The classification of dimorphic basidiomycetes

J.P. Sampaio and R. Bauer



    Class Urediniomycetes

    Class Ustilaginomycetes

    Class Hymenomycetes (Tremellomycetidae)




The current tripartite classification concept of the Basidiomycota is mainly based on sequence analyses of 18S rRNA (Taylor, 1995; Swann and Taylor, 1993; 1995a; 1995b; 1995c). As a consequence of those studies, the classes Urediniomycetes, Ustilaginomycetes and Hymenomycetes have been proposed (Swann and Taylor 1995a). Analysis of the D1/D2 region of the 26S rDNA shows basically the same pattern (Begerow et al., 1997). The data derived from the analysis of cell wall carbohydrates correlate well with the molecular phylogenies (Prillinger et al., 1990; 1991a; 1991b; 1993) and the same holds for the structure of the septal pores and the shape of the spindle pole bodies (McLaughlin et al., 1995a; 1995b and references therein).

  Class Urediniomycetes

This class, as proposed by Swann and Taylor (1995a, 1995b) is characterized by simple septal pores without membrane caps and disc-like spindle pole bodies (Bauer and Oberwinkler 1994, Wells 1994, McLaughlin et al., 1995b). Except for a few species, the basidia are transversally septate. Mannose is the major cell wall carbohydrate, glucose, fucose and rhamnose are the less prevalent neutral sugars and xylose is not present (Prillinger et al., 1991b; 1993).

Within the Urediniomycetes, Swann et al. (1999) defined the subclass Microbotryomycetidae Swann for a monophyletic group (based on 18S rRNA gene sequences) encompassing teleomorphic genera such as Microbotryum Léveillé emend. Vánky, Rhodosporidium Banno, Leucosporidium Fell, Statzell, Hunter & Phaff, Colacogloea Oberwinkler & Bandoni, Kriegeria Bres. and Heterogastridium Oberwinkler & Bauer. This subclass comprises predominantly real or potential mycoparasites having colacosomes (Bauer and Oberwinkler,1991a) and the urediniomycetous smut fungi (Bauer et al.,1997). Fell et al. (2000; 2001) and Scorzetti et al. (2002) have analysed the D1/D2 region of the 26S rDNA of a vast number of (typical) basidiomycetous yeast species and their phylogenetic trees depict a considerable number of anamorphic species of the genera Rhodotorula Harrison and Sporobolomyces Kluyver & van Niel as belonging to the same monophylum. Recently, Sampaio et al. (2003) proposed the order Leucosporidiales Sampaio, Weiss & Bauer for Leucosporidium scottii and closely related sexual and asexual species (Sampaio et al. 2003). In the same report the order Sporidiobolales Sampaio, Weiss & Bauer was erected for Rhodosporidium, Sporidiobolus Nyland and allied asexual taxa. 

In 2001, Swann et al. treated most of the remaining members of the class Urediniomycetes in two additional subclasses, Agaricostilbomycetidae and Urediniomycetidae. Whereas the known members of the Agaricostilbomycetidae are dimorphic, most of the taxa belonging to the Urediniomycetidae are filamentous. Examples of teleomorphic taxa of the Agaricostilbomycetidae are Agaricostilbum Wright, a fungus associated with palms, Chionosphaera Cox, which produces holobasidia, and Kondoa Yamada, Nakagawa & Banno emend. Fonseca, Sampaio & Fell, a genus investigated by Fonseca et al. (2000) who found that it lacks teliospores but forms transversally septate basidia which are able to produce forcibly discharged basidiospores. Based on sequence analyses, the mitosporic genera Kurtzmanomyces Yamada, Itoh, Kawasaki, Banno & Nakase emend. Sampaio, Sterigmatomyces Fell emend. Yamada & Banno and most of the species of Bensingtonia Ingold emend. Nakase & Boekhout belong to the Agaricostilbomycetidae (Fell et al., 2000; 2001; Fonseca et al., 2000; Hamamoto and Nakase, 2000; Scorzetti et al., 2002).

The subclass Urediniomycetidae is mainly composed of plant parasites. The most relevant group of this subclass, in terms of numbers of species and impact in human activities, are the rusts (order Uredinales). Interestingly, taxa with yeast stages are almost absent in the Urediniomycetidae since budding cells are currently known only in species of the genus Septobasidium Pat.

The classification of the Urediniomycetes is far from being settled due, among other factors, to the lack of detailed information for many species. Besides the three subclasses mentioned above, other groups have been detected based on ultrastructural and molecular studies.

One of the unnamed clades includes the sexual genera Naohidea Oberwinkler, Occultifur Oberwinkler, Sakaguchia Yamada, Maeda & Mikata and several asexual species of the genera Rhodotorula and Sporobolomyces. The genera Erythrobasidium Hamamoto, Sugiyama & Komagata and Bannoa Hamamoto belong also to this clade. Erythrobasidium and Bannoa were originally described for sexual species (Hamamoto et al., 1988; 2002) but the postulated basidia of Erytrhobasidium were later interpreted as conidiogenic structures (Sampaio et al., 1999). Bannoa is morphologically and phylogenetically close to Erythrobasidium and also lacks true basidia (Sampaio unpubl.). Takashima et al. (2000) determined the carbohydrate composition for representative taxa of this clade and observed in all cases the absence of fucose, thus providing an additional criterion for the separation of this group in a taxon of its own.

Another distinct clade corresponds to the order Atractiellales sensu Oberwinkler and Bauer (1989). The taxa in this order do not have a yeast stage and belong to the genera Atractiella Sacc., Helicogloea Pat., Phleogena Link and Saccoblastia Möller. Molecular studies are scarce for this group but relevant ultrastructural markers like the type of septal pore and the presence of symplechosomes (Oberwinkler and Bauer, 1989, Bauer and Oberwinkler, 1991b) - designated as microscala by McLaughlin (1990) - indicate that those four genera share a close common ancestor.

Recently, Bauer et al. (2003) proposed the order Classiculales Bauer, Begerow, Oberwinkler & Marvanová for urediniomycetous taxa having the uredinalian type of septal pores in combination with tremelloid haustorial cells.

Below, we present an annotated outline of the class Urediniomycetes which attempts to integrate data from several authors, namely Bandoni, Bauer, Fell, Oberwinkler, Swann and Vánky.



Taxa having colacosomes and taxa related to them


Colacosomes of Colacogloea peniophorae. Colacosomes are the dark electron-dense structures in P (hypha of the parasite); H, hypha of Hyphoderma praetermissum (host). Bar = 1 µm; © R. Bauer.

Order Cryptomycocolacales Oberwinkler & Bauer

Family Cryptomycocolacaceae Oberwinkler & Bauer

        Cryptomycocolax Oberwinkler & Bauer

        Colacosiphon Kirschner, Bauer & Oberwinkler

Order Heterogastridiales Oberwinkler & Bauer

Family Heterogastridiaceae Oberwinkler & Bauer

        Heterogastridium Oberwinkler & Bauer

Order Leucosporidiales Sampaio, Weiss & Bauer

        Leucosporidium Fell, Statzell, Hunter & Phaff

        Leucosporidiella Sampaio


        Mastigobasidium Golubev

Order Sporidiobolales Sampaio, Weiss & Bauer

Family Sporidiobolaceae Moore emend. Sampaio, Weiss & Bauer

        Sporidiobolus Nyland

        Sporobolomyces Kluyver & van Niel

        Mitosporic, not monophyletic

        Rhodosporidium Banno

        Rhodotorula Harrison

        Mitosporic, not monophyletic

Teliospores, basidia and basidiospores of Sporidiobolus salmonicolor. Bar = 5 µm; © J.P. Sampaio.

Order Microbotryales Bauer & Oberwinkler

Phytoparasitic members of the Basidiomycota having transversely septate basidia with multiple production of sessile basidiospores and only intercellular hyphae (Bauer et al., 1997).

Family Microbotryaceae MooreMembers of the Microbotryales having poreless septa at maturity (Bauer et al., 1997).

        Bauerago Vánky

        Liroa Ciferri

        Microbotryum Léveillé emend. Vánky

        Approximately 75 species parasitizing dicotyledonous plants

        Sphacelotheca de Bary emend. Langdon et Fullerton

        Zundeliomyces Vánky


Family Ustilentylomataceae Bauer & Oberwinkler

Members of the Microbotryales having simple septal pores (Bauer et al., 1997).

        Aurantiosporium Piepenbring, Vánky & Oberwinkler

        Fulvisporium Vánky

        Ustilentyloma Savile


Septal pore of Ustilentyloma fluitans. Bar = 0.1 µm; © R. Bauer.

Taxa of the Microbotryomycetidae not ascribed to any order

Family Krieglsteineraceae Pouzar

        Krieglsteinera Pouzar


Family Camptobasidiaceae Moore

        Camptobasidium Marvanová & Suberkropp


Atractocolax Kirschner, Bauer & Oberwinkler


Bensingtonia Ingold emend. Nakase & Boekhout

        Mitosporic, not monophyletic


Colacogloea Oberwinkler & Bandoni


Kriegeria Bres.



Order Agaricostilbales Oberwinkler & Bauer

Family Agaricostilbaceae Oberwinkler & Bauer

        Agaricostilbum Wright emend. Wright, Bandoni & Oberwinkler


Family Chionosphaeraceae Oberwinkler & Bandoni

        Chionosphaera Cox

        Stilbum Tode ex Mérat

        Fibulostilbum Seifert & Oberwinkler

Genera of the Agaricostilbomycetidae not ascribed to any order

Bensingtonia Ingold emend. Nakase & Boekhout

Mitosporic, not monophyletic


Kondoa Yamada, Nakagawa & Banno emend. Fonseca, Sampaio & Fell


Kurtzmanomyces Yamada, Itoh, Kawasaki, Banno & Nakase emend. Sampaio



Mycogloea Olive


Sporobolomyces Kluyver & van Niel

Mitosporic, not monophyletic


Sterigmatomyces Fell emend. Yamada & Banno



Spiculogloea Roberts


Zygogloea Roberts



Order Uredinales Arthur

14 families, 164 genera, aprox. 7000 species

Order Septobasidiales Couch ex Donk

Family Septobasidiaceae Maire

        Auriculoscypha Reid and Manimohan

        Coccidiodictyon Oberwinkler

        Ordonia Racib.

        Septobasidium Pat.

        Uredinella Couch


Family Pachnocybaceae Oberwinkler & Bauer

        Pachnocybe Berk.

Order Platygloeales Moore

        Platygloea Schroeter sensu stricto (P. disciformis and related species)


Taxa of the Urediniomycetidae not ascribed to any order

Family Eocronartiaceae Jülich

        Eocronartium Atkinson


Helicobasidium Pat.


Herpobasidium Lind emend. Oberw & Bandoni


Insolibasidium Oberwinkler & Bandoni


Jola Möller


Platycarpa Couch emend. Oberwinkler & Bandoni


Ptechetelium Oberwinkler & Bandoni


Groups of Urediniomycetes for which higher rank classification is incomplete

Order Atractiellales Oberwinkler & Bandoni

Family Hoehnelomycetaceae Jülich

        Atractiella Sacc.


Family Phleogenaceae Weese

        Phleogena Link


Family Helicogloeaceae Jülich

        Helicogloea Pat.

        Saccoblastia Möller


Symplechosomes of Saccoblastia farinacea. Bar = 0.25 µm; © R. Bauer.


Order Classiculales Bauer, Begerow, Oberwinkler & Marvanová

Family Classiculaceae Bauer, Begerow, Oberwinkler & Marvanová

        Classicula Bauer, Begerow, Oberwinkler & Marvanová

        Jaculispora H. J. Huds. & Ingold



Order Cystobasidiales (Bauer & Sampaio, in prep.)


Septal pore and cystosome of Occultifur internus. Bar = 0.2 µm; © R. Bauer.

        Bannoa Hamamoto

        Cystobasidium (Lagerh.) Neuhoff

        Naohidea Oberwinkler

        Occultifur Oberwinkler

        Sakaguchia Yamada, Maeda & Mikata

        Erythrobasidium Hamamoto, Sugiyama & Komagata

        Rhodotorula Harrison

        Mitosporic, not monophyletic

        Sporobolomyces Kluyver & van Niel

        Mitosporic, not monophyletic


Family Mixiaceae Kramer

        Mixia Kramer



  Class Ustilaginomycetes

The Ustilaginomycetes are predominantly plant-parasites and include ca. 1400 species. They can be distinguished from the other basidiomycetes because they have a distinctive cell wall carbohydrate composition with dominance of glucose and absence of xylose (Prillinger et al.,1990; 1993). The type B secondary structure of the 5S rRNA is shared with the Hymenomycetes (Gottschalk and Blanz, 1985) and, like the Urediniomycetes, they lack parenthesomes at the septal pores (Bauer et al., 1997).

Several recent publications have dealt with the classification of the Ustilaginomycetes (Bauer et al., 1997; Bauer et al., 2001; Begerow et al., 1997) and a comprehensive classification scheme based mainly on the characteristics of host-parasite interactions and septal pore apparatus has been proposed. In general, sequence data gives support to the classification proposals listed below, derived mainly from Bauer et al. (2001) and Begerow et al. (2002). The supra-generic classification of mitosporic taxa belonging to the Ustilaginomycetes is also being improved. The order Malasseziales Moore emend. Begerow, Bauer & Boekhout was redefined and presently accommodates the species of Malassezia Baillon, a genus of the Exobasidiomycetidae (Begerow et al., 1997). The genus Pseudozyma Bandoni emend. Boekhout seems to represent the asexual counterpart of Ustilago (Persoon) Roussel, whereas Tilletiopsis Derx ex Derx belongs to the Exobasidiomycetidae but is polyphyletic (Begerow et al., 2000). Sympodiomycopsis paphiopedili Sugiyama, Tokuoka & Komagata and three species of Rhodotorula, namely Rh. bacarum (Buhagiar) Rodrigues de Miranda & Weijman, Rh. hinnulea (Shivas & Rodrigues de Miranda) Rodrigues de Miranda & Weijman and Rh. phylloplana (Shivas & Rodrigues de Miranda) Rodrigues de Miranda & Weijman belong to the order Microstromatales of the Exobasidiomycetidae (Fell et al., 2001; Begerow et al., 2001). The classification system of the Ustilaginomycetes is presented below.

CLASS USTILAGINOMYCETES Bauer, Oberwinkler & Vánky


Members of the Ustilaginomycetes having local interaction zones and pores without membranous bands or caps (Bauer et al., 1997).

Order Entorrhizales Bauer & Oberwinkler

Family Entorrhizaceae Bauer & Oberwinkler

        Entorrhiza Weber


SUB-CLASS USTILAGINOMYCETIDAE Jülich emend. Bauer & Oberwinkler

Members of the Ustilaginomycetes having enlarged interaction zones. Septa have pores with membranous caps or they are poreless at maturity.

Order Urocystales Bauer & Oberwinkler

Family Melanotaeniaceae Begerow, Bauer & Oberwinkler

        Exoteliospora Bauer, Oberwinkler & Vánky

        Melanotaenium de Bary

        Yelsemia Walker


Family Doassansiopsaceae Begerow, Bauer & Oberwinkler

        Doassansiopsis (Setchell) Dietel


Family Urocystaceae Begerow, Bauer & Oberwinkler

        Mundkurella Thirumalachar

        Urocystis Rabenhorst ex Fuckel

        Ustacystis Zundel

        Vankya Ershad

Septal pore of Ustacystis waldsteiniae. Bar = 0.1 µm; © R. Bauer.

Order Ustilaginales Clinton emend. Bauer & Oberwinkler

Family Ustilaginaceae L. Tulasne & C. Tulasne emend. Bauer & Oberwinkler

        Anthracoidea Brefeld

        Cintractia Cornu

        Cintractiella K. B. Boedijn

        Clintamra Cordas & Durán

        Dermatosorus Sawada ex Ling

        Farysia Raciborski

        Farysporium Vánky

        Franzpetrakia Thirumalachar & Pavgi emend. Guo, Vánky & Mordue

        Geminago Vánky et Bauer

        Heterotolyposporium Vánky

        Kuntzeomyces Hennings ex Saccardo & Sydow

        Leucocintractia Piepenbring, Begerow & Oberwinkler

        Macalpinomyces Langdon & Fullerton emend. Vánky

        Melanopsichium Beck

        Moesziomyces Vánky

        Moreaua Liou & Cheng

        Orphanomyces Saville

        Pericladium Passerini emend. Mundkur

        Planetella Saville

        Pseudozyma Bandoni emend. Boekhout


        "Rhodotorula" acheniorum (Buhagiar & Barnett) Rodrigues de Miranda


        Schizonella Schröter

        Sporisorium Ehrenberg. ex Link

        Stegocintractia Piepenbring, Begerow & Oberwinkler

        Testicularia Klotzsch

        Tolyposporium Woronin ex Schröter

        Tranzscheliella Lavrov

        Trichocintractia Piepenbring, Begerow & Oberwinkler

        Uleiella Schröter

        Ustanciosporium Vánky emend. Piepenbring

        Ustilago (Persoon) Roussel

        Websdanea Vánky


Family Glomosporiaceae Cifferi emend. Begerow, Bauer & Oberwinkler

        Glomosporium Kochman

        Thecaphora Fingerhuth emend. Vánky

        Tothiella Vánky


Family Mycosyringaceae Bauer & Oberwinkler

        Mycosyrinx Beck

SUB-CLASS EXOBASIDIOMYCETIDAE Jülich emend. Bauer & Oberwinkler

Members of the Ustilaginomycetes having local interaction zones. Septa have pores with membranous bands or caps or they are poreless at maturity.

Order Malasseziales Moore emend. Begerow, Bauer & Boekhout

        Malassezia Baillon

Order Georgefischeriales Bauer, Begerow & Oberwinkler

Family Georgefischeriaceae Bauer, Begerow & Oberwinkler

        Georgefischeria Thirumalachar & Narasimhan emend Gandhe

        Jamesdicksonia Thirumalachar, Pavgi & Payak


Family Tilletiariaceae Moore

        Phragmotaenium Bauer, Begerow, Nagler & Oberwinkler

        Tilletiaria Bandoni & Johri

        Tilletiopsis flava (Tubaki) Boekhout


        Tilletiopsis fulvescens Gokhale


        Tolyposporella Atkinson


Family Eballistraceae (Bauer, Begerow, Nagler & Oberwinkler)

        Eballistra Bauer, Begerow, Nagler & Oberwinkler

Order Tilletiales Kreisel ex Bauer & Oberwinkler

Family Tilletiaceae L & C Tullasne emend. Bauer et Oberwinker

        Conidiosporomyces Vánky

        Erratomyces Piepenbring & Bauer

        Ingoldiomyces Vánky

        Neovossia Körnicke

        Oberwinkleria Vánky & Bauer

        Tilletia L. & C Tullasne

Order Microstromatales Bauer & Oberwinkler

Family Microstromataceae Jülich

        "Cerinosterus" cyanescens (de Hoog & de Vries) Moore

        Microstroma Niessl

        "Rhodotorula" bacarum (Buhagiar) Rodrigues de Miranda & Weijman


        "Rhodotorula" phylloplana (Shivas & Rodrigues de Miranda) Rodrigues de Miranda & Weijman


        Sympodiomycopsis Sugiyama, Tokuoka & Komagata



Family Volvocisporaceae Begerow, Bauer & Oberwinkler

        Volvocisporium Begerow, Bauer & Oberwinkler


Superorder Exobasidianae Bauer & Oberwinkler

Members of the Exobasididiomycetidae having simple pores and interaction apparatus

Order Entylomatales Bauer & Oberwinkler

Family Entylomataceae Bauer & Oberwinkler

        Entyloma de Bary

Order Doassansiales Bauer & Oberwinkler

Family Melaniellaceae Bauer, Vánky, Begerow & Oberwinkler

        Mellaniella Bauer, Vánky, Begerow & Oberwinkler


Family Doassansiaceae (Azb. & Karat.) Moore emend. Bauer & Oberwinkler

        Burillia Setchell

        Doassansia Cornu

        Doassinga Vánky, Bauer & Begerow

        Heterodoassansia Vánky

        Nannfeldtiomyces Vánky

        Narasimhania Thirumalachar et Pavgi emend. Vánky

        Pseudodermatosorus Vánky

        Pseudodoassansia (Setchell) Vánky

        Pseudotracya Vánky

        Tracya H. & P. Sydow


Family Rhamphosporaceae Bauer & Oberwinkler

        Rhamphospora Cunningham

Order Exobasidiales Hennings emend. Bauer & Oberwinkler

Family Brachybasidiaceae Gäumann

        Brachybasidium Gäumann

        Dicellomyces Olive

        Exobasidiellum Donk

        Kordyana Racib.

        Proliferobasidium Cunningham


Family Exobasidiaceae Hennings

        Exobasidium Woronin

        Muribasidiospora Kamat e & Rajendren


Family Cryptobasidiaceae Malençon ex Donk

        Botryoconis H. & P. Sydow

        Clinoconidium Pat.

        Coniodictyum Hariot & Pat.

        Drepanoconis Schröter & P. Hennings

        Laurobasidium Jülich


Family Graphiolaceae Fischer

        Graphiola Poiteau

        Stylina H. Sydow


Taxa of the Exobasidiomycetidae not ascribed to any family

Ceraceosorus bombacis (Bakshi) Bakshi


Tilletiopsis albescens Gokhale



Tilletiopsis cremea Tubaki



Tilletiopsis lilacina Tubaki



Tilletiopsis minor Nyland



Tilletiopsis pallescens Gokhale



Tilletiopsis washingtonensis Nyland




  Class Hymenomycetes (Tremellomycetidae)

The class Hymenomycetes as defined by Swann and Taylor (1995a; 1995b; 1995c) includes the complex-septate basidiomycetes (i.e. with dolipore), including the gasteromycetes. Moreover, the members of the Hymenomycetes have glucose as the major cell wall carbohydrate component and, at variance with the Urediniomycetes and Ustilaginomycetes, xylose is present. Swann and Taylor (1995b) distinguished two subclasses, viz. Hymenomycetidae and Tremellomycetidae. A few traits other than DNA sequences differentiate the two subclasses. Sacculate membrane elements are normally observed at the vicinity of the dolipore of the Tremellomycetidae (in some cases they are lacking), whereas the members of the Hymenomycetidae have smooth membranous septal pore caps of various types, but not sacculate. In addition, whereas the vast majority of the Tremellomycetidae produces a yeast phase, such ontogenetic stage is not observed in the Hymenomycetidae. Therefore, the present review will deal only with the Tremellomycetidae. The classification system presented below is based mainly on the revisions of Wells (1994) and Wells and Bandoni (2001) which are derived mostly from comparisons of morphological and ultrastructural data. New teleomorphic genera described in Kirschner et al. (2001) and Sampaio et al. (2002) are also included. Although most mitosporic species of the Tremellomycetidae have been investigated in comprehensive sequence analysis studies (Fell et al., 2000; 2001; Scorzetii et al., 2002), several sexual taxa have not yet been studied using the same approaches. It is anticipated that relevant systematic changes will be proposed when such species are investigated by molecular phylogenetic methods.

Septal pore types in the Hymenomycetes. 

Left: Dolipore with perfurated parenthesomes of Schizophyllum commune (Bar = 0.25 µm; © R. Bauer).

Center: Dolipore with continuous parenthesomes of Tulasnella sp. (Bar = 0.25 µm; © R. Bauer).

Left: Dolipore with cup-shaped parenthesomes of Tremella sp. (Bar = 0.2 µm; © R. Bauer).



Order Cystofilobasidiales Boekhout & Fell

        Itersonilia Derx

        "Cryptococcus" aquaticus (Jones & Slooff) Rodrigues de Miranda & Weijman

        "Cryptococcus" huempii (Ramírez & González) Roeijmans, van Eijk & Yarrow

        "Cryptococcus" macerans (Frederiksen) Phaff & Fell

        Cystofilobasidium Oberwinkler & Bandoni

        Mrakia Yamada & Komagata

        Tausonia Bab’eva


        "Trichosporon" pullulans (Lindner) Diddens & Lodder

        Udeniomyces Nakase & Takematsu


        Xanthophyllomyces Golubev

Order Trichosporonales Boekhout & Fell

        "Cryptococcus" curvatus (Diddens & Lodder) Golubev

        "Cryptococcus" humicola (Daszewska) Golubev

        Trichosporon Behrend


Order Filobasidiales Jülich

Family Filobasidiaceae Olive

        Cryptococcus Vuillemin

        Mitosporic, not monophyletic

        Filobasidium Olive

Basidium and basidiospores of Filobasidium floriforme. Bar = 10 µm; © J.P. Sampaio.



Family Carcinomycetaceae Oberwinkler & Bandoni

        Carcinomyces Oberwinkler & Bandoni

        Christiansenia Hauerslev

        Syzygospora Martin

Order Tremellales Rea emend. Bandoni

Family Cuniculitremaceae Sampaio & Kirschner

        Cuniculitrema Sampaio & Kirschner

        Fellomyces Yamada & I. Banno


        Kockovaella Nakase, Banno & Yamada


        Sterigmatosporidium Kraepelin & Schulze



Family Phragmoxenidiaceae Oberwinkler & Bauer

        Phragmoxenidium Oberwinkler

        Phyllogloea Lowy


Family Sirobasidiaceae Möller

        Fibulobasidium Bandoni

        Sirobasidium Lagerh. & Pat.


Family Tremellaceae Fries emend. Bandoni

        Bullera Derx

        Mitosporic, not monophyletic

        Bulleribasidium Sampaio, Weiss & Bauer

        Bulleromyces Boekhout & Fonseca

        Cryptococcus Vuillemin

        Mitosporic, not monophyletic

        Dioszegia Zsolt emend. Takashima, Deak & Nakase

        Filobasidiella Kwon-Chung

        Holtermannia Sacc. & Traverso

        Papiliotrema Sampaio, Weiss & Bauer

        Sirotrema Bandoni

        Tremella Pers.

        Not monophyletic

        Tremellina Bandoni


        Trimorphomyces Bandoni & Oberw.

        Tsuchiyaea Yamada, Kawasaki, Itoh, Banno & Nakase


        Xenolachne Rogers


Family Tetragoniomycetaceae Oberwinkler & Bandoni

        Tetragoniomyces Oberwinkler & Bandoni


Family Rhynchogastremataceae Oberwinkler & Metzler

        Rhynchogastrema Metzler & Oberwinkler


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Sampaio, J.P. and Bauer, R. 2003. The classification of dimorphic basidiomycetes. Dimorphic Basidiomycetes WWW Project.

Version 01, saved 25 March 2003.

José Paulo Sampaio:

Centro de Recursos Microbiológicos, Secção Autónoma de Biotecnologia, Faculdade de Ciências e Tecnologia, Universidade Nova de Lisboa, 2829-516 Caparica, Portugal.

Robert Bauer:

Universität Tübingen, Institut für Biologie I, Lehrstuhl Spezielle Botanik und Mykologie, Auf der Morgenstelle 1, D-72076 Tübingen, Germany


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