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Spatial variation in plant community composition is an important driver of variation in susceptibility to herbivores. In close proximity, certain neighbors can attract or repel herbivores to a focal plant ("associational effects"). Neighboring plants may also compete for resources, modifying their phenotype in ways that affect susceptibility to herbivores. To test whether and how competition contributes to associational effects, we manipulated the sharing of belowground resources among plant neighbors (spotted Joe Pye weed and common boneset) that serve as alternate hosts for an herbivorous beetle. In the field, the beetle Ophraella notata laid more eggs and inflicted more damage on plants of both species that were released from belowground competition with neighbors. Competition also weakened the effects of neighbor identity during field trials, reducing associational susceptibility. When beetles were forced to choose between the two host species in cage trials, competition again reduced beetle use of Joe Pye weed as a secondary host. To test the role of plant traits related to herbivore defense and nutrition, we quantified leaf protein, specific leaf area, and trichomes, and conducted behavioral assays on leaf disks. Beetles did not distinguish between Joe Pye weed treatments at the leaf disk level, and competition did not impact specific leaf area and protein. Trichome density was higher in both species in the preferred treatment. Overall, our results suggest that belowground interactions between plants may mediate the strength of associational effects, as secondary hosts become more attractive when released from competition with primary host plants., (© 2023. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Turnip mosaic virus (TuMV) is a widespread and economically important pathogen in agricultural crops and has the widest known host range in the virus family Potyviridae. While management of the virus and its aphid vectors in agricultural fields decreases virus incidence, many alternative wild hosts for TuMV may serve as source populations for crop infection through spillover. Over thirty years ago, research demonstrated that the introduced brassica, Dame's Rocket (Hesperis matronalis) hosts several viruses, including TuMV. Here, we use both enzyme-linked immunosorbent assays (ELISA) and next generation sequencing to document the frequent infection by TuMV of Dame's Rocket, which is common and widespread in disturbed areas around crop fields in upstate New York. Deep sequencing of multiple tissue types of symptomatic hosts indicate that the infection is systemic and causes diagnostic, visible symptoms. In a common garden experiment using host populations from across upstate New York, we found evidence for genetic tolerance to TuMV infection in H. matronalis. Field surveys show that TuMV prevalence varies across populations, but is generally higher in agricultural areas. Examining disease dynamics in this and other common alternative hosts will enhance our understanding of TuMV epidemiology and, more broadly, virus distribution in wild plants., Competing Interests: Declaration of Competing Interest All authors declare no competing interests., (Copyright © 2022 The Author(s). Published by Elsevier B.V. All rights reserved.)

Our understanding of the ecological interactions between plant viruses, their insect vectors, and their host plants has increased rapidly over the past decade. The suite of viruses known collectively as the yellow dwarf viruses infect an extensive range of cultivated and noncultivated grasses worldwide and is one of the best-studied plant virus systems. The yellow dwarf viruses are ubiquitous in cereal crops, where they can significantly limit yields, and there is growing recognition that they are also ubiquitous in grassland ecosystems, where they can influence community dynamics. Here, we discuss recent research that has explored ( a ) the extent and impact of yellow dwarf viruses in a diversity of plant communities, ( b ) the role of vector behavior in virus transmission, and ( c ) the prospects for impacts of climate change-including rising temperatures, drought, and elevated CO 2 -on the epidemiology of yellow dwarf viruses.

Organic agriculture outperforms conventional agriculture across several sustainability metrics due, in part, to more widespread use of agroecological practices. However, increased entry of large-scale farms into the organic sector has prompted concerns about 'conventionalization' through input substitution, agroecosystem simplification and other changes. We examined this shift in organic agriculture by estimating the use of agroecological practices across farm size and comparing indicators of conventionalization. Results from our national survey of 542 organic fruit and vegetable farmers show that fewer agroecological practices were used on large farms, which also exhibited the greatest degree of conventionalization. Intercropping, insectary plantings and border plantings were at least 1.4 times more likely to be used on small (0.4-39 cropland ha) compared with large (≥405 cropland ha) farms, whereas reduced tillage was less likely and riparian buffers were more likely on small compared with medium (40-404 cropland ha) farms. Because decisions about management practices can drive environmental sustainability outcomes, policy should support small and medium farms that already use agroecological practices while encouraging increased use of agroecological practices on larger farms., (© 2022. The Author(s), under exclusive licence to Springer Nature Limited.)

Predator-prey interactions influence prey traits through both consumptive and non-consumptive effects, and variation in these traits can shape vector-borne disease dynamics. Meta-analysis methods were employed to generate predation effect sizes by different categories of predators and mosquito prey. This analysis showed that multiple families of aquatic predators are effective in consumptively reducing mosquito survival, and that the survival of Aedes , Anopheles , and Culex mosquitoes is negatively impacted by consumptive effects of predators. Mosquito larval size was found to play a more important role in explaining the heterogeneity of consumptive effects from predators than mosquito genus. Mosquito survival and body size were reduced by non-consumptive effects of predators, but development time was not significantly impacted. In addition, Culex vectors demonstrated predator avoidance behavior during oviposition. The results of this meta-analysis suggest that predators limit disease transmission by reducing both vector survival and vector size, and that associations between drought and human West Nile virus cases could be driven by the vector behavior of predator avoidance during oviposition. These findings are likely to be useful to infectious disease modelers who rely on vector traits as predictors of transmission., Competing Interests: MR, CH, ZG, CR, FE, ME, TD, NG, SH, AP, AM No competing interests declared, (© 2022, Russell et al.)

Legumes are used in crop rotations by both large-scale and smallholder farmers alike to increase soil fertility, especially before high-nitrogen-demanding crops such as corn (maize). Legume crop residues and green manures are rich in nitrogen due to mutualistic rhizobia, bacteria that live in their roots and convert atmospheric nitrogen into a biologically available form. Growers can obtain recommendations from local extension offices about how much less inorganic nitrogen fertilizer needs to be added to a subsequent crop following different legume break crops for the predominant soil type (the nitrogen fertilizer replacement value, or NFRV). Due to the intimate relationship between legumes and rhizobia, conditions that affect plant health can also affect the rhizobia and how much nitrogen they provide. We use a combination of empirical data and previously published values to estimate reductions in nitrogen inputs under outbreaks of plant viruses of varying severity. We also use historical fertilizer prices to examine the economic impacts of this lost fertilizer for farmers. We find that fertilizer losses are greatest for crops that fix large amounts of nitrogen, such as clover and alfalfa as opposed to common bean. The economic impact on farmers is controlled by the proportion of plants with viral infections and the price of synthetic fertilizer. In a year of high disease prevalence, attention is normally focused on the yield of the diseased crops. We suggest that farmers growing legumes as break crops should be concerned about yields of subsequent crops as well. Viral diseases can be difficult to diagnose in the field, so the easiest way for farmers to prevent unexpected yield losses in subsequent crops is to test their soil when it is feasible to do so., (© 2020 Ecological Society of America.)

Intraspecific plant diversity can significantly impact insect herbivore populations in natural systems. Yet, its role as an insect pest control strategy in agriculture has received less attention, and little is known about which crop traits are important to herbivores in different landscape contexts. Moreover, empirical economic analyses on the cost-effectiveness of varietal mixtures are lacking. We used varietal mixtures of Brassica oleracea crops on working farms to examine how two metrics of intraspecific crop diversity, varietal richness and number of plant colors (color richness), affect crop damage and the incidence and abundance of two insect pest species: Pieris rapae and Phyllotreta spp. We evaluated the context-dependency of varietal mixtures by sampling early- and late-season plantings of B. oleracea crops in farms across a gradient of landscape composition. We developed crop budgets and used a net present value analysis to assess the impact of varietal mixtures on input and labor costs, crop revenues, and profit. We found context-dependent effects of varietal mixtures on both pests. In early-season plantings, color richness did not affect Phyllotreta spp. populations. However, increasing varietal richness reduced Phyllotreta spp. incidence in simple landscapes dominated by cropland, but this trend was reversed in complex landscapes dominated by natural habitats. In late-season plantings, color richness reduced the incidence and abundance of P. rapae larvae, but only in complex landscapes where their populations were highest. Varietal richness had the same effect on P. rapae larvae as color richness. Unexpectedly, we consistently found lower pest pressure and reduced crop damage in simple landscapes. Although varietal mixtures did not affect crop damage, increasing color richness corresponded with increased profits, due to increased revenue and a marginal reduction in labor and input costs. We demonstrate varietal mixtures can significantly impact pest populations, and this effect can be mediated by intraspecific variation in crop color. However, the strength and direction of these effects vary by season, landscape composition, and pest species. The association between varietal color richness and profitability indicates farmers could design mixtures to enhance economic returns. We recommend additional research on the benefits of intraspecific trait variation for farmers., (© 2020 The Authors. Ecological Applications published by Wiley Periodicals LLC on behalf of Ecological Society of America.)

Many important endemic and emerging diseases are transmitted by vectors that are biting arthropods. The functional traits of vectors can affect pathogen transmission rates directly and also through their effect on vector population dynamics. Increasing empirical evidence shows that vector traits vary significantly across individuals, populations, and environmental conditions, and at time scales relevant to disease transmission dynamics. Here, we review empirical evidence for variation in vector traits and how this trait variation is currently incorporated into mathematical models of vector-borne disease transmission. We argue that mechanistically incorporating trait variation into these models, by explicitly capturing its effects on vector fitness and abundance, can improve the reliability of their predictions in a changing world. We provide a conceptual framework for incorporating trait variation into vector-borne disease transmission models, and highlight key empirical and theoretical challenges. This framework provides a means to conceptualize how traits can be incorporated in vector borne disease systems, and identifies key areas in which trait variation can be explored. Determining when and to what extent it is important to incorporate trait variation into vector borne disease models remains an important, outstanding question., Competing Interests: Conflict of Interest: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

Smallholders throughout sub-Saharan Africa produce legume crops as sources of food, fodder, and cash income, as well as to improve soil fertility. Ethiopian farmers have developed diverse legume varieties that enable adaptation to changing agroecological and sociocultural conditions. However, over the past several decades, as farm sizes declined and extension services promoted new varieties developed by plant breeders, changes in legume diversity have not been monitored. Based on interviews with smallholder farmers (n = 1296), we investigated the status of inter- and intraspecific legume diversity in major production areas of Ethiopia for five food legumes: common bean (Phaseolus vulgaris L.), field pea (Pisum sativum L.), faba bean (Vicia faba L.), groundnut (Arachis hypogaea L.) and fenugreek (Trigonella foenum-graecum L.). Legume species richness increased with altitude, relative household wealth, and land area planted to legumes. The highest numbers of varieties were found for common bean, followed by field pea, faba bean, groundnut and fenugreek. The average number of varieties planted per household was low (ranging from 1 to 2) and often much lower than the number reported in the same community or zone, which ranged from 2 to 18. For three out of the five species, the number of varieties significantly increased with total land area planted to legumes. Most varieties were rare, planted by less than 1/3 of farmers; however, informants accurately named varieties planted by others in the same community, demonstrating awareness of legume diversity at the community level. Given that the ability to plant multiple legume varieties is limited by land size, policies need to strengthen community-level conservation based on the diverse interests and needs of individual households., Competing Interests: The authors have declared that no competing interests exist.

If pathogen species, strains, or clones do not interact, intuition suggests the proportion of coinfected hosts should be the product of the individual prevalences. Independence consequently underpins the wide range of methods for detecting pathogen interactions from cross-sectional survey data. However, the very simplest of epidemiological models challenge the underlying assumption of statistical independence. Even if pathogens do not interact, death of coinfected hosts causes net prevalences of individual pathogens to decrease simultaneously. The induced positive correlation between prevalences means the proportion of coinfected hosts is expected to be higher than multiplication would suggest. By modelling the dynamics of multiple noninteracting pathogens causing chronic infections, we develop a pair of novel tests of interaction that properly account for nonindependence between pathogens causing lifelong infection. Our tests allow us to reinterpret data from previous studies including pathogens of humans, plants, and animals. Our work demonstrates how methods to identify interactions between pathogens can be updated using simple epidemic models., Competing Interests: The authors have declared that no competing interests exist.

The choice of a modeling approach is a critical decision in the modeling process, as it determines the complexity of the model and the phenomena that the model captures. In this paper, we developed an individual-based model (IBM) and compared it to a previously published ordinary differential equation (ODE) model, both developed to describe the same biological system although with slightly different emphases given the underlying assumptions and processes of each modeling approach. We used both models to examine the effect of insect vector life history and behavior traits on the spread of a vector-borne plant virus, and determine how choice of approach affects the results and their biological interpretation. A non-random distribution of insect vectors across plant hosts emerged in the IBM version of the model and was not captured by the ODE. This distribution led simultaneously to a slower-growing vector population and a faster spread of the pathogen among hosts. The IBM model also enabled us to test the effect of potential control measures to slow down virus transmission. We found that removing virus-infected hosts was a more effective strategy for controlling infection than removing vector-infested hosts. Our findings highlight the need to carefully consider possible modeling approaches before constructing a model.

Plants and animals host many microbial symbionts, including both pathogens and mutualists. However, most experimental studies include only one symbiont, and few examine interactions of more than two microbes with their host. Here, we examined whether coinfection with two pathogens causes a synergistic reduction in the benefits that hosts receive from a microbial mutualist. We also measured the effects of a microbial mutualist on the within- and between-host competition between coinfecting pathogens. We manipulated the presence of Clover yellow vein virus (ClYVV), Bean common mosaic virus (BCMV), rhizobia bacteria, and nitrogen fertilizer in common beans (Phaseolus vulgaris). We found asymmetric, context-dependent interactions among the three microbial symbionts and their host. Coinfection with both viruses led to greater than additive negative effects on the amount of nitrogen that plants received from rhizobia. Rhizobia colonization decreased immune signaling in singly infected plants, but not in coinfected plants. Compared to single ClYVV infection, ClYVV reached higher concentrations within hosts coinfected with BCMV, but only in the presence of rhizobia. Coinfection increased BCMV vertical transmission rates for plants without supplemental nitrogen, but overall vertical transmission opportunities were not affected due to reduced seed production. Examining interactions between multiple microbes sharing a host can reveal important insights about nutrient cycling, disease severity, and pathogen epidemiology., (© 2018 by the Ecological Society of America.)

Potato virus Y (PVY) has reemerged as a serious impediment to seed potato production, responsible for reduced yields and tuber quality, as well as the majority of seed lot rejections by certification programs due to excessive virus incidence. This has led to seed shortages, especially in cultivars highly susceptible to infection. While seed certification programs have been effective at managing many virus diseases below economic thresholds, PVY has rapidly evolved in recent decades to become a complex of strains that evade many certification and farm management practices. The evolution of PVY strains is naturally occurring, but several human influences can be linked to the rapid change in PVY populations affecting the potato crop. Here we highlight the recent history and current status of PVY in potatoes and suggest some approaches for managing the virus moving forward., (Published by Elsevier B.V.)

Degradation of freshwater ecosystems and the services they provide is a primary cause of increasing water insecurity, raising the need for integrated solutions to freshwater management. While methods for characterizing the multi-faceted challenges of managing freshwater ecosystems abound, they tend to emphasize either social or ecological dimensions and fall short of being truly integrative. This paper suggests that management for sustainability of freshwater systems needs to consider the linkages between human water uses, freshwater ecosystems and governance. We present a conceptualization of freshwater resources as part of an integrated social-ecological system and propose a set of corresponding indicators to monitor freshwater ecosystem health and to highlight priorities for management. We demonstrate an application of this new framework -the Freshwater Health Index (FHI) - in the Dongjiang River Basin in southern China, where stakeholders are addressing multiple and conflicting freshwater demands. By combining empirical and modeled datasets with surveys to gauge stakeholders' preferences and elicit expert information about governance mechanisms, the FHI helps stakeholders understand the status of freshwater ecosystems in their basin, how ecosystems are being manipulated to enhance or decrease water-related services, and how well the existing water resource management regime is equipped to govern these dynamics over time. This framework helps to operationalize a truly integrated approach to water resource management by recognizing the interplay between governance, stakeholders, freshwater ecosystems and the services they provide., (Copyright © 2018 The Authors. Published by Elsevier B.V. All rights reserved.)

Infections of one host by multiple parasites are common, and several studies have found that the order of parasite invasion can affect both within-host competition and disease severity. However, it is unclear to what extent coinfection timing might be important to consider when modeling parasite impacts on host populations. Using a model system of two viruses infecting barley, we found that simultaneous infections of the two viruses were significantly more damaging to hosts than sequential coinfections. While priority effects were evident in within-host concentrations of sequential coinfections, priority did not influence any parameters (such as virulence or transmission rate) that affect host population dynamics. We built a susceptible-infected model to examine whether the observed difference in coinfection virulence could impact host population dynamics under a range of scenarios. We found that coinfection timing can have an important but context-dependent effect on projected host population dynamics. Studies that examine only simultaneous coinfections could inflate disease impact predictions.

Maize lethal necrosis (MLN) has emerged as a serious threat to food security in sub-Saharan Africa. MLN is caused by coinfection with two viruses, Maize chlorotic mottle virus and a potyvirus, often Sugarcane mosaic virus. To better understand the dynamics of MLN and to provide insight into disease management, we modeled the spread of the viruses causing MLN within and between growing seasons. The model allows for transmission via vectors, soil, and seed, as well as exogenous sources of infection. Following model parameterization, we predict how management affects disease prevalence and crop performance over multiple seasons. Resource-rich farmers with large holdings can achieve good control by combining clean seed and insect control. However, crop rotation is often required to effect full control. Resource-poor farmers with smaller holdings must rely on rotation and roguing, and achieve more limited control. For both types of farmer, unless management is synchronized over large areas, exogenous sources of infection can thwart control. As well as providing practical guidance, our modeling framework is potentially informative for other cropping systems in which coinfection has devastating effects. Our work also emphasizes how mathematical modeling can inform management of an emerging disease even when epidemiological information remains scanty. [Formula: see text] Copyright © 2017 The Author(s). This is an open access article distributed under the CC BY-NC-ND 4.0 International license .

Plant viruses, often spread by arthropod vectors, impact natural and agricultural ecosystems worldwide. Intuitively, the movement behavior and life history of vectors influence pathogen spread, but the relative contribution of each factor has not been examined. Recent research has highlighted the influence of host infection status on vector behavior and life history. Here, we developed a model to explore how vector traits influence the spread of vector-borne plant viruses. We allowed vector life history (growth rate, carrying capacity) and movement behavior (departure and settlement rates) parameters to be conditional on whether the plant host is infected or healthy and whether the vector is viruliferous (carrying the virus) or not. We ran simulations under a wide range of parameter combinations and quantified the fraction of hosts infected over time. We also ran case studies of the model for Barley yellow dwarf virus, a persistently transmitted virus, and for Potato virus Y, a non-persistently transmitted virus. We quantified the relative importance of each parameter on pathogen spread using Latin hypercube sampling with the statistical partial rank correlation coefficient technique. We found two general types of mechanisms in our model that increased the rate of pathogen spread. First, increasing factors such as vector intrinsic growth rate, carrying capacity, and departure rate from hosts (independent of whether these factors were condition-dependent) led to more vectors moving between hosts, which increased pathogen spread. Second, changing condition-dependent factors such as a vector's preference for settling on a host with a different infection status than itself, and vector tendency to leave a host of the same infection status, led to increased contact between hosts and vectors with different infection statuses, which also increased pathogen spread. Overall, our findings suggest that vector population growth rates had the greatest influence on rates of virus spread, but rates of vector dispersal from infected hosts and from hosts of the same infection status were also very important. Our model highlights the importance of simultaneously considering vector life history and behavior to better understand pathogen spread. Although developed for plant viruses, our model could readily be utilized with other vector-borne pathogen systems., (© 2017 by the Ecological Society of America.)

Disease and community ecology share conceptual and theoretical lineages, and there has been a resurgence of interest in strengthening links between these fields. Building on recent syntheses focused on the effects of host community composition on single pathogen systems, we examine pathogen (microparasite) communities using a stochastic metacommunity model as a starting point to bridge community and disease ecology perspectives. Such models incorporate the effects of core community processes, such as ecological drift, selection and dispersal, but have not been extended to incorporate host-pathogen interactions, such as immunosuppression or synergistic mortality, that are central to disease ecology. We use a two-pathogen susceptible-infected (SI) model to fill these gaps in the metacommunity approach; however, SI models can be intractable for examining species-diverse, spatially structured systems. By placing disease into a framework developed for community ecology, our synthesis highlights areas ripe for progress, including a theoretical framework that incorporates host dynamics, spatial structuring and evolutionary processes, as well as the data needed to test the predictions of such a model. Our synthesis points the way for this framework and demonstrates that a deeper understanding of pathogen community dynamics will emerge from approaches working at the interface of disease and community ecology., (© 2015 John Wiley & Sons Ltd/CNRS.)

Disease dilution (reduced disease prevalence with increasing biodiversity) has been described for many different pathogens. Although the mechanisms causing this phenomenon remain unclear, the disassembly of communities to predictable subsets of species, which can be caused by changing climate, land use or invasive species, underlies one important hypothesis. In this case, infection prevalence could reflect the competence of the remaining hosts. To test this hypothesis, we measured local host species abundance and prevalence of four generalist aphid-vectored pathogens (barley and cereal yellow dwarf viruses) in a ubiquitous annual grass host at 10 sites spanning 2000 km along the North American West Coast. In laboratory and field trials, we measured viral infection as well as aphid fecundity and feeding preference on several host species. Virus prevalence increased as local host richness declined. Community disassembly was non-random: ubiquitous hosts dominating species-poor assemblages were among the most competent for vector production and virus transmission. This suggests that non-random biodiversity loss led to increased virus prevalence. Because diversity loss is occurring globally in response to anthropogenic changes, such work can inform medical, agricultural and veterinary disease research by providing insights into the dynamics of pathogens nested within a complex web of environmental forces.

The pathogen and parasite community that inhabits every free-living organism can control host vital rates including lifespan and reproductive output. To date, however, there have been few experiments examining pathogen community assembly replicated at large-enough spatial scales to inform our understanding of pathogen dynamics in natural systems. Pathogen community assembly may be driven by neutral stochastic colonization and extinction events or by niche differentiation that constrains pathogen distributions to particular environmental conditions, hosts, or vectors. Here, we present results from a regionally-replicated experiment investigating the community of barley and cereal yellow dwarf viruses (B/CYDV's) in over 5000 experimentally planted individuals of six grass species along a 700 km latitudinal gradient along the Pacific coast of North America (USA) in response to experimentally manipulated nitrogen and phosphorus supplies. The composition of the virus community varied predictably among hosts and across nutrient-addition treatments, indicating niche differentiation among virus species. There were some concordant responses among the viral species. For example, the prevalence of most viral species increased consistently with perennial grass cover, leading to a 60% increase in the richness of the viral community within individual hosts (i.e., coinfection) in perennial-dominated plots. Furthermore, infection rates of the six host species in the field were highly correlated with vector preferences assessed in laboratory trials. Our results reveal the importance of niche differentiation in structuring virus assemblages. Virus species distributions reflected a combination of local host community composition, host species-specific vector preferences, and virus responses to host nutrition. In addition, our results suggest that heterogeneity among host species in their capacity to attract vectors or support pathogens between growing seasons can lead to positive covariation among virus species.

Research on plant viruses in natural ecosystems has been increasing rapidly over the past decade. This paper reviews recent research on the barley and cereal yellow dwarf viruses (B/CYDVs) in grasslands of the western US, beginning with the evidence that the disease caused by these viruses facilitated the invasion of western US grasslands by European annual grasses. Observational and experimental studies of B/CYDVs were carried out along a latitudinal gradient (33.8-48.8°N) from southern California to southern Canada. The prevalence and community composition of B/CYDVs were assessed over a variety of scales and under a range of biotic and abiotic conditions. The findings indicate that both biotic and abiotic factors are important influences on virus ecology and epidemiology. Introduced annual grasses are high-quality hosts that amplify both virus and vector populations in this system, but our research suggests that endemic perennial grasses are critically important for sustaining virus populations in contemporary grasslands largely composed of introduced species. Experiments indicated that increased phosphorus supply to hosts resulted in greater host biomass and higher virus prevalence. Using experimental exclosures, it was found that the presence of grazing vertebrate herbivores increased the abundance of annual grasses, resulting in increased virus prevalence. The results of these studies suggest that patterns of B/CYDV prevalence and coinfection in western US grasslands are strongly shaped by the interactions of host plants, vectors, vertebrate herbivores, and abiotic drivers including nutrients., (Copyright © 2011 Elsevier B.V. All rights reserved.)

Landscape composition affects ecosystems services, including agricultural pest management. However, relationships between land use and agricultural insects are not well understood, and many complexities remain to be explored. Here we examine whether nonagricultural landscapes can directly suppress agricultural pests, how multiple spatial scales of land use concurrently affect insect populations, and the relationships between regional land use and insect populations. We tracked densities of three specialist corn (Zea mays) pests (Ostrinia nubilalis, European corn borer; Diabrotica virgifera, western corn rootworm; Diabrotica barberi, northern corn rootworm), and two generalist predator lady beetles (Coleomegilla maculata and Propylea quatuordecimpunctata) in field corn and determined their relationships to agricultural land use at three spatial scales (field perimeter, 1-km, and 20-km radius areas). Pest densities were either higher (D. virgifera and D. barberi) or unchanged (O. nubilalis) in landscapes with more corn, while natural enemy densities were either lower (C. maculata) or unchanged (P. quatuordecimpunctata). Results for D. virgifera and D. barberi indicate that decreasing the area of preferred crop in the landscape can directly suppress specialist insect pests. Multiple scales of land use affected populations of D. virgifera and C. maculata, and D. virgifera populations showed strong relationships with regional, 20-km-scale land use. These results suggest that farm planning and government policies aimed at diversifying local and regional agricultural landscapes show promise for increasing biological control and directly suppressing agricultural pests.

Biological invasions occur when plants, animals, or microbes are introduced to a new geographic region, then spread and have negative consequences for the local ecosystem. Across both plant and animal hosts, viruses can play diverse roles in biological invasions. First, viruses can either decrease or increase the impacts of biological invasions by their hosts. Introduced hosts commonly leave behind many viruses from their native ranges, which may allow the hosts to achieve greater fitness and thus dominate in their introduced ranges. Viruses that do infect introduced hosts may reduce invasive host fitness and impacts. However, many viruses that infect introduced hosts also infect native hosts and may have more severe impacts on the native hosts. Second, viruses can also be invasive species themselves. While many viruses are believed to be introduced, it is challenging to differentiate between those that are native and those that are not. Third, many viruses are transmitted by vectors, which can also be introduced to new regions. Introduced vectors can increase virus transmission rates, altering host communities and ecosystems. Further advancing our understanding of the role of viruses in biological invasions will require research that integrates the systematics, biogeography and ecological history of hosts, vectors, and viruses., (Copyright © 2011 Elsevier B.V. All rights reserved.)

Agricultural ecosystems provide humans with food, forage, bioenergy and pharmaceuticals and are essential to human wellbeing. These systems rely on ecosystem services provided by natural ecosystems, including pollination, biological pest control, maintenance of soil structure and fertility, nutrient cycling and hydrological services. Preliminary assessments indicate that the value of these ecosystem services to agriculture is enormous and often underappreciated. Agroecosystems also produce a variety of ecosystem services, such as regulation of soil and water quality, carbon sequestration, support for biodiversity and cultural services. Depending on management practices, agriculture can also be the source of numerous disservices, including loss of wildlife habitat, nutrient runoff, sedimentation of waterways, greenhouse gas emissions, and pesticide poisoning of humans and non-target species. The tradeoffs that may occur between provisioning services and other ecosystem services and disservices should be evaluated in terms of spatial scale, temporal scale and reversibility. As more effective methods for valuing ecosystem services become available, the potential for 'win-win' scenarios increases. Under all scenarios, appropriate agricultural management practices are critical to realizing the benefits of ecosystem services and reducing disservices from agricultural activities.

Host characteristics commonly determine infection risk, but infection can also be mediated by regional- or local-scale variation in the biotic and abiotic environment. Experiments can clarify the relative importance of these factors. We quantified drivers of infection by barley and cereal yellow dwarf viruses (B/CYDV), a group of generalist, vector-borne grass pathogens, at hierarchically nested spatial scales (10(5)-1 m) by planting individuals of six common grass species into five Pacific Coast grassland sites spanning 7 degrees of latitude (> 5000 total hosts) and applying a factorial combination of nitrogen and phosphorus fertilizer. Infection differed most among experimental blocks (10(2)-10(3) m scale), suggesting that local factors control infection risk; infection increased with cover of long-lived hosts and phosphorus, but not nitrogen, fertilization. For B/CYDV, local context more strongly predicts infection risk than host species traits or regional context; such spatially nested experiments can clarify the factors underlying variation in infection risk.

Host-pathogen interactions may be governed by the number of pathogens coexisting within an individual host (i.e., coinfection) and among different hosts, although most sampling in natural systems focuses on the prevalence of single pathogens and/or single hosts. We measured the prevalence of four barley and cereal yellow dwarf viruses (B/CYDVs) in three grass species at 26 natural grasslands along a 2000-km latitudinal gradient in the western United States and Canada. B/CYDVs are aphid-vectored RNA viruses that cause one of the most prevalent of all plant diseases worldwide. Pathogen prevalence and coinfection were uncorrelated, suggesting that different forces likely drive them. Coinfection, the number of viruses in a single infected host (alpha diversity), did not differ among host species but increased roughly twofold across our latitudinal transect. This increase in coinfection corresponded with a decline in among-host pathogen turnover (beta diversity), suggesting that B/CYDVs in northern populations experience less transmission limitation than in southern populations. In contrast to pathogen diversity, pathogen prevalence was a function of host identity as well as biotic and abiotic environmental conditions. Prevalence declined with precipitation and increased with soil nitrate concentration, an important limiting nutrient for hosts and vectors of B/CYDVs. This work demonstrates the need for further studies of processes governing coinfection, and the utility of applying theory developed to explain diversity in communities of free-living organisms to pathogen systems.

The development of crops genetically engineered for pathogen resistance has raised concerns that crop-to-wild gene flow could release wild or weedy relatives from regulation by the pathogens targeted by the transgenes that confer resistance. Investigation of these risks has also raised questions about the impact of gene flow from conventional crops into wild plant populations. Viruses in natural plant populations can play important roles in plant fecundity and competitive interactions. Here, we show that virus-resistance transgenes and conventional crop genes can increase fecundity of wild plants under virus pressure. We asked how gene flow from a cultivated squash (Cucurbita pepo) engineered for virus resistance would affect the fecundity of wild squash (C. pepo) in the presence and absence of virus pressure. A transgenic squash cultivar was crossed and backcrossed with wild C. pepo from Arkansas. Wild C. pepo, transgenic backcross plants, and non-transgenic backcross plants were compared in field plots in Ithaca, New York, USA. The second and third generations of backcrosses (BC2 and BC3) were used in 2002 and 2003, respectively. One-half of the plants were inoculated with zucchini yellow mosaic virus (ZYMV), and one-half of the plants were maintained as healthy controls. Virus pressure dramatically decreased the fecundity of wild C. pepo plants and non-transgenic backcross plants relative to transgenic backcross plants, which showed continued functioning of the virus-resistance transgene. In 2002, non-transgenic backcross fecundity was slightly higher than wild C. pepo fecundity under virus pressure, indicating a possible benefit of conventional crop alleles, but they did not differ in 2003 when fecundity was lower in both groups. We detected no fitness costs of the transgene in the absence of the virus. If viruses play a role in the population dynamics of wild C. pepo, we predict that gene flow from transgenic, virus-resistant squash and, to a much lesser extent, conventionally bred squash would increase C. pepo fecundity. Studies such as this one, in combination with documentation of the probability of crop-to-wild gene flow and surveys of virus incidence in wild populations, can provide a solid basis for environmental risk assessments of crops genetically engineered for virus resistance.

Most species host multiple pathogens, yet field studies rarely examine the processes determining pathogen diversity within a single host or the effects of coinfection on pathogen dynamics in natural systems. Coinfection can affect pathogen transmission and virulence. In turn, coinfection can be regulated within hosts by interactions such as cross-protective immunity or at broader spatial scales via vector distributions. Using a general model, we demonstrate that coinfection by a group of vectored pathogens is highest with abundant generalist vectors and weak cross-protection and coinfection-induced mortality. Using these predictions, we investigate the distribution of five coexisting aphid-vectored, viral pathogens (barley and cereal yellow dwarf luteoviruses and poleroviruses) in a native perennial grass (Elymus glaucus) in both space (700 km) and time (4 years). Observed coinfection rates were much higher than expected at random, suggesting that within-host processes exerted weak effects on within-host pathogen diversity. Covariance among viruses in space and time was highest for viral species sharing a vector. Temporal correlation arose from the synchronous invasion of two viruses transmitted by a shared aphid species. On the basis of our modeling and empirical results, we expect that factors external to individual hosts may affect the coinfection dynamics in other communities hosting vectored pathogens.

Most pathogens exist within complicated food webs of interacting hosts, vectors, competitors, and predators. Although theory has demonstrated a variety of mechanisms by which predation and competition in food webs can indirectly control infection risk in hosts, there have until now been no experimental tests of this theory. We sampled the effect of long-term exclusion of large vertebrate herbivores on the prevalence of infection by a group of aphid-vectored viruses that infect grasses (barley and cereal yellow dwarf viruses) in an oak savannah in central California. We found that pathogen prevalence was approximately 4-fold higher in the presence of consumers than in areas where they were excluded. Vertebrate consumers did not directly alter infection rates by this aphid-vectored pathogen group, but rather increased infection risk by increasing the relative abundance of highly-competent hosts in the grassland community. This large-scale experiment, measuring changes in host abundance and infection risk in response to altered consumption rates, confirms theoretical predictions that consumers can indirectly increase infection risk by altering the composition of whole communities. Most importantly, these results demonstrate that, even in complex natural communities, alterations to food web composition such as consumer invasion or extinction can lead to significant impacts that cascade throughout entire communities, including changes in infection risk.

Plants engage in multiple, simultaneous interactions with other species; some (enemies) reduce and others (mutualists) enhance plant performance. Moreover, effects of different species may not be independent of one another; for example, enemies may compete, reducing their negative impact on a plant. The magnitudes of positive and negative effects, as well as the frequency of interactive effects and whether they tend to enhance or depress plant performance, have never been comprehensively assessed across the many published studies on plant-enemy and plant-mutualist interactions. We performed a meta-analysis of experiments in which two enemies, two mutualists, or an enemy and a mutualist were manipulated factorially. Specifically, we performed a factorial meta-analysis using the log response ratio. We found that the magnitude of (negative) enemy effects was greater than that of (positive) mutualist effects in isolation, but in the presence of other species, the two effects were of comparable magnitude. Hence studies evaluating single-species effects of mutualists may underestimate the true effects found in natural settings, where multiple interactions are the norm and indirect effects are possible. Enemies did not on average influence the effects on plant performance of other enemies, nor did mutualists influence the effects of mutualists. However, these averages mask significant and large, but positive or negative, interactions in individual studies. In contrast, mutualists ameliorated the negative effects of enemies in a manner that benefited plants; this overall effect was driven by interactions between pathogens and belowground mutualists (bacteria and mycorrhizal fungi). The high frequency of significant interactive effects suggests a widespread potential for diffuse rather than pairwise coevolutionary interactions between plants and their enemies and mutualists. Pollinators and mycorrhizal fungi enhanced plant performance more than did bacterial mutualists. In the greenhouse (but not the field), pathogens reduced plant performance more than did herbivores, pathogens were more damaging to herbaceous than to woody plants, and herbivores were more damaging to crop than to non-crop plants (suggesting evolutionary change in plants or herbivores following crop domestication). We discuss how observed differences in effect size might be confounded with methodological differences among studies.

Introduced plant populations lose interactions with enemies, mutualists and competitors from their native ranges, and gain interactions with new species, under new abiotic conditions. From a biogeographical perspective, differences in the assemblage of interacting species, as well as in abiotic conditions, may explain the demographic success of the introduced plant populations relative to conspecifics in their native range. Within invaded communities, the new interactions and conditions experienced by the invader may influence both its demographic success and its effects on native biodiversity. Here, we examine indirect effects involving enemies, mutualists and competitors of introduced plants, and effects of abiotic conditions on biotic interactions. We then synthesize ideas building on Darwin's idea that the kinds of new interactions gained by an introduced population will depend on its relatedness to native populations. This yields a heuristic framework to explain how biotic interactions and abiotic conditions influence invader success. We conclude that species introductions generally alter plants' interactions with enemies, mutualists and competitors, and that there is increasing evidence that these altered interactions jointly influence the success of introduced populations.

In field experiments manipulating generalist pathogens and host community composition, the presence of a highly susceptible reservoir species drove disease dynamics in multiple nonreservoir species, sometimes decreasing their abundance through apparent competition. The dynamics of generalist pathogens in multispecies host communities remain a major frontier for disease ecology. Of particular interest are how host community structure controls pathogen transmission and how disease spread feeds back to influence the host community. Pathogen spillover occurs when epidemics in a host population are driven not by transmission within that population but by transmission from a reservoir population. Here we review examples of spillover in pathogens infecting humans, domesticated animals, and crops, noting that most empirical evidence for spillover results from nonmanipulative, observational studies. We then present results from two field experiments utilizing an experimentally tractable model system of annual wild grasses and a generalist virus, the barley yellow dwarf virus. In these experiments, the presence of a highly susceptible reservoir species, Avena fatua (wild oats), greatly increased pathogen prevalence in several other species. This result demonstrates pathogen spillover and illustrates the crucial role of host community structure in controlling the dynamics of generalist pathogens. Further, pathogen spillover from A. fatua decreased the abundance of two other host species through pathogen-mediated apparent competition. Thus, our results provide experimental support for theoretical predictions of strong feedbacks between host community structure and generalist disease dynamics.

Invasive plant species both threaten native biodiversity and are economically costly, but only a few naturalized species become pests. Here we report broad, quantitative support for two long-standing hypotheses that explain why only some naturalized species have large impacts. The enemy release hypothesis argues that invaders' impacts result from reduced natural enemy attack. The biotic resistance hypothesis argues that interactions with native species, including natural enemies, limit invaders' impacts. We tested these hypotheses for viruses and for rust, smut and powdery mildew fungi that infect 473 plant species naturalized to the United States from Europe. On average, 84% fewer fungi and 24% fewer virus species infect each plant species in its naturalized range than in its native range. In addition, invasive plant species that are more completely released from pathogens are more widely reported as harmful invaders of both agricultural and natural ecosystems. Together, these results strongly support the enemy release hypothesis. Among noxious agricultural weeds, species accumulating more pathogens in their naturalized range are less widely noxious, supporting the biotic resistance hypothesis. Our results indicate that invasive plants' impacts may be a function of both release from and accumulation of natural enemies, including pathogens.

Field experiments were carried out in Nicaragua to examine the influence of plant community diversity, plant density, and host plant quality on the spread of an insect-transmitted plant pathogen. Population levels of the corn leafhopper Dalbulus maidis, which transmits the corn stunt spiroplasma to maize, were monitored in four experimental communities; low-density maize monoculture, high-density maize monoculture, two-species (maize/bean) polyculture, and multispecies (maize/weeds) polyculture. Leafhopper abundance per plant and the incidence of corn stunt were lower in high-density maize monocultures than in low-density monocultures. Increasing plant diversity by intercropping with nonhost species such as beans or weeds also led to lower leafhooper abundance and decreased disease incidence, but the effect was not enhanced as additional nonhost species were added to the community. Manipulating host plant quality by increasing nitrogen fertilization resulted in higher leafhopper densities at higher nitrogen levels. To explore the role of vector movement in disease spread, leafhopper movement rates and emigration were estimated by observing changes in the spatial gradients of leafhopper densities over time. This method of movement analysis requires neither marking the insects nor releasing them at a single point, and thus reduces the extreme disturbance caused by traditional mark-release techniques. The analysis indicated that leafhopper movement rates were lowest in the polycultures. In particular, across-row movement was strikingly inhibited in the bean polyculture. This polyculture also had the highest rates of emigration. These results indicate that plant quality, density, and diversity significantly affect the spread of corn stunt through their effects on the abundance and movement behavior of the corn leafhopper. These factors could be manipulated in a program of cultural control for corn stunt in maize fields in tropical America., (© 1987 by the Ecological Society of America.)