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Ectomycorrhizal lifestyle in fungi: global diversity, distribution, and evolution of phylogenetic lineages

机译：真菌的外生菌根生活方式：系统发生谱系的全球多样性，分布和进化

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摘要

The ectomycorrhizal (EcM) symbiosis involves a large number of plant and fungal taxa worldwide. During studies on EcM diversity, numerous misidentifications, and contradictory reports on EcM status have been published. This review aims to: (1) critically assess the current knowledge of the fungi involved in the EcM by integrating data from axenic synthesis trials, anatomical, molecular, and isotope studies; (2) group these taxa into monophyletic lineages based on molecular sequence data and published phylogenies; (3) investigate the trophic status of sister taxa to EcM lineages; (4) highlight other potentially EcM taxa that lack both information on EcM status and DNA sequence data; (5) recover the main distribution patterns of the EcM fungal lineages in the world. Based on critically examining original reports, EcM lifestyle is proven in 162 fungal genera that are supplemented by two genera based on isotopic evidence and 52 genera based on phylogenetic data. Additionally, 33 genera are highlighted as potentially EcM based on habitat, although their EcM records and DNA sequence data are lacking. Molecular phylogenetic and identification studies suggest that EcM symbiosis has arisen independently and persisted at least 66 times in fungi, in the Basidiomycota, Ascomycota, and Zygomycota. The orders Pezizales, Agaricales, Helotiales, Boletales, and Cantharellales include the largest number of EcM fungal lineages. Regular updates of the EcM lineages and genera therein can be found at the UNITE homepage http://unite.ut.ee/EcM_lineages. The vast majority of EcM fungi evolved from humus and wood saprotrophic ancestors without any obvious reversals. Herbarium records from 11 major biogeographic regions revealed three main patterns in distribution of EcM lineages: (1) Austral; (2) Panglobal; (3) Holarctic (with or without some reports from the Austral or tropical realms). The holarctic regions host the largest number of EcM lineages; none are restricted to a tropical distribution with Dipterocarpaceae and Caesalpiniaceae hosts. We caution that EcM-dominated habitats and hosts in South America, Southeast Asia, Africa, and Australia remain undersampled relative to the north temperate regions. In conclusion, EcM fungi are phylogenetically highly diverse, and molecular surveys particularly in tropical and south temperate habitats are likely to supplement to the present figures. Due to great risk of contamination, future reports on EcM status of previously unstudied taxa should integrate molecular identification tools with axenic synthesis experiments, detailed morphological descriptions, and/or stable isotope investigations. We believe that the introduced lineage concept facilitates design of biogeographical studies and improves our understanding about phylogenetic structure of EcM fungal communities.
机译：外生菌根（EcM）共生涉及全世界大量的植物和真菌类群。在对EcM多样性进行研究期间，已经发表了许多关于EcM状况的误解和相互矛盾的报道。这项审查的目的是：（1）通过整合来自轴生性合成试验，解剖学，分子和同位素研究的数据，严格评估与EcM有关的真菌的当前知识； （2）根据分子序列数据和已发表的系统发育史，将这些分类群分为单系谱系； （3）调查EcM血统的姊妹类群的营养状况； （4）突出显示缺少EcM状态信息和DNA序列数据的其他潜在EcM分类群； （5）恢复世界上EcM真菌谱系的主要分布模式。在严格审查原始报告的基础上，EcM的生活方式在162个真菌属中得到了证明，其中两个基于同位素证据的属和52个基于系统发育数据的属被补充。此外，尽管缺少其EcM记录和DNA序列数据，但根据栖息地将33个属突出显示为潜在EcM。分子系统发育和鉴定研究表明，EcM共生已独立发生，并且在真菌中，担子菌属，子囊菌和合子菌中至少持续66次。 Pezizales，Agaricales，Hellotiales，Boletales和Cantharellales的订单包括数量最多的EcM真菌谱系。可在UNITE主页http://unite.ut.ee/EcM_lineages上找到EcM谱系及其属的定期更新。 EcM真菌的绝大多数是由腐殖质和腐生木的祖先进化而来，没有任何明显的逆转。来自11个主要生物地理区域的植物标本室记录揭示了EcM谱系分布的三种主要模式：（1）南方； （2）Panglobal； （3）狂犬病（有或没有来自南方或热带领域的报道）。恐怖的地区拥有最多的EcM血统。没有一种植物局限于龙脑香科和Ca木科的热带分布。我们告诫，相对于北部温带地区，南美，东南亚，非洲和澳大利亚的以EcM为主的栖息地和寄主仍然采样不足。总之，EcM真菌在系统发育上是高度多样的，尤其是在热带和南部温带生境中的分子调查可能会补充现有数据。由于存在很大的污染风险，有关先前未研究的分类单元的EcM状态的未来报告应将分子鉴定工具与环氧树酯合成实验，详细的形态学描述和/或稳定的同位素研究相结合。我们相信，引入的谱系概念有助于进行生物地理学研究，并增进我们对EcM真菌群落系统发育结构的了解。

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