Assessing the ecological niche and invasion potential of the Asian giant hornet.

The Asian giant hornet (Vespa mandarinia) was recently detected in western British Columbia, Canada and Washington State, United States. V. mandarinia are an invasion concern due to their ability to kill honey bees and affect humans. Here, we used habitat suitability models and dispersal simulations to assess potential invasive spread of V. mandarinia We show V. mandarinia are most likely to establish in areas with warm to cool annual mean temperature, high precipitation, and high human activity. The realized niche of introduced populations is small compared to native populations, suggesting introduced populations could spread into habitats across a broader range of environmental conditions. Dispersal simulations also show that V. mandarinia could rapidly spread throughout western North America without containment. Given its potential negative impacts and capacity for spread, extensive monitoring and eradication efforts throughout western North America are warranted.

Landscape context affects the sustainability of organic farming systems.

Organic agriculture promotes sustainability compared to conventional agriculture. However, the multifunctional sustainability benefits of organic farms might be mediated by landscape context. Assessing how landscape context affects sustainability may aid in targeting organic production to landscapes that promote high biodiversity, crop yields, and profitability. We addressed this using a meta-analysis spanning 60 crop types on six continents that assessed whether landscape context affected biodiversity, yield, and profitability of organic vs. conventional agroecosystems. We considered landscape metrics reflecting landscape composition (percent cropland), compositional heterogeneity (number and diversity of cover types), and configurational heterogeneity (spatial arrangement of cover types) across our study systems. Organic sites had greater biodiversity (34%) and profits (50%) than conventional sites, despite lower yields (18%). Biodiversity gains increased as average crop field size in the landscape increased, suggesting organic farms provide a "refuge" in intensive landscapes. In contrast, as crop field size increased, yield gaps between organic and conventional farms increased and profitability benefits of organic farming decreased. Profitability of organic systems, which we were only able to measure for studies conducted in the United States, varied across landscapes in conjunction with production costs and price premiums, suggesting socioeconomic factors mediated profitability. Our results show biodiversity benefits of organic farming respond differently to landscape context compared to yield and profitability benefits, suggesting these sustainability metrics are decoupled. More broadly, our results show that the ecological, but not the economic, sustainability benefits of organic agriculture are most pronounced in more intensive agricultural landscapes.

Factors affecting virus prevalence in honey bees in the Pacific-Northwest, USA.

Global efforts to assess honey bee health show viruses are major stressors that undermine colony performance. Identifying factors that affect virus incidence, such as management practices and landscape context, could aid in slowing virus transmission. Here we surveyed viruses in honey bees from 86 sites in the Pacific Northwest, USA, and tested effects of regional bee density, movement associated with commercial pollination, julian date, and hive management on virus prevalence. We also explored patterns of virus co-occurrence and spatial autocorrelation to identify whether local transmission was a primary driver of pathogen distribution. Our surveys found widespread prevalence of Deformed wing virus (DWV), Sacbrood virus (SBV), and Black queen cell virus (BQCV). BQCV and SBV were most prolific in commercial apiaries, while Chronic bee paralysis virus (CPBV) was more common in hobbyist apiaries than commercial apiaries. DWV was most common in urban landscapes and was best predicted by mite prevalence and julian date, while the incidence of both SBV and BQCV were best predicted by regional apiary density. We did not find evidence of additional spatial autocorrelation for any viruses, although high co-occurrence suggests parallel transmission patterns. Our results support the importance of mite management in slowing virus spread and suggest that greater bee density increases transmission. Our study provides support that viruses are widespread in honey bees and connects known mechanisms of virus transmission to the distribution of pathogens observed across the Pacific Northwest.

Orchard Management and Landscape Context Mediate the Pear Floral Microbiome.

Crop-associated microbiota are a key factor affecting host health and productivity. Most crops are grown within heterogeneous landscapes, and interactions between management practices and landscape context often affect plant and animal biodiversity in agroecosystems. However, whether these same factors typically affect crop-associated microbiota is less clear. Here, we assessed whether orchard management strategies and landscape context affected bacterial and fungal communities in pear (Pyrus communis) flowers. We found that bacteria and fungi responded differently to management schemes. Organically certified orchards had higher fungal diversity in flowers than conventional or bio-based integrated pest management (IPM) orchards, but organic orchards had the lowest bacterial diversity. Orchard management scheme also best predicted the distribution of several important bacterial and fungal genera that either cause or suppress disease; organic and bio-based IPM best explained the distributions of bacterial and fungal genera, respectively. Moreover, patterns of bacterial and fungal diversity were affected by interactions between management, landscape context, and climate. When examining the similarity of bacterial and fungal communities across sites, both abundance- and taxon-related turnovers were mediated primarily by orchard management scheme and landscape context and, specifically, the amount of land in cultivation. Our study reveals local- and landscape-level drivers of floral microbiome structure in a major fruit crop, providing insights that can inform microbiome management to promote host health and high-yielding quality fruit. IMPORTANCE Proper crop management during bloom is essential for producing disease-free tree fruit. Tree fruits are often grown in heterogeneous landscapes; however, few studies have assessed whether landscape context and crop management affect the floral microbiome, which plays a critical role in shaping plant health and disease tolerance. Such work is key for identification of tactics and/or contexts where beneficial microbes proliferate and pathogenic microbes are limited. Here, we characterize the floral microbiome of pear crops in Washington State, where major production occurs in intermountain valleys and basins with variable elevation and microclimates. Our results show that both local-level (crop management) and landscape-level (habitat types and climate) factors affect floral microbiota but in disparate ways for each kingdom. More broadly, these findings can potentially inform microbiome management in orchards for promotion of host health and high-quality yields.

Responses of pea plants to multiple antagonists are mediated by order of attack and phytohormone crosstalk.

Plants are often attacked by multiple antagonists and traits of the attacking organisms and their order of arrival onto hosts may affect plant defences. However, few studies have assessed how multiple antagonists, and varying attack order, affect plant defence or nutrition. To address this, we assessed defensive and nutritional responses of Pisum sativum plants after attack by a vector herbivore (Acrythosiphon pisum), a nonvector herbivore (Sitona lineatus), and a pathogen (Pea enation mosaic virus, PEMV). We show viruliferous A. pisum induced several antipathogen plant defence signals, but these defences were inhibited by S. lineatus feeding on peas infected with PEMV. In contrast, S. lineatus feeding induced antiherbivore defence signals, and these plant defences were enhanced by PEMV. Sitona lineatus also increased abundance of plant amino acids, but only when they attacked after viruliferous A. pisum. Our results suggest that diverse communities of biotic antagonists alter defence and nutritional traits of plants through complex pathways that depend on the identity of attackers and their order of arrival onto hosts. Moreover, we show interactions among a group of biotic stressors can vary along a spectrum from antagonism to enhancement/synergism based on the identity and order of attackers, and these interactions are mediated by a multitude of phytohormone pathways.

Vector-borne plant pathogens modify top-down and bottom-up effects on insect herbivores.

Ecological theory predicts that host-plant traits affect herbivore population growth rates, which in turn modulates predator-prey interactions. However, while vector-borne plant pathogens often alter traits of both host plants and vectors, a few studies have assessed how pathogens may act as interaction modifiers within tri-trophic food webs. By applying a food web motif framework, we assessed how a vector-borne plant pathogen (Pea-enation mosaic virus, PEMV) modified both bottom-up (plant-herbivore) and top-down (predator-prey) interactions. Specifically, we assessed trophic interactions with PEMV-infectious Acyrthosiphon pisum (pea aphid) vectors compared to non-infectious aphids in a factorial experiment that manipulated predator and plant communities. We show that PEMV altered bi-trophic relationships, whereby on certain plant species, PEMV reduced vector performance but also increased their susceptibility to predators. However, on other plant species, PEMV weakened top-down control or increased vector performance. Our results suggest that vector-borne plant pathogens are important interaction modifiers for plant-herbivore-predator dynamics: host-plant response to viruses can decrease herbivore abundance by reducing herbivore performance, but also increase herbivore abundance by weakening top-down control. Broadly speaking, trophic interactions that regulate herbivore outbreaks appear to be modified for herbivores actively transmitting viruses to host plants. Consequently, management and monitoring of outbreaking herbivores should consider the infection status of focal populations.

Primacy of plants in driving the response of arthropod communities to drought.

Drought threatens arthropod communities worldwide. Water limitation affects the quantity and quality of plants available to herbivores as food, and can also affect higher trophic-level consumers through variability in prey quality and reduced availability of suitable habitats. Our study assessed the response of an arthropod community to water limited wheat (Triticum aestivum L.) in a field setting. We used rainout shelters to exclude precipitation, irrigated raised bed plots to create three levels of water availability, and monitored arthropod community development over 8 weeks. First, we compared arthropod communities in habitats with different levels of water limitation and found that community composition was reliant on the magnitude of the water stress. This difference was largely due to the loss of piercing-sucking herbivores and predators in high-stress environments. Next, we focused on aphids and their natural enemies to investigate the underlying mechanisms driving community responses using structural equation modeling (SEM). Aphid abundance was negatively affected by water limitation, and this response was primarily associated with stress-induced plant physiological changes and not plant biomass or natural enemy abundance. Natural enemy abundance was also reduced in water-limited habitats, but natural enemies responded to plant biomass and not prey availability. These effects were exacerbated as water stress increased. The absence of natural enemy effects on aphids indicates that top-down predation effects were dampened by strong bottom-up effects of plant water limitation. This study revealed the importance of considering water stress intensity when predicting outcomes of droughts for arthropod communities.

Responses to predation risk cues and alarm pheromones affect plant virus transmission by an aphid vector.

Herbivores assess predation risk in their environment by identifying visual, chemical, and tactile predator cues. Detection of predator cues can induce risk-avoidance behaviors in herbivores that affect feeding, dispersal, and host selection in ways that minimize mortality and reproductive costs. For herbivores that transmit plant pathogens, including many aphids, changes in herbivore behavior in response to predator cues may also affect pathogen spread. However, few studies have assessed how aphid behavioral responses to different types of predator cues affect pathogen transmission. Here, we conducted greenhouse experiments to assess whether responses of pea aphids (Acyrthosiphon pisum) to predation risk and alarm pheromone (E-ß-Farnesene), an aphid alarm signal released in response to predation risk, affected transmission of Pea enation mosaic virus (PEMV). We exposed A. pisum individuals to risk cues, and quantified viral titer in aphids and pea (Pisum sativum) host plants across several time periods. We also assessed how A. pisum responses to risk cues affected aphid nutrition, reproduction, and host selection. We show that exposure to predator cues and alarm pheromone significantly reduced PEMV acquisition and inoculation. Although vectors avoided hosts with predator cues, predator cues did not alter vector reproduction or reduce nutrient acquisition. Overall, these results suggest that non-consumptive effects of predators may indirectly decrease the spread of plant pathogens by altering vector behavior in ways that reduce vector competence and pathogen transmission efficiency.

Landscape structure and climate drive population dynamics of an insect vector within intensely managed agroecosystems.

Characterizing factors affecting insect pest populations across variable landscapes is a major challenge for agriculture. In natural ecosystems, insect populations are strongly mediated by landscape and climatic factors. However, it has proven difficult to evaluate if similar factors predict pest dynamics in agroecosystems because control tactics exert strong confounding effects. We addressed this by assessing whether species distribution models could effectively characterize dynamics of an insect pest in intensely managed agroecosystems. Our study used a regional multi-year data set to assess landscape and climatic drivers of potato psyllid (Bactericera cockerelli) populations, which are often subjected to calendar-based insecticide treatments because they transmit pathogens to crops. Despite this, we show that psyllid populations were strongly affected by landscape and climatic factors. Psyllids were more abundant in landscapes with high connectivity, low crop diversity, and large natural areas. Psyllid population dynamics were also mediated by climatic factors, particularly precipitation and humidity. Our results show that many of the same factors that drive insect population dynamics in natural ecosystems can have similar effects in an intensive agroecosystem. More broadly, our study shows that models incorporating landscape and climatic factors can describe pest populations in agroecosystems and may thus promote more sustainable pest management.

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.

Plant-mediated interactions between a vector and a non-vector herbivore promote the spread of a plant virus.

Herbivores that transmit plant pathogens often share hosts with non-vector herbivores. These co-occurring herbivores can affect vector fitness and behaviour through competition and by altering host plant quality. However, few studies have examined how such interactions may both directly and indirectly influence the spread of a plant pathogen. Here, we conducted field and greenhouse trials to assess whether a defoliating herbivore (Sitona lineatus) mediated the spread of a plant pathogen, Pea enation mosaic virus (PEMV), by affecting the fitness and behaviour of Acrythosiphon pisum, the PEMV vector. We observed higher rates of PEMV spread when infectious A. pisum individuals shared hosts with S. lineatus individuals. Using structural equation models, we showed that herbivory from S. lineatus increased A. pisum fitness, which stimulated vector movement and PEMV spread. Moreover, plant susceptibility to PEMV was indirectly enhanced by S. lineatus, which displaced A. pisum individuals to the most susceptible parts of the plant. Subsequent analyses of plant defence genes revealed considerable differences in plant phytohormones associated with anti-herbivore and anti-pathogen defence when S. lineatus was present. Given that vectors interact with non-vector herbivores in natural and managed ecosystems, characterizing how such interactions affect pathogens would greatly enhance our understanding of disease ecology.

Reciprocal plant-mediated interactions between a virus and a non-vector herbivore.

Vector-borne viruses alter many physical and chemical traits of their plant hosts, indirectly affecting the fitness and behavior of vectors in ways that promote virus transmission. However, it is unclear whether viruses induce plant-mediated shifts in the behavior and fitness of non-vector herbivores, or if non-vectors affect the dynamics of vector-borne viruses. Here we evaluated reciprocal interactions between Pea enation mosaic virus (PEMV), a pathogen transmitted by the aphid Acrythosiphon pisum, and a non-vector weevil, Sitona lineatus. In the field, PEMV-infected plants experienced more defoliation from S. lineatus than uninfected plants; behavioral assays similarly showed S. lineatus adults preferred to feed on infected plants. In turn, infectious A. pisum preferred plants damaged by S. lineatus, and S. lineatus herbivory led to increased PEMV titer. These interactions may be mediated by plant phytohormone levels, as S. lineatus induced jasmonic acid, while PEMV induced salicylic acid. Levels of abscisic acid were not affected by attack from either PEMV or S. lineatus alone, but plants challenged by both had elevated levels of this phytohormone. As plant viruses and their vectors often exist in diverse communities, our study highlights the importance of non-vector species in influencing plant pathogens and their vectors through host-mediated effects.

Financial competitiveness of organic agriculture on a global scale.

To promote global food and ecosystem security, several innovative farming systems have been identified that better balance multiple sustainability goals. The most rapidly growing and contentious of these systems is organic agriculture. Whether organic agriculture can continue to expand will likely be determined by whether it is economically competitive with conventional agriculture. Here, we examined the financial performance of organic and conventional agriculture by conducting a meta-analysis of a global dataset spanning 55 crops grown on five continents. When organic premiums were not applied, benefit/cost ratios (-8 to -7%) and net present values (-27 to -23%) of organic agriculture were significantly lower than conventional agriculture. However, when actual premiums were applied, organic agriculture was significantly more profitable (22-35%) and had higher benefit/cost ratios (20-24%) than conventional agriculture. Although premiums were 29-32%, breakeven premiums necessary for organic profits to match conventional profits were only 5-7%, even with organic yields being 10-18% lower. Total costs were not significantly different, but labor costs were significantly higher (7-13%) with organic farming practices. Studies in our meta-analysis accounted for neither environmental costs (negative externalities) nor ecosystem services from good farming practices, which likely favor organic agriculture. With only 1% of the global agricultural land in organic production, our findings suggest that organic agriculture can continue to expand even if premiums decline. Furthermore, with their multiple sustainability benefits, organic farming systems can contribute a larger share in feeding the world.

Restriction of Francisella novicida genetic diversity during infection of the vector midgut.

The genetic diversity of pathogens, and interactions between genotypes, can strongly influence pathogen phenotypes such as transmissibility and virulence. For vector-borne pathogens, both mammalian hosts and arthropod vectors may limit pathogen genotypic diversity (number of unique genotypes circulating in an area) by preventing infection or transmission of particular genotypes. Mammalian hosts often act as "ecological filters" for pathogen diversity, where novel variants are frequently eliminated because of stochastic events or fitness costs. However, whether vectors can serve a similar role in limiting pathogen diversity is less clear. Here we show using Francisella novicida and a natural tick vector of Francisella spp. (Dermacentor andersoni), that the tick vector acted as a stronger ecological filter for pathogen diversity compared to the mammalian host. When both mice and ticks were exposed to mixtures of F. novicida genotypes, significantly fewer genotypes co-colonized ticks compared to mice. In both ticks and mice, increased genotypic diversity negatively affected the recovery of available genotypes. Competition among genotypes contributed to the reduction of diversity during infection of the tick midgut, as genotypes not recovered from tick midguts during mixed genotype infections were recovered from tick midguts during individual genotype infection. Mediated by stochastic and selective forces, pathogen genotype diversity was markedly reduced in the tick. We incorporated our experimental results into a model to demonstrate how vector population dynamics, especially vector-to-host ratio, strongly affected pathogen genotypic diversity in a population over time. Understanding pathogen genotypic population dynamics will aid in identification of the variables that most strongly affect pathogen transmission and disease ecology.

Organic agriculture promotes evenness and natural pest control.

Human activity can degrade ecosystem function by reducing species number (richness) and by skewing the relative abundance of species (evenness). Conservation efforts often focus on restoring or maintaining species number, reflecting the well-known impacts of richness on many ecological processes. In contrast, the ecological effects of disrupted evenness have received far less attention, and developing strategies for restoring evenness remains a conceptual challenge. In farmlands, agricultural pest-management practices often lead to altered food web structure and communities dominated by a few common species, which together contribute to pest outbreaks. Here we show that organic farming methods mitigate this ecological damage by promoting evenness among natural enemies. In field enclosures, very even communities of predator and pathogen biological control agents, typical of organic farms, exerted the strongest pest control and yielded the largest plants. In contrast, pest densities were high and plant biomass was low when enemy evenness was disrupted, as is typical under conventional management. Our results were independent of the numerically dominant predator or pathogen species, and so resulted from evenness itself. Moreover, evenness effects among natural enemy groups were independent and complementary. Our results strengthen the argument that rejuvenation of ecosystem function requires restoration of species evenness, rather than just richness. Organic farming potentially offers a means of returning functional evenness to ecosystems.

Large-scale, spatially-explicit test of the refuge strategy for delaying insecticide resistance.

The refuge strategy is used worldwide to delay the evolution of pest resistance to insecticides that are either sprayed or produced by transgenic Bacillus thuringiensis (Bt) crops. This strategy is based on the idea that refuges of host plants where pests are not exposed to an insecticide promote survival of susceptible pests. Despite widespread adoption of this approach, large-scale tests of the refuge strategy have been problematic. Here we tested the refuge strategy with 8 y of data on refuges and resistance to the insecticide pyriproxyfen in 84 populations of the sweetpotato whitefly (Bemisia tabaci) from cotton fields in central Arizona. We found that spatial variation in resistance to pyriproxyfen within each year was not affected by refuges of melons or alfalfa near cotton fields. However, resistance was negatively associated with the area of cotton refuges and positively associated with the area of cotton treated with pyriproxyfen. A statistical model based on the first 4 y of data, incorporating the spatial distribution of cotton treated and not treated with pyriproxyfen, adequately predicted the spatial variation in resistance observed in the last 4 y of the study, confirming that cotton refuges delayed resistance and treated cotton fields accelerated resistance. By providing a systematic assessment of the effectiveness of refuges and the scale of their effects, the spatially explicit approach applied here could be useful for testing and improving the refuge strategy in other crop-pest systems.

Effects of Neonicotinoids and Crop Rotation for Managing Wireworms in Wheat Crops.

Soil-dwelling insects are severe pests in many agroecosystems. These pests have cryptic life cycles, making sampling difficult and damage hard to anticipate. The management of soil insects is therefore often based on preventative insecticides applied at planting or cultural practices. Wireworms, the subterranean larvae of click beetles (Coleoptera: Elateridae), have re-emerged as problematic pests in cereal crops in the Pacific Northwestern United States. Here, we evaluated two management strategies for wireworms in long-term field experiments: 1) treating spring wheat seed with the neonicotinoid thiamethoxam and 2) replacing continuous spring wheat with a summer fallow and winter wheat rotation. Separate experiments were conducted for two wireworm species--Limonius californicus (Mannerheim) and Limonius infuscatus (Motschulsky). In the experiment with L. californicus, spring wheat yields and economic returns increased by 24-30% with neonicotinoid treatments. In contrast, in the experiment with L. infuscatus, spring wheat yields and economic returns did not increase with neonicotinoids despite an 80% reduction in wireworms. Thus, the usefulness of seed-applied neonicotinoids differed based on the wireworm species present. In experiments with both species, we detected significantly fewer wireworms with a no-till summer fallow and winter wheat rotation compared with continuous spring wheat. This suggests that switching from continuous spring wheat to a winter wheat and summer fallow rotation may aid in wireworm management. More generally, our results show that integrated management of soil-dwelling pests such as wireworms may require both preventative insecticide treatments and cultural practices.

Phytoseiids in Washington commercial apple orchards: biodiversity and factors affecting abundance.

Galendromus occidentalis (Nesbitt) is an important biological control agent of spider mites (Acari: Tetranychidae) in Washington apple orchards. It was thought to be essentially the sole phytoseiid existing in this system, due in part to its resistance to commonly used orchard pesticides, and organophosphates in particular. To test this assumption, we conducted a survey of 102 commercial apple blocks in Washington to characterize the community of phytoseiid species. Seven phytoseiid species were found in our samples; G. occidentalis and Amblydromella caudiglans (Schuster) were found in the greatest abundance. We hypothesized that the gradual shift away from the use of organophosphates in recent decades may have caused the change in phytoseiid community structure. The survey data and information regarding the management, location, and surrounding habitat of each block were used to determine what factors affect phytoseiid abundances. Galendromus occidentalis abundance was positively affected by the use of conventional (vs. organic) spray programs, and the use of the acaricide bifenazate. Amblydromella caudiglans abundance was negatively affected by bifenazate use and positively affected by herbicide strip weediness; it was also less prevalent in 'Golden Delicious' blocks compared to other cultivars. These results indicate that A. caudiglans reaches higher abundances in orchards that lack certain agricultural disturbances, whereas G. occidentalis can survive in more disturbed environments. Surveys of this nature can provide valuable insight to potential drivers of community structure, allowing for the improvement of integrated pest management programs that incorporate conservation of newly recognized biological control agents such as A. caudiglans.

Effects of reproductive interference on the competitive displacement between two invasive whiteflies.

Reproductive interference is one of the major factors mediating species exclusion among insects. The cryptic species Middle East-Asia Minor 1 (MEAM1) and Mediterranean (MED) of the whitefly Bemisia tabaci complex have invaded many parts of the world and often exhibit niche overlap and reproductive interference. However, contrasting patterns of competitive displacement between the two invaders have been observed between regions such as those in USA and China. Understanding the roles of reproductive interference in competitive interactions between populations of the two species in different regions will help unravel other factors related to their invasion. We integrated laboratory population experiments, behavioural observations and simulation modelling to investigate the role of reproductive interference on species exclusion between MEAM1 and MED in China. In mixed cohorts of the two species MEAM1 always excluded MED in a few generations when the initial proportion of MEAM1 was ⩾0.25. Even when the initial proportion of MEAM1 was only 0.10, however, MEAM1 still had a higher probability of excluding MED than that for MED to exclude MEAM1. Importantly, we show that as MEAM1 increased in relative abundance, MED populations became increasingly male-biased. Detailed behavioural observations confirmed that MEAM1 showed a stronger reproductive interference than MED, leading to reduced frequency of copulation and female progeny production in MED. Using simulation modelling, we linked our behavioural observations with exclusion experiments to show that interspecific asymmetric reproductive interference predicts the rate of species exclusion of MED by MEAM1. These findings not only reveal the importance of reproductive interference in the competitive interactions between the two invasive whiteflies as well as the detailed behavioural mechanisms, but also provide a valuable framework against which the effects of other factors mediating species exclusion can be explored.

Effects of Soil Rhizobia Abundance on Interactions between a Vector, Pathogen, and Legume Plant Host.

Soil rhizobia promote nitrogen fixation in legume hosts, maximizing their tolerance to different biotic stressors, plant biomass, crop growth, and yield. While the presence of soil rhizobia is considered beneficial for plants, few studies have assessed whether variation in rhizobia abundance affects the tolerance of legumes to stressors. To address this, we assessed the effects of variable soil rhizobia inoculum concentrations on interactions between a legume host (Pisum sativum), a vector insect (Acyrthosiphon pisum), and a virus (Pea enation mosaic virus, PEMV). We showed that increased rhizobia abundance reduces the inhibitory effects of PEMV on the nodule formation and root growth in 2-week-old plants. However, these trends were reversed in 4-week-old plants. Rhizobia abundance did not affect shoot growth or virus prevalence in 2- or 4-week-old plants. Our results show that rhizobia abundance may indirectly affect legume tolerance to a virus, but effects varied based on plant age. To assess the mechanisms that mediated interactions between rhizobia, plants, aphids, and PEMV, we measured the relative expression of gene transcripts related to plant defense signaling. Rhizobia concentrations did not strongly affect the expression of defense genes associated with phytohormone signaling. Our study shows that an abundance of soil rhizobia may impact a plant's ability to tolerate stressors such as vector-borne pathogens, as well as aid in developing sustainable pest and pathogen management systems for legume crops. More broadly, understanding how variable rhizobia concentrations can optimize legume-rhizobia symbiosis may enhance the productivity of legume crops.