Warmer autumns and winters could reduce honey bee overwintering survival with potential risks for pollination services.

Honey bees and other pollinators are critical for food production and nutritional security but face multiple survival challenges. The effect of climate change on honey bee colony losses is only recently being explored. While correlations between higher winter temperatures and greater colony losses have been noted, the impacts of warmer autumn and winter temperatures on colony population dynamics and age structure as an underlying cause of reduced colony survival have not been examined. Focusing on the Pacific Northwest US, our objectives were to (a) quantify the effect of warmer autumns and winters on honey bee foraging activity, the age structure of the overwintering cluster, and spring colony losses, and (b) evaluate indoor cold storage as a management strategy to mitigate the negative impacts of climate change. We perform simulations using the VARROAPOP population dynamics model driven by future climate projections to address these objectives. Results indicate that expanding geographic areas will have warmer autumns and winters extending honey bee flight times. Our simulations support the hypothesis that late-season flight alters the overwintering colony age structure, skews the population towards older bees, and leads to greater risks of colony failure in the spring. Management intervention by moving colonies to cold storage facilities for overwintering has the potential to reduce honey bee colony losses. However, critical gaps remain in how to optimize winter management strategies to improve the survival of overwintering colonies in different locations and conditions. It is imperative that we bridge the gaps to sustain honey bees and the beekeeping industry and ensure food and nutritional security.

Turning a Pest into a Natural Enemy: Removing Earwigs from Stone Fruit and Releasing Them in Pome Fruit Enhances Pest Control.

The European earwig Forficula auricularia (L.) (Dermaptera: Forficulidae) is an omnivorous insect that is considered a minor pest of stone fruit and a key predator of pests in pome fruit orchards. In many pome fruit orchards, earwigs are absent or in low abundance due to broad-spectrum spray programs and the slow recolonization rate of earwigs. Orchards in transition to organic or "selective" conventional programs often struggle to achieve effective levels of biological control, and thus, may benefit from inoculating earwigs to expedite their re-establishment. In a two-year study, we evaluated the potential for mass trapping earwigs from stone fruit using rolled cardboard traps to reduce fruit damage and provide earwigs for augmentation in pome fruit. We also tested whether a single mass release or five releases (on alternating weeks) of the same total number of earwigs in apples and pears reduced pests relative to plots where no releases occurred. Mass trapping did not decrease earwig abundance or substantially reduce fruit damage in stone fruit orchards. However, trapping was an efficient method for providing earwigs for augmentation. Earwig abundances were only increased in orchards where earwigs were previously low or absent; however, multiple orchards with varying prior levels of earwigs exhibited reductions in key pests (woolly apple aphid and pear psylla). For some other pests evaluated, plots with mass releases of earwigs had a slight trend in overall lower pest density when compared with control plots. A strategy for moving earwigs out of stone fruit orchards and into pome fruit orchards could be an effective method for augmenting orchard predator populations.

Bacterial Endosymbionts Identified From Leafhopper (Hemiptera: Cicadellidae) Vectors of Phytoplasmas.

Insects often harbor bacterial endosymbionts that provide them with nutritional benefit or with protection against natural enemies, plant defenses, insecticides, and abiotic stresses. Certain endosymbionts may also alter acquisition and transmission of plant pathogens by insect vectors. We identified bacterial endosymbionts from four leafhopper vectors (Hemiptera: Cicadellidae) of 'Candidatus Phytoplasma' species by direct sequencing 16S rDNA and confirmed endosymbiont presence and identity by species-specific conventional PCR. We examined three vectors of Ca. Phytoplasma pruni, causal agent of cherry X-disease [Colladonus geminatus (Van Duzee), Colladonus montanus reductus (Van Duzee), Euscelidius variegatus (Kirschbaum)] - and a vector of Ca. Phytoplasma trifolii, the causal agent of potato purple top disease [Circulifer tenellus (Baker)]. Direct sequencing of 16S identified the two obligate endosymbionts of leafhoppers, 'Ca. Sulcia' and 'Ca. Nasuia', which are known to produce essential amino acids lacking in the leafhoppers' phloem sap diet. About 57% of C. geminatus also harbored endosymbiotic Rickettsia. We identified 'Ca. Yamatotoia cicadellidicola' in Euscelidius variegatus, providing just the second host record for this endosymbiont. Circulifer tenellus harbored the facultative endosymbiont Wolbachia, although the average infection rate was only 13% and all males were Wolbachia-uninfected. A significantly greater percentage of Wolbachia-infected Ci. tenellus adults than uninfected adults carried Ca. P. trifolii, suggesting that Wolbachia may increase this insect's ability to tolerate or acquire this pathogen. Results of our study provide a foundation for continued work on interactions between leafhoppers, bacterial endosymbionts, and phytoplasma.

A trait-based framework for predicting foodborne pathogen risk from wild birds.

Recent foodborne illness outbreaks have heightened pressures on growers to deter wildlife from farms, jeopardizing conservation efforts. However, it remains unclear which species, particularly birds, pose the greatest risk to food safety. Using >11,000 pathogen tests and 1565 bird surveys covering 139 bird species from across the western United States, we examined the importance of 11 traits in mediating wild bird risk to food safety. We tested whether traits associated with pathogen exposure (e.g., habitat associations, movement, and foraging strategy) and pace-of-life (clutch size and generation length) mediated foodborne pathogen prevalence and proclivities to enter farm fields and defecate on crops. Campylobacter spp. were the most prevalent enteric pathogen (8.0%), while Salmonella and Shiga-toxin producing Escherichia coli (STEC) were rare (0.46% and 0.22% prevalence, respectively). We found that several traits related to pathogen exposure predicted pathogen prevalence. Specifically, Campylobacter and STEC-associated virulence genes were more often detected in species associated with cattle feedlots and bird feeders, respectively. Campylobacter was also more prevalent in species that consumed plants and had longer generation lengths. We found that species associated with feedlots were more likely to enter fields and defecate on crops. Our results indicated that canopy-foraging insectivores were less likely to deposit foodborne pathogens on crops, suggesting growers may be able to promote pest-eating birds and birds of conservation concern (e.g., via nest boxes) without necessarily compromising food safety. As such, promoting insectivorous birds may represent a win-win-win for bird conservation, crop production, and food safety. Collectively, our results suggest that separating crop production from livestock farming may be the best way to lower food safety risks from birds. More broadly, our trait-based framework suggests a path forward for co-managing wildlife conservation and food safety risks in farmlands by providing a strategy for holistically evaluating the food safety risks of wild animals, including under-studied species.

Coevolution, diversification and alternative states in two-trophic communities.

Single-trait eco-evolutionary models of arms races between consumers and their resource species often show inhibition rather than promotion of community diversification. In contrast, modelling arms races involving multiple traits, we found that arms races can promote diversification when trade-off costs among traits make simultaneous investment in multiple traits either more beneficial or more costly. Coevolution between resource and consumer species generates an adaptive landscape for each, with the configuration giving predictable suites of consumer and resource species. Nonetheless, the adaptive landscape contains multiple alternative stable states, and which stable community is reached depends on small stochastic differences occurring along evolutionary pathways. Our results may solve a puzzling conflict between eco-evolutionary theory that predicts community diversification via consumer-resource interactions will be rare, and empirical research that has uncovered real cases. Furthermore, our results suggest that these real cases might be just a subset of alternative stable communities.

Conventional Soil Management May Promote Nutrients That Lure an Insect Pest to a Toxic Crop.

Slow and consistent nutrient release by organic fertilizers can improve plant nutrient balance and defenses, leading to herbivore avoidance of organically managed crops in favor of conventional crops with weaker defenses. We propose that this relative attraction to conventional plants, coupled with the use of genetically modified, insecticidal crops (Bt), has created an unintentional attract-and-kill system. We sought to determine whether Bt and non-Bt corn Zea mays L. plants grown in soil collected from five paired organic and conventional fields differed in attractiveness to European corn borer [Ostrinia nubilalis (Hübner)] moths, by conducting ovipositional choice and flight tunnel assays. We then examined the mechanisms driving the observed differences in attraction by comparing soil nutrient profiles, soil microbial activity, plant nutrition, and plant volatile profiles. Finally, we assessed whether European corn borer abundance near corn fields differed based on soil management. European corn borer preferred plants grown in conventional soil but did not discriminate between Bt and non-Bt corn. Organic management and more alkaline soil were associated with an increased soil magnesium:potassium ratio, which increased plant magnesium, and were linked to reduced European corn borer oviposition. There was an inconsistent trend for higher European corn borer moth activity near conventional fields. Our results extend the mineral balance hypothesis describing conventional plant preference by showing that it can also improve attraction to plants with genetically inserted toxins. Unintentional attract (to conventional) and (Bt) kill is a plausible scenario for pest declines in response to Bt corn adoption, but this effect may be obscured by variation in other management practices and landscape characteristics.

Small Molecules in the Venom of the Scorpion Hormurus waigiensis.

Despite scorpion stings posing a significant public health issue in particular regions of the world, certain aspects of scorpion venom chemistry remain poorly described. Although there has been extensive research into the identity and activity of scorpion venom peptides, non-peptide small molecules present in the venom have received comparatively little attention. Small molecules can have important functions within venoms; for example, in some spider species the main toxic components of the venom are acylpolyamines. Other molecules can have auxiliary effects that facilitate envenomation, such as purines with hypotensive properties utilised by snakes. In this study, we investigated some non-peptide small molecule constituents of Hormurus waigiensis venom using LC/MS, reversed-phase HPLC, and NMR spectroscopy. We identified adenosine, adenosine monophosphate (AMP), and citric acid within the venom, with low quantities of the amino acids glutamic acid and aspartic acid also being present. Purine nucleosides such as adenosine play important auxiliary functions in snake venoms when injected alongside other venom toxins, and they may have a similar role within H. waigiensis venom. Further research on these and other small molecules in scorpion venoms may elucidate their roles in prey capture and predator defence, and gaining a greater understanding of how scorpion venom components act in combination could allow for the development of improved first aid.

Plant evolution can mediate negative effects from honey bees on wild pollinators.

Pollinators are introduced to agroecosystems to provide pollination services. Introductions of managed pollinators often promote ecosystem services, but it remains largely unknown whether they also affect evolutionary mutualisms between wild pollinators and plants.Here, we developed a model to assess effects of managed honey bees on mutualisms between plants and wild pollinators. Our model tracked how interactions among wild pollinators and honey bees affected pollinator and plant populations.We show that when managed honey bees have a competitive advantage over wild pollinators, or a greater carrying capacity, the honey bees displace the wild pollinator. This leads to reduced plant density because plants benefit less by visits from honey bees than wild pollinators that coevolved with the plants.As wild pollinators are displaced, plants evolve by increasing investment in traits that are attractive for honey bees but not wild pollinators. This evolutionary switch promotes wild pollinator displacement. However, higher mutualism investment costs by the plant to the honey bee can promote pollinator coexistence.Our results show plant evolution can promote displacement of wild pollinators by managed honey bees, while limited plant evolution may lead to pollinator coexistence. More broadly, effects of honey bees on wild pollinators in agroecosystems, and effects on ecosystem services, may depend on the capacity of plant populations to evolve.

Highly diversified crop-livestock farming systems reshape wild bird communities.

Agricultural intensification is a leading threat to bird conservation. Highly diversified farming systems that integrate livestock and crop production might promote a diversity of habitats useful to native birds foraging across otherwise-simplified landscapes. At the same time, these features might be attractive to nonnative birds linked to a broad range of disservices to both crop and livestock production. We evaluated the influence of crop-livestock integration on wild bird richness and density along a north-south transect spanning the U.S. West Coast. We surveyed birds on 52 farms that grew primarily mixed vegetables and fruits alone or integrated livestock into production. Crop-livestock systems harbored higher native bird density and richness relative to crop-only farms, a benefit more pronounced on farms embedded in nonnatural landscapes. Crop-livestock systems bolstered native insectivores linked to the suppression of agricultural pest insects but did not bolster native granivores that may be more likely to damage crops. Crop-livestock systems also significantly increased the density of nonnative birds, primarily European Starlings (Sturnus vulgaris) and House Sparrows (Passer domesticus) that may compete with native birds for resources. Models supported a small, positive correlation between nonnative density and overall native bird density as well as between nonnative density and native granivore density. Relative to crop-only farms, on average, crop-livestock systems exhibited 1.5 times higher patch richness, 2.4 times higher density of farm structures, 7.3 times smaller field sizes, 2.4 times greater integration of woody crops, and 5.3 times greater integration of pasture/hay habitat on farm. Wild birds may have responded to this habitat diversity and/or associated food resources. Individual farm factors had significantly lower predictive power than farming system alone (change in C statistic information criterion (ΔCIC) = 80.2), suggesting crop-livestock systems may impact wild birds through a suite of factors that change with system conversion. Collectively, our findings suggest that farms that integrate livestock and crop production can attract robust native bird communities, especially within landscapes devoted to intensified food production. However, additional work is needed to demonstrate persistent farm bird communities through time, ecophysiological benefits to birds foraging on these farms, and net effects of both native and nonnative wild birds in agroecosystems.

A novel application of the Price equation reveals that landscape diversity promotes the response of bees to regionally rare plant species.

Bees are ecosystem service providers that are globally threatened by losses of plant diversity. However, effects of multi-species floral displays on bees in agro-ecosystems with variable landscape context remain poorly understood, hindering pollinator conservation tactics. We addressed this knowledge gap through a novel application of the modified Price equation to evaluate responses of bees to diverse floral communities on 36 farms in Washington, USA, over 3 years. We found that floral richness, not floral identity, was the best predictor of floral visits by bees. However, the benefits of regionally rare floral species (i.e. plants found at relatively few sites) were only fully realised when farms were embedded in diverse landscapes. Our analysis used the modified Price equation to demonstrate that plant diversity, rather than specific plant species, promotes pollinator visitation, and that diverse landscapes promote the response of pollinators to regionally rare plant species.

Environmental impacts of abrasive blasting of transmission towers in protected areas.

In Australia, and other parts of the world, tower infrastructure in electricity transmission networks are nearing the end of their asset life. In changing economic, political and regulatory environments Transmission Network Service Providers are implementing new approaches to asset management and reinvestment, such as refurbishment to extend the life of existing assets, instead of replacement. As part of these refurbishment efforts, abrasive blasting and recoating is being employed to remove corrosion and extend the life of steel electricity transmission towers. New controls and procedures have been developed to manage the most likely impacts associated with the abrasive blasting of transmission towers. However, little or no data have been available on the environmental impacts of abrasive blasting or the effectiveness of management procedures currently being used to mitigate potential adverse environmental impacts.We conducted an integrated study on the impacts of abrasive blasting, which brought together on-site research; modelling; and controlled laboratory trials. The study was undertaken during a transmission tower refurbishment project within the World Heritage listed Wet Tropics Region in Queensland, Australia. Measured metal deposition around towers due to blasting, was primarily as large particles (>PM10) at 12-30 m from the tower. Soil concentrations of metals were highest under towers, with a small number of samples showing elevated zinc at 12-30 m. The presence of spent abrasive media and dust on the geofabric material used under the towers and up to 15 m from the tower base, as part of control measures used to contain the abrasive products, indicates that deposition also occurs between 0 and 12 m from the tower.The potential impacts of the abrasive blasting technique on plants and invertebrates appear to be low. Five species of tropical rainforest tree seedlings exposed to abrasive blasting dust at worst-case levels had no negative impact on physiological performance or plant health. This research will assist Transmission Network Service Providers and other operators of corroded linear infrastructure to plan and implement mitigating management actions and procedures during abrasive blasting projects and assist regulators and the community to better understand the impacts of the practice.

Honey Bee (Apis mellifera) Pollen Foraging Reflects Benefits Dependent on Individual Infection Status.

Parasites often modify host foraging behavior, for example, by spurring changes to nutrient intake ratios or triggering self-medication. The gut parasite, Nosema ceranae, increases energy needs of the European or Western honey bee (Apis mellifera), but little is known about how infection affects foraging behavior. We used a combination of experiments and observations of caged and free-flying individual bees and hives to determine how N. ceranae affects honey bee foraging behavior. In an experiment with caged bees, we found that infected bees with access to a high-quality pollen were more likely to survive than infected bees with access to a lower quality pollen or no pollen. Non-infected bees showed no difference in survival with pollen quality. We then tested free-flying bees in an arena of artificial flowers and found that pollen foraging bees chose pollen commensurate with their infection status; twice as many infected bees selected the higher quality pollen than the lower quality pollen, while healthy bees showed no preference between pollen types. However, healthy and infected bees visited sucrose and pollen flowers in the same proportions. Among hive-level observations, we found no significant correlations between N. ceranae infection intensity in the hive and the proportion of bees returning with pollen. Our results indicate that N. ceranae-infected bees benefit from increased pollen quality and will selectively forage for higher quality while foraging for pollen, but infection status does not lead to increased pollen foraging at either the individual or hive levels.

Native turncoats and indirect facilitation of species invasions.

At local scales, native species can resist invasion by feeding on and competing with would-be invasive species. However, this relationship tends to break down or reverse at larger scales. Here, we consider the role of native species as indirect facilitators of invasion and their potential role in this diversity-driven 'invasion paradox'. We coin the term 'native turncoats' to describe native facilitators of non-native species and identify eight ways they may indirectly facilitate species invasion. Some are commonly documented, while others, such as indirect interactions within competitive communities, are largely undocumented in an invasion context. Therefore, we use models to evaluate the likelihood that these competitive interactions influence invasions. We find that native turncoat effects increase with the number of resources and native species. Furthermore, our findings suggest the existence, abundance and effectiveness of native turncoats in a community could greatly influence invasion success at large scales.

Trade-offs for butterfly alpha and beta diversity in human-modified landscapes and tropical rainforests.

The accelerating expansion of human populations and associated economic activity across the globe have made maintaining large, intact natural areas increasingly challenging. The difficulty of preserving large intact landscapes in the presence of growing human populations has led to a growing emphasis on landscape approaches to biodiversity conservation with a complementary strategy focused on improving conservation in human-modified landscapes. This, in turn, is leading to intense debate about the effectiveness of biodiversity conservation in human-modified landscapes and approaches to better support biodiversity in those landscapes. Here, we compared butterfly abundance, alpha richness, and beta diversity in human-modified landscapes (urban, sugarcane) and natural, forested areas to assess the conservation value of human-modified landscapes within the Wet Tropics bioregion of Australia. We used fruit-baited traps to sample butterflies and analyzed abundance and species richness in respective land uses over a one-year period. We also evaluated turnover and spatial variance components of beta diversity to determine the extent of change in temporal and spatial variation in community composition. Forests supported the largest numbers of butterflies, but were lowest in each, alpha species richness, beta turnover, and the spatial beta diversity. Sugarcane supported higher species richness, demonstrating the potential for conservation at local scales in human-modified landscapes. In contrast, beta diversity was highest in urban areas, likely driven by spatial and temporal variation in plant composition within the urban landscapes. Thus, while improving conservation on human-modified landscapes may improve local alpha richness, conserving variation in natural vegetation is critical for maintaining high beta diversity.

Biological control agents in the Anthropocene: current risks and future options.

Global climate change is often expected to disrupt biological control. Predicting the effects of climate change on biological control, and identifying natural enemies that will thrive in future climate scenarios, is thus essential to ensure agricultural sustainability. To promote biological control under climate change, land managers should prioritise the conservation of natural enemy diversity to ensure some effective natural enemies are always present despite often-unpredictable climate scenarios. In addition, ecophysiological and habitat domain models should be combined to predict climate change-induced shifts in predation by diverse predator communities. Finally, insights from land managers during extreme weather events, such as droughts and heat waves, may be invaluable in the effort to identify key biological control agents for future scenarios.

Changes in predator exposure, but not in diet, induce phenotypic plasticity in scorpion venom.

Animals embedded between trophic levels must simultaneously balance pressures to deter predators and acquire resources. Venomous animals may use venom toxins to mediate both pressures, and thus changes in this balance may alter the composition of venoms. Basic theory suggests that greater exposure to a predator should induce a larger proportion of defensive venom components relative to offensive venom components, while increases in arms races with prey will elicit the reverse. Alternatively, reducing the need for venom expenditure for food acquisition, for example because of an increase in scavenging, may reduce the production of offensive venom components. Here, we investigated changes in scorpion venom composition using a mesocosm experiment where we manipulated scorpions' exposure to a surrogate vertebrate predator and live and dead prey. After six weeks, scorpions exposed to surrogate predators exhibited significantly different venom chemistry compared with naive scorpions. This change included a relative increase in some compounds toxic to vertebrate cells and a relative decrease in some compounds effective against their invertebrate prey. Our findings provide, to our knowledge, the first evidence for adaptive plasticity in venom composition. These changes in venom composition may increase the stability of food webs involving venomous animals.

A spatial theory for emergent multiple predator-prey interactions in food webs.

Predator-prey interaction is inherently spatial because animals move through landscapes to search for and consume food resources and to avoid being consumed by other species. The spatial nature of species interactions necessitates integrating spatial processes into food web theory and evaluating how predators combine to impact their prey. Here, we present a spatial modeling approach that examines emergent multiple predator effects on prey within landscapes. The modeling is inspired by the habitat domain concept derived from empirical synthesis of spatial movement and interactions studies. Because these principles are motivated by synthesis of short-term experiments, it remains uncertain whether spatial contingency principles hold in dynamical systems. We address this uncertainty by formulating dynamical systems models, guided by core habitat domain principles, to examine long-term multiple predator-prey spatial dynamics. To describe habitat domains, we use classical niche concepts describing resource utilization distributions, and assume species interactions emerge from the degree of overlap between species. The analytical results generally align with those from empirical synthesis and present a theoretical framework capable of demonstrating multiple predator effects that does not depend on the small spatial or temporal scales typical of mesocosm experiments, and help bridge between empirical experiments and long-term dynamics in natural systems.

Does Biodiversity-Ecosystem Function Literature Neglect Tropical Ecosystems?

Current evidence suggests that there is a positive relationship between biodiversity and ecosystem functioning, but few studies have addressed tropical ecosystems where the highest levels of biodiversity occur. We develop two hypotheses for the implications of generalizing from temperate studies to tropical ecosystems, and discuss the need for more tropical research.

Increased pollinator habitat enhances cacao fruit set and predator conservation.

The unique benefits of wild pollinators to the productivity of agricultural crops have become increasingly recognized in recent decades. However, declines in populations of wild pollinator species, largely driven by the conversion of natural habitat to agricultural land and broad-spectrum pesticide use often lead reductions in the provision of pollination services and crop production. With growing evidence that targeted pollinator conservation improves crop yield and/or quality, particularly for pollination specialist crops, efforts are increasing to substitute agriculturally intensive practices with those that alleviate some of the negative impacts of agriculture on pollinators and the pollination services they provide, in part through the provision of suitable pollinator habitat. Further, similarities between the responses of some pollinators and predators to habitat management suggest that efforts to conserve pollinators may also encourage predator densities. We evaluated the effects of one habitat management practice, the addition of cacao fruit husks to a monoculture cacao farm, on the provision of pollination services and the densities of two groups of entomophagous predators. We also evaluated the impacts of cacao fruit husk addition on pollen limitation, by crossing this habitat manipulation with pollen supplementation treatments. The addition of cacao fruit husks increased the number of fruits per tree and along with hand pollination treatments, increased final yields indicating a promotion of the pollination ecosystem service provided by the specialist pollinators, midges. We also found that cacao fruit husk addition increased the densities of two predator groups, spiders and skinks. Further, the conservation of these predators did not inhibit pollination through pollinator capture or deterrence. The findings show that, with moderate habitat management, both pollinator and predator conservation can be compatible goals within a highly specialized plant-pollinator system. The effectiveness of this habitat manipulation may be attributable to the increased availability of alternative habitat and food resources for both pollinators and predators. The results exemplify a win-win relationship between agricultural production and biological conservation, whereby agricultural practices to support vital pollinators and pollination services can increase production as well as support species conservation.