Ectomycorrhizal fungal communities in ice-age relict forests of Pinus pumila on nine mountains correspond to summer temperature

Ectomycorrhizal (ECM) fungi are critical symbionts of major forest trees, and their communities are affected by various environmental factors including temperature. However, previous knowledge concerning temperature effects does not exclude the effects of host species and coexisting plants, which usually change with temperature, and should be rigorously tested under the same vegetation type. Herein we examined ECM fungal communities in ice-age relict forests dominated by a single host species (Pinus pumila) distributed on nine mountains across >1000 km in Japan. Direct sequencing of rDNA ITS regions identified 154 ECM fungal species from 4134 ECM root-tip samples. Gradient analyses revealed a large contribution of temperature, especially summer temperature, to ECM fungal communities. Additionally, we explored global sequence records of each fungal species to infer its potential temperature niche, and used it to estimate the temperature of the observed communities. The estimated temperature was significantly correlated with the actual temperature of the research sites, especially in summer seasons, indicating inherent temperature niches of the fungal components could determine their distribution among the sites. These results indicate that temperature is still a significant determinant in structuring ECM fungal communities after excluding the effects of host species and coexisting plants. The results also imply that the rising temperature under global warming may have been affecting soil microbes unnoticeably, while such microbial community change may have been contributing to the resilience of the same vegetation.