Plant Species Loss Affects Life-History Traits of Aphids and Their Parasitoids

The consequences of plant species loss are rarely assessed in a multi-trophic context and especially effects on life-history traits of organisms at higher trophic levels have remained largely unstudied. We used a grassland biodiversity experiment and measured the effects of two components of plant diversity, plant species richness and the presence of nitrogen-fixing legumes, on several life-history traits of naturally colonizing aphids and their primary and secondary parasitoids in the field. We found that, irrespective of aphid species identity, the proportion of winged aphid morphs decreased with increasing plant species richness, which was correlated with decreasing host plant biomass. Similarly, emergence proportions of parasitoids decreased with increasing plant species richness. Both, emergence proportions and proportions of female parasitoids were lower in plots with legumes, where host plants had increased nitrogen concentrations. This effect of legume presence could indicate that aphids were better defended against parasitoids in high-nitrogen environments. Body mass of emerged individuals of the two most abundant primary parasitoid species was, however, higher in plots with legumes, suggesting that once parasitoids could overcome aphid defenses, they could profit from larger or more nutritious hosts. Our study demonstrates that cascading effects of plant species loss on higher trophic levels such as aphids, parasitoids and secondary parasitoids begin with changed life-history traits of these insects. Thus, life-history traits of organisms at higher trophic levels may be useful indicators of bottom-up effects of plant diversity on the biodiversity of consumers.     

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.     

Effects of plant sex on insect abundance across three trophic levels in the perennial shrub Buddleja cordata

Although there is a growing interest in the effects of intra‐specific plant genetic variation on species interactions, the effects of plant sex, an important axis of genetic variation, have been less studied. In addition, previous work investigating plant sex effects on species interactions has frequently focused on bitrophic interactions (e.g., herbivory), usually ignoring plant sex effects on higher trophic levels (i.e., natural enemies). Here, we investigated the effects of plant sex on herbivore abundance and that of their natural enemies associated with the dioecious shrub Buddleja cordata Kunth (Scrophulariaceae). Furthermore, we measured a subset of plant traits frequently involved in herbivore resistance and the potentially underlying plant sex effects. To this end, we recorded the abundances of a specialist leaf‐chewing caterpillar [Acronyctodes mexicanaria Walker (Lepidoptera: Geometridae)] throughout an entire growing season. We also recorded information about the caterpillar’s parasitoids, as well as leaf water content, phenolic compounds, phosphorus, and nitrogen for male and female plants of B. cordata. Plant sex did not significantly influence caterpillar abundance but did have an effect on natural enemies, with parasitoid abundance being 2.4‐fold greater on female than on male plants. The effect of plant sex on parasitoids remained significant after accounting for caterpillar abundance, suggesting that it was underlain by a trait‐mediated (rather than density‐mediated) mechanism. Finally, we found that male plants had a higher concentration of phenolic compounds (other traits did not differ between plant sexes). These results provide valuable evidence for the extended effects of plant sex on the third trophic level and point at plant traits potentially mediating such effects.     

Quantifying the impact of above‐ and belowground higher trophic levels on plant and herbivore performance by modeling1

Growing empirical evidence suggests that aboveground and belowground multitrophic communities interact. However, investigations that comprehensively explore the impacts of above‐ and belowground third and higher trophic level organisms on plant and herbivore performance are thus far lacking. We tested the hypotheses that above‐ and belowground higher trophic level organisms as well as decomposers affect plant and herbivore performance and that these effects cross the soil–surface boundary. We used a well‐validated simulation model that is individual‐based for aboveground trophic levels such as shoot herbivores, parasitoids, and hyperparasitoids while considering belowground herbivores and their antagonists at the population level. We simulated greenhouse experiments by removing trophic levels and decomposers from the simulations in a factorial design. Decomposers and above‐ and belowground third trophic levels affected plant and herbivore mortality, root biomass, and to a lesser extent shoot biomass. We also tested the effect of gradual modifications of the interactions between different trophic level organisms with a sensitivity analysis. Shoot and root biomass were highly sensitive to the impact of the fourth trophic level. We found effects that cross the soil surface, such as aboveground herbivores and parasitoids affecting root biomass and belowground herbivores influencing aboveground herbivore mortality. We conclude that higher trophic level organisms and decomposers can strongly influence plant and herbivore performance. We propose that our modelling framework can be used in future applications to quantitatively explore the possible outcomes of complex above‐ and belowground multitrophic interactions under a range of environmental conditions and species compositions.     

Tree diversity and functional leaf traits drive herbivore‐associated microbiomes in subtropical China

Herbivorous insects acquire microorganisms from host plants or soil, but it remains unclear how the diversity and functional composition of host plants contribute to structuring herbivore microbiomes. Within a controlled tree diversity setting, we used DNA metabarcoding of 16S rRNA to assess the contribution of Lepidoptera species and their local environment (particularly, tree diversity, host tree species, and leaf traits) to the composition of associated bacterial communities. In total, we obtained 7,909 bacterial OTUs from 634 caterpillar individuals comprising 146 species. Tree diversity was found to drive the diversity of caterpillar‐associated bacteria both directly and indirectly via effects on caterpillar communities, and tree diversity was a stronger predictor of bacterial diversity than diversity of caterpillars. Leaf toughness and dry matter content were important traits of the host plant determining bacterial species composition, while leaf calcium and potassium concentration influenced bacterial richness. Our study reveals previously unknown linkages between trees and their characteristics, herbivore insects, and their associated microbes, which contributes to developing a more nuanced understanding of functional dependencies between herbivores and their environment, and has implications for the consequences of plant diversity loss for trophic interactions.     

Multitrophic effects of experimental changes in plant diversity on cavity-nesting bees, wasps, and their parasitoids

Plant diversity changes can impact the abundance, diversity, and functioning of species at higher trophic levels. We used an experimental gradient in grassland plant diversity ranging from 1 to 16 plant species to study multitrophic interactions among plants, cavity-nesting bees and wasps, and their natural enemies, and analysed brood cell density, insect diversity (species richness), and bee and wasp community similarity over two consecutive years. The bee and wasp communities were more similar among the high (16 species) diversity plots than among plots of the lower diversity levels (up to 8 species), and a more similar community of bees and wasps resulted in a more similar community of their parasitoids. Plant diversity, which was closely related to flower diversity, positively and indirectly affected bee diversity and the diversity of their parasitoids via increasing brood cell density of bees. Increasing plant diversity directly led to higher wasp diversity. Parasitism rates of bees and wasps (hosts) were not affected by plant diversity, but increased with the diversity of their respective parasitoids. Decreases in parasitism rates of bees arose from increasing brood cell density of bees (hosts), whereas decreasing parasitism rates of wasps arose from increasing wasp diversity (hosts). In conclusion, decreases in plant diversity propagated through different trophic levels: from plants to insect hosts to their parasitoids, decreasing density and diversity. The positive relationship between plant diversity and the community similarity of higher trophic levels indicates a community-stabilising effect of high plant diversity.     

Describing a multitrophic plant-herbivore-parasitoid system at four spatial scales

Herbivore-parasitoid interactions must be studied using a multitrophic and multispecies approach. The strength and direction of multiple effects through trophic levels may change across spatial scales. In this work, we use the herbaceous plant Ruellia nudiflora, its moth herbivore Tripudia quadrifera, and several parasitoid morphospecies that feed on the herbivore to answer the following questions: Do herbivore and parasitoid attack levels vary depending on the spatial scale considered? With which plant characteristics are the parasitoid and the herbivore associated? Do parasitoid morphospecies vary in the magnitude of their positive indirect effect on plant reproduction? We evaluated three approximations of herbivore and parasitoid abundance (raw numbers, ratios, and attack rates) at four spatial scales: regional (three different regions which differ in terms of abiotic and biotic characteristics); population (i.e. four populations within each region); patch (four 1 m² plots in each population); and plant level (using a number of plant characteristics). Finally, we determined whether parasitoids have a positive indirect effect on plant reproductive success (seed number). Herbivore and parasitoid numbers differed at three of the spatial scales considered. However, herbivore/fruit ratio and attack rates did not differ at the population level. Parasitoid/host ratio and attack rates did not differ at any scale, although there was a tendency of a higher attack in one region. At the plant level, herbivore and parasitoid abundances were related to different plant traits, varying the importance and the direction (positive or negative) of those traits. In addition, only one parasitoid species (Bracon sp.) had a positive effect on plant fitness saving up to 20% of the seeds in a fruit. These results underline the importance of knowing the scales that are relevant to organisms at different trophic levels and distinguish between the specific effects of species.     

Elevated air temperature alters an old-field insect community in a multifactor climate change experiment

To address how multiple, interacting climate drivers may affect plant–insect community associations, we sampled insects that naturally colonized a constructed old‐field plant community grown for over 2 years under simultaneous CO₂, temperature, and water manipulation. Insects were sampled using a combination of sticky traps and vacuum sampling, identified to morphospecies and the insect community with respect to abundance, richness, and evenness quantified. Individuals were assigned to four broad feeding guilds in order to examine potential trophic level effects. Although there were occasional effects of CO₂ and water treatment, the effects of warming on the insect community were large and consistent. Warming significantly increased Order Thysanoptera abundance and reduced overall morphospecies richness and evenness. Nonmetric multidimensional scaling found that only temperature affected insect community composition, while a Sørensen similarity index showed less correspondence in the insect community between temperature treatments compared with CO₂ or soil water treatments. Within the herbivore guild, elevated temperature significantly reduced richness and evenness. Corresponding reductions of diversity measures at higher trophic levels (i.e. parasitoids), along with the finding that herbivore richness was a significant predictor of parasitoid richness, suggest trophic‐level effects within the insect community. When the most abundant species were considered in temperature treatments, a small number of species increased in abundance at elevated temperature, while others declined compared with ambient temperature. Effects of temperature in the dominant insects demonstrated that treatment effects were limited to a relatively small number of morphospecies. Observed effects of elevated CO₂ concentration on whole‐community foliar N concentration did not result in any effect on herbivores, which are probably the most susceptible guild to changes in plant nutritional quality. These results demonstrate that climatic warming may alter certain insect communities via effects on insect species most responsive to a higher temperature, contributing to a change in community structure.     

Traits across trophic levels interact to influence parasitoid establishment in biological control releases

A central goal in ecology is to predict what governs a species’ ability to establish in a new environment. One mechanism driving establishment success is individual species’ traits, but the role of trait combinations among interacting species across different trophic levels is less clear. Deliberate or accidental species additions to existing communities provide opportunities to study larger scale patterns of establishment success. Biological control introductions are especially valuable because they contain data on both the successfully established and unestablished species. Here, we used a recent dataset of importation biological control introductions to explore how life‐history traits of 132 parasitoid species and their herbivorous hosts interact to affect parasitoid establishment. We find that of five parasitoid and herbivore traits investigated, one parasitoid trait—host range—weakly predicts parasitoid establishment; parasitoids with higher levels of phylogenetic specialization have higher establishment success, though the effect is marginal. In addition, parasitoids are more likely to establish when their herbivore host has had a shorter residence time. Interestingly, we do not corroborate earlier findings that gregarious parasitoids and endoparasitoids are more likely to establish. Most importantly, we find that life‐history traits of the parasitoid species and their hosts can interact to influence establishment. Specifically, parasitoids with broader host ranges are more likely to establish when the herbivore they have been released to control is also more of a generalist. These results provide insight into how multiple species’ traits and their interactions, both within and across trophic levels, can influence establishment of species of higher trophic levels.     

Plant diversity and the stability of foodwebs

Insect outbreaks in forest and agriculture monocultures led Charles Elton to propose, a half-century ago, that higher plant diversity stabilized animal foodweb dynamics in natural ecosystems. We tested this hypothesis by studying arthropod community dynamics in a long-term experimental manipulation of grassland plant species diversity. Over the course of a decade, we found that higher plant diversity increased the stability (i.e. lowered year-to-year variability) of a diverse (>700 species) arthropod community across trophic levels. As the number of plant species increased, the stability of both herbivore and predator species richness and of total herbivore abundance increased. The underlying mechanisms driving these diversity-stability relationships were plant diversity, via effects on primary productivity and plant community stability, and portfolio effects. Taken together, our results show that higher plant diversity provides more temporally consistent food and habitat resources to arthropod foodwebs. Consequently, actively managing for high plant diversity may have stronger than expected benefits for increasing animal diversity and controlling pest outbreaks.     

Herbivore species richness, composition and community structure mediate predator richness effects and top-down control of herbivore biomass

Changes in predator species richness can have important consequences for ecosystem functioning at multiple trophic levels, but these effects are variable and depend on the ecological context in addition to the properties of predators themselves. Here, we report an experimental study to test how species identity, community attributes, and community structure at the herbivore level moderate the effects of predator richness on ecosystem functioning. Using mesocosms containing predatory insects and aphid prey, we independently manipulated species richness at both predator and herbivore trophic levels. Community structure was also manipulated by changing the distribution of herbivore species across two plant species. Predator species richness and herbivore species richness were found to negatively interact to influence predator biomass accumulation, an effect which is hypothesised to be due to the breakdown of functional complementarity among predators in species-rich herbivore assemblages. The strength of predator suppression of herbivore biomass decreased as herbivore species richness and distribution across host plants increased, and positive predator richness effects on herbivore biomass suppression were only observed in herbivore assemblages of relatively low productivity. In summary, the study shows that the species richness, productivity and host plant distribution of prey communities can all moderate the general influence of predators and the emergence of predator species richness effects on ecosystem functioning.     

Effects of pepper (Capsicum chinense) genotypic diversity on insect herbivores

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Cascade effects of crop species richness on the diversity of pest insects and their natural enemies

Understanding how plant species richness influences the diversity of herbivorous and predatory/parasitic arthropods is central to community ecology.We explore the effects of crop species richness on the diversity of pest insects and their natural enemies.Using data from a four-year experiment with five levels of crop species richness,we found that crop species richness significantly affected the pest species richness,but there were no significant effects on richness of the pests’natural enemies.In contrast,the species richness of pest insects significantly affected their natural enemies.These findings suggest a cascade effect where trophic interactions are strong between adjacent trophic levels,while the interactions between connected but nonadjacent trophic levels are weakened by the intermediate trophic level.High crop species richness resulted in a more stable arthropod community compared with communities in monoculture crops.Our results highlight the complicated cross-trophic interactions and the crucial role of crop diversity in the food webs of agro-ecosystems.     

Patch size effects are more important than genetic diversity for plant–herbivore interactions in Brassica crops

1. Considerable evidence suggests that the diversity within plant communities may strongly affect the strength of species interactions, but the majority of studies only considered interspecific diversity. 2. This paper examines the effect of intraspecific genetic diversity within Brassica fields on two Brassica specialists, cabbage root fly, and diamondback moth, and on a parasitoid attacking diamondback moths. Genetic diversity was manipulated both in a replacement and an additive design. 3. Both herbivore densities and parasitism rates were higher in smaller plots, with limited responses to increased within‐plot diversity. All species showed variable densities across genotypes, and preference hierarchies were species specific. 4. Responses to plot size in root flies scaled with the diameter‐to‐area ratio, suggesting that patch detectability affected local density, whereas responses by diamondback moths and parasitoids deviated from this ratio. These species differences could be traced to differences in the residence time within patches, where diamondback moths typically spend longer and more variable time periods in patches than root flies. 5. The lack of response to genetic diversity by both herbivores suggests that egg‐laying rates are affected by decisions on the plant and not by attraction from a distance, neither to the plant itself nor the patch. Patterns of differential attack may then be due to different acceptability for studied genotypes. 6. Future theories on insect responses to spatial heterogeneity should focus on species traits and how traits interact with information landscapes in the field.     

Plant diversity induces shifts in the functional structure and diversity across trophic levels

Changes to primary producer diversity can cascade up to consumers and affect ecosystem processes. Although the effect of producer diversity on higher trophic groups have been studied, these studies often quantify taxonomy‐based measures of biodiversity, like species richness, which do not necessarily reflect the functioning of these communities. In this study, we assess how plant species richness affects the functional composition and diversity of higher trophic levels and discuss how this might affect ecosystem processes, such as herbivory, predation and decomposition. Based on six different consumer traits, we examined the functional composition of arthropod communities sampled in experimental plots that differed in plant species richness. The two components we focused on were functional variation in the consumer community structure (functional structure) and functional diversity, expressed as functional richness, evenness and divergence. We found a consistent positive effect of plant species richness on the functional richness of herbivores, carnivores, and omnivores, but not decomposers, and contrasting patterns for functional evenness and divergence. Increasing plant species richness shifted the omnivore community to more predatory and less mobile species, and the herbivore community to more specialized and smaller species. This was accompanied by a shift towards more species occurring in the vegetation than in the ground layer. Our study shows that plant species richness strongly affects the functional structure and diversity of aboveground arthropod communities. The observed shifts in body size (herbivores), specialization (herbivores), and feeding mode (omnivores) together with changes in the functional diversity may underlie previously observed increases in herbivory and predation in plant communities of higher diversity.     

Negative per capita effects of two invasive plants, Lythrum salicaria and Phalaris arundinacea, on the moth diversity of wetland communities

Invasive plants have been shown to negatively affect the diversity of plant communities. However, little is known about the effect of invasive plants on the diversity at other trophic levels. In this study, we examine the per capita effects of two invasive plants, purple loosestrife (Lythrum salicaria) and reed canary grass (Phalaris arundinacea), on moth diversity in wetland communities at 20 sites in the Pacific Northwest, USA. Prior studies document that increasing abundance of these two plant species decreases the diversity of plant communities. We predicted that this reduction in plant diversity would result in reduced herbivore diversity. Four measurements were used to quantify diversity: species richness (S), community evenness (J), Brillouin's index (H) and Simpson's index (D). We identified 162 plant species and 156 moth species across the 20 wetland sites. The number of moth species was positively correlated with the number of plant species. In addition, invasive plant abundance was negatively correlated with species richness of the moth community (linear relationship), and the effect was similar for both invasive plant species. However, no relationship was found between invasive plant abundance and the three other measures of moth diversity (J, H, D) which included moth abundance in their calculation. We conclude that species richness within, and among, trophic levels is adversely affected by these two invasive wetland plant species.     

Functional trait diversity across trophic levels determines herbivore impact on plant community biomass

Understanding the consequences of trophic interactions for ecosystem functioning is challenging, as contrasting effects of species and functional diversity can be expected across trophic levels. We experimentally manipulated functional identity and diversity of grassland insect herbivores and tested their impact on plant community biomass. Herbivore resource acquisition traits, i.e. mandible strength and the diversity of mandibular traits, had more important effects on plant biomass than body size. Higher herbivore functional diversity increased overall impact on plant biomass due to feeding niche complementarity. Higher plant functional diversity limited biomass pre‐emption by herbivores. The functional diversity within and across trophic levels therefore regulates the impact of functionally contrasting consumers on primary producers. By experimentally manipulating the functional diversity across trophic levels, our study illustrates how trait‐based approaches constitute a promising way to tackle existing links between trophic interactions and ecosystem functioning.     

Host-based genotype variation in insects revisited

Elucidation of the genetic variability of a model insect species, the grain aphid, Sitobion avenae (Fabricius), a predominantly asexual herbivore within the temperate agro-ecosystem tested, was initiated using molecular DNA markers (RAPDs). This revealed genetic profiles that appeared related to host adaptation at the specific level amongst the natural populations colonizing different grasses and cereals (Poaceae) within the same geographic location. These profiles were recorded either as 'specialist' genotypes found on specific grasses, or as 'generalist' genotypes colonizing several host types including cultivated cereals or native grasses. These findings are compared with analogous systems found amongst insect species, including at a higher trophic level, i.e. interactions between hymenopterous aphid parasitoids. As the aphids and their respective plant hosts occur in the same geographical region at the same time, this appears to be a rare example of the evolutionary transition leading to sympatric speciation in insects. Hence, this study highlights the importance of understanding not only the demographic parameters to genetic diversity, but also the more intricate correlation of genetic diversity to host types in agricultural environments.     

Herbivore species identity rather than diversity of the non‐host community determines foraging behaviour of the parasitoid wasp Cotesia glomerata

Extensive research has been conducted to reveal how species diversity affects ecosystem functions and services. Yet, consequences of diversity loss for ecosystems as a whole as well as for single community members are still difficult to predict. Arthropod communities typically are species‐rich, and their species interactions, such as those between herbivores and their predators or parasitoids, may be particularly sensitive to changes in community composition. Parasitoids forage for herbivorous hosts by using herbivore‐induced plant volatiles (indirect cues) and cues produced by their host (direct cues). However, in addition to hosts, non‐suitable herbivores are present in a parasitoid's environment which may complicate the foraging process for the parasitoid. Therefore, ecosystem changes in the diversity of herbivores may affect the foraging efficiency of parasitoids. The effect of herbivore diversity may be mediated by either species numbers per se, by specific species traits, or by both. To investigate how diversity and identity of non‐host herbivores influence the behaviour of parasitoids, we created environments with different levels of non‐host diversity. On individual plants in these environments, we complemented host herbivores with 1–4 non‐host herbivore species. We subsequently studied the behaviour of the gregarious endoparasitoid Cotesia glomerata L. (Hymenoptera: Braconidae) while foraging for its gregarious host Pieris brassicae L. (Lepidoptera: Pieridae). Neither non‐host species diversity nor non‐host identity influenced the preference of the parasitoid for herbivore‐infested plants. However, after landing on the plant, non‐host species identity did affect parasitoid behaviour, whereas non‐host diversity did not. One of the non‐host species, Trichoplusia ni Hübner (Lepidoptera: Noctuidae), reduced the time the parasitoid spent on the plant as well as the number of hosts it parasitized. We conclude that non‐host herbivore species identity has a larger influence on C. glomerata foraging behaviour than non‐host species diversity. Our study shows the importance of species identity over species diversity in a multitrophic interaction of plants, herbivores, and parasitoids.     

Extended larval development time for aphid parasitoids in the presence of plant endosymbionts

Abstract 1. Variation in plant chemistry does not only mediate interactions between plants and herbivores but also those between herbivores and their natural enemies, and plants and natural enemies. 2. Endophytic fungi complete their whole life cycle within the host plant’s tissue and are associated with a large diversity of plant species. Endophytes of the genus Neotyphodium alter the chemistry of the host plant by producing herbivore toxic alkaloids. 3. Here we asked whether the endophyte‐tolerant aphid species Metopolophium festucae could be defended against its parasitoid Aphidius ervi when feeding on endophyte‐infected plants. In a laboratory experiment, we compared life‐history traits of A. ervi when exposed to hosts on endophyte‐infected or endophyte‐free Lolium perenne. 4. The presence of endophytes significantly increased larval and pupal development times, but did not affect the mortality of immature parasitoids or the longevity of the adults. Although the number of parasitoid mummies tended to be reduced on endophyte‐infected plants, the number of emerging parasitoids did not differ significantly between the two treatments. 5. This shows that the metabolism of individual aphids feeding on infected plants may be changed and help in the defence against parasitoids. An increase in parasitoid development time should ultimately reduce the population growth of A. ervi. Therefore, endophyte presence may represent an advantage for endophyte‐tolerant aphid species through extended parasitoid development and its effect on parasitoid population dynamics.