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Species diversity, updated classification and divergence times of the phylum Mucoromycota

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Zygomycetes are phylogenetically early diverging, ecologically diverse, industrially valuable, agriculturally beneficial, and clinically pathogenic fungi. Although new phyla and subphyla have been constantly established to accommodate specific members and a subkingdom Mucoromyceta, comprising Calcarisporiellomycota, Glomeromycota, Mortierellomycota and Mucoromycota, was erected to unite core zygomycetous fungi, phylogenetic relationships within phyla have not been well resolved. Taking account of the information of monophyly and divergence time estimated from ITS and LSU rDNA sequences, the present study updates the classification framework of the phylum Mucoromycota from the class down to the generic rank: three classes, three orders, 20 families (including five new families Circinellaceae, Protomycocladaceae, Rhizomucoraceae, Syzygitaceae and Thermomucoraceae) and 64 genera. The taxonomic hierarchy was calibrated with estimated divergence times: phylum earlier than 617 Mya, classes and orders earlier than 547 Mya, families earlier than 199 Mya, and genera earlier than 12 Mya. Along with this outline, all genera of Mucoromycota are annotated and 58 new species are described. In addition, three new combinations are proposed. In this study, we update the taxonomic backbone of the phylum Mucoromycota and reinforce its phylogeny. We also contribute numerous new taxa and enrich the diversity of Mucoromycota.

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Data availability

All sequences have been deposited at GenBank database (Table 1 and S1).


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We thank the following scholars for their help with sampling and culture isolation: Yu-Chuan Bai and Peng-Cheng Deng (North Minzu University), Meng Zhou, Hong-Ming Zhou, Shun Liu, Xiao-Wu Man, Chang-Ge Song and Tai-Min Xu (Beijing Forestry University), Ru-Yong Zheng, He Huang, Hong-Mei Liu, Ya-Ning Wang, Jing Yang, Mao-Qiang He, Yue Zhang, Xing-Chun Li, Jie Li, Xiao-Ling Liu, Gui-Qing Chen, Feng-Yan Bai, Xue-Wei Wang (Prof.), Xue-Wei Wang (Ph.D. student), Xin-Yu Zhu and Long Wang (Institute of Microbiology, Chinese Academy of Chinese), Xiao Ju (Nanjing Agricultural University), Zhi-Kang Zhang (Ludong University), Ze Liu, Xiang Sun and Qi-Ming Wang (Hebei University), Zhong-Dong Yu (Northwest A&F University), Yan-Yan Long (Guangxi Academy of Agricultural Sciences), Nian-Kai Zeng (Hainan Medical University), Ze-Fen Yu and Min Qiao (Yunnan University), Pei-Wu Cui (Hunan University of Chinese Medicine), Jun-Feng Liang (Chinese Academy of Forestry), Yan-Feng Hang (Guizhou University), Wang-Qiu Deng (Guangdong Institute of Microbiology), Pu Liu, Li-Ying Ren (Jilin Agricultural University), Biao Xu (Tarim University), Rafael F. Castañeda-Ruiz (Academia de Ciencias de Cuba, Cuba), Kerstin Voigt (Friedrich Schiller University Jena, Germany), Manfred Binder (Clark University, USA), and Timothy Y. James (Michigan University, USA), Nicolas Corradi and Mathu Malar C (University of Ottawa, USA), and Francis Michel Martin (Head of Department at French National Institute for Agriculture, Food, and Environment, France).


This study was supported by the National Natural Science Foundation of China (Grant Nos. 31970009, 32170012 and 32000010), the Second Tibetan Plateau Scientific Expedition and Research Program (STEP, Grant No. 2019QZKK0503), and the Third Xinjiang Scientific Expedition and Research Program (STEP, Grant No. 2021XJKK0505).

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Authors and Affiliations



Heng Zhao is responsible for culture isolations, identifications and descriptions, data analyses, drafting and editing the manuscript; Yu-Cheng Dai and Xiao-Yong Liu for funding acquisition; Yuan Yuan, Yu-Cheng Dai and Xiao-Yong Liu conceiving this study, and improving the manuscript; Yong Nie, Tong-Kai Zong, Ke Wang, Mei-Lin Lv, Yu-Jin Cui, Ablat·Tohtirjap, Jia-Jia Chen, Chang-Lin Zhao, Fang Wu and Bao-Kai Cui contributed to the sample collection and data analyses.

Corresponding authors

Correspondence to Yuan Yuan, Yu-Cheng Dai or Xiao-Yong Liu.

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All authors declare no conflict of interest.

Additional information

Handling Editor: Jian-Kui Liu.

Supplementary Information

Below is the link to the electronic supplementary material.

Fig. S1

. Species information concerning publication years, type’ origins, and first record time in China. (XLSX 33 kb)

Fig. S1

. A Maximum Likelihood phylogenetic tree illustrating relationships within subkingdom Mucoromyceta based on ITS and LSU rDNA sequences, with Neurospora crassa as outgroup. New taxa proposed in the present study are in red. Maximum Likelihood bootstrap values (≥50%) / Bayesian Inference (BI) Posterior Probabilities (≥0.80) are indicated along branches. Taxa are highlighted in different colors. A scale bar in the upper left indicates substitutions per site. (PDF 92 kb)

Supplementary file3 (DOCX 34 kb)

Supplementary file4 (DOCX 25 kb)

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Zhao, H., Nie, Y., Zong, TK. et al. Species diversity, updated classification and divergence times of the phylum Mucoromycota. Fungal Diversity 123, 49–157 (2023).

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