Effects of management outweigh effects of plant diversity on restored animal communities in tallgrass prairies.

A primary goal of ecological restoration is to increase biodiversity in degraded ecosystems. However, the success of restoration ecology is often assessed by measuring the response of a single functional group or trophic level to restoration, without considering how restoration affects multitrophic interactions that shape biodiversity. An ecosystem-wide approach to restoration is therefore necessary to understand whether animal responses to restoration, such as changes in biodiversity, are facilitated by changes in plant communities (plant-driven effects) or disturbance and succession resulting from restoration activities (management-driven effects). Furthermore, most restoration ecology studies focus on how restoration alters taxonomic diversity, while less attention is paid to the response of functional and phylogenetic diversity in restored ecosystems. Here, we compared the strength of plant-driven and management-driven effects of restoration on four animal communities (ground beetles, dung beetles, snakes, and small mammals) in a chronosequence of restored tallgrass prairie, where sites varied in management history (prescribed fire and bison reintroduction). Our analyses indicate that management-driven effects on animal communities were six-times stronger than effects mediated through changes in plant biodiversity. Additionally, we demonstrate that restoration can simultaneously have positive and negative effects on biodiversity through different pathways, which may help reconcile variation in restoration outcomes. Furthermore, animal taxonomic and phylogenetic diversity responded differently to restoration, suggesting that restoration plans might benefit from considering multiple dimensions of animal biodiversity. We conclude that metrics of plant diversity alone may not be adequate to assess the success of restoration in reassembling functional ecosystems.

Flowering plant composition shapes pathogen infection intensity and reproduction in bumble bee colonies.

Pathogens pose significant threats to pollinator health and food security. Pollinators can transmit diseases during foraging, but the consequences of plant species composition for infection is unknown. In agroecosystems, flowering strips or hedgerows are often used to augment pollinator habitat. We used canola as a focal crop in tents and manipulated flowering strip composition using plant species we had previously shown to result in higher or lower bee infection in short-term trials. We also manipulated initial colony infection to assess impacts on foraging behavior. Flowering strips using high-infection plant species nearly doubled bumble bee colony infection intensity compared to low-infection plant species, with intermediate infection in canola-only tents. Both infection treatment and flowering strips reduced visits to canola, but we saw no evidence that infection treatment shifted foraging preferences. Although high-infection flowering strips increased colony infection intensity, colony reproduction was improved with any flowering strips compared to canola alone. Effects of flowering strips on colony reproduction were explained by nectar availability, but effects of flowering strips on infection intensity were not. Thus, flowering strips benefited colony reproduction by adding floral resources, but certain plant species also come with a risk of increased pathogen infection intensity.

Reintroduced grazers and prescribed fire effects on beetle assemblage structure and function in restored grasslands.

Ecological restoration seeks to reestablish functioning ecosystems, but planning and evaluation often focus on taxonomic community structure and neglect consumers and their functional roles. The functional trait composition of insect assemblages, which make up the majority of animal diversity in many systems, can reveal how they are affected by restoration management and the consequences for ecosystem function. We sampled ground beetle (Coleoptera: Carabidae) assemblages in restored tallgrass prairies varying in management with prescribed fire and reintroduced American bison (Bison bison) to describe their taxonomic and functional trait structure. We also measured seed and arthropod predation to relate management, beetle assemblage characteristics, and function, and to test if function is maximized by trait diversity, dominant trait values, or beetle abundance. Beetle assemblages primarily varied with restoration age, declining over time in richness and both taxonomic and functional diversity, but bison presence also influenced taxonomic composition. Prescribed fire reduced seed predation in summer and arthropod predation in fall. Although seed predation was unrelated to beetle assemblages, arthropod predation was greater in sites with higher abundances of carnivorous ground beetles. The relatively weak impacts of fire and bison on functional assemblage structure is a promising sign that these management disturbances, aimed at supporting a diverse native plant community, are not detrimental to beetle assemblages. The significance of reduced predator function following prescribed fire will depend on the restoration context and whether seed or arthropod predation relates to management goals.

Introduced ecological engineers drive behavioral changes of grasshoppers, consequently linking to its abundance in two grassland plant communities.

Introduced ecosystem engineers are expected to have extensive ecological impacts on a broad range of resident biota by altering the physical-chemical structure of ecosystems. Livestock that are potentially important introduced ecosystem engineers in grassland systems could create and/or modify habitats for native plant-dwelling insects. Yet, there is little knowledge of how insects respond to engineering effects of introduced livestock. To bridge this gap, we tested how domestic sheep affects the behavior and abundance of a native grasshopper Euchorthippus unicolor at both low (11.8 ± 0.4 plant species per plot) and high (19.8 ± 0.5 plant species per plot) diversity sites. Results found grasshoppers shifted their resting and feeding locations from the upper to the intermediate or low layers of vegetation, and fed on more plants species following livestock engineering effects. In the low plant diversity habitats, grazing caused grasshoppers to increase switching frequency, spend more time searching for host plants, and reduce time spent feeding, but had opposite effects on all the three behaviors in the high-diversity habitats. Moreover, grazing engineering effects on behavioral changes of grasshoppers were potentially related to their abundance. Overall, this study highlights native insect species' behavior and abundance in responses to introduced ecological engineers, and suggests that ecosystem engineers of non-native species have strong and important impacts extending beyond their often most obvious and frequently documented direct ecological effects.

Soil microbial community composition in tallgrass prairie restorations converge with remnants across a 27-year chronosequence.

Restoration and management of natural ecosystems is a critical strategy in mitigating global biodiversity loss. This is exemplified in the American Midwest by efforts aimed at reclaiming historical grasslands lost to high-yield agriculture. While restorations traditionally take the form of plant reintroduction and management, advances in microbial analyses suggest that soil communities could be indicators restoration success. However, current understanding of key microbial taxa and functional activities in both natural and restored ecosystems is limited. Here, we investigated the impact of nearly 30 years of carefully managed restoration on soil microbial communities at the Nachusa Grasslands in northern Illinois, USA. We characterized bacterial and archaeal communities in a chronosequence of restored tallgrass prairies ranging from 1 to 27 years old across a growing season and compared them to communities in pre-restoration agricultural fields and remnant prairies. Results indicate that older restorations harboured communities statistically distinct from newer restorations. These communities converged toward those in local prairie remnants, suggesting that plant-focussed restoration has yielded soil bacterial communities reflective of a successful restoration. Recovery of microbial clades within the Verrucomicrobia and Acidobacteria are an important feature of this convergence, and these groups could be targeted for future soil-focussed, bottom-up restoration studies.

Attracting mutualists and antagonists: plant trait variation explains the distribution of specialist floral herbivores and pollinators on crops and wild gourds.

UNLABELLED: • PREMISE OF THE STUDY: Floral traits play important roles in pollinator attraction and defense against floral herbivory. However, plants may experience trade-offs between conspicuousness to pollinators and herbivore attraction. Comparative studies provide an excellent framework to examine the role of multiple traits shaping mutualist and antagonist interactions.• METHODS: To assess whether putative defensive and attractive traits predict species interactions, we grew 20 different Cucurbitaceae species and varieties in the field to measure interactions with pollinators and herbivores and in the greenhouse to assess trait variation. Cucurbits are characterized by the production of cucurbitacins, bitter nonvolatile terpenoids that are effective against generalist herbivores but can attract specialist beetles. We determined whether plant traits such as cucurbitacins predict herbivore resistance and pollinator attraction using an information-theoretic approach.• KEY RESULTS: Mutualists and floral antagonists were attracted to the same cucurbit varieties once they flowered. However, rather than cucurbitacin concentration, we found that the size of the flower and volatile emissions of floral sesquiterpenoids explained both pollinator and floral herbivore visitation preference across cucurbit taxa. This pattern held across cucurbit taxa and within the Cucurbita genus.• CONCLUSIONS: Surprisingly, floral sesquiterpenoid volatiles, which are associated with direct defense, indirect defense, and attraction, rather than defense traits such as cucurbitacins, appeared to drive interactions with both pollinators and floral herbivores across cucurbit taxa. Identifying the relevant plant traits for attraction and deterrence is important in this economically valuable crop, particularly if pollinators and floral herbivores use the same plant traits as cues.

Functional consequences of animal community changes in managed grasslands: An application of the CAFE approach.

In the midst of an ongoing biodiversity crisis, much research has focused on species losses and their impacts on ecosystem functioning. The functional consequences (ecosystem response) of shifts in communities are shaped not only by changes in species richness, but also by compositional shifts that result from species losses and gains. Species differ in their contribution to ecosystem functioning, so species identity underlies the consequences of species losses and gains on ecosystem functions. Such research is critical to better predict the impact of disturbances on communities and ecosystems. We used the "Community Assembly and the Functioning of Ecosystems" (CAFE) approach, a modification of the Price equation to understand the functional consequences and relative effects of richness and composition changes in small nonvolant mammal and dung beetle communities as a result of two common disturbances in North American prairie restorations, prescribed fire and the reintroduction of large grazing mammals. Previous research in this system has shown dung beetles are critically important decomposers, while small mammals modulate much energy in prairie food webs. We found that dung beetle communities were more responsive to bison reintroduction and prescribed fires than small nonvolant mammals. Dung beetle richness increased after bison reintroduction, with higher dung beetle community biomass resulting from changes in remaining species (context-dependent component) rather than species turnover (richness components); prescribed fire caused a minor increase in dung beetle biomass for the same reason. For small mammals, bison reintroduction reduced energy transfer through the loss of species, while prescribed fire had little impact on either small mammal richness or energy transfer. The CAFE approach demonstrates how bison reintroduction controls small nonvolant mammal communities by increasing prairie food web complexity, and increases dung beetle populations with possible benefits for soil health through dung mineralization and soil bioturbation. Prescribed fires, however, have little effect on small mammals and dung beetles, suggesting a resilience to fire. These findings illustrate the key role of re-establishing historical disturbance regimes when restoring endangered prairie ecosystems and their ecological function.

Linking agricultural practices, mycorrhizal fungi, and traits mediating plant-insect interactions.

Agricultural management has profound effects on soil communities. Activities such as fertilizer inputs can modify the composition of arbuscular mycorrhizal fungi (AMF) communities, which form important symbioses with the roots of most crop plants. Intensive conventional agricultural management may select for less mutualistic AMF with reduced benefits to host plants compared to organic management, but these differences are poorly understood. AMF are generally evaluated based on their direct growth effects on plants. However, mycorrhizal colonization also may alter plant traits such as tissue nutrients, defensive chemistry, or floral traits, which mediate important plant-insect interactions like herbivory and pollination. To determine the effect of AMF from different farming practices on plant performance and traits that putatively mediate species interactions, we performed a greenhouse study by inoculating Cucumis sativus (cucumber, Cucurbitaceae) with AMF from conventional farms, organic farms, and a commercial AMF inoculum. We measured growth and a suite of plant traits hypothesized to be important predictors of herbivore resistance and pollinator attraction. Several leaf and root traits and flower production were significantly affected by AMF inoculum. Both conventional and organic AMF reduced leaf P content but increased Na content compared to control and commercial AMF. Leaf defenses were unaffected by AMF treatments, but conventional AMF increased root cucurbitacin C, the primary defensive chemical of C. sativus, compared to organic AMF. These effects may have important consequences for herbivore preference and population dynamics. AMF from both organic and conventional farms decreased flower production relative to commercial and control treatments, which may reduce pollinator attraction and plant reproduction. AMF from both farm types also reduced seed germination, but effects on plant growth were limited. Our results suggest that studies only considering AMF effects on growth may overlook changes in plant traits that have the potential to influence interactions, and hence yield, on farms. Given the effects of AMF on plant traits documented here, and the great importance of both herbivores and pollinators to wild and cultivated plants, we advocate for comprehensive assessments of mycorrhizal effects in complex community contexts, with the aim of incorporating multispecies interactions both above and below the soil surface.

Interactions among predators and the cascading effects of vertebrate insectivores on arthropod communities and plants.

Theory on trophic interactions predicts that predators increase plant biomass by feeding on herbivores, an indirect interaction called a trophic cascade. Theory also predicts that predators feeding on predators, or intraguild predation, will weaken trophic cascades. Although past syntheses have confirmed cascading effects of terrestrial arthropod predators, we lack a comprehensive analysis for vertebrate insectivores-which by virtue of their body size and feeding habits are often top predators in these systems-and of how intraguild predation mediates trophic cascade strength. We report here on a meta-analysis of 113 experiments documenting the effects of insectivorous birds, bats, or lizards on predaceous arthropods, herbivorous arthropods, and plants. Although vertebrate insectivores fed as intraguild predators, strongly reducing predaceous arthropods (38%), they nevertheless suppressed herbivores (39%), indirectly reduced plant damage (40%), and increased plant biomass (14%). Furthermore, effects of vertebrate insectivores on predatory and herbivorous arthropods were positively correlated. Effects were strongest on arthropods and plants in communities with abundant predaceous arthropods and strong intraguild predation, but weak in communities depauperate in arthropod predators and intraguild predation. The naturally occurring ratio of arthropod predators relative to herbivores varied tremendously among the studied communities, and the skew to predators increased with site primary productivity and in trees relative to shrubs. Although intraguild predation among arthropod predators has been shown to weaken herbivore suppression, we find this paradigm does not extend to vertebrate insectivores in these communities. Instead, vertebrate intraguild preda-tion is associated with strengthened trophic cascades, and insectivores function as dominant predators in terrestrial plant-arthropod communities.

Restoration age and reintroduced bison may shape soil bacterial communities in restored tallgrass prairies.

Knowledge of how habitat restoration shapes soil microbial communities often is limited despite their critical roles in ecosystem function. Soil community diversity and composition change after restoration, but the trajectory of these successional changes may be influenced by disturbances imposed for habitat management. We studied soil bacterial communities in a restored tallgrass prairie chronosequence for >6 years to document how diversity and composition changed with age, management through fire, and grazing by reintroduced bison, and in comparison to pre-restoration agricultural fields and remnant prairies. Soil C:N increased with restoration age and bison, and soil pH first increased and then declined with age, although bison weakened this pattern. Bacterial richness and diversity followed a similar hump-shaped pattern as soil pH, such that the oldest restorations approached the low diversity of remnant prairies. ß-diversity patterns indicated that composition in older restorations with bison resembled bison-free sites, but over time they became more distinct. In contrast, younger restorations with bison maintained unique compositions throughout the study, suggesting bison disturbances may cause a different successional trajectory. We used a novel random forest approach to identify taxa that indicate these differences, finding that they were frequently associated with bacteria that respond to grazing in other grasslands.

Taxonomic and functional restoration of tallgrass prairie soil microbial communities in comparison to remnant and agricultural soils.

Restoring ecosystems requires the re-establishment of diverse soil microbial communities that drive critical ecosystem functions. In grasslands, restoration and management require the application of disturbances like fire and grazing. Disturbances can shape microbial taxonomic composition and potentially functional composition as well. We characterized taxonomic and functional gene composition of soil communities using whole genome shotgun metagenomic sequencing to determine how restored soil communities differed from pre-restoration agricultural soils and original remnant soils, how management affects soil microbes, and whether restoration and management affect the number of microbial genes associated with carbohydrate degradation. We found distinct differences in both taxonomic and functional diversity and composition among restored, remnant, and agricultural soils. Remnant soils had low taxonomic and functional richness and diversity, as well as distinct composition, indicating that restoration of agricultural soils does not re-create soil microbial communities that match remnants. Prescribed fire management increased functional diversity, which also was higher in more recently planted restorations. Finally, restored and post-fire soils included high abundances of genes encoding cellulose-degrading enzymes, so restorations and their ongoing management can potentially support functions important in carbon cycling.

Herbivory reduces plant interactions with above- and belowground antagonists and mutualists.

Herbivores affect plants through direct effects, such as tissue damage, and through indirect effects that alter species interactions. Interactions may be positive or negative, so indirect effects have the potential to enhance or lessen the net impacts of herbivores. Despite the ubiquity of these interactions, the indirect pathways are considerably less understood than the direct effects of herbivores, and multiple indirect pathways are rarely studied simultaneously. We placed herbivore effects in a comprehensive community context by studying how herbivory influences plant interactions with antagonists and mutualists both aboveground and belowground. We manipulated early-season aboveground herbivore damage to Cucumis sativus (cucumber, Cucurbitaceae) and measured interactions with subsequent aboveground herbivores, root-feeding herbivores, pollinators, and arbuscular mycorrhizal fungi (AMF). We quantified plant growth and reproduction and used an enhanced pollination treatment to determine if plants were pollen limited. Increased herbivory reduced interactions with both antagonists and mutualists. Plants with high levels of early herbivory were significantly less likely to suffer leaf damage later in the summer and tended to be less attacked by root herbivores. Herbivory also reduced pollinator visitation, likely due to fewer and smaller flowers, and reduced AMF colonization. The net effect of herbivory on plant growth and reproduction was strongly negative, but lower fruit and seed production were not due to reduced pollinator visits, because reproduction was not pollen limited. Although herbivores influenced interactions between plants and other organisms, these effects appear to be weaker than the direct negative effects of early-season tissue loss.

Winter predation by insectivorous birds and consequences for arthropods and plants in summer.

Top-down effects of predators can have important consequences for ecosystems. Insectivorous birds frequently have strong predation effects on herbivores and other arthropods, as well as indirect effects on herbivores' host plants. Diet studies have shown that birds in temperate ecosystems consume arthropods in winter as well as in summer, but experimental studies of bird predation effects have not attempted to quantitatively separate winter predation impacts from those in summer. To understand if winter foraging by insectivorous birds has consequences for arthropods or plants, we performed a meta-analysis of published bird exclusion studies in temperate forest and shrubland habitats. We categorized 85 studies from 41 publications by whether birds were excluded year-round or only in summer, and analyzed arthropod and plant response variables. We also performed a manipulative field experiment in which we used a factorial design to exclude birds from Quercus velutina Lam. saplings in winter and summer, and censused arthropods and herbivore damage in the following growing season. In the meta-analysis, birds had stronger negative effects on herbivores in studies that included winter exclusion, and this effect was not due to study duration. However, this greater predation effect did not translate to a greater impact on plant damage or growth. In the field experiment, winter exclusion did not influence herbivore abundance or their impacts on plants. We have shown that winter feeding by temperate insectivorous birds can have important consequences for insect herbivore populations, but the strength of these effects may vary considerably among ecosystems. A full understanding of the ecological roles of insectivorous birds will require explicit consideration of their foraging in the non-growing season, and we make recommendations for how future studies can address this.

The effect of natural disturbances on forest biodiversity: an ecological synthesis.

Disturbances alter biodiversity via their specific characteristics, including severity and extent in the landscape, which act at different temporal and spatial scales. Biodiversity response to disturbance also depends on the community characteristics and habitat requirements of species. Untangling the mechanistic interplay of these factors has guided disturbance ecology for decades, generating mixed scientific evidence of biodiversity responses to disturbance. Understanding the impact of natural disturbances on biodiversity is increasingly important due to human-induced changes in natural disturbance regimes. In many areas, major natural forest disturbances, such as wildfires, windstorms, and insect outbreaks, are becoming more frequent, intense, severe, and widespread due to climate change and land-use change. Conversely, the suppression of natural disturbances threatens disturbance-dependent biota. Using a meta-analytic approach, we analysed a global data set (with most sampling concentrated in temperate and boreal secondary forests) of species assemblages of 26 taxonomic groups, including plants, animals, and fungi collected from forests affected by wildfires, windstorms, and insect outbreaks. The overall effect of natural disturbances on α-diversity did not differ significantly from zero, but some taxonomic groups responded positively to disturbance, while others tended to respond negatively. Disturbance was beneficial for taxonomic groups preferring conditions associated with open canopies (e.g. hymenopterans and hoverflies), whereas ground-dwelling groups and/or groups typically associated with shady conditions (e.g. epigeic lichens and mycorrhizal fungi) were more likely to be negatively impacted by disturbance. Across all taxonomic groups, the highest α-diversity in disturbed forest patches occurred under moderate disturbance severity, i.e. with approximately 55% of trees killed by disturbance. We further extended our meta-analysis by applying a unified diversity concept based on Hill numbers to estimate α-diversity changes in different taxonomic groups across a gradient of disturbance severity measured at the stand scale and incorporating other disturbance features. We found that disturbance severity negatively affected diversity for Hill number q = 0 but not for q = 1 and q = 2, indicating that diversity-disturbance relationships are shaped by species relative abundances. Our synthesis of α-diversity was extended by a synthesis of disturbance-induced change in species assemblages, and revealed that disturbance changes the ß-diversity of multiple taxonomic groups, including some groups that were not affected at the α-diversity level (birds and woody plants). Finally, we used mixed rarefaction/extrapolation to estimate biodiversity change as a function of the proportion of forests that were disturbed, i.e. the disturbance extent measured at the landscape scale. The comparison of intact and naturally disturbed forests revealed that both types of forests provide habitat for unique species assemblages, whereas species diversity in the mixture of disturbed and undisturbed forests peaked at intermediate values of disturbance extent in the simulated landscape. Hence, the relationship between α-diversity and disturbance severity in disturbed forest stands was strikingly similar to the relationship between species richness and disturbance extent in a landscape consisting of both disturbed and undisturbed forest habitats. This result suggests that both moderate disturbance severity and moderate disturbance extent support the highest levels of biodiversity in contemporary forest landscapes.

Termite sensitivity to temperature affects global wood decay rates.

Deadwood is a large global carbon store with its store size partially determined by biotic decay. Microbial wood decay rates are known to respond to changing temperature and precipitation. Termites are also important decomposers in the tropics but are less well studied. An understanding of their climate sensitivities is needed to estimate climate change effects on wood carbon pools. Using data from 133 sites spanning six continents, we found that termite wood discovery and consumption were highly sensitive to temperature (with decay increasing >6.8 times per 10°C increase in temperature)-even more so than microbes. Termite decay effects were greatest in tropical seasonal forests, tropical savannas, and subtropical deserts. With tropicalization (i.e., warming shifts to tropical climates), termite wood decay will likely increase as termites access more of Earth's surface.

Leaf quality, predators, and stochastic processes in the assembly of a diverse herbivore community.

Ecological communities are structured by both deterministic, niche-based processes and stochastic processes such as dispersal. A pressing issue in ecology is to determine when and for which organisms each of these types of processes is important in community assembly. The roles of deterministic and stochastic processes have been studied for a variety of communities, but very few researchers have addressed their contribution to insect herbivore community structure. Insect herbivore niches are often described as largely shaped by the antagonistic pressures of predation and host plant defenses. However host plants are frequently discrete patches of habitat, and their spatial arrangement can affect herbivore dispersal patterns. We studied the roles of predation, host plant quality, and host spatial proximity for the assembly of a diverse insect herbivore community on Quercus alba (white oak) across two growing seasons. We examined abundances of feeding guilds to determine if ecologically similar species responded similarly to variation in niches. Most guilds responded similarly to leaf quality, preferring high-nitrogen, low-tannin host plants, particularly late in the growing season, while bird predation had little impact on herbivore abundance. The communities on the high-quality plants tended to be larger and, in some cases, have greater species richness. We analyzed community composition by correlating indices of community similarity with predator presence, leaf quality similarity, and host plant proximity. Birds did not affect community composition. Community similarity was significantly associated with distance between host plants and uncorrelated with leaf quality similarity. Thus although leaf quality significantly affected the total abundance of herbivores on a host plant, in some cases leading to increased species richness, dispersal limitation may weaken this relationship. The species composition of these communities may be driven by stochastic processes rather than variation in host plant characteristics or differential predation by insectivorous birds.

Light environment and the impacts of foliage quality on herbivorous insect attack and bird predation.

Theory predicts that variation in plant traits will modify both the direct interactions between plants and herbivores and the indirect impacts of predators of those herbivores. Light has strong effects on leaf quality, so the impacts of herbivores and predators may differ between plants grown in sun and shade. However, past experiments have often been unable to separate the effects of light environment on plant traits and herbivory from direct effects on herbivores and predators. We first manipulated light availability in an open habitat using a shade cloth pre-treatment to produce oak saplings with different leaf qualities. Leaves on plants exposed to high light were thicker and tougher and had lower nitrogen and water contents, and higher carbon and phenolic contents than leaves on plants under a shade cloth. Then, in the main experiment, we moved all plants to a common shade environment where bird predators were excluded in a factorial design. We measured insect herbivore abundance and leaf damage. Herbivores were significantly more abundant and caused greater leaf damage on sun trees, although these leaf characteristics are usually associated with low-quality food. Bird exclusion did not change herbivore abundance but did increase leaf damage. Contrary to our predictions, the effects of birds did not differ between trees grown in sun and shade conditions. Thus, differences in effects of predators on herbivores and plants between light habitats, when observed, might be due to variation in predator abundance and not bottom-up effects of host plant quality.

Effects of above- and belowground herbivory on growth, pollination, and reproduction in cucumber.

Plants experience unique challenges due to simultaneous life in two spheres, above- and belowground. Interactions with other organisms on one side of the soil surface may have impacts that extend across this boundary. Although our understanding of plant-herbivore interactions is derived largely from studies of leaf herbivory, belowground root herbivores may affect plant fitness directly or by altering interactions with other organisms, such as pollinators. In this study, we investigated the effects of leaf herbivory, root herbivory, and pollination on plant growth, subsequent leaf herbivory, flower production, pollinator attraction, and reproduction in cucumber (Cucumis sativus). We manipulated leaf and root herbivory with striped cucumber beetle (Acalymma vittatum) adults and larvae, respectively, and manipulated pollination with supplemental pollen. Both enhanced leaf and root herbivory reduced plant growth, and leaf herbivory reduced subsequent leaf damage. Plants with enhanced root herbivory produced 35% fewer female flowers, while leaf herbivory had no effect on flower production. While leaf herbivory reduced the time that honey bees spent probing flowers by 29%, probing times on root-damaged plants were over twice as long as those on control plants. Root herbivory increased pollen limitation for seed production in spite of increased honey bee preference for plants with root damage. Leaf damage and hand-pollination treatments had no effect on fruit production, but plants with enhanced root damage produced 38% fewer fruits that were 25% lighter than those on control plants. Despite the positive effect of belowground damage on honey bee visitation, root herbivory had a stronger negative effect on plant reproduction than leaf herbivory. These results demonstrate that the often-overlooked effects of belowground herbivores may have profound effects on plant performance.

Disturbance-Induced Trophic Niche Shifts In Ground Beetles (Coleoptera: Carabidae) In Restored Grasslands.

Ecosystem restoration is a critical component of land management, countering the loss of native biodiversity. Restoration efforts are enhanced by reintroducing naturally occurring ecosystem processes, including disturbances that may impact species characteristics such as niche position or niche size. In grasslands, grazing and fire affect plant diversity and habitat complexity, which potentially influence insect dietary behaviors and thus their contributions to functions like seed and arthropod predation. Using carbon and nitrogen stable isotopes, we characterized variation in the dietary niche of six ground beetle species (Coleoptera: Carabidae) in response to grazing by reintroduced bison and prescribed fire disturbances in twenty tallgrass prairies. Management disturbances did not affect activity density for most beetle species and mean trophic position was mostly unaffected. However, five of six species exhibited increased trophic niche area and breadth with disturbances, indicating a switch to a more generalist diet that incorporated a wider range of food items. The combination of bison and fire impacts may increase vegetation patchiness and heterogeneity, driving these diet changes. Morphological traits and microhabitat preferences might mediate response to disturbances and the resulting heterogeneity. Combining prescribed fire and grazing, which increases plant diversity and vegetation structural diversity, may help beetle communities establish over time and support the ecological functions to which these insects contribute.