Abstract
BackgroundEctomycorrhizae (ECM) are symbioses formed by polyphyletic assemblages of fungi (mostly Agaricomycetes) and plants (mostly Pinaceae and angiosperms in the rosid clade). Efforts to reconstruct the evolution of the ECM habit in Agaricomycetes have yielded vastly different results, ranging from scenarios with many relatively recent origins of the symbiosis and no reversals to the free-living condition; a single ancient origin of ECM and many subsequent transitions to the free-living condition; or multiple gains and losses of the association. To test the plausibility of these scenarios, we performed Bayesian relaxed molecular clock analyses including fungi, plants, and other eukaryotes, based on the principle that a symbiosis cannot evolve prior to the origin of both partners. As we were primarily interested in the relative ages of the plants and fungi, we did not attempt to calibrate the molecular clock using the very limited fossil record of Agaricomycetes.ResultsTopologically constrained and unconstrained analyses suggest that the root node of the Agaricomycetes is much older than either the rosids or Pinaceae. The Agaricomycetidae, a large clade containing the Agaricales and Boletales (collectively representing 70% of Agaricomycetes), is also significantly older than the rosids. The relative age of Agaricomycetidae and Pinaceae, however, is sensitive to tree topology, and the inclusion or exclusion of the gnetophyte Welwitschia mirabilis.ConclusionThe ancestor of the Agaricomycetes could not have been an ECM species because it existed long before any of its potential hosts. Within more derived clades of Agaricomycetes, there have been at least eight independent origins of ECM associations involving angiosperms, and at least six to eight origins of associations with gymnosperms. The first ECM symbioses may have involved Pinaceae, which are older than rosids, but several major clades of Agaricomycetes, such as the Boletales and Russulales, are young enough to have been plesiomorphically associated with either rosids or Pinaceae, suggesting that some contemporary ECM partnerships could be of very ancient origin.
Highlights
Ectomycorrhizae (ECM) are symbioses formed by polyphyletic assemblages of fungi and plants
An unconstrained relaxed molecular clock analysis returned a topology that is largely consistent with prior multilocus phylogenies of fungi and other eukaryotes, with several exceptions: 1) the Polyporales is placed as the sister group of the Russulales; 2) the opisthokonts are paraphyletic, with Dictyostelium as the sister group of the rhodophyte-viridiplantae clade; 3)Amborella and Nymphaea form a clade that is the sister group to the remaining angiosperms; and, 4) gnetophytes are placed as the sister group of the rest of the seed plants (Figure 2)
To assess the impact of the topology on relative age estimates, we performed a pair of constrained analyses, which reflect a consensus phylogeny based on multilocus analyses of fungi [4,10,26], plants [23,24,25,27,28,29], and other eukaryotes [30,31]
Summary
Ectomycorrhizae (ECM) are symbioses formed by polyphyletic assemblages of fungi (mostly Agaricomycetes) and plants (mostly Pinaceae and angiosperms in the rosid clade). This study focuses on the evolution of the ECM habit in the Agaricomycotina (Basidiomycota), which contains the vast majority of ECM-forming fungi. Most studies on this subject have used phylogenetic approaches coupled with ancestral state reconstruction (ASR) [1,2,3,4,5]. We use molecular clock analyses to estimate the relative ages of major clades of Agaricomycotina and their potential plant hosts. We focus on key nodes in the fungal phylogeny that subtend clades containing ECM and saprotrophic taxa, and ask whether it is plausible that the ancestors at these nodes could have been partners in ancient ECM associations
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