J.P. Sampaio and R. Bauer

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).

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.

        Cryptomycocolax Oberwinkler & Bauer

        Colacosiphon Kirschner, Bauer & Oberwinkler

        Leucosporidiella Sampaio

        Mitosporic

        Sporidiobolus Nyland

        Sporobolomyces Kluyver & van Niel

        Mitosporic, not monophyletic

        Rhodotorula Harrison

        Mitosporic, not monophyletic

        Liroa Ciferri

        Microbotryum Léveillé emend. Vánky

        Approximately 75 species parasitizing dicotyledonous plants

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.

        Camptobasidium Marvanová & Suberkropp

Atractocolax Kirschner, Bauer & Oberwinkler

Bensingtonia Ingold emend. Nakase & Boekhout

        Agaricostilbum Wright emend. Wright, Bandoni & Oberwinkler

        Chionosphaera Cox

        Stilbum Tode ex Mérat

Bensingtonia Ingold emend. Nakase & Boekhout

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

Mycogloea Olive

Sporobolomyces Kluyver & van Niel

Sterigmatomyces Fell emend. Yamada & Banno

Spiculogloea Roberts

Zygogloea Roberts

14 families, 164 genera, aprox. 7000 species

        Coccidiodictyon Oberwinkler

        Ordonia Racib.

        Septobasidium Pat.

        Uredinella Couch

Herpobasidium Lind emend. Oberw & Bandoni

Insolibasidium Oberwinkler & Bandoni

Jola Möller

Platycarpa Couch emend. Oberwinkler & Bandoni

Ptechetelium Oberwinkler & Bandoni

        Saccoblastia Möller

        Classicula Bauer, Begerow, Oberwinkler & Marvanová

        Jaculispora H. J. Huds. & Ingold

        Cystobasidium (Lagerh.) Neuhoff

        Naohidea Oberwinkler

        Occultifur Oberwinkler

        Sakaguchia Yamada, Maeda & Mikata

        Erythrobasidium Hamamoto, Sugiyama & Komagata

        Rhodotorula Harrison

        Sporobolomyces Kluyver & van Niel

        Mixia Kramer

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.

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

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

        Exoteliospora Bauer, Oberwinkler & Vánky

        Melanotaenium de Bary

        Yelsemia Walker

        Mundkurella Thirumalachar

        Urocystis Rabenhorst ex Fuckel

        Ustacystis Zundel

        Vankya Ershad

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

        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

        Mitosporic

        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

        Glomosporium Kochman

        Thecaphora Fingerhuth emend. Vánky

        Tothiella Vánky

        Mycosyrinx Beck

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

        Georgefischeria Thirumalachar & Narasimhan emend Gandhe

        Phragmotaenium Bauer, Begerow, Nagler & Oberwinkler

        Tilletiaria Bandoni & Johri

        Tilletiopsis flava (Tubaki) Boekhout

        Mitosporic

        Mitosporic

        Eballistra Bauer, Begerow, Nagler & Oberwinkler

        Conidiosporomyces Vánky

        Erratomyces Piepenbring & Bauer

        Ingoldiomyces Vánky

        Neovossia Körnicke

        Oberwinkleria Vánky & Bauer

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

        Mitosporic

        Sympodiomycopsis Sugiyama, Tokuoka & Komagata

        Mitosporic

Members of the Exobasididiomycetidae having simple pores and interaction apparatus

        Entyloma de Bary

        Mellaniella Bauer, Vánky, Begerow & 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

        Rhamphospora Cunningham

        Brachybasidium Gäumann

        Dicellomyces Olive

        Exobasidiellum Donk

        Kordyana Racib.

        Proliferobasidium Cunningham

        Exobasidium Woronin

        Muribasidiospora Kamat e & Rajendren

        Botryoconis H. & P. Sydow

        Clinoconidium Pat.

        Coniodictyum Hariot & Pat.

        Drepanoconis Schröter & P. Hennings

        Laurobasidium Jülich

        Graphiola Poiteau

        Stylina H. Sydow

Ceraceosorus bombacis (Bakshi) Bakshi

Tilletiopsis albescens Gokhale

Mitosporic

Mitosporic

Mitosporic

Mitosporic

Mitosporic

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).

        "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

        Mitosporic

        "Trichosporon" pullulans (Lindner) Diddens & Lodder

        Udeniomyces Nakase & Takematsu

        Mitosporic

        Xanthophyllomyces Golubev

        "Cryptococcus" curvatus (Diddens & Lodder) Golubev

        "Cryptococcus" humicola (Daszewska) Golubev

        Trichosporon Behrend

        Mitosporic

        Cryptococcus Vuillemin

        Mitosporic, not monophyletic

        Filobasidium Olive

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

        Carcinomyces Oberwinkler & Bandoni

        Christiansenia Hauerslev

        Syzygospora Martin

        Cuniculitrema Sampaio & Kirschner

        Fellomyces Yamada & I. Banno

        Mitosporic

        Kockovaella Nakase, Banno & Yamada

        Mitosporic

        Sterigmatosporidium Kraepelin & Schulze

        Mitosporic

        Phragmoxenidium Oberwinkler

        Phyllogloea Lowy

        Fibulobasidium Bandoni

        Sirobasidium Lagerh. & Pat.

        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

        Trimorphomyces Bandoni & Oberw.

        Tsuchiyaea Yamada, Kawasaki, Itoh, Banno & Nakase

        Mitosporic       

        Xenolachne Rogers

        Tetragoniomyces Oberwinkler & Bandoni

Bauer, R. and Oberwinkler, F. 1991a. The colacosomes: new structures at the host-parasite interface of a mycoparasitic basidiomycete. Botanica Acta 104: 53-57.

Bauer, R. and Oberwinkler, F. 1991b. The symplechosome: a unique cell organelle of some basidiomycetes. Botanica Acta 104: 93-97.

Bauer, R. and Oberwinkler, F. 1994. Meiosis, septal pore architecture and systematic position of the heterobasidiomycetous fern parasite Herpobasidium filicinum. Canadian Journal of Botany 72: 1229-1242.

Bauer, R., Oberwinkler, F. and Vánky, K. 1997. Ultrastructural markers and systematics in smut fungi and allied taxa. Canadian Journal of Botany. 75: 1273-1314.

Bauer, R., Begerow, D, Oberwinkler, F. Piepenbring, M. and Berbee, ML. 2001. Ustilaginomycetes In The Mycota – vol VII, Systematics and Evolution, D.J. McLaughlin, E.G. McLaughlin and P.A. Lemke (eds.), Springer-Verlag, Berlin, pp 58-83.

Bauer, R., Begerow, D., Oberwinkler, F. and Marvanová, L. 2003. Classicula: the teleomorph of Naiadella fluitans. Mycologia (in press).

Begerow, D., Bauer, R. and Oberwinkler, F. 1997. Phylogenetic studies on nuclear large subunit ribosomal DNA sequences of smut fungi and related taxa. Canadian Journal Botany 75: 2045-2056.

Begerow, D., Bauer, R. and Boekhout, T. 2000. Phylogenetic placements of ustilaginomycetous anamorphs as deduced from nuclear LSU rDNA sequences. Mycological Research 104: 53-60.

Begerow, D., Bauer, R. and Oberwinkler, F. 2001. Muribasidiospora: Microstromatales or Exobasidiales? Mycological Research 105: 798-810.

Begerow, D., Bauer, R. and Oberwinkler, F. 2002. The Exobasidiales: an evolutionary hypothesis Mycological Progress 1: 187-199.

Fell, J.W., Boekhout, T., Fonseca, A., Scorzetti, G. and Statzell-Tallman, A. 2000. Biodiversity and systematics of basidiomycetous yeasts as determined by large-subunit rDNA D1/D2 domain sequence analysis. International Journal of Systematic and Evolutionary Microbiology 50: 1351-1371.

Fell, J.W., Boekhout, T., Fonseca, A., Sampaio, J.P. 2001. Basidiomycetous yeasts. In The Mycota – vol VII, Systematics and Evolution, D.J. McLaughlin, E.G. McLaughlin and P.A. Lemke (eds.), Springer-Verlag, Berlin, pp 3-35.

Fonseca, A., Sampaio, J.P., Inácio, I. and Fell, J.W. 2000. Emendation of the basidiomycetous yeast genus Kondoa and the description of Kondoa aeria sp. nov. Antonie van Leeuwenhoek 77: 293-302.

Gottschalk, M. and Blanz, P.A. 1985. Untersuchungen an 5S ribosomalen Ribonucleinsäuren als Beitrag zur Klärung von Systematik und Phylogenie der Basidiomyceten. Zeitschrift für Mycologie 51: 205-243.

Hamamoto, M. and Nakase, T. 2000. Phylogenetic analysis of the ballistoconidium-forming yeast genus Sporobolomyces based on 18S rDNA sequences. International Journal of Systematic and Evolutionary Microbiology 50: 1373-1380.

Hamamoto, M., Sugiyama, J. and Komagata, K. 1988. Transfer of Rhodotorula hasegawae to a new basidiomycetous genus Erythrobasidium as Erythrobasidium hasegawae comb. nov. Journal of General and Applied Microbiology 34: 279-287.

Hamamoto, M., Thanh, V.N. and Nakase, T. 2002. Bannoa hahajimensis gen. nov., sp. nov., and three related anamorphs, Sporobolomyces bischofiae sp. nov., Sporobolomyces ogasawarensis sp. nov. and Sporobolomyces syzygii sp. nov., yeasts isolated from plants in Japan. International Journal of Systematic and Evolutionary Microbiology 52: 1023-1032.

Kirschner, R., Sampaio, J.P., Gadanho, M., Weiss, M. and Oberwinkler, F. 2001. Cuniculitrema polymorpha (Tremellales, gen. nov. and sp. nov.), a heterobasidiomycete vectored by bark beetles, which is the teleomorph of Sterigmatosporidium polymorphum. Antonie van Leeuwenhoek 80: 149-161.

McLaughlin, D.J. 1990. A new cytoplasmic structure in the basidiomycete Helicogloea: the microscala. Experimental Mycology 14: 331-338.

McLaughlin, D.J., Berres, M.E. and Szabo, L.J. 1995a. Molecules and morphology in basidiomycete phylogeny. Canadian Journal of Botany 73: 684-692.

McLaughlin, D.J., Frieders, E.M. and Lü, H. 1995b. A microscopist’s view of heterobasidiomycete phylogeny. Studies in Mycology 38: 91-109.

Oberwinkler, Fand Bauer, R. 1989. The systematics of gasteroid, auricularioid Heterobasidiomycetes. Sydowia 41: 224-256.

Prillinger, H., Dörfler, Ch., Laaser, G. e Hauska, G. 1990. Ein Beitrag zur Systematik und Entwicklungsbiologie Höherer Pilze. Hefe-Typen der Basidiomyceten. Teil III:Ustilago-Typ. Zeitschrift für Mykologie 56: 251-278.Prillinger, H., Laaser, G., Dörfler, Ch., e Ziegler, K. 1991a. Ein Beitrag zur Systematik und Entwicklungsbiologie Höherer Pilze. Hefe-Typen der Basidiomyceten. Teil IV: Dacrymyces-Typ, Tremella-Typ. Sydowia 53: 170-218.

Prillinger, H., Deml, G., Dörfler, Ch., Laaser, G. e Lockau, W. 1991b. Ein Beitrag zur Systematik und Entwicklungsbiologie Höherer Pilze. Hefe-Typen der Basidiomyceten. Teil II: Microbotryum-Typ. Botanica Acta 104: 5-17.

Prillinger, H., Oberwinkler, F., Umile, C., Tlachac, K., Bauer, R., Dörfler, C. e Taufratzhofer, E. 1993. Analysis of cell wall carbohydrates (neutral sugars) from ascomycetous and basidiomycetous yeasts with and without derivatization. Journal of General and Applied Microbiology 39: 1-34.

Sampaio, J.P., Bauer, R., Begerow, D. andOberwinkler, F. 1999. Occultifur externus sp. nov., a new species of simple-pored auricularioid heterobasidiomycete from plant litter in Portugal. Mycologia 91: 1094-1101.

Sampaio, J.P., Weiß, M., Gadanho, M. and Bauer, R. 2002. New taxa in the Tremellales: Bulleribasidium oberjochense gen. et sp. nov., Papiliotrema bandonii gen. et sp. nov. and Fibulobasidium murrhardtense sp. nov. Mycologia 94: 873-887.

Sampaio, J.P., Gadanho, M., Bauer, R. and Weiß, M. 2003. Taxonomic studies in the Microbotryomycetidae: Leucosporidium golubevii sp. nov., Leucosporidiella gen. nov. and the new orders Leucosporidiales and Sporidiobolales. Mycological Progress (in press).

Scorzetti, G., Fell, J.W., Fonseca, A. and Statzell-Tallman, A. 2002. Systematics of basidiomycetous yeasts: a comparison of large subunit D1/D2 and internal transcribed spacer rDNA regions. FEMS Yeast Research 2: 495-517.

Swann, E.C. and Taylor, J.W. 1993. Higher taxa of basidiomycetes: an 18S rRNA gene perspective. Mycologia 85: 923-936.

Swann, E.C. and Taylor, J.W. 1995a. Phylogenetic perspectives on basidiomycete systematics: evidence from the 18S rRNA gene. Canadian Journal of Botany 73: 649-1433.

Swann, E.C. and Taylor, J.W. 1995b. Phylogenetic diversity of yeast-producing basidiomycetes. Mycological Research 99: 1205-1210.

Swann, E.C. and Taylor, J.W. 1995c. Toward a phylogenetic systematics of the Basidiomycota: integrating yeasts and filamentous basidiomycetes using 18S rRNA gene sequences. Studies in Mycology 38: 147-161.

Swann, E.C. Frieders, E.M. and McLaughlin, D.J. 1999. Microbotryum, Kriegeria and the changing paradigm in basidiomycete classification. Mycologia 91: 51-66.Swann, E.C. Frieders, E.M. and McLaughlin, D.J. 2001. Urediniomycetes In The Mycota – vol VII, Systematics and Evolution, D.J. McLaughlin, E.G. McLaughlin and P.A. Lemke (eds.), Springer-Verlag, Berlin, pp 37-56

Takashima, M., Hamamoto, M. and Nakase, T. 2000. Taxonomic significance of fucose in the class Urediniomycetes: distribution of fucose in cell wall and phylogeny of urediniomycetous yeasts. Systematic and Applied Microbiology 23: 63-70.

Taylor, J.W. 1995. Molecular phylogenetic classification of fungi. Archives of Medical Research 26: 307-314.

Wells, K. 1994. Jelly fungi, then and now! Mycologia 86: 18-48.

Wells, K. and Bandoni, R.J. 2001. Heterobasidiomycetes. In The Mycota – vol VII, Systematics and Evolution, D.J. McLaughlin, E.G. McLaughlin and P.A. Lemke (eds.), Springer-Verlag, Berlin, pp 85-120.