Heterogeneous landscape promotes distinct microbial communities in an imperiled scrub ecosystem.

Habitat heterogeneity is a key driver of biodiversity of macroorganisms, yet how heterogeneity structures belowground microbial communities is not well understood. Importantly, belowground microbial communities may respond to any number of abiotic, biotic, and spatial drivers found in heterogeneous environments. Here, we examine potential drivers of prokaryotic and fungal communities in soils across the heterogenous landscape of the imperiled Florida scrub, a pyrogenic ecosystem where slight differences in elevation lead to large changes in water and nutrient availability and vegetation composition. We employ a comprehensive, large-scale sampling design to characterize the communities of prokaryotes and fungi associated with three habitat types and two soil depths (crust and subterranean) to evaluate (i) differences in microbial communities across these heterogeneous habitats, (ii) the relative roles of abiotic, biotic, and spatial drivers in shaping community structure, and (iii) the distribution of fungal guilds across these habitats. We sequenced soils from 40 complete replicates of habitat × soil depth combinations and sequenced the prokaryotic 16S and fungal internal transcribed spacer (ITS) regions using Illumina MiSeq. Habitat heterogeneity generated distinct communities of soil prokaryotes and fungi. Spatial distance played a role in structuring crust communities, whereas subterranean microbial communities were primarily structured by the shrub community, whose roots they presumably interacted with. This result helps to explain the unexpected transition we observed between arbuscular mycorrhiza-dominated soils at low-elevation habitats to ectomycorrhiza-dominated soils at high-elevation habitats. Our results challenge previous notions of environmental determinism of microbial communities and generate new hypotheses regarding symbiotic relationships across heterogeneous environments.

Allelopathy-selected microbiomes mitigate chemical inhibition of plant performance.

Allelopathy is a common and important stressor that shapes plant communities and can alter soil microbiomes, yet little is known about the direct effects of allelochemical addition on bacterial and fungal communities or the potential for allelochemical-selected microbiomes to mediate plant performance responses, especially in habitats naturally structured by allelopathy. Here, we present the first community-wide investigation of microbial mediation of allelochemical effects on plant performance by testing how allelopathy affects soil microbiome structure and how these microbial changes impact germination and productivity across 13 plant species. The soil microbiome exhibited significant changes to 'core' bacterial and fungal taxa, bacterial composition, abundance of functionally important bacterial and fungal taxa, and predicted bacterial functional genes after the addition of the dominant allelochemical native to this habitat. Furthermore, plant performance was mediated by the allelochemical-selected microbiome, with allelopathic inhibition of plant productivity moderately mitigated by the microbiome. Through our findings, we present a potential framework to understand the strength of plant-microbial interactions in the presence of environmental stressors, in which frequency of the ecological stress may be a key predictor of microbiome-mediation strength.

What kills the virtually immortal palms of the Florida scrub?

PREMISE: Life span varies greatly across plants, with some species being capable of extreme longevity. Yet even long-lived individuals are susceptible to climatic events, fire, and other challenges. We examined rare mortality events and their causes in two long-lived palmettos over four decades. METHODS: We monitored the survival of the clonal Serenoa repens and non-clonal, Florida-endemic Sabal etonia from 1981 to 2022 in four habitats along an elevational gradient within the globally imperiled Florida scrub ecosystem. We considered several challenges to palmetto survival, including extreme fires, shading due to lack of fire, droughts, periods of high precipitation, and possible pathogens. RESULTS: Survival of palmettos was remarkably high, and mortality was infrequent (Serenoa: cumulative, 5.7%; annualized, 0%-0.68%; Sabal: cumulative, 3.5%; annualized, 0%-0.43%). Mortality was highest in higher-elevation habitats with greater soil drainage, and smaller palmettos were more likely to die. When subjected to extreme fire, Serenoa suffered greater mortality than Sabal. Mortality in long-unburned habitats with increased shading rivaled that which occurred with extreme fire. There was no evidence of mortality due to lethal bronzing palm disease. CONCLUSIONS: Both palmettos had exceptionally low mortality rates, which, coupled with earlier work showing slow rates of transition from seedling to adult and remarkable adult longevity, suggest notably low rates of population turnover. Observed mortality in long-unburned habitats suggests the importance of fire-management planning with prescription burning. Lengthy age to reproduction and/or dependency on clonal propagation limits migration or genetic adaptation to altered conditions caused by climate change.

Microbiome-mediated response to pulse fire disturbance outweighs the effects of fire legacy on plant performance.

Fire plays a major role in structuring plant communities across the globe. Interactions with soil microbes impact plant fitness, scaling up to influence plant populations and distributions. Here we present the first factorial manipulation of both fire and soil microbiome presence to investigate their interactive effects on plant performance across a suite of plant species with varying life history traits. We conducted fully factorial experiments on 11 species from the Florida scrub ecosystem to test plant performance responses to soils with varying fire histories (36 soil sources), the presence/absence of a microbiome, and exposure to an experimental burn. Results revealed interactive 'pulse' effects between fire and the soil microbiome on plant performance. On average, post-fire soil microbiomes strongly reduced plant productivity compared to unburned or sterilized soils. Interestingly, longer-term fire 'legacy' effects had minor impacts on plant performance and were unrelated to soil microbiomes. While pulse fire effects on plant-microbiome interactions are short-term, they could have long-term consequences for plant communities by establishing differential microbiome-mediated priority effects during post-disturbance succession. The prominence of pulse fire effects on plant-microbe interactions has even greater import due to expected increases in fire disturbances resulting from anthropogenic climate change.

Community of Bark and Ambrosia Beetles (Coleoptera: Curculionidae: Scolytinae and Platypodinae) Infesting Brazilian Peppertree Treated With Herbicide and the Volatile Tree Response.

Brazilian peppertree, Schinus terebinthifolia Raddi (Anacardiaceae), is one of the most invasive weeds of natural and agricultural areas of Florida, Hawaii, and Texas (USA). Herbicides are the main tool used to manage populations of this weed. Faunal inventories of the insects associated with invasive populations of the weed have mostly listed leaf-feeding phytophagous, pollinator, or predacious species. Among these, bark and ambrosia beetles were collected only once from S. terebinthifolia in the invaded range and there are no reports from the native range. A diverse assemblage of bark and ambrosia beetles, many well-known economic pests of ornamentals, was reared from S. terebinthifolia bolts collected at a restoration site in Florida that had been treated with herbicide (triclopyr ester). A similar collection of beetles was captured on ethanol-baited sticky traps. No beetles emerged from bolts of untreated trees, almost none emerged from those wounded with a machete (3.1% of total), whereas nearly all the beetles collected emerged from bolts that had been treated with herbicide (62.3%) or the combination wounded + herbicide (34.6%). Ethanol was detected from the herbicide and wound + herbicide-treated bolts suggesting this was the attractive kairomone. Abundant amounts of other volatiles were collected from all bolts, especially from the wounded treatment, but no association was detected between volatile emissions and beetle infestation. Further studies are needed to determine whether invasive populations of S. terebinthifolia treated with herbicides constitute reservoirs for pest bark and ambrosia beetles that may spill over onto neighboring ornamental hosts.

Localized Induced Defenses Limit Gall Formation by Eriophyid Mite Against Invasive Lygodium microphyllum (Schizaeales: Lygodiaceae).

A potential barrier to the establishment of weed biological control agents is interference from other management tactics that induce plant defenses. Methods that suppress the weed such as feeding by other biological control agents or mechanical removal are especially disposed to inducing plant defenses and potentially limiting agent establishment. Here, we focused on the invasive weed Lygodium microphyllum (Cav.) R. Br. (Schizaeales: Lygodiaceae, Old World climbing fern) and one of its biological control agents, the mite Floracarus perrepae Knihinicki and Boczek (Acariformes: Eriophyidae). We experimentally induced plant defenses in potted plants via damage or application of jasmonic acid, a hormone typically involved in plant defenses, and measured the responses of the mite in a screenhouse. Localized damage to the pinnae (e.g., leaflets) via cutting or larval feeding from a second biological control agent, Neomusotima conspurcatalis (Warren) (Lepidoptera; Crambidae), reduced F. perrepae gall formation, but not the number of mites per gall. In contrast, damage to rachises (e.g., stems) did not affect galling, likely because plant defense responses were not systemic. Application of jasmonic acid reduced gall formation but not the numbers of mites within galls. Taken together, we found that localized damage interfered with gall formation but not within-gall reproduction. However, these effects on the mite from induced plant defenses are likely short-lived, and therefore interference between management tactics is unlikely to affect F. perrepae establishment and performance.

Water availability modifies productivity response to biodiversity and nitrogen in long-term grassland experiments.

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.

Environmental stress destabilizes microbial networks.

Environmental stress is increasing worldwide, yet we lack a clear picture of how stress disrupts the stability of microbial communities and the ecosystem services they provide. Here, we present the first evidence that naturally-occurring microbiomes display network properties characteristic of unstable communities when under persistent stress. By assessing changes in diversity and structure of soil microbiomes along 40 replicate stress gradients (elevation/water availability gradients) in the Florida scrub ecosystem, we show that: (1) prokaryotic and fungal diversity decline in high stress, and (2) two network properties of stable microbial communities-modularity and negative:positive cohesion-have a clear negative relationship with environmental stress, explaining 51-78% of their variation. Interestingly, pathogenic taxa/functional guilds decreased in relative abundance along the stress gradient, while oligotrophs and mutualists increased, suggesting that the shift in negative:positive cohesion could result from decreasing negative:positive biotic interactions consistent with the predictions of the Stress Gradient Hypothesis. Given the crucial role microbiomes play in ecosystem functions, our results suggest that, by limiting the compartmentalization of microbial associations and creating communities dominated by positive associations, increasing stress in the Anthropocene could destabilize microbiomes and undermine their ecosystem services.

Colonization by Biological Control Agents on Post-Fire Regrowth of Invasive Lygodium microphyllum (Lygodiaceae).

Integration of biological control with other management tactics such as prescribed burning is often important for successful invasive weed control. A critical step in this integration is determining whether the agent can colonize postburn growth of the weed. Here, we investigated postburn colonization by biological control agents on regrowth of the invasive vine Lygodium microphyllum (Cav.) R. Br. (Lygodiaceae, Old World climbing fern) in Florida. We monitored regrowth and subsequent colonization of two agents already established in Florida-the gall-inducing mite Floracarus perrepae Knihinicki and Boczek (Acariformes: Eriophyidae) and the foliage-feeding moth Neomusotima conspurcatalis Warren (Lepidoptera: Crambidae)-following three prescribed burns. We provide the first report of natural colonization by the F. perrepae mite and N. conspurcatalis moth on postburn L. microphyllum regrowth, and this colonization typically began 5-9 mo postburn. Furthermore, we report that L. microphyllum can recover to prefire levels of percent cover in as little as 5 mo. Our findings indicate that biological control of L. microphyllum has the potential to be integrated with prescribed burns.

Do plant-microbe interactions support the Stress Gradient Hypothesis?

The Stress Gradient Hypothesis (SGH), which predicts increasing ratios of facilitative:competitive interactions with increasing stress, has long been a guiding framework for conceptualizing plant-plant interactions. Recently, there has been a growing recognition of the roles of microbes in mitigating or exacerbating environmental stress for their plant hosts. As such, we might predict, based on the SGH, that beneficial microbial effects on plant performance should be positively associated with stress. Specifically, we hypothesized that support for the SGH would depend on the host plant's habitat specialization such that species that specialize in high stress habitats and thus likely coevolved with the resident microbes would exhibit stronger support for the SGH than non-specialist plant species. We further hypothesized that support for the SGH would vary with germination frequency, since boosting germination of low-frequency germinators is one effective means by which microbes can benefit plant species performance. Here, we explore whether plant-microbial interactions support the SGH using 12 plant species native to the Florida rosemary scrub. We conducted factorial experiments that manipulated the presence of microbes in nine soils collected along an elevational stress gradient, and recorded germination frequency and biomass. Microbes increased the germination frequency of four species, all of which had relatively low germination rates. Furthermore, we found support for the SGH in nearly one-half of the species examined, with soil microbes facilitating germination with increasing stress for 5 of the 12 species tested, and none of the species exhibiting the opposite trend. Support for the SGH was not predicted by either the plant hosts' habitat specialization or germination frequency. In contrast to germination, biomass results showed little support for the SGH, with four of 12 species refuting and one species supporting SGH predictions. Taken together, our study documents that interactions between the soil microbial community and plant species along a stress gradient can support the SGH, but emphasizes that these effects are life-history-stage dependent. This work also identifies a common mechanism (germination facilitation) by which microbes can benefit plant species in stressful habitats.

Use of standardized methods to improve extinction-risk classification.

Standardized classification methods based on quantifiable risk metrics are critical for evaluating extinction threats because they increase objectivity, consistency, and transparency of listing decisions. Yet, in the United States, neither federal nor state agencies use standardized methods for listing species for legal protection, which could put listing decisions at odds with the magnitude of the risk. We used a recently developed set of quantitative risk metrics for California herpetofauna as a case study to highlight discrepancies in listing decisions made without standardized methods. We also combined such quantitative metrics with classification tree analysis to attempt to increase the transparency of previous listing decisions by identifying the criteria that had inherently been given the most weight. Federally listed herpetofauna in California scored significantly higher on the risk-metric spectrum than those not federally listed, whereas state-listed species did not score any higher than species that were not state listed. Based on classification trees, state endemism was the most important predictor of listing status at the state level and distribution trend (decline in a species' range size) and population trend (decline in a species' abundance at localized sites) were the most important predictors at the federal level. Our results emphasize the need for governing bodies to adopt standardized methods for assessing conservation risk that are based on quantitative criteria. Such methods allow decision makers to identify criteria inherently given the most weight in determining listing status, thus increasing the transparency of previous listing decisions, and produce an unbiased comparison of conservation threat across all species to promote consistency, efficiency, and effectiveness of the listing process.

Uso de Métodos Estandarizados para Mejorar la Clasificación del Riesgo de Extinción Resumen Los métodos estandarizados de clasificación basados en medidas cuantificables del riesgo de extinción son sumamente importantes para evaluar las amenazas de extinción ya que incrementan la objetividad, consistencia y transparencia de las decisiones de listado. Aún así, en los Estados Unidos, ni las agencias federales ni las estatales usan métodos estandarizados para enlistar a las especies para su protección legal, lo que podría poner en discrepancia a las decisiones de listado con la magnitud del riesgo. Usamos un conjunto de medidas cuantitativas del riesgo, desarrollado recientemente para la herpetofauna de California, como un estudio de caso que nos permitiera resaltar las discrepancias en las decisiones de listado hechas sin métodos estandarizados. También combinamos dichas medidas cuantitativas con un análisis de árbol de clasificación para intentar incrementar la transparencia de las decisiones de listado previas al identificar los criterios a los cuales se les había otorgado mayor peso inherentemente. La herpetofauna de California que se encontraba enlistada a nivel federal tuvo un puntaje significativamente más alto en el espectro de la medida del riesgo que aquellas especies que no estaban enlistadas, mientras que las especies enlistadas a nivel estatal no tuvieron un puntaje más alto que aquellas especies que no estaban enlistadas a nivel estatal. Con base en los árboles de clasificación, el endemismo estatal fue el indicador más importante del estado de listado a nivel estatal y tanto la tendencia de distribución (declinación del tamaño de la extensión de una especie) y como la tendencia poblacional (declinación de la abundancia de una especie en sitios localizados) fueron los indicadores más importantes a nivel federal. Nuestros resultados enfatizan la necesidad que tienen los cuerpos de gobierno de adoptar los métodos estandarizados que están basados en criterios cuantitativos para la evaluación del riesgo de conservación. Dichos métodos permiten que quienes toman las decisiones identifiquen los criterios a los cuales se les otorga inherentemente el mayor peso al determinar el estado de listado, lo que incrementa la transparencia de las decisiones previas de listado, y produce una comparación sin sesgos de la amenaza de conservación en todas las especies para promover la regularidad, eficiencia y efectividad de los procesos de listado.

Soil Microbiomes Underlie Population Persistence of an Endangered Plant Species.

Microbiomes can dramatically alter individual plant performance, yet how these effects influence higher-order processes is not well resolved. In particular, little is known about how microbiome effects on individual plants alter plant population dynamics, a question critical to imperiled species conservation. Here we integrate bioassays, multidecadal demographic data, and integral projection modeling to determine how the presence of the natural soil microbiome underlies plant population dynamics. Simulations indicated that the presence of soil microbiomes boosted population growth rates (λ) of the endangered Hypericum cumulicola by 13% on average, the difference between population growth versus decline in 76% of patches. The greatest benefit (47% increase in λ) occurred in low-nutrient, high-elevation habitats, suggesting that the soil microbiome may help expand H. cumulicola's distribution to include these stressful habitats. Our results demonstrate that soil microbiomes can significantly affect plant population growth and persistence and support the incorporation of soil microbiomes into conservation planning.

Testing for loss of Epichloë and non-epichloid symbionts under altered rainfall regimes.

PREMISE: Microbial symbionts can buffer plant hosts from environmental change. Therefore, understanding how global change factors alter the associations between hosts and their microbial symbionts may improve predictions of future changes in host population dynamics and microbial diversity. Here, we investigated how one global change factor, precipitation, affected the maintenance or loss of symbiotic fungal endophytes in a C3 grass host. Specifically, we examined the distinct responses of Epichloë (vertically transmitted and systemic) and non-epichloid endophytes (typically horizontally transmitted and localized) by considering (1) how precipitation altered associations with Epichloë and non-epichloid endophytic taxa across host ontogeny, and (2) interactive effects of water availability and Epichloë on early seedling life history stages. METHODS: We manipulated the presence of Epichloë amarillans in American beachgrass (Ammophila breviligulata) in a multiyear field experiment that imposed three precipitation regimes (ambient or ±30% rainfall). In laboratory assays, we investigated the interactive effects of water availability and Epichloë on seed viability and germination. RESULTS: Reduced precipitation decreased the incidence of Epichloë in leaves in the final sampling period, but had no effect on associations with non-epichloid taxa. Epichloë reduced the incidence of non-epichloid endophytes, including systemic p-endophytes, in seeds. Laboratory assays suggested that association with Epichloë is likely maintained, in part, due to increased seed viability and germination regardless of water availability. CONCLUSIONS: Our study empirically demonstrates several pathways for plant symbionts to be lost or maintained across host ontogeny and suggests that reductions in precipitation can drive the loss of a plant's microbial symbionts.

Wind speed predicts population dynamics of the eriophyid mite Floracarus perrepae on invasive Old World climbing fern (Lygodium microphyllum) in a shade house colony.

Lygodium microphyllum is one of the most noxious invasive plants in Florida, USA, smothering native vegetation in cypress swamps, pine flatwoods, and Everglades tree islands and altering fire regimes. The eriophyid mite Floracarus perrepae was introduced from Australia to help control L. microphyllum infestations. While F. perrepae exhibits high population growth rates in its native range, its population dynamics in Florida are unknown, particularly the dynamics that occur within the leaf roll galls the mite induces on the margins of leaves. Here, we monitored a shade house colony of F. perrepae in south Florida for 2 years to identify seasonal patterns and potential climate drivers of within-gall mite density. Gall dissections of mite-infested colony plants were performed monthly. Mite density within galls exhibited two cycles per year: a strong cycle that boomed in spring and busted in summer, and a weak cycle that moderately increased mite density in fall and declined in winter. Climate variables, particularly those related to wind speed, were positively associated with higher mite density. Our study sheds light on the within-gall dynamics of F. perrepae and suggests that the highest within-gall mite densities occur in the spring and fall.

Microbial mitigation-exacerbation continuum: a novel framework for microbiome effects on hosts in the face of stress.

A key challenge to understanding microbiomes and their role in ecological processes is contextualizing their effects on host organisms, particularly when faced with environmental stress. One influential theory, the Stress Gradient Hypothesis, might predict that the frequency of positive interactions increases with stressful conditions such that microbial taxa would mitigate harmful effects on host performance. Yet, equally plausible is that microbial taxa could exacerbate these effects. Here, we introduce the Mitigation-Exacerbation Continuum as a novel framework to conceptualize microbial mediation of stress. We (1) use this continuum to quantify microbial mediation of stress for six plant species and (2) test the association between these continuum values and natural species' abundance. We factorially manipulated a common stress (allelopathy) and the presence of soil microbes to quantify microbial effects in benign and stressed environments for two critical early life-history metrics, seed germination and seedling biomass. Although we found evidence of both mitigation and exacerbation among the six species, exacerbation was more common. Across species, the degree of microbial-mediated effects on germination explained >80% of the variation of natural field abundances. Our results suggest a critical role of soil microbes in mediating plant stress responses, and a potential microbial mechanism underlying species abundance.

Disentangling environmental and host sources of fungal endophyte communities in an experimental beachgrass study.

Disentangling the ecological factors that contribute to the assembly of the microbial symbiont communities within eukaryotic hosts is an ongoing challenge. Broadly speaking, symbiont propagules arrive either from external sources in the environment or from internal sources within the same host individual. To understand the relative importance of these propagule sources to symbiont community assembly, we characterized symbiotic fungal endophyte communities within the roots of three species of beachgrass in a field experiment. We manipulated two aspects of the external environment, successional habitat and physical disturbance. To determine the role of internal sources of propagules for endophyte community assembly, we used beachgrass individuals with different pre-existing endophyte communities. Endophyte species richness and community composition were characterized using culture-based and next-generation sequencing approaches. Our results showed that external propagule sources associated with successional habitat, but not disturbance, were particularly important for colonization of most endophytic taxa. In contrast, internal propagule sources played a minor role for most endophytic taxa but were important for colonization by the dominant taxon Microdochium bolleyi. Our findings highlight the power of manipulative field experiments to link symbiont community assembly to its underlying ecological processes, and to ultimately improve predictions of symbiont community assembly across environments.

Quantifying the associations between fungal endophytes and biocontrol-induced herbivory of invasive purple loosestrife (Lythrum salicaria L.).

Fungal endophytes are one of several groups of heterotrophic organisms that associate with living plants. The net effects of these groups of organisms on each other and ultimately on their host plants depend in part on how they facilitate or antagonize one another. In this study we quantified the associations between endophyte communities and herbivory induced by a biological control in the invasive Lythrum salicaria at various spatial scales using a culture-based approach. We found positive associations between herbivory damage and endophyte isolation frequency and richness at the site level and weak, positive associations at the leaf level. Herbivory damage was more strongly influenced by processes at the site level than were endophyte isolation frequency and community structure, which were influenced by processes at the plant and leaf levels. Furthermore, endophytic taxa found in low herbivory sites were nested subsets of those taxa found at high herbivory sites. Our findings suggest that endophyte communities of L. salicaria are associated with, and potentially facilitated by, biocontrol-induced herbivory. Quantifying the associations between heterotrophic groups ultimately may lead to a clearer understanding of their complex interactions with plants.

Draft Genome Sequence of Microdochium bolleyi, a Dark Septate Fungal Endophyte of Beach Grass.

Here, we present the genome sequence of the dark septate fungal endophyte Microdochium bolleyi (Ascomycota, Sordariomycetes, Xylariales). The assembled genome size was 38.84 Mbp and consisted of 173 scaffolds and 13,177 predicted genes.

Plant Host Species and Geographic Distance Affect the Structure of Aboveground Fungal Symbiont Communities, and Environmental Filtering Affects Belowground Communities in a Coastal Dune Ecosystem.

Microbial symbionts inhabit tissues of all plants and animals. Their community composition depends largely on two ecological processes: (1) filtering by abiotic conditions and host species determining the environments that symbionts are able to colonize and (2) dispersal-limitation determining the pool of symbionts available to colonize a given host and community spatial structure. In plants, the above- and belowground tissues represent such distinct habitats for symbionts that we expect different effects of filtering and spatial structuring on their symbiont communities. In this study, we characterized above- and belowground communities of fungal endophytes--fungi living asymptomatically within plants--to understand the contributions of filtering and spatial structure to endophyte community composition. We used a culture-based approach to characterize endophytes growing in leaves and roots of three species of coastal beachgrasses in dunes of the USA Pacific Northwest. For leaves, endophyte isolation frequency and OTU richness depended primarily on plant host species. In comparison, for roots, both isolation frequency and OTU richness increased from the nutrient-poor front of the dune to the higher-nutrient backdune. Endophyte community composition in leaves exhibited a distance-decay relationship across the region. In a laboratory assay, faster growth rates and lower spore production were more often associated with leaf- than root-inhabiting endophytes. Overall, our results reveal a greater importance of biotic filtering by host species and dispersal-limitation over regional geographic distances for aboveground leaf endophyte communities and stronger effects of abiotic environmental filtering and locally patchy distributions for belowground root endophyte communities.

Invasive congeners differ in successional impacts across space and time.

Invasive species can alter the succession of ecological communities because they are often adapted to the disturbed conditions that initiate succession. The extent to which this occurs may depend on how widely they are distributed across environmental gradients and how long they persist over the course of succession. We focus on plant communities of the USA Pacific Northwest coastal dunes, where disturbance is characterized by changes in sediment supply, and the plant community is dominated by two introduced grasses--the long-established Ammophila arenaria and the currently invading A. breviligulata. Previous studies showed that A. breviligulata has replaced A. arenaria and reduced community diversity. We hypothesize that this is largely due to A. breviligulata occupying a wider distribution across spatial environmental gradients and persisting in later-successional habitat than A. arenaria. We used multi-decadal chronosequences and a resurvey study spanning 2 decades to characterize distributions of both species across space and time, and investigated how these distributions were associated with changes in the plant community. The invading A. breviligulata persisted longer and occupied a wider spatial distribution across the dune, and this corresponded with a reduction in plant species richness and native cover. Furthermore, backdunes previously dominated by A. arenaria switched to being dominated by A. breviligulata, forest, or developed land over a 23-yr period. Ammophila breviligulata likely invades by displacing A. arenaria, and reduces plant diversity by maintaining its dominance into later successional backdunes. Our results suggest distinct roles in succession, with A. arenaria playing a more classically facilitative role and A. breviligulata a more inhibitory role. Differential abilities of closely-related invasive species to persist through time and occupy heterogeneous environments allows for distinct impacts on communities during succession.