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Exobasidiomycetes

Ceraceosorales Doassansiales Entylomatales Exobasidiales Georgefischeriales Microstromatales Tilletiales

Ustilaginomycetes

Urocystales Ustilaginales

Malasseziomycetes

Malassezioales

Moniliellomycetes

Moniliellales

The Ustilaginomycotina
Ustilaginomycotina
is a subdivision within the division Basidiomycota
Basidiomycota
of the kingdom Fungi. It consists of the classes Ustilaginomycetes
Ustilaginomycetes
and Exobasidiomycetes,[1] and in 2014 the subdivision was reclassified and the two additional classes Malasseziomycetes and Moniliellomycetes added.[2][3] The name was first published by Doweld in 2001; Bauer and colleagues later published it in 2006 as an isonym.[4] Ustilagomycotina and Agaricomycotina
Agaricomycotina
are considered to be sister groups, and they are in turn sister groups to the subdivision Pucciniomycotina.[4] Ustilaginomycotina
Ustilaginomycotina
comprises 115 genera with more than 1700 species.[4][5] The subdivision is mostly plant parasites on vascular plants, and the distribution of the subdivision is therefore restricted to the distribution of the host. The group is also called the true smut fungi because of the production of teliospores. The name smut is still used as a term since it circumscribes the organization and life cycle of Ustilaginomycotina, but it is not a taxonomic term. Ustilaginomycotina
Ustilaginomycotina
has some of the best known and studied genera of plant parasites like Ustilago
Ustilago
and Tilletia
Tilletia
and it is also of great economic importance.[6][7]

Contents

1 Morphology

1.1 Defining Features

2 Classification

2.1 Historic Classification 2.2 Modern Classification

3 Ecology

3.1 Distribution 3.2 Role in the Environment

4 Life Cycle 5 References 6 External links

Morphology[edit] Ustilaginomycotina
Ustilaginomycotina
is morphologically a highly diverse group. It consists of two states: anamorphic yeast state and teleomorphic filamentous smut state.[7] These two states in the life cycle can look very different. Defining Features[edit] The Ustilaginomycotina
Ustilaginomycotina
has different ultrastructural morphologies that defines the subdivision and some of them are also used to delimit the classes in the subdivision. Cellular interaction The cellular interactions are referring to the interaction between the hyphae and the host plant cells. These zones provide a useful ultrastructural character for delimiting the classes in Ustilaginomycotina.[1][8] The host-interactions zones can either be local interaction zones and then characterizing the Exobasidiomycetes, or enlarged interaction zones characterizing the Ustilaginomycetes. Cellular composition Ustilaginomycotina
Ustilaginomycotina
has a distinctive cell wall composition consistent of mostly glucose and absence of xylose. This character separates the Ustilaginomycotina
Ustilaginomycotina
from Pucciniomycotina
Pucciniomycotina
and Agaricomycotina.[9] Septal pores The architecture of the septal pores plays an important part in delimiting the subdivision in Basidiomycotina. In contrast to the Pucciniomycotina, the Ustilaginomycotina
Ustilaginomycotina
has a septal pore with a membrane cap or it is poreless. It does not have a dolipore or parenthesome as the Agaricomycotina
Agaricomycotina
do.[1][10] 5S rRNA In 1985 Gottschalk and Blanz did a study about the 5s ribosomal RNA and distinguished two types of structures in the Basidiomycota. These two types was named the type A secondary structure and the type B secondary structure of the 5S rRNA. This is a useful tool for delimiting between the subdivisions. The Ustilaginomycetina has the type B secondary structure and they share this character with the Agaricomycotina, and it separates them from the Pucciniomycotina
Pucciniomycotina
which has the type A secondary structure.[11] Classification[edit] Historic Classification[edit] In 1847 Tulasne and Tulasne divided the so-called smut fungi into two groups called Ustilaginacceae and Tilletiaceae.[12] Traditionally morphological characters of the basidia was used for the delimitation of the smut fungi group, but after a thorough investigation of the ultrastructural characters the group was revised.[1][4] With the use of electron microscopy, Bauer et al. 1997 found two separates lines of the smut fungi, namely the Microbotryales (which is now moved to Puzziniomycotina) and the Ustilaginomycetes. Within the Ustilaginomycetes
Ustilaginomycetes
they identified three linages: the Entorrhizomycetidea, Ustilaginomycetidae and the Exobasidiomycetidae.[1][13] Modern Classification[edit] Ustilaginomycotina
Ustilaginomycotina
consists of both anamorphic yeast species and teleomorphic filamentous smuts.[1][7] These two states can look very different and is not always easy to connect. The two different states has therefore produced different names on the same species. With the use of both morphological characters and molecular data, the subdivision is now reclassified and many of the species has been renamed by the 'One Fungus
Fungus
= One Name’ principle.' [2] By looking at ultrastructural morphological structures like host-parasite interactions, the septal pore apparatus [1][10] and molecular sequence analysis a new classification was proposed. Here Ustilaginomycotina
Ustilaginomycotina
consisted of the three classes Ustilaginomycetes, Exobasidiomycetes
Exobasidiomycetes
and Entorrhizomycetes.[7] But, the presence of Entorrhizomycetes was questioned and are now considered as incertae sedis in the Basidiomycota.[5] In 2014 a new multiple gene sequence analysis showed that two additional classes, forming two independent deep lineages should be recognized as classes in the subdivision: Malasseziomycetes and Moniliellomycetes.[3] Ecology[edit] Distribution[edit] Since Ustilaginomycotina
Ustilaginomycotina
is mostly plant parasites, the group is restricted to the host species of vascular plants, and mainly on angiosperms and monocots.[1][14] This encompasses a geographical distribution in both tropical, temperate and arctic regions. Most species are highly host-specific and this may be a product of coevolution with different angiosperm lineages. This is supported by studies that shows that some monophyletic lineages in the Ustilaginomycotina
Ustilaginomycotina
are restricted to monophyletic lineages in the angiosperms. But not all taxa in Ustilaginomycotina
Ustilaginomycotina
are host-specific, some have a broad host range and others have also made a host jump to other vascular plants and not only monocots in the angiosperms.[14][15] Role in the Environment[edit] Ustilaginomycotina
Ustilaginomycotina
have an array of plant pathogens, and some are parasitizing on economically important species like wheat, barley and corn. In some cases the yield loss is minimal, in other the crops has to be quarantined. Some of the galls produced by the smuts is considered as a delicacy in some parts of the world. Malassezia lineages also causes harm on human skin. Ustilago
Ustilago
maydis Tilletia Malassezia Life Cycle[edit]

Life cycle of Ustilago
Ustilago
maydis

The life cycle of the subdivision is dimorphic and it consists of two phases in the life cycle. One saprobic haploid phase and a parasitic (biotrophic) dikaryotic phase. The saprobic phase is initiated by the production of haploid yeasts, which fuses with another spore and produce the n+n hyphae which will infect the host. The infection happens with the production of a structure called appressoria, which is a specialized cell that is used to penetrate the host cuticle. Inside the host, the fungi will produce hyphae and another specialized structure called haustoria. This will take nutrition from the plant, and is a parasitic feature. Almost all of the Ustilaginomycotina
Ustilaginomycotina
will then sporulate inside the host, and this happens with the spore becoming thick-walled and will separate, now called a teliospore. The teliospore is the most conspicuous part of the individual and represent the smut syndrome. This teliospore is a specialized resting spore that can survive outside their host. The teliospore is released from the host, and it will produce a diploid basidium and the cycle starts over. Species from the subdivision obtained from the nature will often be in the yeast-like state.[6][14][15] References[edit]

^ a b c d e f g h Bauer, R., Oberwinkler, F. and Vánky, K. (1997). " Ultrastructural
Ultrastructural
markers and systematics in smut fungi and allied taxa". Canadian Journal of Botany. 75: 1273–1314. doi:10.1139/b97-842. CS1 maint: Multiple names: authors list (link) ^ a b Wang, QW, Begerow, D., Groenewald, M., Liu, XZ., Theelen, T., Bai, F. Y., Boekhout, T. (2014). "Multigene phylogeny and taxonomic revision of yeasts and related fungi in the Ustilaginomycotina". Studies in Mycology. 81: 55–83. doi:10.1016/j.simyco.2015.10.004. CS1 maint: Multiple names: authors list (link) ^ a b Wang QM, Theelen B, Groenewald M (2014). "Moniliellomycetes and Malasseziomycetes, two new classes in Ustilaginomycotina". Persoonia. 33: 41–47. doi:10.3767/003158514x682313.  ^ a b c d Bauer, R., Begerow, JP., Samp, M., Weiß, M., Oberwinkler F. (2006). "The simple-septate basidiomycetes: a synopsis". Mycological Progress. 5 (1): 41–66. doi:10.1007/s11557-006-0502-0. CS1 maint: Multiple names: authors list (link) ^ a b Hibbett, D.S., Binder, M., Bischoff, J.F., Blackwell, M., Cannon, P.F., Eriksson, O.E., Huhndorf, S., James, T., Kirk, P.M., Lücking, R., Lumbsch, H.T., Lutzoni, F., Matheny, P.B., McLaughlin, D.J., Powell, M.J., Redhead, S., Schoch, C.L., Spatafora, J.W., Stalpers, J.A., Vilgalys, R., Aime, M.C., Aptroot, A., Bauer, R., Begerow, D., Benny, G.L., Castlebury, L.A., Crous, P.W., Dai, Y.-C., Gams, W., Geiser, D.M., Griffith, G.W., Gueidan, C., Hawksworth, D.L., Hestmark, G., Hosaka, K., Humber, R.A., Hyde, K.D., Ironside, J.E., Kõljalg, U., Kurtzman, C.P., Larsson, K.-H., Lichtwardt, R., Longcore, J., Miadlikowska, J., Miller, A., Moncalvo, J.-M., Mozley-Standridge, S., Oberwinkler, F., Parmasto, E., Reeb, V., Rogers, J.D., Roux, C., Ryvarden, L., Sampaio, J.P., Schüßler, A., Sugiyama, J., Thorn, R.G., Tibell, L., Untereiner, W.A., Walker, C., Wang, Z., Weir, A., Weiss, M., White, M.M., Winka, K., Yao, Y.-J. & Zhang, N. (2007). "A higher-level phylogenetic classification of the Fungi". Mycological Research. 111: 509–547. CS1 maint: Multiple names: authors list (link) ^ a b Bauer, R., Begerow, D., Oberwinkler, F. (2008). " Ustilaginomycotina
Ustilaginomycotina
R. Bauer, Begerow, J. P. Samp., M. Weiß & Oberw. 2006. The true smut fungi". The Tree of Life Web Project. CS1 maint: Multiple names: authors list (link) ^ a b c d Begerow, D., Stol,l M., Bauer, R. (2006). "A phylogenetic hypothesis of Ustilaginomycotina
Ustilaginomycotina
based on multiple gene analyses and morphological data". Mycologia. 98: 906–916. doi:10.3852/mycologia.98.6.906. CS1 maint: Multiple names: authors list (link) ^ Bauer, R., Mendgen, K. and Oberwinkler, F. (1995). "Cellular interaction of the smut fungus Ustacystis waldsteiniae". Canadian Journal of Botany. 73: 867–883. doi:10.1139/b95-095. CS1 maint: Multiple names: authors list (link) ^ Prillinger, H., Oberwinkler, F., Umile, C., Tlachac, K., Bauer, R., Dörfler, C. and 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. doi:10.2323/jgam.39.1. CS1 maint: Multiple names: authors list (link) ^ a b Bauer, R., Mendgen, K. and Oberwinkler, F. (1995). "Septal pore apparatus of the smut Ustacystis waldsteiniae". Mycologia. 87: 18–24. doi:10.2307/3760941. CS1 maint: Multiple names: authors list (link) ^ Gottschalk, M. & Blanz, P.A. (1985). "Untersuchungen an 5S ribosomalen Ribonucleinsäuren als Beitrag zur Klärung von Systematik und Phylogenie der Basidiomyceten". Zeitschrift für Mykologie. 51: 205–243.  ^ Tulasne, L.; Tulasne, C. (1847). "Mémoire sur les Ustilaginées comparées Uredinées". Annales des Sciences Naturelles. Botanique. 3 (7): 12–127.  ^ Vánky, K. (2003). "Cintractiellaceae fam. nov. (Ustilaginomycetes)". Fungal Diversity. 13: 167–173.  ^ a b c Begerow, D., Göker, M., Lutz, M., Stoll, M. (2004). "On the evolution of smut fungi on their hosts". Frontiers in Basidiomycote Mycology (PDF). pp. 81–98. CS1 maint: Multiple names: authors list (link) ^ a b Begerow. D., Schafer, AM., Kellner, R., Oberwinkel, D., Bauer, R. (2014). The Mycota, Vol. VII, Part A: systematics and evolution. Ustilaginomycotina
Ustilaginomycotina
(2nd ed.). Berlin, Germany: Springer-Verlag. pp. 295–329. CS1 maint: Multiple names: authors list (link)

External links[edit]

Tree of Life Ustilaginomycotina

v t e

Opisthokont: True fungi classification, fungal orders

Domain Archaea Bacteria Eukaryota (Supergroup Plant Hacrobia Heterokont Alveolata Rhizaria Excavata Amoebozoa Opisthokonta

Animal Fungi)

Dikarya

Ascomycota (sac fungi)

Pezizomycotina

Leotiomyceta

Dothideomyceta

Coniocybomycetes Lichinomycetes Arthoniomycetes Dothideomycetes Eurotiomycetes Lecanoromycetes

Sordariomyceta

Xylonomycetes Geoglossomycetes Leotiomycetes Laboulbeniomycetes Sordariomycetes

Other

Orbiliomycetes Pezizomycetes

Saccharomycotina

Saccharomycetes

Taphrinomycotina

Archaeorhizomycetes Neolectomycetes Pneumocystidomycetes Schizosaccharomycetes Taphrinomycetes

Basidiomycota (with basidia)

Pucciniomycotina

Tritirachiomycetes Mixiomycetes Agaricostilbomycetes Cystobasidiomycetes Microbotryomycetes Classiculomycetes Cryptomycocolacomycetes Atractiellomycetes Pucciniomycetes

Ustilaginomycotina

Monilielliomycetes Malasseziomycetes Ustilaginomycetes Exobasidiomycetes

Agaricomycotina

Hymenomycete

Dacrymycetales Agaricomycetes

Other

Wallemiomycetes Bartheletiomycetes Tremellomycetes

Entorrhizomycota

Entorrhizomycetes

Glomeromycota

Glomeromycetes

Zygomycota (paraphyletic)

Mucoromycotina

Mortierellomycetes Mucoromycetes

Kickxellomycotina

Zoopagomycetes Kickxellomycetes

Entomophthoromycotina

Neozygitomycetes Basidiobolomycetes Entomophthoromycetes

Zoosporic fungi (paraphyletic)

Olpidiomycota

Olpidiomycetes

Blastocladiomycota

Blastocladiomycetes

Chytridiomycota

Neocallimastigomycetes Hyaloraphidiomycetes Monoblepharidomycetes Chytridiomycetes

Fungal phyla are underlined. See also: fungi imperfecti (polyphyletic group).

Taxon identifiers

Wd: Q1204312 EoL: 3020735 Fungorum: 90778 ITIS: 936294 MycoBank:

.