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Most organisms face the problem of foraging and maintaining growth while avoiding predators. Typical animal responses to predator exposure include reduced feeding, elevated metabolism, and altered development rate, all of which can be beneficial in the presence of predators but detrimental in their absence. How then do animals balance growth and predator avoidance? In a series of field and greenhouse experiments, we document that the tobacco hornworm caterpillar, Manduca sexta, reduced feeding by 30–40% owing to the risk of predation by stink bugs, but developed more rapidly and gained the same mass as unthreatened caterpillars. Assimilation efficiency, extraction of nitrogen from food, and percent body lipid content all increased during the initial phase (1-3 d) of predation risk, indicating that enhanced nutritional physiology allows caterpillars to compensate when threatened. However, we report physiological costs of predation risk, including altered body composition (decreased glycogen) and reductions in assimilation efficiency later in development. Our findings indicate that hornworm caterpillars use temporally dynamic compensatory mechanisms that ameliorate the trade-off between predator avoidance and growth in the short term, deferring costs to a period when they are less vulnerable to predation.

Introduced plants may be important foraging resources for honey bees and wild pollinators, but how often and why pollinators visit introduced plants across an entire plant community is not well understood. Understanding the importance of introduced plants for pollinators could help guide management of these plants and conservation of pollinator habitat. We assessed how floral abundance and pollinator preference influence pollinator visitation rate and diversity on 30 introduced versus 24 native plants in central New York. Honey bees visited introduced and native plants at similar rates regardless of floral abundance. In contrast, as floral abundance increased, wild pollinator visitation rate decreased more strongly for introduced plants than native plants. Introduced plants as a group and native plants as a group did not differ in bee diversity or preference, but honey bees and wild pollinators preferred different plant species. As a case study, we then focused on knapweed (Centaurea spp.), an introduced plant that was the most preferred plant by honey bees, and that beekeepers value as a late‐summer foraging resource. We compared the extent to which honey bees versus wild pollinators visited knapweed relative to coflowering plants, and we quantified knapweed pollen and nectar collection by honey bees across 22 New York apiaries. Honey bees visited knapweed more frequently than coflowering plants and at a similar rate as all wild pollinators combined. All apiaries contained knapweed pollen in nectar, 86% of apiaries contained knapweed pollen in bee bread, and knapweed was sometimes a main pollen or nectar source for honey bees in late summer. Our results suggest that because of diverging responses to floral abundance and preferences for different plants, honey bees and wild pollinators differ in their use of introduced plants. Depending on the plant and its abundance, removing an introduced plant may impact honey bees more than wild pollinators. We found that because of diverging responses to floral abundance and preferences for different plants, honey bees and wild pollinators varied in their use of introduced plants. In particular, knapweed (Centaurea spp.)—an introduced plant valued by beekeepers—was most often visited and preferred by honey bees but not wild pollinators. Our results suggest that when considering pollinator use of introduced plant species, it is critical to distinguish between honey bees—the dominant managed pollinator in the United States—and wild pollinators, as removing introduced plants may benefit one group of pollinators at a cost to the other.

Plants may benefit from limiting the community of generalist floral visitors if the species that remain are more effective pollinators and less effective pollenivores. Plants can reduce access to pollen through altered floral cues or morphological structures, but can also reduce consumption through direct pollen defenses. We observed that Eucera (Peponapis) pruinosa , a specialist bee on Cucurbita plants, collected pure loads of pollen while generalist honey bees and bumble bees collected negligible amounts of cucurbit pollen, even though all groups of bees visited these flowers. Cucurbit flowers have no morphological adaptations to limit pollen collection by bees, thus we assessed their potential for physical, nutritional, and chemical pollen traits that might act as defenses to limit pollen loss to generalist pollinators. Bumble bee ( Bombus impatiens ) microcolonies experienced reduced pollen consumption, mortality, and reproduction as well as increased stress responses when exposed to nutritional and mechanical pollen defenses. These bees also experienced physiological effects of these defenses in the form of hindgut expansion and gut melanization. Chemical defenses alone increased the area of gut melanization in larger bees and induced possible compensatory feeding. Together, these results suggest that generalist bumble bees avoid collecting cucurbit pollen due to the physiological costs of physical and chemical pollen defenses.

Reports of pollinator declines have prompted efforts to understand contributing factors and protect vulnerable species. While pathogens can be widespread in bee communities, less is known about factors shaping pathogen prevalence among species. Functional traits are often used to predict susceptibility to stressors, including pathogens, in other species-rich communities. Here, we evaluated the relationship between bee functional traits (body size, phenology, nesting location, sociality, and foraging choice) and prevalence of trypanosomes, neogregarines, and the microsporidian Nosema ceranae in wild bee communities. For the most abundant bee species in our system, Bombus impatiens , we also evaluated the relationship between intra-specific size variation and pathogen prevalence. A trait-based model fit the neogregarine prevalence data better than a taxa-based model, while the taxonomic model provided a better model fit for N. ceranae prevalence, and there was no marked difference between the models for trypanosome prevalence. We found that Augochlorella aurata was more likely to harbor trypanosomes than many other bee taxa. Similarly, we found that bigger bees and those with peak activity later in the season were less likely to harbor trypanosomes, though the effect of size was largely driven by A. aurata . We found no clear intra-specific size patterns for pathogen prevalence in B. impatiens . These results indicate that functional traits are not always better than taxonomic affinity in predicting pathogen prevalence, but can help to explain prevalence depending on the pathogen in species-rich bee communities.

Honey bees are commonly co-exposed to pesticides during crop pollination, including the fungicide captan and neonicotinoid insecticide thiamethoxam. We assessed the impact of exposure to these two pesticides individually and in combination, at a range of field-realistic doses. In laboratory assays, mortality of larvae treated with captan was 80–90% greater than controls, dose-independent, and similar to mortality from the lowest dose of thiamethoxam. There was evidence of synergism (i.e., a non-additive response) from captan-thiamethoxam co-exposure at the highest dose of thiamethoxam, but not at lower doses. In the field, we exposed whole colonies to the lowest doses used in the laboratory. Exposure to captan and thiamethoxam individually and in combination resulted in minimal impacts on population growth or colony mortality, and there was no evidence of synergism or antagonism. These results suggest captan and thiamethoxam are each acutely toxic to immature honey bees, but whole colonies can potentially compensate for detrimental effects, at least at the low doses used in our field trial, or that methodological differences of the field experiment impacted results (e.g., dilution of treatments with natural pollen). If compensation occurred, further work is needed to assess how it occurred, potentially via increased queen egg laying, and whether short-term compensation leads to long-term costs. Further work is also needed for other crop pollinators that lack the social detoxification capabilities of honey bee colonies and may be less resilient to pesticides.

Herbivores are squeezed between the two omnipresent threats of variable food quality and natural enemy attack, but these two factors are not independent of one another. The mechanisms by which organisms navigate the dual challenges of foraging while avoiding predation are poorly understood. We tested the effects of plant defense and predation risk on herbivory in an assemblage of leaf-chewing insects on Solanum lycopersicum (tomato) that included two Solanaceae specialists (Manduca sexta and Leptinotarsa decemlineata) and one generalist (Trichoplusia ni). Defenses were altered using genetic manipulations of the jasmonate phytohormonal cascade, whereas predation risk was assessed by exposing herbivores to cues from the predaceous stink bug, Podisus maculiventris. Predation risk reduced herbivore food intake by an average of 29% relative to predator-free controls. Interestingly, this predator-mediated impact on foraging behavior largely attenuated when quantified in terms of individual growth rate. Only one of the three species experienced lower body weight under predation risk and the magnitude of this effect was small (17% reduction) compared with effects on foraging behavior. Manduca sexta larvae, compensated for their predator-induced reduction in food intake by more effectively converting leaf tissue to body mass. They also had higher whole-body lipid content when exposed to predators, suggesting that individuals convert energy to storage forms to draw upon when risk subsides. In accordance with expectations based on insect diet breadth, plant defenses tended to have a stronger impact on consumption and growth in the generalist than the two specialists. These data both confirm the ecological significance of predators in the foraging behavior of herbivorous prey and demonstrate how sophisticated compensatory mechanisms allow foragers to partially offset the detrimental effects of reduced food intake. The fact that these mechanisms operated across a wide range of plant resistance phenotypes suggests that compensation is not always constrained by reduced food quality.

Biodiversity loss is proceeding at an unprecedented rate, yet we lack a thorough understanding of the consequences of losing diversity at different scales. While species diversity is known to impact community and ecosystem processes, genotypic diversity is assumed to have relatively smaller effects. Nonetheless, a few recent studies suggest that genotypic diversity may have quantitatively similar ecological consequences compared to species diversity. Here we show that increasing either genotypic diversity of common evening primrose ( Oenothera biennis ) or species diversity of old-field plant species resulted in nearly equivalent increases (∼∼17%%) in aboveground primary production. The predominant mechanism explaining this effect, niche complementarity, was similar for each type of diversity. Arthropod species richness also increased with both types of plant diversity, but the mechanisms leading to this effect differed: abundance-driven accumulation of arthropod species was important in plant genotypic polycultures, whereas resource specialization was important in plant species polycultures. Thus, similar increases in primary productivity differentially impacted higher trophic levels in response to each type of plant diversity. These results highlight important ecological similarities and differences between genotypic and species diversity and suggest that genotypic diversity may play a larger role in community and ecosystem processes than previously realized.

Concern over insect declines has increased attention on the effects of pesticide residues on native insects. We collected strawberry (target) and common milkweed (non-target) foliage and flowers on two small Central New York farms, within crop fields as well as field margins, and analyzed the tissues for pesticide residues of 94 agrochemicals. We found quantifiable levels of 13 fungicides, herbicides, and insecticides, mostly at low concentrations (typically less than 200 ppb where detected), and more often on strawberry than milkweed. We generally found higher pesticide residue levels early in the season (June vs. July) and on leaves compared to flowers. Residue levels in fields did not differ strongly from margins but pesticide drift may have left low-level residues on milkweed leaves and flowers in margins. Given that non-target effects of fungicides are understudied, we selected two prevalent fungicides found in this study (cyprodinil and difenoconazole) and used them in laboratory assays to assess impacts on early instar monarch ( Danaus plexippus ) caterpillar feeding, growth, and growth efficiency on common milkweed, along with two other milkweed ( Asclepias ) species. We tested the hypothesis that exposure to pesticides may be most impactful on poor quality host plants. For both fungicides, exposure at the highest doses (>100,000 ppb) reduced feeding, with the strongest effect on Asclepias asperula , the lowest quality host plant. Effects on caterpillar growth were similarly negative and consistent across host plant species. Finally, effects of fungicides on gross growth efficiency of caterpillars were negative, but dependent on the fungicide. Effects of cyprodinil were stronger than difenoconazole, but at realistically low concentrations there was little effect of either fungicide. Nonetheless, higher concentrations of these chemicals, approximating those experienced directly after fungicide application, may impact non-target species. The observed interaction of fungicides with host plant species highlights the importance of considering resource quality in the assessment of non-target effects of pesticides.

Sensitivity analysis of ecological and epidemiological models is often used to identify the best targets of opportunity for control or management, for example, which subpopulations should be prioritised for vaccination or protection. However, effectiveness of a management action depends not just on which system component is targeted but also on when an action is taken. Traditional methods of model sensitivity analysis apply to time‐invariant parameter perturbations, which limits their ability to address questions about the timing of management interventions. A semianalytic method for performing time‐dependent sensitivity analysis (TDSA) has been recently introduced to address this need (Ng et al., in press). However, some of the steps typically require substantial time and effort. We have developed an R package, tdsa , that automates all steps required for TDSA by using numerical rather than analytic differentiation of model terms to evaluate the sensitivity equations. By avoiding analytic differentiation, the package substantially reduces user effort and risk of human error and increases the generality of TDSA by making it possible to analyse nonparametric or equation‐free models. The results of TDSA may be sensitive to assumptions of the system model. By reducing the effort needed to perform TDSA, the tdsa package facilitates comparison of results under different model assumptions, hence allowing for more robust conclusions. We illustrate this using an example involving the transmission of Morogoro virus in the Natal multimammate mouse, under different assumptions about how the contact rate depends on mouse population density. By automating most of the mathematical and programming steps of the semianalytical approach, the tdsa package makes TDSA accessible to a wide range of users with varying quantitative expertise.

Several fungal plant pathogens induce 'pseudoflowers' on their hosts to facilitate insect-mediated transmission of gametes and spores. When spores must be transmitted to host flowers to complete the fungal life cycle, we predict that pseudoflowers should evolve traits that mimic flowers and attract the most effective vectors in the flower-visiting community. We quantified insect visitation to flowers, healthy leaves and leaves infected with Monilinia vaccinii-corymbosi (Mvc), the causative agent of mummy berry disease of blueberry. We developed a nested PCR assay for detecting Mvc spores on bees, flies and other potential insect vectors. We also collected volatiles from blueberry flowers, healthy leaves and leaves infected with Mvc, and experimentally manipulated specific pathogen-induced volatiles to assess attractiveness to potential vectors. Bees and flies accounted for the majority of contacts with flowers, leaves infected with Mvc and healthy leaves. Flowers were contacted most often, while there was no difference between bee or fly contacts with healthy and infected leaves. While bees contacted flowers more often than flies, flies contacted infected leaves more often than bees. Bees were more likely to have Mvc spores on their bodies than flies, suggesting that bees may be more effective vectors than flies for transmitting Mvc spores to flowers. Leaves infected with Mvc had volatile profiles distinct from healthy leaves but similar to flowers. Two volatiles produced by flowers and infected leaves, cinnamyl alcohol and cinnamic aldehyde, were attractive to bees, while no volatiles manipulated were attractive to flies or any other insects. These results suggest that Mvc infection of leaves induces mimicry of floral volatiles, and that transmission occurs primarily via bees, which had the highest likelihood of carrying Mvc spores and visited flowers most frequently.