora through plant‐mediated indirect interactions. Our field surveys, genetic analyses and community‐manipulation experiments show that herbivore community composition determines the degree of herbivore‐induced regrowth of willows (Salicaceae), which in turn, promotes the divergent evolution of feeding preference in the leaf beetle from exclusive preference for new leaves to a lack of preference among leaf‐age types. Regrowth intensity depends both on the differential response of willows to different herbivore species and the integration of those herbivore species in the community. Because herbivore‐induced regrowth involves phenological changes in new leaf production, leaf beetle populations develop divergent feeding preferences according to local regrowth intensity. Therefore, herbivore community composition shapes the selection regime for leaf beetle evolution through trait‐mediated indirect interactions.     

Consequences of variation in plant defense for biodiversity at higher trophic levels

Antagonistic interactions between insect herbivores and plants impose selection on plants to defend themselves against these attackers. Although selection on plant defense traits has typically been studied for pairwise plant-attacker interactions, other community members of plant-based food webs are unavoidably affected by these traits as well. A plant trait might, for example, affect parasitoids and predators feeding on the herbivore. Consequently, defensive plant traits structure the diversity and composition of the complex community associated with the plant, and communities as a whole also feed back to selection on plant traits. Here, we review recent developments in our understanding of how plant defense traits structure insect communities and discuss how molecular mechanisms might drive community-wide effects.     

Effects of phylogeny,leaf traits,and the altitudinal distribution of host plants on herbivore assemblages on congeneric Acer species

Historical, niche-based, and stochastic processes have been proposed as the mechanisms that drive community assembly. In plant–herbivore systems, these processes can correspond to phylogeny, leaf traits, and the distribution of host plants, respectively. Although patterns of herbivore assemblages among plant species have been repeatedly examined, the effects of these factors among co-occurring congeneric host plant species have rarely been studied. Our aim was to reveal the process of community assembly for herbivores by investigating the effects of phylogeny, leaf traits, and the altitudinal distribution of closely related host plants of the genus Acer. We sampled leaf functional traits for 30 Acer species in Japan. Using a newly constructed phylogeny, we determined that three of the six measured leaf traits (leaf thickness, C/N ratio, and condensed tannin content) showed a phylogenetic signal. In a field study, we sampled herbivore communities on 14 Acer species within an elevation gradient and examined relationships between herbivore assemblages and host plants. We found that herbivore assemblages were significantly correlated with phylogeny, leaf traits, phylogenetic signals, and the altitudinal distribution of host plants. Our results indicate that the interaction between historical and current ecological processes shapes herbivore community assemblages.     

Differential effects of host plant hybridization on herbivore community structure and grazing pressure on forest canopies

Interspecific hybridization in plants is known to have ecological effects on associated organisms. We examined the differences in insect herbivore community structure and grazing pressure on tree canopy leaves among natural hybrids and their parental oak species. We measured leaf traits, herbivore community structure, and grazing pressure on leaves of two oak species, Quercus crispula and Q. dentata, and their hybrids. The concentration of nitrogen in canopy leaves was greater in hybrids and in Q. dentata than in Q. crispula. The concentration of total phenolics was lower in hybrids than in Q. crispula. The concentration of condensed tannin was greater in hybrids than in Q. crispula. Relative herbivore abundance and species richness were greater on oak hybrids than on either parental species; herbivore species diversity and composition on hybrids were close to those on Q. crispula. Herbivore grazing pressure was lower on hybrids and Q. dentata than on Q. crispula. There was a negative correlation between herbivore grazing pressure and leaf nitrogen, suggesting that interspecific variation among oak taxa in herbivore pressure may be explained by leaf nitrogen; variation in herbivore community structure among oak taxa is likely to be controlled by polygenic leaf traits. Differing responses of (1) herbivore community structure and (2) herbivore grazing pressure to host plant hybridization may play important roles in regulating herbivore biodiversity in cool‐temperate forest canopies.     

Population variation in plant traits associated with ant attraction and herbivory in Chamaecrista fasciculata (Fabaceae)

The benefits of ant–plant–herbivore interactions for the plant depend on the abundance of ants and herbivores and the selective pressures these arthropods exert. In plants bearing extrafloral nectaries (EFN), different mean trait values may be selected for by different populations in response to local herbivore pressure, ultimately resulting in the evolution of differences in plant traits that attract ants as defensive agents against herbivory. To determine if variation in traits that mediate ant–plant interactions reflect herbivore selective pressures, we quantified intra- and inter-population variation in plant traits for eight populations of the EFN-bearing annual Chamaecrista fasciculata (Michx.) (Fabaceae). Censuses in rural and urban areas of Missouri and Illinois (USA) showed population differences in ant attendance and herbivore pressure. Seeds were collected from each population, and plants were grown in a common greenhouse environment to measure sugar production, nectar volume and composition, EFN size and time of emergence, leaf pubescence, and leaf quality throughout plant development. Populations varied mainly in terms of nectary size, sugar production, and nectar volume, but to a lesser degree in leaf pubescence. Populations of C. fasciculata within urban areas (low in insect abundance) had small nectaries and the lowest nectar production. There was a positive correlation across populations between herbivore density and leaf damage by those herbivores on the one hand and sugar production and nectar volume on the other. These results, in conjunction with lack of evidence for maternally based environmental effects, suggest that population differences in herbivore damage have promoted differential evolution of EFN-related traits among populations. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Inbreeding in horsenettle influences herbivore resistance

Abstract 1. Plant traits (e.g. nutrition, allelochemistry) are an important determinant of the feeding preferences and performance of insect herbivores. Recent evidence suggests that plant inbreeding can affect plant–insect interactions by impacting host‐plant quality and resistance to herbivory. 2. The effect of inbreeding on host‐plant quality for, and resistance against, the tobacco hornworm, Manduca sexta L., was assessed in the wild solanaceous weed horsenettle, Solanum carolinense L. Caterpillar preference, relative growth rate (RGR), total leaf consumption (TC), and per cent total nitrogen in leaves were examined using selfed and outcrossed progeny of eight maternal plants. 3. Inbreeding significantly influenced insect preference, with caterpillars preferring leaf discs from selfed versus outcrossed plants. There was also a breeding effect for RGR and TC, with both higher on selfed plants. No breeding effect for per cent total nitrogen was observed. 4. The results of this study indicate that inbreeding decreased resistance against the tobacco hornworm, but did not affect plant quality. Decreased plant resistance will likely alter interactions with the herbivore community and could also have important consequences for plant–herbivore–natural enemy interactions.     

Eco-evolutionary dynamics in herbivorous insect communities mediated by induced plant responses

It is increasingly recognized that the ecology of communities and evolution of species within communities are interdependent, and researchers have been paying attention to this rapidly emerging field of research, i.e., through studies on eco-evolutionary dynamics. Most of the studies on eco-evolutionary dynamics have been concerned with direct trophic interactions. However, community ecologists have shown that trait-mediated indirect effects play an important role in shaping the structure of natural communities. In particular, in terrestrial plant–insect systems, indirect effects mediated through herbivore-induced plant responses are common and have a great impact on the structure of herbivore communities. This review describes eco-evolutionary dynamics in herbivorous insect communities, and specifically focuses on the key role of herbivore-induced plant responses in eco-evolutionary dynamics. First, I review studies on the evolution of herbivore traits relevant to plant induction and discuss evolution in a community context mediated by induced plant responses. Second, I highlight how intraspecific genetic variation or evolution in herbivore traits can influence herbivore community structure. Finally, I propose the hypothetical model that induced plant responses supports eco-evolutionary feedback in herbivore communities. In this review, I argue that the application of the indirect interaction web approaches into studies on eco-evolutionary will provide profound insights into understanding of mechanisms of the generation and maintenance of biodiversity.     

Plant phylogeny drives arboreal caterpillar assemblages across the Holarctic

1. Assemblages of insect herbivores are structured by plant traits such as nutrient content, secondary metabolites, physical traits, and phenology. Many of these traits are phylogenetically conserved, implying a decrease in trait similarity with increasing phylogenetic distance of the host plant taxa. Thus, a metric of phylogenetic distances and relationships can be considered a proxy for phylogenetically conserved plant traits and used to predict variation in herbivorous insect assemblages among co‐occurring plant species.
2. Using a Holarctic dataset of exposed‐feeding and shelter‐building caterpillars, we aimed at showing how phylogenetic relationships among host plants explain compositional changes and characteristics of herbivore assemblages.
3. Our plant–caterpillar network data derived from plot‐based samplings at three different continents included >28,000 individual caterpillar–plant interactions. We tested whether increasing phylogenetic distance of the host plants leads to a decrease in caterpillar assemblage overlap. We further investigated to what degree phylogenetic isolation of a host tree species within the local community explains abundance, density, richness, and mean specialization of its associated caterpillar assemblage.
4. The overlap of caterpillar assemblages decreased with increasing phylogenetic distance among the host tree species. Phylogenetic isolation of a host plant within the local plant community was correlated with lower richness and mean specialization of the associated caterpillar assemblages. Phylogenetic isolation had no effect on caterpillar abundance or density. The effects of plant phylogeny were consistent across exposed‐feeding and shelter‐building caterpillars.
5. Our study reveals that distance metrics obtained from host plant phylogeny are useful predictors to explain compositional turnover among hosts and host‐specific variations in richness and mean specialization of associated insect herbivore assemblages in temperate broadleaf forests. As phylogenetic information of plant communities is becoming increasingly available, further large‐scale studies are needed to investigate to what degree plant phylogeny structures herbivore assemblages in other biomes and ecosystems.

     

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.     

桂林岩溶石山青冈群落植物功能性状的尺度变化与关联

研究植物功能性状在不同尺度的变异和关联,对于揭示植物对环境的适应策略和群落构建规律具有重要意义。以岩溶石山青冈群落为研究对象,测量了研究区内20个样方74种木本植物的叶面积、比叶面积和木材密度3个功能性状值,利用性状梯度分析法分析了3个性状在群落内部(α组分)及群落间(β组分)的变异格局及相关性。结果表明:(1)群落内3个植物功能性状的α值范围均大于β值范围,即物种相对于共生物种性状值的变化大于沿着群落平均性状梯度的变化。(2)植物功能性状比叶面积的种内差异引起的变化小于群落水平。(3)叶面积与比叶面积、比叶面积与木材密度、叶面积与木材密度的β组分相关性均最强,而α组分间无相关性或相关性较弱,即叶面积与比叶面积、比叶面积与木材密度、叶面积与木材密度两两性状间的相关性在群落间的依赖程度比群落内共生物种的依赖性要强,暗示物种在群落内和群落间采取不同的生态策略来适应环境。     

Linking plant genes to insect communities: Identifying the genetic bases of plant traits and community composition

Community genetics aims to understand the effects of intraspecific genetic variation on community composition and diversity, thereby connecting community ecology with evolutionary biology. Thus far, research has shown that plant genetics can underlie variation in the composition of associated communities (e.g., insects, lichen and endophytes), and those communities can therefore be considered as extended phenotypes. This work, however, has been conducted primarily at the plant genotype level and has not identified the key underlying genes. To address this gap, we used genome‐wide association mapping with a population of 445 aspen (Populus tremuloides) genets to identify the genes governing variation in plant traits (defence chemistry, bud phenology, leaf morphology, growth) and insect community composition. We found 49 significant SNP associations in 13 Populus genes that are correlated with chemical defence compounds and insect community traits. Most notably, we identified an early nodulin‐like protein that was associated with insect community diversity and the abundance of interacting foundation species (ants and aphids). These findings support the concept that particular plant traits are the mechanistic link between plant genes and the composition of associated insect communities. In putting the “genes” into “genes to ecosystems ecology”, this work enhances understanding of the molecular genetic mechanisms that underlie plant–insect associations and the consequences thereof for the structure of ecological communities.     

Community Functional Responses to Soil and Climate at Multiple Spatial Scales: When Does Intraspecific Variation Matter?

Despite increasing evidence of the importance of intraspecific trait variation in plant communities, its role in community trait responses to environmental variation, particularly along broad-scale climatic gradients, is poorly understood. We analyzed functional trait variation among early-successional herbaceous plant communities (old fields) across a 1200-km latitudinal extent in eastern North America, focusing on four traits: vegetative height, leaf area, specific leaf area (SLA), and leaf dry matter content (LDMC). We determined the contributions of species turnover and intraspecific variation to between-site functional dissimilarity at multiple spatial scales and community trait responses to edaphic and climatic factors. Among-site variation in community mean trait values and community trait responses to the environment were generated by a combination of species turnover and intraspecific variation, with species turnover making a greater contribution for all traits. The relative importance of intraspecific variation decreased with increasing geographic and environmental distance between sites for SLA and leaf area. Intraspecific variation was most important for responses of vegetative height and responses to edaphic compared to climatic factors. Individual species displayed strong trait responses to environmental factors in many cases, but these responses were highly variable among species and did not usually scale up to the community level. These findings provide new insights into the role of intraspecific trait variation in plant communities and the factors controlling its relative importance. The contribution of intraspecific variation to community trait responses was greatest at fine spatial scales and along edaphic gradients, while species turnover dominated at broad spatial scales and along climatic gradients.     

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.     

Community-level relaxation of plant defenses against herbivores at high elevation

Ecological gradients shape the evolution of plant functional traits through variation in climate, abiotic factors, and biological processes. While recent observations highlight elevation clines in functional trait variation, several traits vary independently of others and vary in species-specific manner, thereby hindering community-level generalization. In this study, we examined whether whole plant communities at different elevations independently converge to elevation-specific trait values. Using a combination of field-derived trait quantification and a literature survey, we analyzed eight functional traits relating to plant palatability [specific leaf area (SLA), leaf dry matter content, carbon-to-nitrogen ratio], and defenses (spines, trichomes, silica, and the relative importance of phenolic- or non-phenolic-based chemical defenses in plants) of plants growing along several mountain transects in Switzerland, and computed community-level means for each trait. We observed a general decline of most defense traits in communities at high elevations. The individual traits’ patterns underlined a concerted decline of the physico-chemical defense syndrome at high elevation, and low SLA values at high elevation also indicate investment in the production of leaves with slow turnover rates. In addition, we found that high-elevation plant trait values tend to converge, while at low elevation, to diverge within communities. Our data suggest that community-level defense syndromes along elevation gradients are influenced by ecological filtering. Lower herbivore pressures select for reduced plant defenses, while harsh environmental conditions select for investment in tolerance-related traits. Secondly, since species are highly variable in their functional trait space within communities, niche-partitioning theory might explain which variabilities of the defense functional space are selected within communities, particularly at low elevation.     

Importance of species interactions to community heritability: a genetic basis to trophic-level interactions

Recent community genetics studies have shown that specific genotypes of a host plant support distinct arthropod communities. Building upon these findings, we examined the hypothesis that a trophic community consisting of cottonwood trees, a galling herbivore and avian predators could also be related to the genetics of the host tree. We found genetic correlations among phytochemistry of individual tree genotypes, the density of a galling herbivore, and the intensity of avian predation on these herbivores. We detected significant broad-sense heritability of these interactions that range from H      = 0.70 to 0.83. The genetic basis of these interactions tended to increase across trophic levels suggesting that small genetic changes in the cottonwood phenotype could have major consequences at higher trophic levels affecting species interactions and energy flow. These findings show a heritable basis to trophic-level interactions indicating that there is a significant genetic basis to community composition and energy flow that is predictable by plant genotype. Our data clearly link plant genetics to patterns of avian foraging and show that species interactions are important components of community heritability and ecosystem processes. Overall, these data support the hypothesis that evolution of plant traits can alter trophic-level interactions and community composition.     

Evolution in plant populations as a driver of ecological changes in arthropod communities

Heritable variation in traits can have wide-ranging impacts on species interactions, but the effects that ongoing evolution has on the temporal ecological dynamics of communities are not well understood. Here, we identify three conditions that, if experimentally satisfied, support the hypothesis that evolution by natural selection can drive ecological changes in communities. These conditions are: (i) a focal population exhibits genetic variation in a trait(s), (ii) there is measurable directional selection on the trait(s), and (iii) the trait(s) under selection affects variation in a community variable(s). When these conditions are met, we expect evolution by natural selection to cause ecological changes in the community. We tested these conditions in a field experiment examining the interactions between a native plant (Oenothera biennis) and its associated arthropod community (more than 90 spp.). Oenothera biennis exhibited genetic variation in several plant traits and there was directional selection on plant biomass, life-history strategy (annual versus biennial reproduction) and herbivore resistance. Genetically based variation in biomass and life-history strategy consistently affected the abundance of common arthropod species, total arthropod abundance and arthropod species richness. Using two modelling approaches, we show that evolution by natural selection in large O. biennis populations is predicted to cause changes in the abundance of individual arthropod species, increases in the total abundance of arthropods and a decline in the number of arthropod species. In small O. biennis populations, genetic drift is predicted to swamp out the effects of selection, making the evolution of plant populations unpredictable. In short, evolution by natural selection can play an important role in affecting the dynamics of communities, but these effects depend on several ecological factors. The framework presented here is general and can be applied to other systems to examine the community-level effects of ongoing evolution.     

The influence of range‐wide plant genetic variation on soil invertebrate communities

Plant genetic variation can have far‐reaching effects on associated communities and ecosystems. Heritable variation in ecologically relevant plant traits is often non‐randomly distributed across a species’ range and can exhibit geographic clines. In the event of range expansions and migration, previously unfamiliar genotypes may have large impacts on resident communities and ecosystems due to the introduction of novel and heritable phenotypic variation. Here we test the hypothesis that geographic origin of a focal plant genotype has effects on belowground invertebrate communities using a common garden field experiment. We sampled soil invertebrates from 103 Oenothera biennis genotypes, which were collected from across the species’ range and planted into a common garden field experiment at the northern range limit. We enumerated 24 000 individuals from 190 morphospecies and found that the diversity, abundance, and composition of soil invertebrate communities varied greatly among plant genotypes. Despite strong effects of plant genotype, we found few genetic correlations between plant traits and soil invertebrate community variables. However, herbivore damage was strongly related to variation in the soil invertebrate community. Geographic origin of plant genotypes had at most a weak effect on belowground communities. We speculate that predicting the extended effects of population movement on associated communities will require detailed knowledge of the trait variation occurring within focal species across particular environmental gradients.     

Plant genetics shapes inquiline community structure across spatial scales

Recent research in community genetics has examined the effects of intraspecific genetic variation on species diversity in local communities. However, communities can be structured by a combination of both local and regional processes and to date, few community genetics studies have examined whether the effects of instraspecific genetic variation are consistent across levels of diversity. In this study, we ask whether host-plant genetic variation structures communities of arthropod inquilines within distinct habitat patches – rosette leaf galls on tall goldenrod ( Solidago altissima ). We found that genetic variation determined inquiline diversity at both local and regional spatial scales, but that trophic-level responses varied independently of one another. This result suggests that herbivores and predators likely respond to heritable plant traits at different spatial scales. Together, our results show that incorporating spatial scale is essential for predicting the effects of genetically variable traits on different trophic levels and levels of diversity within the communities that depend on host plants.