Aquatic plant management can help mitigate the ecological, social, and economic harms caused by nuisance plants but can also have unintentional negative effects on native aquatic plants. The characteristics of management regimes, such as frequency and extent, may modify the impacts on native plant communities. Long-term and large-scale monitoring can span a range of management frequencies and extents, as well as habitat conditions, providing insights on native plant responses to management. We used a dataset of aquatic plant management and plant surveys collected from 373 waterbodies over 22 years in Florida, USA, to assess the effects of management on native plant communities. Because plant surveys were collected through nonstandardized approaches (i.e., meanders), we first evaluated the surveys relative to point-intercept sampling. We then evaluated the effects of management frequency (percentage of years managed) and extent (percentage of waterbody area managed), with respect to different targets and methods, on native plants. Management targets included Hydrilla verticillata (L.f) Royle subsp. verticillata, the floating plants Pontederia crassipes and Pistia stratiotes (grouped together), and all other managed plants (both native and non-native) grouped together (e.g., Cyperus blepharoleptos, Typha spp.). The meander approach estimated higher native plant richness and occurrences than point-intercept sampling, but there were large discrepancies in abundance estimates. Based on meander surveys and complimentary management records, native plant richness and occurrences were positively associated with more frequent management of H. verticillata and other plants. Native plant richness and occurrences were negatively associated with more frequent floating plant management and more extensive management with herbicides. Overall, we demonstrated that frequency and extent are important characteristics of management to consider when evaluating the effects of management on native plant taxa. Adjustments to management frequency and extent that maintain effective control of aquatic plants may also allow managers to maintain or even promote native plant diversity.

Decomposition of plant litter, facilitated primarily by microbial decomposers, plays a critical role in biogeochemical cycling and ecosystem function. The rate of litter decomposition can determine its environmental impact, where accelerated decomposition alters the timing and rate of nutrient release and may promote nutrient leaching, whereas slowed decomposition can result in litter accumulation, which impacts seedling recruitment, fire regimes, perennation of microbial communities, and slows nutrient release. Mutualistic endophytes are known to slow litter decomposition, but less is known about the impact that plant pathogens, present in diseased litter, have on decomposition rates. We compared litter decomposition of the invasive annual grass Microstegium vimineum with Bipolaris leaf spot symptoms, a fungal disease, to litter without symptoms of the disease in a year-long common garden experiment. We found leaf tissue with disease symptoms decomposed later in the year compared to litter without symptoms. By summer, 54% of leaf tissue from healthy sites remained compared to 80% of leaf material from diseased litter. Fungal infection did not impact the lignin or C:N content of the litter. There were significant differences in fungal community composition between infected and healthy litter at the start of the experiment that persisted until the end of summer. Disease epidemics prior to senescence contributed to the persistence of infected tissue, which could slow the return of nutrients to the environmental pool and promote the survival and dispersal of pathogen inoculum the following season.

Most species can be coinfected by multiple pathogens that may interact through shared resources (i.e., resource competition) or the host immune system (i.e., apparent competition). Community theory developed for free-living organisms suggests that coinfecting pathogens can persist if they satisfy the mutual invasion criterion of coexistence, establishing infections in hosts that are already infected. Furthermore, the likelihood of coexistence may depend on host nutrition which can affect shared resources and host immunity. Here, we apply the novel approach of combining a dynamical model and experimental mutual invasibility trials to explore the effects of host nutrient supply on the coexistence of two viral plant pathogens. We focus on among-pathogen interactions mediated by shared resources. First, we used a model to generate hypotheses about how nitrogen (N) and phosphorus (P) supply rates affect the ability of two plant viruses to invade established infections of the other virus. Then, we experimentally manipulated the N and P supplied to oats (Avena sativa) in a growth chamber experiment and tested mutual invasion of two RNA viral pathogens, BYDV-PAV and CYDV-RPV. Nutrient supplies ranged from rates that barely kept hosts alive up to high, but sub-toxic, rates. Model simulations suggested that the viruses were more likely to invade established infections either when they could replicate at lower N and P concentrations or when plant N and P concentrations increased due to a combination of nutrient supply rates and resident virus nutrient use. In the experiment, each virus successfully invaded hosts infected by the other and had consistent growth rates across N and P supply rates. Our results suggest that BYDV-PAV and CYDV-RPV can coexist across a wide range environmental nutrient supply, which is consistent with the high levels of co-occurrence of these two viruses in field populations.

Managing invasive species with prevention and early-detection strategies can avert severe ecological and economic impacts. Horizon scanning, an evidence-based process combining risk screening and consensus building to identify threats, has become a valuable tool for prioritizing invasive species management and prevention. We assembled a working group of experts from academic, government, and nonprofit agencies and organizations, and conducted a multi-taxa horizon scan for Florida, USA, the first of its kind in North America. Our primary objectives were to identify high-risk species and their introduction pathways, to detail the magnitude and mechanism of potential impacts, and, more broadly, to demonstrate the utility of horizon scanning. As a means to facilitate future horizon scans, we document the process used to generate the list of taxa for screening. We evaluated 460 taxa for their potential to arrive, establish, and cause negative ecological and socioeconomic impacts, and identified 40 potential invaders, including alewife, zebra mussel, crab-eating macaque, and red swamp crayfish. Vertebrates and aquatic invertebrates posed the greatest invasion threat, over half of the high-risk taxa were omnivores, and there was high confidence in the scoring of high-risk taxa. Common arrival pathways were ballast water, biofouling of vessels, and escape from the pet/aquarium/horticulture trade. Competition, predation, and damage to agriculture/forestry/aquaculture were common impact mechanisms. We recommend full risk analysis for the high-risk taxa; increased surveillance at Florida's ports, state borders, and high-risk pathways; and periodic review and revision of the list. Few horizon scans detail the comprehensive methodology (including list-building), certainty estimates for all scoring categories and the final score, detailed pathways, and the magnitude and mechanism of impact. Providing this information can further inform prevention efforts and can be efficiently replicated in other regions. Moreover, harmonizing methodology can facilitate data sharing and enhance interpretation of results for stakeholders and the general public.

Invasive plants, which cause substantial economic and ecological impacts, acquire both pathogens and beneficial microbes in their introduced ranges. Communities of fungal endophytes are known to mediate impacts of pathogens on plant fitness but few studies have examined the temporal dynamics of fungal communities on invasive plants. The annual grass Microstegium vimineum, an invader of forests and riparian areas throughout the eastern United States, experiences annual epidemics of disease caused by Bipolaris pathogens. Our objective was to characterize the dynamics of foliar fungal communities on M. vimineum over a growing season during a foliar disease epidemic. First, we asked how the fungal community in the phyllosphere changed over 2 months that corresponded with increasing disease severity. Second, we experimentally suppressed disease with fungicide in half of the plots and asked how the treatment affected fungal community diversity and composition. We found increasingly diverse foliar fungal communities and substantial changes in community composition between timepoints using high-throughput amplicon sequencing of the internal transcribed spacer 2 region. Monthly fungicide application caused shifts in fungal community composition relative to control samples. Fungicide application increased diversity at the late-season timepoint, suggesting that it suppressed dominant fungicide sensitive taxa and allowed other fungal taxa to flourish. These results raise new questions regarding the roles of putative endophytes found in the phyllosphere given the limited number of pathogens known to cause disease on M. vimineum in its invasive range.

Food has long been known to perform dual functions of nutrition and medicine, but mounting evidence suggests that complex host-pathogen dynamics can emerge along continuous resource gradients. Empirical examples of nonmonotonic responses of infection with increasing host resources (e.g., low prevalence at low and high resource supply but high prevalence at intermediate resources) have been documented across the tree of life, but these dynamics, when observed, often are interpreted as nonintuitive, idiosyncratic features of pathogen and host biology. Here, by developing generalized versions of existing models of resource dependence for within- and among-host infection dynamics, we provide a synthetic view of nonmonotonic infection dynamics. We demonstrate that where resources jointly impact two (or more) processes (e.g., growth, defense, transmission, mortality, predation), nonmonotonic infection dynamics, including alternative states, can emerge across a continuous resource supply gradient. We review the few empirical examples that concurrently measured resource effects on multiple rates and pair this with a wide range of examples in which resource dependence of multiple rates could generate nonmonotonic infection outcomes under realistic conditions. This review and generalized framework highlight the likely generality of such resource effects in natural systems and point to opportunities ripe for future empirical and theoretical work.

Climate change is increasing the frequency and intensity of extreme events like drought and flooding, which threaten to amplify other global change drivers such as species invasion. We investigate the effect of wet and dry extreme precipitation regimes on invasive species' abundances in northern tallgrass prairies. Because soil moisture is a key determinant of prairie composition, theory and evidence suggest drought conditions will hinder invasion, whereas wetter conditions will enhance invasion. To test this hypothesis, we explored the effect of precipitation on invasive plant species abundance from 2010 to 2019 in 25 managed prairies using observations from 267 transects comprising 6675 plots throughout western Minnesota, USA. We estimated how increases in the number of extremely wet or dry months in a year altered overall invasive species abundance and the abundance of the highly invasive grasses Poa pratensis and Bromus inermis. We found that a greater occurrence of abnormally wet months increased invasive species abundance but found mixed evidence that abnormally dry conditions hindered invasion. Further, more moderately wet and dry months reduced native grass abundance. Together, these results suggest that more frequent extremely wet months may intensify invasive dominance and that dry months may not counterbalance these trends. Given the considerable uncertainty still surrounding the interactive effects of climate change and invasion on native plant communities, this research represents an important step toward quantifying the complex influence of precipitation extremes on invasion dynamics in managed ecosystems of critical conservation concern.

Early detection and eradication of invasive plants are more cost-effective than managing well-established invasive plant populations and their impacts. However, there is high uncertainty around which taxa are likely to become invasive in a given area. Horizon scanning that combines a data-driven approach with rapid risk assessment and consensus building among experts can help identify invasion threats. We performed a horizon scan of potential invasive plant threats to Florida, USA—a state with a high influx of introduced species, conditions that are generally favorable for plant establishment, and a history of negative impacts from invasive plants. We began with an initial list of 2128 non-native plant taxa that are known invaders or crop pests. We built on previous invasive species horizon scans by developing data-based criteria to prioritize 100 taxa for rapid risk assessment. The semi-automated prioritization process included selecting taxa “on the horizon” (i.e., not yet in the target location and not on a noxious weed list) with climate matching, naturalization history, “weediness” record, and global commonness. We derived overall invasion risk scores with rapid risk assessment by evaluating the likelihood of each of the taxa arriving, establishing, and having an impact in Florida. Then, following a consensus-building discussion, we identified six plant taxa as high risk, with overall risk scores ranging from 75 to 100 out of a possible 125. The six taxa are globally distributed, easily transported to new areas, found in regions with climates similar to Florida’s, and can impact native plant communities, human health, or agriculture. Finally, we evaluated our initial and final lists for potential biases. Assessors tended to assign higher risk scores to taxa that had more available information. In addition, we identified biases towards four plant families and certain geographical regions of origin. Our horizon scan approach identified taxa conforming to metrics of high invasion risk and used a methodology refined for plants that can be applied to other locations.

Host nutrient supply can mediate host-pathogen and pathogen-pathogen interactions. In terrestrial systems, plant nutrient supply is mediated by soil microbes, suggesting a potential role of soil microbes in plant diseases beyond soil-borne pathogens and induced plant defenses. Long-term nitrogen (N) enrichment can shift pathogenic and nonpathogenic soil microbial community composition and function, but it is unclear if these shifts affect plant-pathogen and pathogen-pathogen interactions. In a growth chamber experiment, we tested the effect of long-term N enrichment on infection by Barley Yellow Dwarf Virus (BYDV-PAV) and Cereal Yellow Dwarf Virus (CYDV-RPV), aphid-vectored RNA viruses, in a grass host. We inoculated sterilized growing medium with soil collected from a long-term N enrichment experiment (ambient, low, and high N soil treatments) to isolate effects mediated by the soil microbial community. We crossed soil treatments with a N supply treatment (low, high) and virus inoculation treatment (mock-, singly-, and co-inoculated) to evaluate the effects of long-term N enrichment on plant-pathogen and pathogen-pathogen interactions, as mediated by N availability. We measured the proportion of plants infected (i.e., incidence), plant biomass, and leaf chlorophyll content. BYDV-PAV incidence (0.96) declined with low N soil (to 0.46), high N supply (to 0.61), and co-inoculation (to 0.32). Low N soil mediated the effect of N supply on BYDV-PAV: instead of N supply reducing BYDV-PAV incidence, the incidence increased. Additionally, ambient and low N soil ameliorated the negative effect of co-inoculation on BYDV-PAV incidence. BYDV-PAV infection only reduced chlorophyll when plants were grown with low N supply and ambient N soil. There were no significant effects of long-term N soil on CYDV-RPV incidence. Soil inoculant with different levels of long-term N enrichment had different effects on host-pathogen and pathogen-pathogen interactions, suggesting that shifts in soil microbial communities with long-term N enrichment may mediate disease dynamics.

This publication describes why invasive species are an important issue in Florida and how nonnative species become invasive by moving through five stages of human-aided biological invasion. Written by Deah Lieurance, Amy Kendig, and Christina Romagosa, and published by the UF/IFAS Agronomy Department, July 2022.

Invasive species impact ecosystems through their large abundances and strong per capita effects. Enemies can regulate abundances and per capita effects, but are notably absent for many new invaders. However, invaders acquire enemies over time and as they spread; processes hypothesized to mitigate negative invader impacts by reducing abundance or per capita effects. Alternatively, properties of invaders or acquired enemies, such as an enemy’s ability to attack multiple species, may hinder enemy mitigation of invader impacts. We used field experiments to evaluate disease mitigation of invader impacts using the invasive grass Microstegium vimineum , which hosts an emerging fungal disease, and a native grass competitor, Elymus virginicus . We manipulated competition through density gradients of each plant species, and we reduced ambient foliar diseases with fungicide and autoclaving. We then modeled long-term population dynamics with field-estimated parameters. In the field, disease did not reduce invader abundance or per capita effects. The invader amplified disease on itself and the competitor, and disease reduced invader and competitor fitness components (e.g., germination). The dynamical model predicted that disease impacts on the competitor are greater than on the invader, such that disease will reduce invader abundance by 18%, and competitor abundance by 88%, over time. Our study suggests that enemies acquired by invaders will not necessarily mitigate invader impacts if the invader amplifies the enemy and the enemy attacks and suppresses competitor species. ### Competing Interest Statement The authors have declared no competing interest.

Plant litter can alter ecosystems and promote plant invasions by altering resource availability, depositing phytotoxins, and transmitting microorganisms to living plants. Transmission of microorganisms from invasive plant litter to live plants may gain importance as invasive plants, which often escape pathogens upon introduction to a new range, acquire new pathogens over time. It is unclear, however, if invasive plant litter affects native plant communities by promoting disease. Microstegium vimineum is an invasive grass that suppresses native populations, in part through litter production, and has acquired new fungal leaf spot diseases since its introduction to the United States. In a greenhouse experiment, we evaluated how M. vimineum litter and its pathogens mediated competition with the native grass Elymus virginicus. M. vimineum litter promoted disease on E. virginicus and suppressed establishment and biomass of both species. Litter had stronger negative effects on E. virginicus than M. vimineum, increasing the relative biomass of M. vimineum. Live plant competition reduced biomass of both species and live M. vimineum increased disease incidence on E. virginicus. Altogether, invasive grass litter suppressed both species, ultimately favoring the invasive species in competition, and increased disease incidence on the native species.

Community structure depends jointly on species' responses to, and effects on, environmental factors. Many such factors, including detritus, are studied in ecosystem ecology. Detritus in terrestrial ecosystems is dominated by plant litter (nonliving organic material), which, in addition to its role in material cycling, can act as a niche factor modulating interactions among plants. Litter thus links traditional community and ecosystem processes, which are often studied separately. We explore this connection using population dynamics models of two plant species and a litter pool. We first find conditions determining the outcome of interactions between these species, highlighting the role that litter plays and the role of broader ecosystem parameters, such as decomposition rate. Species trade-offs in tolerance to direct competition and litter-based interference competition allow for coexistence, provided the litter-tolerant species produces more litter at the population level; otherwise, priority effects may result. When species coexist, litter-mediated interactions between plants disrupt the traditional relationship between biomass accumulation and decomposition. Increasing decomposition rate may have no effect on standing litter density and, in some cases, may even increase litter load. These results illustrate how ecosystem variables can influence community outcomes that then feed back to influence the ecosystem.

Viral dynamics within plant hosts can be important for understanding plant disease prevalence and impacts. However, few mathematical modeling efforts aim to characterize within-plant viral dynamics. In this paper, we derive a simple system of delay differential equations that describes the spread of infection throughout the plant by barley and cereal yellow dwarf viruses via the cell-to-cell mechanism. By incorporating ratio-dependent incidence function and logistic growth of the healthy cells, the model can capture a wide range of biologically relevant phenomena via the disease-free, endemic, mutual extinction steady states, and a stable periodic orbit. We show that when the basic reproduction number is less than $ 1 $ ($ R_0 < 1 $), the disease-free steady state is asymptotically stable. When $ R_0>1 $, the dynamics either converge to the endemic equilibrium or enter a periodic orbit. Using a ratio-dependent transformation, we show that if the infection rate is very high relative to the growth rate of healthy cells, then the system collapses to the mutual extinction steady state. Numerical and bifurcation simulations are provided to demonstrate our theoretical results. Finally, we carry out parameter estimation using experimental data to characterize the effects of varying nutrients on the dynamics of the system. Our parameter estimates suggest that varying the nutrient supply of nitrogen and phosphorous can alter the dynamics of the infection in plants, specifically reducing the rate of viral production and the rate of infection in certain cases.

Diversity and nitrogen addition have positive relationships with plant productivity, yet climate-induced changes in water availability threaten to upend these established relationships. Using long-term data from three experiments in a mesic grassland (ranging from 17 to 34 yr of data), we tested how the effects of species richness and nitrogen addition on community-level plant productivity changed as a function of annual fluctuations in water availability using growing season precipitation and the Standardized Precipitation-Evapotranspiration Index (SPEI). While results varied across experiments, our findings demonstrate that water availability can magnify the positive effects of both biodiversity and nitrogen addition on productivity. These results suggest that productivity responses to anthropogenic species diversity loss and increasing nitrogen deposition could depend on precipitation regimes, highlighting the importance of testing interactions between multiple global change drivers.

Early detection and eradication of invasive plants are more cost-effective than managing well-established invasive plant populations and their impacts. However, there is high uncertainty around which taxa are likely to become invasive in a given area. Horizon scanning, which pairs rapid risk assessment with consensus building among experts, can help identify invasion threats. We performed a horizon scan of potential invasive plant threats to Florida, USA—a state with a high influx of introduced species, conditions that are favorable for plant establishment, and a history of negative impacts from invasive plants. We began with a list of 2128 non-native plant species and subspecies that are crop pests or invasive somewhere in the world and used publicly available data to prioritize 100 taxa for rapid risk assessment. We derived overall invasion risk scores by evaluating the likelihood and certainty of each of the 100 taxa arriving, establishing, and having an impact in Florida. Through the rapid risk assessments and a consensus-building discussion, we identified six plant taxa with high overall risk scores ranging from 75 to 100 out of a possible 125. The six taxa are globally distributed, easily transported to new areas, found in regions with climates similar to Florida’s, and can impact native plant communities, human health, or agriculture. We recommend more thorough risk assessments for each of these six species and, if appropriate, policy and management actions to limit invasive plant introduction and establishment in Florida.

Infectious diseases and invasive species can be strong drivers of biological systems that may interact to shift plant community composition. For example, disease can modify resource competition between invasive and native species. Invasive species tend to interact with a diversity of native species, and it is unclear how native species differ in response to disease-mediated competition with invasive species. Here, we quantified the biomass responses of three native North American grass species (Dichanthelium clandestinum, Elymus virginicus, and Eragrostis spectabilis) to disease-mediated competition with the non-native invasive grass Microstegium vimineum. The foliar fungal pathogen Bipolaris gigantea has recently emerged in Microstegium populations, causing a leaf spot disease that reduces Microstegium biomass and seed production. In a greenhouse experiment, we examined the effects of B. gigantea inoculation on two components of competitive ability for each native species: growth in the absence of competition and biomass responses to increasing densities of Microstegium. Bipolaris gigantea inoculation affected each of the three native species in unique ways, by increasing (Dichanthelium), decreasing (Elymus), or not changing (Eragrostis) their growth in the absence of competition relative to mock inoculation. Bipolaris gigantea inoculation did not, however, affect Microstegium biomass or mediate the effect of Microstegium density on native plant biomass. Thus, B. gigantea had species-specific effects on native plant competition with Microstegium through species-specific biomass responses to B. gigantea inoculation, but not through modified responses to Microstegium density. Our results suggest that disease may uniquely modify competitive interactions between invasive and native plants for different native plant species.

Summary Nutrient supply rates to hosts can mediate host–pathogen interactions. In terrestrial systems, nutrient supply to plants is mediated by soil microbes, suggesting a potential indirect effect of soil microbes on plant–pathogen interactions. Soil microbes also may affect plant pathogens by inducing plant defenses. We tested the role of soil microbes, nitrogen supply to plant hosts, and co-inoculation on infection by aphid-vectored RNA viruses, Barley Yellow Dwarf Virus (BYDV-PAV) and Cereal Yellow Dwarf Virus (CYDV-RPV), in a grass host grown in soil microbes collected from a long-term nitrogen enrichment experiment. BYDV-PAV incidence declined with high nitrogen supply, co-inoculation, or presence of soil microbes exposed to long-term low nitrogen enrichment. However, when combined, the negative effects of these treatments were sub-additive: nitrogen and co-inoculation did not reduce BYDV-PAV incidence in plants grown with the soil microbes. While soil microbes impacted leaf chlorophyll, they did not alter biomass or CYDV-RPV incidence. Soil microbes mediated the effects of nitrogen supply and co-inoculation on infection incidence and the effects of infection on host symptoms. Thus, soil microbial communities can indirectly control disease dynamics, altering the effects of nitrogen enrichment on plant–pathogen and pathogen–pathogen interactions in terrestrial systems.

The densities of highly competent plant hosts (i.e. those that are susceptible to and successfully transmit a pathogen) may shape pathogen community composition and disease severity, altering disease risk and impacts. Life history and evolutionary history can influence host competence; longer lived species tend to be better defended than shorter lived species and pathogens adapt to infect species with which they have longer evolutionary histories. It is unclear, however, how the densities of species that differ in competence due to life and evolutionary histories affect plant pathogen community composition and disease severity. We examined foliar fungal pathogens of two host groups in a California grassland: native perennial and non-native annual grasses. We first characterized pathogen community composition and disease severity of the two host groups to approximate differences in competence. We then used observational and manipulated gradients of native perennial and non-native annual grass densities to assess the effects of each host group on pathogen community composition and disease severity in 1-m(2)plots. Native perennial and non-native annual grasses hosted distinct pathogen communities but shared generalist pathogens. Native perennial grasses experienced 26% higher disease severity than non-native annuals. Only the observational gradient of native perennial grass density affected disease severity; there were no other significant relationships between host group density and either disease severity or pathogen community composition. Synthesis. The life and evolutionary histories of grasses likely influence their competence for different pathogen species, exemplified by distinct pathogen communities and differences in disease severity. However, there was limited evidence that the density of either host group affected pathogen community composition or disease severity. Therefore, competence for different pathogens likely shapes pathogen community composition and disease severity but may not interact with host density to alter disease risk and impacts at small scales.