Ustilaginomycotina

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Ustilaginomycotina
Ustilago maydis de 2.jpg
Ustilago maydis
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Fungi
Division: Basidiomycota
Subdivision: Ustilaginomycotina
Doweld (2001)
Classes/orders

Exobasidiomycetes

Ceraceosorales
Doassansiales
Entylomatales
Exobasidiales
Georgefischeriales
Microstromatales
Tilletiales

Ustilaginomycetes

Urocystales
Ustilaginales

Malasseziomycetes

Malasseziales

Monilielliomycetes

Moniliellales

Peribolosporomycetes

Peribolosporales

The Ustilaginomycotina is a subdivision within the division Basidiomycota of the kingdom Fungi. It consists of the classes Ustilaginomycetes and Exobasidiomycetes, [1] and in 2014 the subdivision was reclassified and the two additional classes Malasseziomycetes and Monilielliomycetes 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 are considered to be sister groups, and they are in turn sister groups to the subdivision Pucciniomycotina. [4]

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 has some of the best known and studied genera of plant parasites like Ustilago and Tilletia and it is also of great economic importance. [6] [7]

Morphology

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

The 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 has a distinctive cell wall composition consistent of mostly glucose and absence of xylose. This character separates the Ustilaginomycotina from 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 has a septal pore with a membrane cap or it is poreless. It does not have a dolipore or parenthesome as the 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 which has the type A secondary structure. [11]

Classification

Historic classification

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 Pucciniomycotina) and the Ustilaginomycetes. Within the Ustilaginomycetes they identified three lineages: the Entorrhizomycetidea, Ustilaginomycetidae and the Exobasidiomycetidae. [1] [13]

Modern classification

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 = 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 consisted of the three classes Ustilaginomycetes, 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 Monilielliomycetes. [3]

Ecology

Distribution

Since 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 are restricted to monophyletic lineages in the angiosperms. But not all taxa in 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

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 maydis

Tilletia

Malassezia

Life cycle

Life cycle of Ustilago maydis 03 03 07 life cycle of Ustilago maydis on corn, Ustilaginales Basidiomycota (M. Piepenbring).png
Life cycle of 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 an appressorium, which is generated by a specialized cell 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 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]

Related Research Articles

<span class="mw-page-title-main">Basidiomycota</span> Division of fungi

Basidiomycota is one of two large divisions that, together with the Ascomycota, constitute the subkingdom Dikarya within the kingdom Fungi. Members are known as basidiomycetes. More specifically, Basidiomycota includes these groups: agarics, puffballs, stinkhorns, bracket fungi, other polypores, jelly fungi, boletes, chanterelles, earth stars, smuts, bunts, rusts, mirror yeasts, and Cryptococcus, the human pathogenic yeast.

<span class="mw-page-title-main">Smut (fungus)</span> Reproductive structure of fungi

The smuts are multicellular fungi characterized by their large numbers of teliospores. The smuts get their name from a Germanic word for 'dirt' because of their dark, thick-walled, and dust-like teliospores. They are mostly Ustilaginomycetes and comprise seven of the 15 orders of the subphylum. Most described smuts belong to two orders, Ustilaginales and Tilletiales. The smuts are normally grouped with the other basidiomycetes because of their commonalities concerning sexual reproduction.

<span class="mw-page-title-main">Ustilaginomycetes</span> Class of fungi

Ustilaginomycetes is the class of true smut fungi. They are plant parasites with about 1400 recognised species in 70 genera. They have a simple septum with a septal pore cap, this is different from Agaricomycotina which has a dolipore septum with parenthoesome. The group is monophyletic.

<span class="mw-page-title-main">Ustilaginales</span> Order of fungi

The Ustilaginales are an order of fungi within the class Ustilaginomycetes. The order contained 8 families, 49 genera, and 851 species in 2008.

<span class="mw-page-title-main">Pucciniomycotina</span> Subdivision of fungi

Pucciniomycotina is a subdivision of fungus within the division Basidiomycota. The subdivision contains 10 classes, 21 orders, and 38 families. Over 8400 species of Pucciniomycotina have been described - more than 8% of all described fungi. The subdivision is considered a sister group to Ustilaginomycotina and Agaricomycotina, which may share the basal lineage of Basidiomycota, although this is uncertain due to low support for placement between the three groups. The group was known as Urediniomycetes until 2006, when it was elevated from a class to a subdivision and named after the largest order in the group, Pucciniales.

<span class="mw-page-title-main">Exobasidiomycetes</span> Class of fungi

The Exobasidiomycetes are a class of fungi sometimes associated with the abnormal outgrowths of plant tissues known as galls. The class includes Exobasidium camelliae Shirai, the camellia leaf gall and Exobasidium vaccinii Erikss, the leaf and flower gall. There are eight orders in the Exobasidiomycetes, including the Ceraceosorales, Doassansiales, Entylomatales, Exobasidiales, Georgefischeriales, Malasseziales, Microstromatales and the Tilletiales. Four of the eight orders include smut fungi. The families Ceraceosoraceae and Malasseziaceae were formally validated in 2009 for the orders Ceraceosorales and Malasseziales, respectively.

The Doassansiales are an order of fungi in the class Exobasidiomycetes. The order consist of three families: the Doassansiaceae, the Melaniellaceae, and the Rhamphosporaceae.

<span class="mw-page-title-main">Entylomatales</span> Order of fungi

The Entylomatales are an order of smut fungi in the class Exobasidiomycetes. A monotypic order, it consists of a single family, the Entylomataceae. Both the family and order were circumscribed in 1997.

<span class="mw-page-title-main">Georgefischeriales</span> Order of fungi

The Georgefischeriales are an order of smut fungi in the class Exobasidiomycetes. The order consists of four families, the Eballistraceae, the Georgefischeriaceae, the Gjaerumiaceae, and the Tilletiariaceae.

<span class="mw-page-title-main">Tilletiales</span> Order of fungi

The Tilletiales are an order of smut fungi in the class Exobasidiomycetes. It is a monotypic order, consisting of a single family, the Tilletiaceae, which contains seven genera. The roughly 150 species in the Tilletiales all infect hosts of the grass family, except for species of Erratomyces, which occur on legumes.

<span class="mw-page-title-main">Entorrhizomycetes</span> Class of fungi

Entorrhizomycetes is the sole class in the phylum Entorrhizomycota, within the Fungi subkingdom Dikarya along with Basidiomycota and Ascomycota. It contains three genera and is a small group of teliosporic root parasites that form galls on plants in the Juncaceae (rush) and Cyperaceae (sedge) families. Prior to 2015 this phylum was placed under the subdivision Ustilaginomycotina. A 2015 study did a "comprehensive five-gene analyses" of Entorrhiza and concluded that the former class Entorrhizomycetes is possibly either a close sister group to the rest of Dikarya or Basidiomycota.

<span class="mw-page-title-main">Agaricostilbomycetes</span> Class of fungi

The Agaricostilbomycetes are a class of fungi in the subdivision Pucciniomycotina of the Basidiomycota. The class consists of a single order, six families, and 15 genera. Its type genus, Agaricostilbum was originally placed in Ascomycota, and later, Agaricomycotina, before being placed in Pucinniomycotina.

<span class="mw-page-title-main">Pucciniomycetes</span> Class of fungi

Pucciniomycetes is a diverse class of fungi in the subphylum Pucciniomycotina of phylum Basidiomycota. The class contains 5 orders, 21 families, 190 genera, and approximately 8,016 species. It has been estimated that this class contains about one third of all teleomorphic basidiomycetes. Pucciniomycetes contains many economically important plant pathogenic fungal rusts; the order Pucciniales is the largest clade in this class, representing approximately 7,000 species.

<span class="mw-page-title-main">Kriegeriales</span> Order of fungi

The Kriegeriales are an order of fungi in the subdivision Pucciniomycotina. Most species are known only from their yeast states and can be found in a variety of habitats, ranging from arctic waters to tropical ferns. Hyphal states produce auricularioid basidia.

Robert Bauer was a German mycologist, specialising in rust (Uredinales) and smut (Ustilaginomycetes) fungi.

<i>Doassansiopsis</i> Genus of fungi

Doassansiopsis is a genus of smut fungi belonging to the monotypic family DoassansiopsidaceaeBegerow, R.Bauer & Oberw., 1998, within the class Ustilaginomycetes and order Urocystidales.

<span class="mw-page-title-main">Urocystidales</span> Order of fungi

The Urocystidales are an order of fungi within the class Ustilaginomycetes. The order contains 6 families and about 400 genera. They are a sister order to Ustilaginales.

<span class="mw-page-title-main">Doassansiaceae</span> Family of fungi

The Doassansiaceae are a family of fungi in the division Basidiomycota and order of Doassansiales. The family contains 11 genera and about 58 species. They have a widespread distribution. Doassansiaceae is also known and classified as a smut fungi.

The Melaniellaceae are a family of fungi in the division Basidiomycota and order of Doassansiales. The family contains 1 genera and 2 species. They have a distribution in south and south-east Asia.

The Spiculogloeomycetes are a class of fungi in the subdivision Pucciniomycotina of the Basidiomycota. The class consists of a single order, the Spiculogloeales, together with an additional, unassigned genus, Meniscomyces. Many species are currently known only from their yeast states. Species in the genus Spiculogloea form hyphal states that produce auricularioid basidia and are parasitic on other fungi.

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