Trajectories and tipping points of piñon-juniper woodlands after fire and thinning.

Piñon-juniper (PJ) woodlands are a dominant community type across the Intermountain West, comprising over a million acres and experiencing critical effects from increasing wildfire. Large PJ mortality and regeneration failure after catastrophic wildfire have elevated concerns about the long-term viability of PJ woodlands. Thinning is increasingly used to safeguard forests from fire and in an attempt to increase climate resilience. We have only a limited understanding of how fire and thinning will affect the structure and function of PJ ecosystems. Here, we examined vegetation structure, microclimate conditions, and PJ regeneration dynamics following ~20 years post-fire and thinning treatments. We found that burned areas had undergone a state shift that did not show signs of returning to their previous state. This shift was characterized by (1) distinct plant community composition dominated by grasses; (2) a lack of PJ recruitment; (3) a decrease in the sizes of interspaces in between plants; (4) lower abundance of late successional biological soil crusts; (5) lower mean and minimum daily soil moisture values; (6) lower minimum daily vapor pressure deficit; and (7) higher photosynthetically active radiation. Thinning created distinct plant communities and served as an intermediate between intact and burned communities. More intensive thinning decreased PJ recruitment and late successional biocrust cover. Our results indicate that fire has the potential to create drier and more stressful microsite conditions, and that, in the absence of active management following fire, there may be shifts to persistent ecological states dominated by grasses. Additionally, more intensive thinning had a larger impact on community structure and recruitment than less intensive thinning, suggesting that careful consideration of goals could help avoid unintended consequences. While our results indicate the vulnerability of PJ ecosystems to fire, they also highlight management actions that could be adapted to create conditions that promote PJ re-establishment.

Is there tree senescence? The fecundity evidence.

Despite its importance for forest regeneration, food webs, and human economies, changes in tree fecundity with tree size and age remain largely unknown. The allometric increase with tree diameter assumed in ecological models would substantially overestimate seed contributions from large trees if fecundity eventually declines with size. Current estimates are dominated by overrepresentation of small trees in regression models. We combined global fecundity data, including a substantial representation of large trees. We compared size-fecundity relationships against traditional allometric scaling with diameter and two models based on crown architecture. All allometric models fail to describe the declining rate of increase in fecundity with diameter found for 80% of 597 species in our analysis. The strong evidence of declining fecundity, beyond what can be explained by crown architectural change, is consistent with physiological decline. A downward revision of projected fecundity of large trees can improve the next generation of forest dynamic models.

Sequential hybridization may have facilitated ecological transitions in the Southwestern pinyon pine syngameon.

Multispecies interbreeding networks, or syngameons, have been increasingly reported in natural systems. However, the formation, structure, and maintenance of syngameons have received little attention. Through gene flow, syngameons can increase genetic diversity, facilitate the colonization of new environments, and contribute to hybrid speciation. In this study, we evaluated the history, patterns, and consequences of hybridization in a pinyon pine syngameon using morphological and genomic data to assess genetic structure, demographic history, and geographic and climatic data to determine niche differentiation. We demonstrated that Pinus edulis, a dominant species in the Southwestern US and a barometer of climate change, is a core participant in the syngameon, involved in the formation of two drought-adapted hybrid lineages including the parapatric and taxonomically controversial fallax-type. We found that species remain morphologically and genetically distinct at range cores, maintaining species boundaries while undergoing extensive gene flow in areas of sympatry at range peripheries. Our study shows that sequential hybridization may have caused relatively rapid speciation and facilitated the colonization of different niches, resulting in the rapid formation of two new lineages. Participation in the syngameon may allow adaptive traits to be introgressed across species barriers and provide the changes needed to survive future climate scenarios.

Limits of thermal and hydrological tolerance in a foundation tree species (Populus fremontii) in the desert southwestern United States.

Populus fremontii is among the most dominant, and ecologically important riparian tree species in the western United States and can thrive in hyper-arid riparian corridors. Yet, P. fremontii forests have rapidly declined over the last decade, particularly in places where temperatures sometimes exceed 50°C. We evaluated high temperature tolerance of leaf metabolism, leaf thermoregulation, and leaf hydraulic function in eight P. fremontii populations spanning a 5.3°C mean annual temperature gradient in a well-watered common garden, and at source locations throughout the lower Colorado River Basin. Two major results emerged. First, despite having an exceptionally high Tcrit (the temperature at which Photosystem II is disrupted) relative to other tree taxa, recent heat waves exceeded Tcrit , requiring evaporative leaf cooling to maintain leaf-to-air thermal safety margins. Second, in midsummer, genotypes from the warmest locations maintained lower midday leaf temperatures, a higher midday stomatal conductance, and maintained turgor pressure at lower water potentials than genotypes from more temperate locations. Taken together, results suggest that under well-watered conditions, P. fremontii can regulate leaf temperature below Tcrit along the warm edge of its distribution. Nevertheless, reduced Colorado River flows threaten to lower water tables below levels needed for evaporative cooling during episodic heat waves.

Globally, tree fecundity exceeds productivity gradients.

Lack of tree fecundity data across climatic gradients precludes the analysis of how seed supply contributes to global variation in forest regeneration and biotic interactions responsible for biodiversity. A global synthesis of raw seedproduction data shows a 250-fold increase in seed abundance from cold-dry to warm-wet climates, driven primarily by a 100-fold increase in seed production for a given tree size. The modest (threefold) increase in forest productivity across the same climate gradient cannot explain the magnitudes of these trends. The increase in seeds per tree can arise from adaptive evolution driven by intense species interactions or from the direct effects of a warm, moist climate on tree fecundity. Either way, the massive differences in seed supply ramify through food webs potentially explaining a disproportionate role for species interactions in the wet tropics.

Evidence of climate-driven selection on tree traits and trait plasticity across the climatic range of a riparian foundation species.

Selection on quantitative traits by heterogeneous climatic conditions can lead to substantial trait variation across a species range. In the context of rapidly changing environments, however, it is equally important to understand selection on trait plasticity. To evaluate the role of selection in driving divergences in traits and their associated plasticities within a widespread species, we compared molecular and quantitative trait variation in Populus fremontii (Fremont cottonwood), a foundation riparian distributed throughout Arizona. Using SNP data and genotypes from 16 populations reciprocally planted in three common gardens, we first performed QST -FST analyses to detect selection on traits and trait plasticity. We then explored the environmental drivers of selection using trait-climate and plasticity-climate regressions. Three major findings emerged: (1) There was significant genetic variation in traits expressed in each of the common gardens and in the phenotypic plasticity of traits across gardens, both of which were heritable. (2) Based on QST -FST comparisons, there was evidence of selection in all traits measured; however, this result varied from no effect in one garden to highly significant in another, indicating that detection of past selection is environmentally dependent. We also found strong evidence of divergent selection on plasticity across environments for two traits. (3) Traits and/or their plasticity were often correlated with population source climate (R2 up to .77 and .66, respectively). These results suggest that steep climate gradients across the Southwest have played a major role in shaping the evolution of divergent phenotypic responses in populations and genotypes now experiencing climate change.

Tradeoffs between leaf cooling and hydraulic safety in a dominant arid land riparian tree species.

Leaf carbon gain optimization in hot environments requires balancing leaf thermoregulation with avoiding excessive water loss via transpiration and hydraulic failure. The tradeoffs between leaf thermoregulation and transpirational water loss can determine the ecological consequences of heat waves that are increasing in frequency and intensity. We evaluated leaf thermoregulation strategies in warm- (>40°C maximum summer temperature) and cool-adapted (<40°C maximum summer temperature) genotypes of the foundation tree species, Populus fremontii, using a common garden near the mid-elevational point of its distribution. We measured leaf temperatures and assessed three modes of leaf thermoregulation: leaf morphology, midday canopy stomatal conductance and stomatal sensitivity to vapour pressure deficit. Data were used to parameterize a leaf energy balance model to estimate contrasts in midday leaf temperature in warm- and cool-adapted genotypes. Warm-adapted genotypes had 39% smaller leaves and 38% higher midday stomatal conductance, reflecting a 3.8°C cooler mean leaf temperature than cool-adapted genotypes. Leaf temperatures modelled over the warmest months were on average 1.1°C cooler in warm- relative to cool-adapted genotypes. Results show that plants adapted to warm environments are predisposed to tightly regulate leaf temperatures during heat waves, potentially at an increased risk of hydraulic failure.

Invasive and native grasses exert negative plant-soil feedbacks on the woody shrub Artemisia tridentata.

Displacement of diverse native plant communities by low-diversity invasive communities is a global problem. In the western United States, the displacement of sagebrush-dominated communities by cheatgrass has increased since the 1920s. Restoration outcomes are poor, potentially due to soil alteration by cheatgrass. We explored the poorly understood role of plant-soil feedbacks in the dominance of cheatgrass in a greenhouse study where uninvaded sagebrush soils were conditioned with either cheatgrass, a native bunchgrass or sagebrush. Sagebrush seedlings were grown in the soils that remained following the removal of conditioning plants. We expected cheatgrass to strongly suppress sagebrush due to a change in belowground microbial communities, conspecifics to have neutral effects and the native bunchgrass to have intermediate effects as it coevolved with sagebrush but belongs to a different functional group. We assessed the effects of conditioning on sagebrush growth, tissue nutrients, and carbon allocation. We also characterized the abundance, diversity and community composition of root microbial associates. Cheatgrass strongly suppressed sagebrush growth at high and low conditioning densities, the native bunchgrass showed suppression at high conditioning densities only and conspecific effects were neutral. Tissue nutrients, amount of root colonization by soil fungi or root microbial community composition were not associated with these plant-soil feedbacks. Although we did not identify the precise mechanism, our results provide key evidence that rapid soil alteration by cheatgrass results in negative plant-soil feedbacks on sagebrush growth. These feedbacks likely contribute to cheatgrass dominance and the poor success of sagebrush restoration.

Repeated Genetic and Adaptive Phenotypic Divergence across Tidal Elevation in a Foundation Plant Species.

AbstractMicrogeographic genetic divergence can create fine-scale trait variation. When such divergence occurs within foundation species, then it might impact community structure and ecosystem function and cause other cascading ecological effects. We tested for parallel microgeographic trait and genetic divergence in Spartina alterniflora, a foundation species that dominates salt marshes of the US Atlantic and Gulf coasts. Spartina is characterized by tall-form (1-2 m) plants at lower tidal elevations and short-form (<0.5 m) plants at higher tidal elevations, yet whether this trait variation reflects plastic and/or genetically differentiated responses to these environmental conditions remains unclear. In the greenhouse, seedlings raised from tall-form plants grew taller than those from short-form plants, indicating a heritable difference in height. When we reciprocally transplanted seedlings back into the field for a growing season, composite fitness (survivorship and seed production) and key plant traits (plant height and biomass allocation) differed interactively across origin and transplant zones in a manner indicative of local adaptation. Further, a survey of single nucleotide polymorphisms revealed repeated, independent genetic differentiation between tall- and short-form Spartina at five of six tested marshes across the native range. The observed parallel, microgeographic genetic differentiation in Spartina likely underpins marsh health and functioning and provides an underappreciated mechanism that might increase capacity of marshes to adapt to rising sea levels.

Plant response to fungal root endophytes varies by host genotype in the foundation species Spartina alterniflora.

PREMISE: Root-associated fungi provide a wide range of functions for their host plants, including nutrient provisioning, pathogen protection, and stress alleviation. In so doing, they can markedly influence host-plant structural and physiological traits, although the degree to which these effects vary within particular plant host species is not well understood. METHODS: We conducted a 7-month common-garden inoculation experiment to test the potential effects of a marine fungus (Lulwoana sp.) on the phenotypic traits of different genotypes of the host, the salt marsh plant species Spartina alterniflora. Lulwoana belongs to the dark septate endophytes (DSE), a polyphyletic group of fungi that are commonly found colonizing healthy plant roots, though their ecological role remains unclear. RESULTS: We documented significant impacts of Lulwoana on S. alterniflora morphology, biomass, and biomass allocation. For most traits in our study, these impacts varied significantly in direction and/or magnitude across S. alterniflora genotypes. Effects that were consistent across genotype were generally negative. Plant response was not predicted by the percentage of roots colonized, consistent with findings that dark septate endophytes do not necessarily influence plant growth responses through direct contact with roots. CONCLUSIONS: The observed changes in stem height, biomass, and biomass allocation have important effects on plant competitive ability, growth, and fitness, suggesting that plant-fungal interactions have community and ecosystem level effects in salt marshes.

Tree genetics defines fungal partner communities that may confer drought tolerance.

Plant genetic variation and soil microorganisms are individually known to influence plant responses to climate change, but the interactive effects of these two factors are largely unknown. Using long-term observational studies in the field and common garden and greenhouse experiments of a foundation tree species (Pinus edulis) and its mutualistic ectomycorrhizal fungal (EMF) associates, we show that EMF community composition is under strong plant genetic control. Seedlings acquire the EMF community of their seed source trees (drought tolerant vs. drought intolerant), even when exposed to inoculum from the alternate tree type. Drought-tolerant trees had 25% higher growth and a third the mortality of drought-intolerant trees over the course of 10 y of drought in the wild, traits that were also observed in their seedlings in a common garden. Inoculation experiments show that EMF communities are critical to drought tolerance. Drought-tolerant and drought-intolerant seedlings grew similarly when provided sterile EMF inoculum, but drought-tolerant seedlings grew 25% larger than drought-intolerant seedlings under dry conditions when each seedling type developed its distinct EMF community. This demonstration that particular combinations of plant genotype and mutualistic EMF communities improve the survival and growth of trees with drought is especially important, given the vulnerability of forests around the world to the warming and drying conditions predicted for the future.

Common garden experiments disentangle plant genetic and environmental contributions to ectomycorrhizal fungal community structure.

The interactions among climate change, plant genetic variation and fungal mutualists are poorly understood, but probably important to plant survival under drought. We examined these interactions by studying the ectomycorrhizal fungal (EMF) communities of pinyon pine seedlings (Pinus edulis) planted in a wildland ecosystem experiencing two decades of climate change-related drought. We established a common garden containing P. edulis seedlings of known maternal lineages (drought tolerant, DT; drought intolerant, DI), manipulated soil moisture and measured EMF community structure and seedling growth. Three findings emerged: EMF community composition differed at the phylum level between DT and DI seedlings, and diversity was two-fold greater in DT than in DI seedlings. EMF communities of DT seedlings did not shift with water treatment and were dominated by an ascomycete, Geopora sp. By contrast, DI seedlings shifted to basidiomycete dominance with increased moisture, demonstrating a lineage by environment interaction. DT seedlings grew larger than DI seedlings in high (28%) and low (50%) watering treatments. These results show that inherited plant traits strongly influence microbial communities, interacting with drought to affect seedling performance. These interactions and their potential feedback effects may influence the success of trees, such as P. edulis, in future climates.

Legacy effects of tree mortality mediated by ectomycorrhizal fungal communities.

Successive droughts have resulted in extensive tree mortality in the southwestern United States. Recovery of these areas is dependent on the survival and recruitment of young trees. For trees that rely on ectomycorrhizal fungi (EMF) for survival and growth, changes in soil fungal communities following tree mortality could negatively affect seedling establishment. We used tree-focused and stand-scale measurements to examine the impact of pinyon pine mortality on the performance of surviving juvenile trees and the potential for mutualism limitation of seedling establishment via altered EMF communities. Mature pinyon mortality did not affect the survival of juvenile pinyons, but increased their growth. At both tree and stand scales, high pinyon mortality had no effect on the abundance of EMF inocula, but led to altered EMF community composition including increased abundance of Geopora and reduced abundance of Tuber. Seedling biomass was strongly positively associated with Tuber abundance, suggesting that reductions in this genus with pinyon mortality could have negative consequences for establishing seedlings. These findings suggest that whereas mature pinyon mortality led to competitive release for established juvenile pinyons, changes in EMF community composition with mortality could limit successful seedling establishment and growth in high-mortality sites.

Beyond ICOM8: perspectives on advances in mycorrhizal research from 2015 to 2017.

This report reviews important advances in mycorrhizal research that occurred during the past 2 years. We highlight major advancements both within and across levels of biological organization and describe areas where greater integration has led to unique insights. Particularly active areas of research include exploration of the mechanisms underpinning the development of the mycorrhizal symbiosis, the mycorrhizal microbiome, comparisons among types of mycorrhizas from molecular to ecosystem scales, the extent and function of mycorrhizal networks and enhanced understanding of the role of mycorrhizas in carbon dynamics from local to global scales. The top-tier scientific journals have acknowledged mycorrhizas to be complex adaptive systems that play key roles in the development of communities and ecosystem processes. Understanding the mechanisms driving these large-scale effects requires integration of knowledge across scales of biological organization.

Accounting for local adaptation in ectomycorrhizas: a call to track geographical origin of plants, fungi, and soils in experiments.

Local adaptation, the differential success of genotypes in their native versus foreign environments, can influence ecological and evolutionary processes, yet its importance is difficult to estimate because it has not been widely studied, particularly in the context of interspecific interactions. Interactions between ectomycorrhizal (EM) fungi and their host plants could serve as model system for investigations of local adaptation because they are widespread and affect plant responses to both biotic and abiotic selection pressures. Furthermore, because EM fungi cycle nutrients and mediate energy flow into food webs, their local adaptation may be critical in sustaining ecological function. Despite their ecological importance and an extensive literature on their relationships with plants, the vast majority of experiments on EM symbioses fail to report critical information needed to assess local adaptation: the geographic origin of the plant, fungal inocula, and soil substrate used in the experiment. These omissions limit the utility of such studies and restrict our understanding of EM ecology and evolution. Here, we illustrate the potential importance of local adaptation in EM relationships and call for consistent reporting of the geographic origin of plant, soil, and fungi as an important step towards a better understanding of the ecology and evolution of EM symbioses.

Plant species differ in early seedling growth and tissue nutrient responses to arbuscular and ectomycorrhizal fungi.

Experiments with plant species that can host both arbuscular mycorrhizal fungi (AMF) and ectomycorrhizal fungi (EMF) are important to separating the roles of fungal type and plant species and understanding the influence of the types of symbioses on plant growth and nutrient acquisition. We examined the effects of mycorrhizal fungal type on the growth and tissue nutrient content of two tree species (Eucalyptus grandis and Quercus costaricensis) grown under four nutrient treatments (combinations of low versus high nitrogen (N) and phosphorus (P) with different N:P ratios) in the greenhouse. Trees were inoculated with unidentified field mixtures of AMF or EMF species cultivated on root fragments of AMF- or EMF-specific bait plants. In E. grandis, inoculation with both AMF and EMF positively affected belowground plant dry weight and negatively affected aboveground dry weight, while only inoculation with AMF increased tissue nutrient content. Conversely, Q. costaricensis dry weight and nutrient content did not differ significantly among inoculation treatments, potentially due to its dependence on cotyledon reserves for growth. Mineral nutrition of both tree species differed with the ratio of N to P applied while growth did not. Our results demonstrate that both tree species' characteristics and the soil nutrient environment can affect how AMF and EMF interact with their host plants. This research highlights the importance of mycorrhizal fungal-tree-soil interactions during early seedling growth and suggests that differences between AMF and EMF associations may be crucial to understanding forest ecosystem functioning.

Home-field advantage? evidence of local adaptation among plants, soil, and arbuscular mycorrhizal fungi through meta-analysis.

BACKGROUND: Local adaptation, the differential success of genotypes in their native versus foreign environment, arises from various evolutionary processes, but the importance of concurrent abiotic and biotic factors as drivers of local adaptation has only recently been investigated. Local adaptation to biotic interactions may be particularly important for plants, as they associate with microbial symbionts that can significantly affect their fitness and may enable rapid evolution. The arbuscular mycorrhizal (AM) symbiosis is ideal for investigations of local adaptation because it is globally widespread among most plant taxa and can significantly affect plant growth and fitness. Using meta-analysis on 1170 studies (from 139 papers), we investigated the potential for local adaptation to shape plant growth responses to arbuscular mycorrhizal inoculation. RESULTS: The magnitude and direction for mean effect size of mycorrhizal inoculation on host biomass depended on the geographic origin of the soil and symbiotic partners. Sympatric combinations of plants, AM fungi, and soil yielded large increases in host biomass compared to when all three components were allopatric. The origin of either the fungi or the plant relative to the soil was important for explaining the effect of AM inoculation on plant biomass. If plant and soil were sympatric but allopatric to the fungus, the positive effect of AM inoculation was much greater than when all three components were allopatric, suggesting potential local adaptation of the plant to the soil; however, if fungus and soil were sympatric (but allopatric to the plant) the effect of AM inoculation was indistinct from that of any allopatric combinations, indicating maladaptation of the fungus to the soil. CONCLUSIONS: This study underscores the potential to detect local adaptation for mycorrhizal relationships across a broad swath of the literature. Geographic origin of plants relative to the origin of AM fungal communities and soil is important for describing the effect of mycorrhizal inoculation on plant biomass, suggesting that local adaptation represents a powerful factor for the establishment of novel combinations of fungi, plants, and soils. These results highlight the need for subsequent investigations of local adaptation in the mycorrhizal symbiosis and emphasize the importance of routinely considering the origin of plant, soil, and fungal components.

Genetics-based interactions among plants, pathogens, and herbivores define arthropod community structure.

Plant resistance to pathogens or insect herbivores is common, but its potential for indirectly influencing plant-associated communities is poorly known. Here, we test whether pathogens' indirect effects on arthropod communities and herbivory depend on plant resistance to pathogens and/or herbivores, and address the overarching interacting foundation species hypothesis that genetics-based interactions among a few highly interactive species can structure a much larger community. In a manipulative field experiment using replicated genotypes of two Populus species and their interspecific hybrids, we found that genetic variation in plant resistance to both pathogens and insect herbivores modulated the strength of pathogens' indirect effects on arthropod communities and insect herbivory. First, due in part to the pathogens' differential impacts on leaf biomass among the two Populus species and the hybrids, the pathogen most strongly impacted arthropod community composition, richness, and abundance on the pathogen-susceptible tree species. Second, we found similar patterns comparing pathogen-susceptible and pathogen-resistant genotypes within species. Third, within a plant species, pathogens caused a fivefold greater reduction in herbivory on insect-herbivore-susceptible plant genotypes than on herbivore-resistant genotypes, demonstrating that the pathogen-herbivore interaction is genotype dependent. We conclude that interactions among plants, pathogens, and herbivores can structure multitrophic communities, supporting the interacting foundation species hypothesis. Because these interactions are genetically based, evolutionary changes in genetic resistance could result in ecological changes in associated communities, which may in turn feed back to affect plant fitness.

Plant genetics and interspecific competitive interactions determine ectomycorrhizal fungal community responses to climate change.

Although the importance of plant-associated microbes is increasingly recognized, little is known about the biotic and abiotic factors that determine the composition of that microbiome. We examined the influence of plant genetic variation, and two stressors, one biotic and one abiotic, on the ectomycorrhizal (EM) fungal community of a dominant tree species, Pinus edulis. During three periods across 16 years that varied in drought severity, we sampled the EM fungal communities of a wild stand of P. edulis in which genetically based resistance and susceptibility to insect herbivory was linked with drought tolerance and the abundance of competing shrubs. We found that the EM fungal communities of insect-susceptible trees remained relatively constant as climate dried, while those of insect-resistant trees shifted significantly, providing evidence of a genotype by environment interaction. Shrub removal altered the EM fungal communities of insect-resistant trees, but not insect-susceptible trees, also a genotype by environment interaction. The change in the EM fungal community of insect-resistant trees following shrub removal was associated with greater shoot growth, evidence of competitive release. However, shrub removal had a 7-fold greater positive effect on the shoot growth of insect-susceptible trees than insect-resistant trees when shrub density was taken into account. Insect-susceptible trees had higher growth than insect-resistant trees, consistent with the hypothesis that the EM fungi associated with susceptible trees were superior mutualists. These complex, genetic-based interactions among species (tree-shrub-herbivore-fungus) argue that the ultimate impacts of climate change are both ecological and evolutionary.

Tree genotype and genetically based growth traits structure twig endophyte communities.

PREMISE OF THE STUDY: Fungal endophytes asymptomatically inhabit plant tissues where they have mutualistic, parasitic, or commensal relationships with their hosts. Although plant-fungal interactions at the genotype scale have broad ecological and evolutionary implications, the sensitivity of endophytes in woody tissues to differences among plant genotypes is poorly understood. We hypothesize that (1) endophyte communities in Populus angustifolia (Salicaceae) twigs vary among tree genotypes, (2) endophyte variation is linked to quantitative tree traits, and (3) tree genotype influences interspecific fungal interactions. METHODS: Endophytes were isolated from twigs of replicated P. angustifolia genotypes in a common garden and characterized with PCR-RFLP and DNA sequencing. Twig length and diameter, aboveground tree biomass, and condensed tannins were also quantified. KEY RESULTS: (1) Aspects of fungal community structure, including composition and total isolation frequency (i.e., abundance), varied among genotypes. (2) Aboveground biomass and twig diameter were positively associated with isolation frequency and covaried with composition, whereas twig length and condensed tannin concentration were not significantly correlated to endophytes. (3) Fungal co-occurrence patterns suggested negative species interactions, but the presence of significant co-occurrences was genotype dependent. CONCLUSIONS: The species is often assumed to be the most important ecological unit; however, these results indicate that genetically based trait variation within a species can influence an important community of associated organisms. Given the dominance of plants as primary producers and the ubiquity of endophytes, the effect of host genetic variation on endophytes has fundamental implications for our understanding of terrestrial ecosystems.