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Symbiotic fungi commonly increase plant acquisition of soil nutrients. Because prior work has focused on root fungi, we examined how leaf endophytes (Epichloe) influenced plant responses to fertilization and altered plant traits that may cascade to food webs and ecosystem processes. We manipulated endophyte presence/absence in two populations of Trisetum spicatum, a wild relative of oat, under a 2 × 2 addition of soil nitrogen (N) and phosphorus (P) in the greenhouse. Endophyte symbiosis altered how plant biomass responded to soil N and how plant traits responded to soil P. Endophytes boosted the biomass gains from N-fertilization in one population. Plants from a second population had weak benefits of symbiosis, but the endophyte altered plant traits, by increasing specific leaf area under P-fertilization, root diameter under low P, and concentration of the fungal alkaloid AcAP under N fertilization. Endophyte presence suppressed the typically observed increase in root hair density in response to soil P limitation. Under low P, symbiotic plants from both populations had improved forage quality relative to symbiont-free plants, although N-fertilization had a larger effect size on forage quality than did symbiosis. Finally, the two populations differed in production of fungal alkaloids, which generally increased in response to fertilization. Predicting how microbial symbionts mediate plant acquisition of nutrients requires understanding how much their effects vary among plant and endophyte genotypes. Here, the magnitude and direction of leaf symbionts’ effects on plant yield and traits varied between populations and with soil nutrient availability.

Epichloë species (Clavicipitaceae, Ascomycota) are endophytic symbionts of many cool-season grasses. Many interactions between Epichloë and their host grasses contribute to plant growth promotion, protection from many pathogens and insect pests, and tolerance to drought stress. Resistance to insect herbivores by endophytes associated with Hordeum species has been previously shown to vary depending on the endophyte-grass-insect combination. We explored the genetic and chemotypic diversity of endophytes present in wild Hordeum species. We analyzed seeds of Hordeum bogdanii, H. brevisubulatum, and H. comosum obtained from the US Department of Agriculture's (USDA) National Plant Germplasm System (NPGS), of which some have been reported as endophyte-infected. Using polymerase chain reaction (PCR) with primers specific to Epichloë species, we were able to identify endophytes in seeds from 17 of the 56 Plant Introduction (PI) lines, of which only 9 lines yielded viable seed. Phylogenetic analyses of housekeeping, alkaloid biosynthesis, and mating type genes suggest that the endophytes of the infected PI lines separate into five taxa: Epichloë bromicola, Epichloë tembladerae, and three unnamed interspecific hybrid species. One PI line contained an endophyte that is considered a new taxonomic group, Epichloë sp. HboTG-3 (H. bogdanii Taxonomic Group 3). Phylogenetic analyses of the interspecific hybrid endophytes from H. bogdanii and H. brevisubulatum indicate that these taxa all have an E. bromicola allele but the second allele varies. We verified in planta alkaloid production from the five genotypes yielding viable seed. Morphological characteristics of the isolates from the viable Hordeum species were analyzed for their features in culture and in planta. In the latter, we observed epiphyllous growth and in some cases sporulation on leaves of infected plants.

Aim Fungal symbionts are ubiquitous in plants and can mitigate abiotic stressors associated with climate change. Predicting fungal symbiont distributions under future climates first requires knowledge of current distributions and their potential drivers. Location We documented colonization by fungal symbionts in perennial, cool-season grasses along altitudinal gradients in the Rocky Mountains of Colorado, USA. Methods Across seven replicate altitudinal gradients, spanning c. 1400 vertical meters, we scored fungal colonization for 46 grass species. We documented altitudinal clines in colonization by both above-ground and below-ground fungal symbionts for the first time, including localized foliar endophytes (LFE) and systemic endophytes (epichloae) in leaves and arbuscular mycorrhizal fungi (AMF) and dark septate endophytes (DSE) in roots. For a subset of 16 well-sampled grass species, we used model selection procedures to evaluate the relative importance of geography, edaphic factors and host plant identity. We also assessed the influence of host phylogenetic relatedness and colonization by co-infecting fungi. Results Levels of fungal colonization varied strongly with host plant identity, but the effects of particular host species were not consistent across fungal groups. In addition to the influence of host identity, epichloae colonization declined with elevation and varied with geography (latitude/longitude) and edaphic factors. Geography and collection date were important predictors of LFE colonization, with higher colonization later in the growing season. Colonization estimates for the obligately plant-associated fungi (epichloae, AMF) were phylogenetically conserved across the grass supertree. Positive correlations between AMF and DSE, which remained even after accounting for host plant relatedness, suggested possible synergisms between these fungal groups. Main conclusions Our survey showed greater host specificity in patterns of fungal colonization than prior reports and revealed that different fungal symbiont groups do not share similar drivers. Conserving plant–fungal symbioses under future climates may require unique strategies for different plant species and fungal symbiont types.

Few studies have tested whether mutualisms may affect species distributions by altering the niches of partner species. We show that a fungal endophyte is associated with a shift in the soil moisture niche of its host plant relative to a co-occurring, endophyte-free congener. The endophyte appeared to initially restrict its host's distribution to wetter microsites before positively affecting its growth, suggesting the value of considering symbiont effects at different partner life stages. Our study identifies a symbiotic relationship as a potential mechanism facilitating the coexistence of two species, suggesting that symbiont effects on host niche may have community-level consequences., Mutualisms can play important roles in influencing species coexistence and determining community composition. However, few studies have tested whether such interactions can affect species distributions by altering the niches of partner species. In subalpine meadows of the Rocky Mountains, USA, we explored whether the presence of a fungal endophyte (genus Epichloë) may shift the niche of its partner plant, marsh bluegrass (Poa leptocoma) relative to a closely related but endophyte-free grass species, nodding bluegrass (Poa reflexa). Using observations and a 3-year field experiment, we tested two questions: (i) Do P. leptocoma and P. reflexa occupy different ecological niches? and (ii) Does endophyte presence affect the relative fitness of P. leptocoma versus P. reflexa in the putative niches of these grass species? The two species were less likely to co-occur than expected by chance. Specifically, P. leptocoma grew closer to water sources and in wetter soils than P. reflexa, and also had higher root colonization by mycorrhizal fungi. Endophyte-symbiotic P. leptocoma seeds germinated with greater frequency in P. leptocoma niches relative to P. reflexa niches, whereas neither endophyte-free (experimentally removed) P. leptocoma seeds nor P. reflexa seeds showed differential germination between the two niche types. Thus, endophyte presence constrained the germination and early survival of host plants to microsites occupied by P. leptocoma. However, endophyte-symbiotic P. leptocoma ultimately showed greater growth than endophyte-free plants across all microsites, indicating a net benefit of the symbiosis at this life history stage. Differential effects of endophyte symbiosis on different host life history stages may thus contribute to niche partitioning between the two congeneric plant species. Our study therefore identifies a symbiotic relationship as a potential mechanism facilitating the coexistence of two species, suggesting that symbiont effects on host niche may have community-level consequences.