title: Phylogenetic classification of yeasts and related taxa within Pucciniomycotina.
abstract: Most small genera containing yeast species in the Pucciniomycotina (Basidiomycota, Fungi) are monophyletic, whereas larger genera including Bensingtonia, Rhodosporidium, Rhodotorula, Sporidiobolus and Sporobolomyces are polyphyletic. With the implementation of the "One Fungus = One Name" nomenclatural principle these polyphyletic genera were revised. Nine genera, namely Bannoa, Cystobasidiopsis, Colacogloea, Kondoa, Erythrobasidium, Rhodotorula, Sporobolomyces, Sakaguchia and Sterigmatomyces, were emended to include anamorphic and teleomorphic species based on the results obtained by a multi-gene phylogenetic analysis, phylogenetic network analyses, branch length-based methods, as well as morphological, physiological and biochemical comparisons. A new class Spiculogloeomycetes is proposed to accommodate the order Spiculogloeales. The new families Buckleyzymaceae with Buckleyzyma gen. nov., Chrysozymaceae with Chrysozyma gen. nov., Microsporomycetaceae with Microsporomyces gen. nov., Ruineniaceae with Ruinenia gen. nov., Symmetrosporaceae with Symmetrospora gen. nov., Colacogloeaceae and Sakaguchiaceae are proposed. The new genera Bannozyma, Buckleyzyma, Fellozyma, Hamamotoa, Hasegawazyma, Jianyunia, Rhodosporidiobolus, Oberwinklerozyma, Phenoliferia, Pseudobensingtonia, Pseudohyphozyma, Sampaiozyma, Slooffia, Spencerozyma, Trigonosporomyces, Udeniozyma, Vonarxula, Yamadamyces and Yunzhangia are proposed to accommodate species segregated from the genera Bensingtonia, Rhodosporidium, Rhodotorula, Sporidiobolus and Sporobolomyces. Ballistosporomyces is emended and reintroduced to include three Sporobolomyces species of the sasicola clade. A total of 111 new combinations are proposed in this study.
authors.affiliation: State Key Laboratory of Mycology, Institute of Microbiology, Chinese Academy of Sciences, Beijing 100101, China.
authors.forename: Q-M
authors.initials: QM
authors.lastname: Wang
authors.affiliation: Leibniz Institute DSMZ - German Collection of Microorganisms and Cell Cultures, Braunschweig, Germany.
authors.forename: A M
authors.initials: AM
authors.lastname: Yurkov
authors.affiliation: Leibniz Institute DSMZ - German Collection of Microorganisms and Cell Cultures, Braunschweig, Germany.
authors.forename: M
authors.initials: M
authors.lastname: Göker
authors.affiliation: Science & Education, The Field Museum, 1400 S. Lake Shore Drive, Chicago, IL 60605, USA.
authors.forename: H T
authors.initials: HT
authors.lastname: Lumbsch
authors.affiliation: Science & Education, The Field Museum, 1400 S. Lake Shore Drive, Chicago, IL 60605, USA.
authors.forename: S D
authors.initials: SD
authors.lastname: Leavitt
authors.affiliation: CBS Fungal Biodiversity Centre (CBS-KNAW), Uppsalalaan 8, 3584 CT Utrecht, The Netherlands.
authors.forename: M
authors.initials: M
authors.lastname: Groenewald
authors.affiliation: CBS Fungal Biodiversity Centre (CBS-KNAW), Uppsalalaan 8, 3584 CT Utrecht, The Netherlands.
authors.forename: B
authors.initials: B
authors.lastname: Theelen
authors.affiliation: State Key Laboratory of Mycology, Institute of Microbiology, Chinese Academy of Sciences, Beijing 100101, China.
authors.forename: X-Z
authors.initials: XZ
authors.lastname: Liu
authors.affiliation: State Key Laboratory of Mycology, Institute of Microbiology, Chinese Academy of Sciences, Beijing 100101, China; CBS Fungal Biodiversity Centre (CBS-KNAW), Uppsalalaan 8, 3584 CT Utrecht, The Netherlands; Shanghai Key Laboratory of Molecular Medical Mycology, Changzheng Hospital, Second Military Medical University, Shanghai, China.
authors.forename: T
authors.initials: T
authors.lastname: Boekhout
authors.affiliation: State Key Laboratory of Mycology, Institute of Microbiology, Chinese Academy of Sciences, Beijing 100101, China; CBS Fungal Biodiversity Centre (CBS-KNAW), Uppsalalaan 8, 3584 CT Utrecht, The Netherlands.
authors.forename: F-Y
authors.initials: FY
authors.lastname: Bai
keywords: B. aurantiaca (Saito) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; B. kluyveri-nielii (van der Walt) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; B. ogasawarensis (Hamam., Thanh & Nakase) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; B. phyllomatis (van der Walt & Y. Yamada) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; B. salicina (B.N. Johri & Bandoni) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; B. syzygii (Hamam., Thanh & Nakase) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; B. taupoensis (Hamam. & Nakase) F.Y. Bai, Q.M. Wang, M. Groenew. & Boekhout; B. yamatoana (Nakase, M. Suzuki & M. Itoh) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Ballistosporomycessasicola (Nakase & M. Suzuki) F.Y. Bai, Q.M. Wang, M. Groenew. & Boekhout; Bannoabischofiae (Hamam., Thanh & Nakase) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Bannozyma Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Bannozymaarctica (Vishniac & M. Takash.) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Buckleyzyma Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Buckleyzymaarmeniaca (R.G. Shivas & Rodr. Mir.) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Buckleyzymaceae Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; C. diffluens (Ruinen) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; C. eucalyptica (C.H. Pohl, M.S. Smit & Albertyn) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; C. falcata (Nakase, M. Itoh & M. Suzuki) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; C. foliorum (Ruinen) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; C. griseoflava (Nakase & M. Suzuki) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; C. lophatheri (Nakase, Tsuzuki, F.L. Lee, Jindam. & M. Takash.) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; C. philyla (van der Walt, Klift & D.B. Scott) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; C. retinophila (Thanh, M.S. Smit, Moleleki & Fell) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; C. terpenoidalis (Thanh, M.S. Smit, Moleleki & Fell) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Chrysozyma Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Chrysozymaceae Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Chrysozymafushanensis (Nakase, F.L. Lee & M. Takash.) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Colacogloeaceae Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Colacogloeacycloclastica (Thanh, M.S. Smit, Moleleki & Fell) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Cystobasidiopsislactophilus (Nakase, M. Itoh, M. Suzuki & Bandoni) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Cystobasidiumportillonense (F. Laich, I. Vaca & R. Chávez) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; E. yunnanense (F.Y. Bai, M. Takash., Hamam. & Nakase) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Erythrobasidiumelongatum (R.G. Shivas & Rodr. Mir.) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Fellozyma Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Fellozymainositophila (Nakase & M. Suzuki) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Fungi; GMYC approach; H. singularis (Phaff & do Carmo-Sousa) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Hamamotoa Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Hamamotoalignophila (Dill, C. Ramírez & González) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Hasegawazyma Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Hasegawazymalactosa (Hasegawa) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Jianyunia Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Jianyuniasakaguchii (Sugita, M. Takash., Hamam. & Nakase) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; K. miscanthi (Nakase & M. Suzuki) Q.M. Wang, M. Groenew., F.Y. Bai & Boekhout; K. phyllada (van der Walt & Y. Yamada) Q.M. Wang, M. Groenew., F.Y. Bai & Boekhout; K. sorbi (F.Y. Bai & Q.M. Wang) Q.M. Wang, M. Groenew., F.Y. Bai & Boekhout; K. subrosea (Nakase & M. Suzuki) Q.M. Wang, M. Groenew., F.Y. Bai & Boekhout; K. thailandica (Fungsin, Hamam. & Nakase ) Q.M. Wang, M. Groenew., F.Y. Bai & Boekhout; K. yuccicola (Nakase & M. Suzuki) Q.M. Wang, M. Groenew., F.Y. Bai & Boekhout; Kondoachangbaiensis (F.Y. Bai & Q.M. Wang) Q.M. Wang, M. Groenew., F.Y. Bai & Boekhout; M. magnisporus (Nakase, Tsuzuki, F.L. Lee, Sugita, Jindam. & M. Takash.) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; M. orientis (Pohl, M.S. Smit & Albertyn) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; M. pini (Pohl, M.S. Smit & Albertyn) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Microsporomyces Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Microsporomycesbloemfonteinensis (Pohl, M.S. Smit & Albertyn) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Microsporomycetaceae Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Molecular phylogeny; O. straminea (Golubev & Scorzetti) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; O. yarrowii (Á. Fonseca & van Uden) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Oberwinklerozyma Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Oberwinklerozymasilvestris (Golubev & Scorzetti) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; P. buffonii (C. Ramírez) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; P. corallina (N. Furuya & M. Takash.) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; P. dimennae (Hamam. & Nakase) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; P. glacialis (Margesin & J.P. Samp.) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; P. himalayensis (Shivaji, Bhadra & Rao) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; P. linderae (Nakase, M. Takash. & Hamam.) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; P. musae (M. Takash., S.O. Suh & Nakase) F.Y. Bai, Q.M. Wang, M. Groenew. & Boekhout; P. novozealandica (Hamam. & Nakase) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; P. producta (N. Furuya & M. Takash.) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; P. psychrophila (Margesin & J.P. Samp.) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; P. pustula (Buhagiar) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; P. subbrunnea (Nakase & M. Suzuki) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Phenoliferia Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Phenoliferiapsychrophenolica (Margesin & J.P. Samp.) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Phyllozyma Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Phyllozymacoprosmicola (Hamam. & Nakase) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Phylogenetic rank boundary optimisation; Pseudobensingtonia F.Y. Bai, Q.M. Wang, M. Groenew. & Boekhout; Pseudobensingtoniaingoldii (Nakase & Itoh.) F.Y. Bai, Q.M. Wang, M. Groenew. & Boekhout; Pseudohyphozyma Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Pseudohyphozymabogoriensis (Deinema) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Pucciniomycotina; R. azoricus (J.P. Samp. & Gadanho) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; R. babjevae (Golubev) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; R. colostri (T. Castelli) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; R. diobovata (S.Y. Newell & I.L. Hunter) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; R. diospyroris (Nakase, Tsuzuki, F.L. Lee, Jindam. & M. Takash.) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; R. dracophylli (Hamam. & Nakase) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; R. kratochvilovae (Hamam., Sugiy. & Komag.) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; R. lusitaniae (Á. Fonseca & J.P. Samp.) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; R. microsporus (Higham ex Fell, Blatt & Statzell) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; R. nylandii (M. Takash. & Nakase) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; R. odoratus (J.P. Samp., Á. Fonseca & Valério) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; R. paludigena (Fell & Tallman) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; R. poonsookiae (M. Takash. & Nakase) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; R. pyrrosiae (Nakase, Tsuzuki, F.L. Lee, Jindam. & M. Takash.) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; R. rubra (Nakase, Oakada & Sugiy.) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; R. ruineniae (Holzschu, Tredick & Phaff) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; R. sphaerocarpa (S.Y. Newell & Fell) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; R. toruloides (I. Banno) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Rhodosporidiobolus Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Rhodosporidiobolus fluvialis (Fell, Kurtzman, Tallman & J.D. Buck) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Rhodotorulaalborubescens (Derx) Q.M. Wang, F.Y. Bai, Groenew. & Boekhout; Ruinenia Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Ruineniaceae Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Ruineniaclavata (F.Y. Bai & Q.M. Wang) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; S. oryzae (F.Y. Bai & Y.M. Cai) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; S. foliicola (R.G. Shivas & Rodr. Mir.) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; S. gracilis (Derx) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; S. johnsonii (Nyland) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; S. lamellibrachiae (Nagah., Hamam., Nakase & Horikoshi) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; S. marina (Phaff, Mrak & Williams) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; S. meli (Libkind, van Broock & J.P. Samp.) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; S. novozelandica (W.B. Kendr. & X.D. Gong) F.Y. Bai, Q.M. Wang, Groenewald & Boekhout; S. pilati (F.H. Jacob, Faure-Raynaud & Berton) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; S. pulcherrima (J.E. Wright) F.Y. Bai, Q.M. Wang, Groenewald & Boekhout; S. symmetrica (F.Y. Bai & Q.M. Wang) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; S. tsugae (Phaff & do Carmo-Sousa) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; S. vanillica (J.P. Samp.) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; S. vermiculata (M. Takash. & Nakase) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Sakaguchiaceae Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Sakaguchiacladiensis (Fell, Statzell & Scorzetti) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Sampaiozyma Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Sampaiozymaingeniosa (Di Menna) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Slooffia Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Slooffia cresolica (Middelhoven & Spaaij) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Spencerozyma Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Spencerozymacrocea (Shifrine & Phaff) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Spiculogloeomycetes Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Sporobolomyceslongiusculus (Libkind, van Broock & J.P. Samp.) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Sterigmatomyceshyphaenes (Har. & Pat.) F.Y. Bai, Q.M. Wang, Groenewald & Boekhout; Symmetrospora Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Symmetrosporaceae Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Symmetrosporacoprosmae (Hamam. & Nakase) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Taxonomy; Trigonosporomyces Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Trigonosporomyceshylophilus (van der Walt, van der Klift & D.B. Scott) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Udeniozyma Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Udeniozymaferulica (J.P. Samp. & van Uden) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Ustilentylomagraminis (Rodr. Mir. & Diem) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Vonarxula Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Vonarxulajavanica (Ruinen) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Y. sonckii (Hopsu-Havu, Tunnela & Yarrow) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Yamadamyces Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Yamadamycesrosulatus (Golubev & Scorzetti) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Yeasts; Yunzhangia Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout; Yunzhangiaauriculariae (Nakase) Q.M. Wang, F.Y. Bai, M. Groenew. & Boekhout