Tania N. Kim

Trophic cascades in agricultural landscapes: indirect effects of landscape composition on crop yield

The strength and prevalence of trophic cascades, defined as positive, indirect effects of natural enemies (predatory and parasitic arthropods) on plants, is highly variable in agroecosystems. This variation may in part be due to the spatial or landscape context in which hese trophic cascades occur. In 2011 and 2012, we conducted a natural enemy exclusion experiment in soybean fields along a gradient of landscape composition across southern Wisconsin and Michigan, USA. We used structural equation modeling to ask (1) whether natural enemies influence biocontrol of soybean aphids (SBA) and soybean yield and (2) whether landscape effects on natural enemies influence the strength of the trophic cascades. We found that natural enemies (NE) suppressed aphid populations in both years of our study, and, in 2011, the yield of soybean plants exposed to natural enemies was 37% higher than the yield of plants with aphid populations protected from natural enemies. The strength of the :rophic cascade was also influenced by landscape context. We found that landscapes with a higher proportion of soybean and higher diversity habitats resulted in more NE, fewer aphids, and, in some cases, a trend toward greater soybean yield. These results indicate that landscape context is important for understanding spatial variability in biocontrol and yield, but other factors, such as environmental variability and compensatory growth, might overwhelm the beneficial effects of biocontrol on crop yield.

Plant neighborhood effects on herbivory: damage is both density and frequency dependent

Neighboring plants can affect the likelihood that a focal plant is attacked by herbivores. Both the density of conspecific neighbors (resource concentration or dilution effects) and the relative density of heterospecific neighbors (associational effects or effects of neighbor frequency) within the local neighborhood can affect herbivore load and plant damage. Understanding how these neighborhood effects influence processes such as plant competition or natural selection on plant resistance traits will require knowing how both plant density and frequency affect damage, but previous studies have generally confounded density and frequency effects. In this study, we independently manipulated the absolute density and frequency (i.e., relative density) of two plant species (Solanum carolinense and Solidago altissima) to characterize neighborhood composition effects on S. carolinense damage by herbivores, providing the first picture of how both density and frequency of neighbors influence damage in a single system. We found both a positive effect of S. carolinense density on S. carolinense damage (a resource concentration effect) and a nonlinear effect of S. altissima frequency on S. carolinense damage (associational susceptibility). If these types of patterns are common in nature, future studies seeking to understand neighborhood effects on damage need to incorporate both density and frequency effects and capture any nonlinear effects by selecting a range of values rather than focusing on only a pair of densities or frequencies. This type of data on neighborhood effects will allow us to understand the contribution of neighborhood effects to population-level processes such as competition, the evolution of plant resistance to herbivores, and yield gains in agricultural crop mixtures.

Insect herbivores change the outcome of plant competition through both inter- and intraspecific processes.

Insect herbivores can affect plant abundance and community composition, and theory suggests that herbivores influence plant communities by altering interspecific interactions among plants. Because the outcome of interspecific interactions is influenced by the per capita competitive ability of plants, density dependence, and intrinsic rates of increase, measuring herbivore effects on all these processes is necessary to understand the mechanisms by which herbivores influence plant communities. We fit alternative competition models to data from a response surface experiment conducted over four years to examine how herbivores affected the outcome of competition between two perennial plants, Solidago altissima and Solanum carolinense. Within a growing season, herbivores reduced S. carolinense plant size but did not affect the size of S. altissima, which exhibited compensatory growth. Across seasons, herbivores did not affect S. carolinense density or biomass but reduced both the density and population growth of S. altissima. The best-fit models indicated that the effects of herbivores varied with year. In some years, herbivores increased the per capita competitive effect of S. altissima on S. carolinense; in other years, herbivores influenced the intrinsic rate of increase of S. altissima. We examined possible herbivore effects on the longer-term outcome of competition (over the time scale of a typical old-field habitat), using simulations based on the best-fit models. In the absence of herbivores, plant coexistence was observed. In the presence of herbivores, S. carolinense was excluded by S. altissima in 72.3% of the simulations. We demonstrate that herbivores can influence the outcome of competition through changes in both per capita competitive effects and intrinsic rates of increase. We discuss the implications of these results for ecological succession and biocontrol.

The direct and indirect effects of fire on the assembly of insect herbivore communities: examples from the Florida scrub habitat.

Disturbance is a major source of spatial and temporal heterogeneity. In fire-maintained systems, disturbance by fire is often used as a management tool to increase biological diversity, restore degraded habitats, and reduce pest outbreaks. Much attention has been given to how plant communities recover from fire, but relatively few studies have examined post-fire responses of higher order species, such as insect herbivores. Because dynamic feedbacks occur between plants and their consumers, which can in turn influence the response of the entire ecosystem, incorporating higher trophic level responses into our understanding of the effects of fire is essential. In this study, we used structural equation modeling (SEM) to tease apart the direct and indirect effects of fire on insect herbivore assemblages found on three common oak species in the Florida scrub (Quercus inopina, Q. chapmanii, and Q. geminata). We investigated how fire affected herbivore abundance, richness, and community composition both directly and indirectly through environmental heterogeneity at different spatial scales (e.g., leaf quality, plant architecture, and habitat structure). We also investigated how seasonality and landscape heterogeneity influenced post-fire responses of insect herbivores and whether fire effects on herbivore assemblages varied among different host plants. Our general findings were that fire effects were (1) largely indirect, mediated through habitat structure (although direct fire effects were observed on Q. inopina herbivores), (2) non-linear through time due to self-thinning processes occurring in the scrub habitat, and (3) varied according to herbivore assemblage as a result of differences in the composition of species in each herbivore community. To the best of our knowledge, this is the first comprehensive study to examine how fire influences the assembly of insect herbivore communities through both direct and indirect pathways and at multiple spatial scales.

Selective manipulation of a non-dominant plant and its herbivores affects an old-field plant community

Competition and herbivory can interact to influence the recovery of plant communities from disturbance. Previous attention has focused on the roles of dominant plant species in structuring plant communities, leaving the roles of subordinate species often overlooked. In this study, we examined how manipulating the density of a subordinate plant species, Solanum carolinense, and its insect herbivores influenced an old-field plant community in northern Florida following a disturbance. Five years following the disturbance, the initial densities of S. carolinense planted at the start of the experiment negatively influenced total plant cover and species diversity, and the cover of some grasses (e.g., Paspalum urvillei) and forbs (e.g., Rubus trivalis). Selectively removing herbivores from S. carolinense increased S. carolinense abundance (both stem densities and cover), increased the total cover of plants in the surrounding plant community, and affected plant community composition. Some plant species increased (e.g., Digitaria ciliaris, Solidago altissima) and others decreased (e.g., Paspalum notatum, Cynodon dactylon) in cover in response to herbivore removal. Herbivore effects on plant community metrics did not depend on S. carolinense density (no significant herbivory by density interaction), suggesting that even at low densities, a reduction of S. carolinense herbivores can influence the rest of the plant community. The recovery of the plant community was context dependent, depending on site- and plot-level differences in underlying environmental conditions and pre-disturbance plant community composition. We demonstrate that the density of and herbivory on a single subordinate plant species can affect the structure of an entire plant community.

Harvesting biofuel grasslands has mixed effects on natural enemy communities and no effects on biocontrol services

Summary Perennial bioenergy systems, such as switchgrass and restored prairies, are alternatives to commonly used annual monocultures such as maize. Perennial systems require lower chemical input, provide greater ecosystem services such as carbon storage, greenhouse gas mitigation and support greater biodiversity of beneficial insects. However, biomass harvest will be necessary in managing these perennial systems for bioenergy production, and it is unclear how repeated harvesting might affect ecosystem services. In this study, we examined how repeated production-scale harvesting of diverse perennial grasslands influences vegetation structure, natural enemy communities (arthropod predators and parasitoids), and natural biocontrol services in two states (Wisconsin and Michigan, USA) over multiple years. We found that repeated biomass harvest reduced litter biomass and increased bare ground cover. Some natural enemy groups, such as ground-dwelling arthropods, decreased in abundance with harvest, whereas others such as foliar-dwelling arthropods increased in abundance. The disparity in responses is likely due to how different taxonomic groups utilize vegetation and differences in dispersal abilities. At the community level, biomass harvest altered community composition, increased total arthropod abundance and decreased evenness but did not influence species richness, diversity or biocontrol services. Harvest effects varied with time, diminishing in strength both within the season (for total abundance and evenness), across seasons (for evenness) or were consistent throughout the duration of the study (for community composition). Greater functional redundancy and compensatory responses of the different taxonomic groups may have buffered against the potentially negative effects of harvest on biocontrol services. Synthesis and applications. Our results show that in the short-term, repeated harvesting of perennial grasslands (when insect activity is low) consistently altered vegetation structure but had mixed effects on natural enemy communities and no discernible effects on biocontrol services. However, the long-term effects of repeated harvesting on vegetation structure, natural enemies and other arthropod-derived ecosystem services such as pollination and decomposition remain largely unknown.

Contrasting Foraging Patterns: Testing Resource-Concentration and Dilution Effects with Pollinators and Seed Predators.

Resource concentration effects occur when high resource density patches attract and support more foragers than low density patches. In contrast, resource dilution effects can occur if high density patches support fewer consumers. In this study, we examined the foraging rates of pollinators and seed predators on two perennial plant species (Rudbeckia triloba and Verbena stricta) as functions of resource density. Specifically, we examined whether resource-dense patches (densities of flower and seeds on individual plants) resulted in greater visitation and seed removal rates, respectively. We also examined whether foraging rates were context-dependent by conducting the study in two sites that varied in resource densities. For pollinators, we found negative relationships between the density of flowers per plant and visitation rates, suggesting dilution effects. For seed predators, we found positive relationships consistent with concentration effects. Saturation effects and differences in foraging behaviors might explain the opposite relationships; most of the seed predators were ants (recruitment-based foragers), and pollinators were mostly solitary foragers. We also found that foraging rates were site-dependent, possibly due to site-level differences in resource abundance and consumer densities. These results suggest that these two plant species may benefit from producing as many flowers as possible, given high levels of pollination and low seed predation.