Lineages of ectomycorrhizal fungi revisited: Foraging strategies and novel lineages revealed by sequences from belowground

In the fungal kingdom, the ectomycorrhizal (EcM) symbiosis has evolved independently in multiple groups that are referred to as lineages. A growing number of molecular studies in the fields of mycology, ecology, soil science, and microbiology generate vast amounts of sequence data from fungi in their natural habitats, particularly from soil and roots. However, as the number and diversity of sequences has increased, it has become increasingly difficult to accurately identify the fungal species in these samples and to determine their trophic modes. In particular, there has been significant controversy regarding which fungal groups form ectomycorrhizas, the morphological “exploration types” that these fungi form on roots, and the ecological strategies that they use to obtain nutrients. To address this problem, we have synthesized the phylogenetic and taxonomic breadth of EcM fungi by using the wealth of accumulated sequence data. We also compile available information about exploration types of 143 genera of EcM fungi (including 67 new reports) that can be tentatively used to help infer the ecological strategies of different fungal groups. Phylogenetic analyses of ribosomal DNA ITS and LSU sequences enabled us to recognize 20 novel lineages of EcM fungi. Most of these are rare and have a limited distribution. Five new lineages occur exclusively in tropical and subtropical habitats. Altogether 46 fungal genera were added to the list of EcM fungal taxa and we anticipate that this number will continue to grow rapidly as taxonomic works segregate species-rich genera into smaller, monophyletic units. Three genera were removed from the list of EcM groups due to refined taxonomic and phylogenetic information. In all, we suggest that EcM symbiosis has arisen independently in 78-82 fungal lineages that comprise 251-256 genera. The EcM fungal diversity of tropical and southern temperate ecosystems remains significantly understudied and we expect that these regions are most likely to reveal additional EcM taxa.