Ground beetle movement is deterred by habitat edges: a mark-release-recapture study on the effectiveness of border crops in an agricultural landscape.

Border crops can increase beneficial insect biodiversity within agricultural fields by supplementing insects with food and nesting resources. However, the effectiveness of border crops relies on insect movement between adjacent habitats and some insects might consider habitat boundaries as barriers. Therefore, understanding insect movement between habitats is needed to determine the effectiveness of border crops for ecosystem services such as pest control within agricultural habitats. Our objective was to compare ground beetle (Coleoptera: Carabidae) movement across soybean plots that were bordered by corn and grassland habitat to determine whether habitat boundaries were considered barriers of movement to predatory beetles. Using a grid of pitfall traps within these habitats, we conducted a mark, release, and recapture experiment to track and evaluate ground beetle movement patterns. We found that ground beetles stayed in the habitat of their release and that movement between habitats, despite the type of bordering habitat or type of edge, was uncommon. We also found that long-distance movement was rare as most beetles moved less than 5 m (regardless of release or recaptured habitat) and movement was perpendicular to habitat edges. These results suggest that any edge habitat, including agricultural-agricultural boundaries and natural-agricultural boundaries, are likely barriers to ground beetle movement. Therefore, in order for border crops to be effective in pest management by ground beetles, making habitat edges more permeable, especially using techniques such as edge softening, could promote cross-habitat movement and ultimately contribute to natural pest control in agricultural systems.

Microbial vectoring capacity by internal- and external-infesting stored product insects after varying dispersal periods between novel food patches: An underestimated risk.

Understanding the ability of internal- and external-infesting stored product insects to vector microbes is important for estimating the relative risk that insects pose to postharvest commodities as they move between habitat patches and in the landscape. Thus, the aim of the current study was to evaluate and compare the microbial growth in novel food patches at different dispersal periods by different populations of Sitophilus oryzae (e.g., internal-infesting) and Lasioderma serricorne (e.g., external-infesting). Adults of both species collected from laboratory colonies or field-captured populations were either placed immediately in a novel food patch, or given a dispersal period of 24 or 72 h in a sterilized environment before entering a surrogate food patch. Vectored microbes in new food patches were imaged after 3 or 5 days of foraging, and microbial growth was processed using ImageJ while fungal species were identified through sequencing the ITS4/5 ribosomal subunit. We found that increasing dispersal time resulted in multiple-fold reductions in microbial growth surrogate food patches by L. serricorne but not S. oryzae. This was likely attributable to higher mobility by S. oryzae than L. serricorne. A total of 20 morphospecies were identified from 13 genera among the 59 sequences, with a total of 23% and 16% classified as Aspergillus and Penicillium spp. Our data suggest that there is a persistent risk of microbial contamination by both species, which has important food safety implications at food facilities.

Differentiating spillover: an examination of cross-habitat movement in ecology spillover in ecology.

Organisms that immigrate into a recipient habitat generate a movement pattern that affects local population dynamics and the environment. Spillover is the pattern of unidirectional movement from a donor habitat to a different, adjacent recipient habitat. However, ecological definitions are often generalized to include any cross-habitat movement, which limits within- and cross-discipline collaboration. To assess spillover nomenclature, we reviewed 337 studies within the agriculture, disease, fisheries and habitat fragmentation disciplines. Each study's definition of spillover and the methods used were analysed. We identified four descriptors (movement, habitat type and arrangement, and effect) used that differentiate spillover from other cross-habitat movement patterns (dispersal, foray loops and edge movement). Studies often define spillover as movement (45%) but rarely measure it as such (4%), particularly in disease and habitat fragmentation disciplines. Consequently, 98% of studies could not distinguish linear from returning movement out of a donor habitat, which can overestimate movement distance. Overall, few studies (12%) included methods that matched their own definition, revealing a distinct mismatch. Because theory shows that long-term impacts of the different movement patterns can vary, differentiating spillover from other movement patterns is necessary for effective long-term and inter-disciplinary management of organisms that use heterogeneous landscapes.

Short-term spatial dispersion patterns between the larger grain borer and the maize weevil in grain columns.

The invasive larger grain borer (Prostephanus truncatus) and the maize weevil (Sitophilus zeamais) co-occur in many regions of the world. While competition between these 2 species has been studied extensively, there is little information on spatial dispersion patterns in bulk storage of grain. To evaluate potential overlap in realized niche, we evaluated the short-term spatial dispersion behavior of P. truncatus and S. zeamais in monolayers of maize alone or together for 1 day compared to 7 days. We evaluated competition under three different densities, namely 10-20, 75-150, and 150-300 insects/kg for P. truncatus and S. zeamais. The monolayers were equally divided into 24 zones to track location the abundance of insects and damage to maize. We found that both species generally aggregated together and were correlated to the same location as heterospecifics. After 1 day, most of the insects for both species were near the top of the monolayer, but by 7 days, most individuals were at the bottom of the monolayers. In monolayers, when alone, P. truncatus created a clear path of destruction to the bottom of the monolayer, but when S. zeamais was present, damage was lessened and shifted upwards in the grain column. In an olfactometer assay, P. truncatus preferred maize odors, while S. zeamais exhibited no preference among maize, conspecifics, and heterospecifics. In evaluating relative emissions, each of these treatments emitted unique odors but with significant overlap. These data may improve targeting of chemical control tactics by identifying the position of these insects in the grain mass.

Attraction, mobility, and preference by Lasioderma serricorne (Coleoptera: Ptinidae) to microbially-mediated volatile emissions by two species of fungi in stored grain.

Insects and microbes are known to interact in a variety of ways at food facilities, compounding damage. However, little research has explicated how specific common fungal species affect the behavior of the cosmopolitan secondary stored product pest, Lasioderma serricorne. Enhanced knowledge about attraction to microbially-produced volatile organic compounds (MVOCs) may be used to manipulate insect behavior. Aspergillus flavus and Fusarium verticillioides are two common, widespread pre- and postharvest fungi on small cereals that produce aflatoxins and fumonisins, respectively, while directly competing with each other for nutrients. Our goals were to (1) characterize the volatile emissions from grain inoculated by A. flavus or F. verticillioides derived from the cuticle of L. serricorne compared to uninoculated and sanitized grain, and (2) understand how MVOCs from each fungal species affects mobility, attraction, and preference by L. serricorne. Headspace collection revealed that the F. verticillioides- and A. flavus-inoculated grain produced significantly different volatiles compared to sanitized grain or the positive control. Changes in MVOC emissions affected close-range foraging during an Ethovision movement assay, with a greater frequency of entering and spending time in a small zone with kernels inoculated with A. flavus compared to other treatments. In the release-recapture assay, MVOCs were found to be attractive to L. serricorne at longer distances in commercial pitfall traps. There was no preference shown among semiochemical stimuli in a still-air, four-way olfactometer. Overall, our study suggests that MVOCs are important for close- and long-range orientation of L. serricorne during foraging, and that MVOCs may have the potential for inclusion in behaviorally-based tactics for this species.

Grain Inoculated with Different Growth Stages of the Fungus, Aspergillus flavus, Affect the Close-Range Foraging Behavior by a Primary Stored Product Pest, Sitophilus oryzae (Coleoptera: Curculionidae).

Although some research has investigated the interactions among stored product insects and microbes, little research has examined how specific fungal life stages affect volatile emissions in grain and linked it to the behavior of Sitophilus oryzae, the cosmopolitan rice weevil. Thus, our goals were to 1) isolate, culture, and identify two fungal life stages of Aspergillus flavus, 2) characterize the volatile emissions from grain inoculated by each fungal morphotype, and 3) understand how microbially-produced volatile organic compounds (MVOCs) from each fungal morphotype affect foraging, attraction, and preference by S. oryzae. We hypothesized that the headspace blends would be unique among our treatments and that this will lead to preferential mobility by S. oryzae among treatments. Using headspace collection coupled with GC-MS, we found the sexual life stage of A. flavus had the most unique emissions of MVOCs compared to the other semiochemical treatments. This translated to a higher interaction with kernels containing grain with the A. flavus sexual life stage, as well as a higher cumulative time spent in those zones by S. oryzae in a video-tracking assay in comparison to the asexual life stage. While fungal cues were important for foraging at close-range, the release-recapture assay indicated that grain volatiles were more important for attraction at longer distances. There was no significant preference between grain and MVOCs in a four-way olfactometer. Overall, this study enhances our understanding of how fungal cues affect the close and longer range foraging ecology of a primarily stored product insect.

Native Flowering Border Crops Attract High Pollinator Abundance and Diversity, Providing Growers the Opportunity to Enhance Pollination Services.

Over the past century, habitat loss from agricultural intensification has contributed to pollinator decline. One way to mitigate the harmful effects of agricultural intensification is through the re-introduction of native flowering plants as border strips that provide supplemental floral and nesting resources to pollinators. However, border crop species vary in bloom period and flower densities, and are thus likely to attract different suites of pollinator species. Resulting differences in pollinator community composition are likely to affect their ability to provide pollination services to adjacent crop habitat. To address these issues, we implemented a two-year study on the impact of different flowering border crops on pollinator abundance, richness, and community composition. We also examined which crop features (bloom duration, number of flowers, floral area) were most likely to affect pollinator densities. We found that native flowering plant border crops of diverse prairie mix and monocultures of silflower (Silphium integrifolium Michx.) and cup plant (Silphium perfoliatum L.) attracted the highest abundance and species richness of bees and pollinator groups combined, while alfalfa (Medicago sativa L.) attracted the highest lepidopteran abundance and species richness. We also found a significant, positive relationship between pollinator abundance and floral resource amount and bloom duration. These findings offer valuable insight into the impacts of different land management strategies on different pollinator groups, and thus provide landowners with management options for attracting specific pollinator groups and species.

Microbial Volatile Organic Compounds from Tempered and Incubated Grain Mediate Attraction by a Primary but Not Secondary Stored Product Insect Pest in Wheat.

There has been a dearth of research elucidating the behavioral effect of microbially-produced volatile organic compounds on insects in postharvest agriculture. Demonstrating attraction to MVOC's by stored product insects would provide an additional source of unique behaviorally-relevant stimuli to protect postharvest commodities at food facilities. Here, we assessed the behavioral response of a primary (Rhyzopertha dominica) and secondary (Tribolium castaneum) grain pest to bouquets of volatiles produced by whole wheat that were untempered, or tempered to 12%, 15%, or 19% grain moisture and incubated for 9, 18, or 27 days. We hypothesized that MVOC's may be more important for the secondary feeder because they signal that otherwise unusable, intact grains have become susceptible by weakening of the bran. However, contrary to our expectations, we found that the primary feeder, R. dominica, but not T. castaneum was attracted to MVOC's in a wind tunnel experiment, and in a release-recapture assay using commercial traps baited with grain treatments. Increasing grain moisture resulted in elevated grain damage detected by near-infrared spectroscopy and resulted in small but significant differences in the blend of volatiles emitted by treatments detected by gas chromatography coupled with mass spectrometry (GC-MS). In sequencing the microbial community on the grain, we found a diversity of fungi, suggesting that an assemblage was responsible for emissions. We conclude that R. dominica is attracted to a broader suite of MVOC's than T. castaneum, and that our work highlights the importance of understanding insect-microbe interactions in the postharvest agricultural supply chain.

A Systematic Review of the Behavioral Responses by Stored-Product Arthropods to Individual or Blends of Microbially Produced Volatile Cues.

Microbes are ubiquitous and play important ecological roles in a variety of habitats. While research has been largely focused on arthropods and microbes separately in the post-harvest supply chain, less attention has been paid to their interactions with each other. Up to this point, there has been no attempt to systematically describe the patterns of behavioral responses by stored-product insects to microbially produced volatile organic compounds (MVOCs). Thus, our aims were to evaluate whether stored-product arthropods were primarily and significantly attracted, repelled, or had a net neutral effect (e.g., unaffected or mixed) by MVOCs presented as (1) complex headspace blends or (2) single constituents and known mixtures. In total, we found 43 articles that contained 384 sets of tests with different combinations of methodology and/or qualitative findings, describing the behavioral responses of 24 stored-product arthropod species from two classes, four orders, and 14 families to 58 individual microbial compounds and the complex headspace blends from at least 78 microbial taxa. A total of five and four stored-product arthropod species were significantly attracted and repelled by MVOCs across odor sources, respectively, while 13 were unaffected or exhibited mixed effects. We summarize the biases in the literature, including that the majority of tests have occurred in the laboratory with a limited subset of methodology and has largely only assessed the preference of adult arthropods. Finally, we identify foundational hypotheses for the roles that MVOCs play for stored-product arthropods as well as gaps in research and future directions, while highlighting that the behavioral responses to MVOCs are complex, context-, and taxon-dependent, which warrants further investigation.

Species traits elucidate crop pest response to landscape composition: a global analysis.

Recent synthesis studies have shown inconsistent responses of crop pests to landscape composition, imposing a fundamental limit to our capacity to design sustainable crop protection strategies to reduce yield losses caused by insect pests. Using a global dataset composed of 5242 observations encompassing 48 agricultural pest species and 26 crop species, we tested the role of pest traits (exotic status, host breadth and habitat breadth) and environmental context (crop type, range in landscape gradient and climate) in modifying the pest response to increasing semi-natural habitats in the surrounding landscape. For natives, increasing semi-natural habitats decreased the abundance of pests that exploit only crop habitats or that are highly polyphagous. On the contrary, populations of exotic pests increased with an increasing cover of semi-natural habitats. These effects might be related to changes in host plants and other resources across the landscapes and/or to modified top-down control by natural enemies. The range of the landscape gradient explored and climate did not affect pests, while crop type modified the response of pests to landscape composition. Although species traits and environmental context helped in explaining some of the variability in pest response to landscape composition, the observed large interspecific differences suggest that a portfolio of strategies must be considered and implemented for the effective control of rapidly changing communities of crop pests in agroecosystems.

How plant neighborhood composition influences herbivory: Testing four mechanisms of associational resistance and susceptibility.

Neighboring plants can decrease or increase each other's likelihood of damage from herbivores through associational resistance or susceptibility, respectively. Associational effects (AE) can transpire through changes in herbivore or plant traits that affect herbivore movement, densities, and feeding behaviors to ultimately affect plant damage. While much work has focused on understanding the mechanisms that underlie associational effects, we know little about how these mechanisms are influenced by neighborhood composition, i.e., plant density or relative frequency which is necessary to make predictions about when AE should occur in nature. Using a series of field and greenhouse experiments, I examined how plant density and relative frequency affected plant damage to Solanum carolinense and four mechanisms that underlie AE; (i) accumulation of insect herbivores and arthropod predators, (ii) microclimate conditions, (iii) plant resistance, and (iv) specialist herbivore preference. I found a positive relationship between S. carolinense damage and the relative frequency of a non-focal neighbor (Solidago altissima) and all four AE mechanisms were influenced by one or multiple neighborhood components. Frequency-dependence in S. carolinense damage is most likely due to greater generalist herbivore load on S. carolinense (through spillover from S. altissima) with microclimate variables, herbivore preference, predation pressures, and plant resistance having relatively weaker effects. Associational effects may have long-term consequences for these two plant species during plant succession and understanding context-dependent herbivory has insect pest management implication for other plant species in agriculture and forestry.

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.

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.

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.

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.