Entorrhiza ist eine systematisch isoliert stehende Gattung der Ständerpilze (Basidiomycota), die eine eigene Abteilung Entorrhizomycota bildet.[1]
Entorrhiza sind Pflanzenparasiten. Sie entwickeln sich in den Wurzeln von Cyperaceae und Juncaceae und führen hier zu Gallbildung. Sie bilden in den Zellen Knäuel von septierten Hyphen. Ihre Teliosporen bilden sie innerhalb der lebenden Wirtszelle am Ende der Hyphen. Die Teliosporen teilen sich in vier Zellen mit kreuzförmig stehenden Septen und bilden so eine Phragmobasidie. Die Keimung der Teliosporen erfolgt durch die Bildung von vier Keimschläuchen.
Die Septalporen sind nicht von Membrankappen umgeben, bilden jedoch einen Doliporus.
Die Entorrhizomycetes wurden einige Zeit zu den Ustilaginomycotina gestellt, von Hibbett et al. 2007 aber als incertae sedis ohne Zugehörigkeit zu einer Unterabteilung direkt den Basidiomycota zugeordnet.[2] Sie bestehen aus einer einzigen Gattung.[3] Robert Bauer und Kollegen stellten schließlich die eigene Abteilung Entorrhizomycota auf, basierend auf molekularbiologischen Untersuchungen und auch der einzigartigen Merkmalen hinsichtlich der Ultrastruktur und Reproduktion.[1] Dabei werden die Entorrhizomycota entweder als Schwestergruppe der Dikarya oder der Ständerpilze angesehen.[1]
Der Index Fungorum listet folgende Arten:[4]
Entorrhiza ist eine systematisch isoliert stehende Gattung der Ständerpilze (Basidiomycota), die eine eigene Abteilung Entorrhizomycota bildet.
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[2] 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.
Taxonomy based on the work of Wijayawardene et al. 2019.[4]
All members of Entorrhizomycetes are obligate parasites on the roots of plants.[7] Sori are produced as galls on the roots of hosts. Galls are tubercular with a globoid, irregular or elongated shape and are composed of vascular bundles, parenchymatous cells and fungal mycelium.[6] Younger segments of the galls are pale in color whilst older segments turn brown.[2] Mycelium consists of dikaryotic and septate hyphae with fibrillate walls that lack clamp connections. Initially, the mycelium grows intercellularily before producing coiled intracellular hyphae terminating in globose cells that detach and develop into teliospores.[2] Teliospores germinate into tetrads through internal septation, and each tetrad compartment produce hyphae that terminate in sigmoid propagules.[2] Bauer et al. noted that young teliospores have two nuclei, older teliospores have only one nucleus, and each tetrad compartment has one nucelus each. This indicates that karyogamy and meiosis occurs in the teliospore.[2] It has been observed that teliospores are liberated when the host plant dies and the galls disintegrate,[6] and that the number of galls is higher in waterlogged soils compared to well-drained soils.[8] These observations might support the hypothesis that entorrhizomycetes disperse through soil moisture.[2]
Both Talbotiomyces and Juncorrhiza are segregate taxa from Entorrhiza sensu lato.[6][7] Entorrhiza sensu stricto is diagnosed by teliospores with longitudinally ridged or cerebriform ornamentation and infecting plants belonging to Cyperaceae, whilst Juncorrhiza is diagnosed by teliospores with verrucose-tuberculate ornamentation and infecting plants belonging to Juncaceae.[7] Talbotiomyces is distinguished from species in Entorrhizales by hyphal septa with simple pores that lack caps or membranes (species in Entorrhizales have dolipores that lack caps or membranes) and infecting plants belonging to Caryophyllales.[6][7]
Molecular phylogeny place Entorrhizomycetes as either a sister group to Basidiomycota or a sister group to Dikarya as a whole. Entorrhizomycetes share many traits with basidiomycetes such as dikaryotic vegetative mycelium, fibrillate cell walls, hyphal septa with a tripartite profile, and similarities in the spindle pole body.[2] Bauer et al. speculated that the teliospore tetrad in entorrhizomycetes might represent the ancestral state of dikaryan meiosporangia. This is based on the observation that the septa in the tetrads have pores, and that the tetrad compartments germinate into hyphae terminating in propagules. The basidial cells separated by pored septa in basidiomycete phragmobasidia represent meiospores that in turn release vegetative propagules (that are usually characterised as basidiospores).[2] It is possible that an ancestral structure similar to the teliospore tetrad evolved into phragmobasidia which in turn evolved into holobasidia on multiple occasions during the transition from water-dispersal to air-dispersal. If entorrhizomycetes are sister to Dikarya, it is also possible that the teliospore tetrad is homologous to the meiospore tetrads of early-diverging ascomycetes.[2]
The stem age of the Entorrhizomycota has been estimated to approximately 560 Mya during the late Neoproterozoic era. Divergence between Talbotiomycetales and Entorrhizales is estimated to approximately 50 Mya, and divergence between Entorrhiza and Juncorrhiza is estimated to approximately 42 Mya. Both Entorrhiza and Juncorrhiza underwent a major radiation during the Oligocene and Miocene epochs. Given that these divergence estimates are incongruent or only slightly congruent with the estimated stem ages of the host plant lineages, and incongruence in the co-phylogeny between Entorrhizales and host plants, host-shift speciation is more likely to have occurred than co-speciation during these divergences and the radiation of Entorrhizales.[7]
Entorrhizomycetes have much lower number of species and more limited host range than their estimated age would indicate. One possible explanation is that many lineages have gone extinct along with their hosts during mass extinction events in the past. Another explanation is that much of the diversity in this phylum remains undiscovered.[2] The latter explanation is supported by the fact that host plants don't show any aboveground symptoms of infection,[7] and there might be species that don't cause galls on their hosts.[2]
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.
Entorrhizomycetes Begerow et al. – klasa podstawczaków (Basidiomycota). Typem nomenklatorycznym jest Entorrhiza[1]. W tzw. "systemie Adla 2012" klad Entorrhizales wyprowadzony jest bezpośrednio z kladu Basidiomycota[2].
Do klasy Entorrhizomycetes należą grzyby fitopasożytnicze rozwijające się wewnątrz komórek żywiciela[3].
Rodzina Entorrhizaceae i rząd Entorrhizales zostały utworzone przez Roberta Bauera i Franza Oberwinklera w artykule Ultrastructural markers and systematics in smut fungi and allied taxa opublikowanym w „Canadian Journal of Botany” z 1997. Klasę Entorrhizomycetes po raz pierwszy wyodrębnili Dominik Begerow, Matthias Stoll i Robert Bauer w artykule A phylogenetic hypothesis of Ustilaginomycotina based on multiple gene analyses and morphological data opublikowanym w „Mycologia” z 2006:
Entorrhizomycetes Begerow, Stoll & R. Bauer class. nov. (= Entorrhizomycetidae R. Bauer & Oberw., in Bauer et al 1997)
Fungi phytoparasitici hyphis glomeratis septatis intracellularibus teliosporas terminales procreantibus.
Phytoparasitic fungi forming intracellular septate hyphal coils with terminal teliospores.Klasa ta jest zaliczana według kodeksu Index Fungorum do typu Basidiomycota. Należą do niej[4]:
Entorrhizomycetes Begerow et al. – klasa podstawczaków (Basidiomycota). Typem nomenklatorycznym jest Entorrhiza. W tzw. "systemie Adla 2012" klad Entorrhizales wyprowadzony jest bezpośrednio z kladu Basidiomycota.
Entorrhizomycetes é uma classe de fungos do filo Basidiomycota. Contém uma única ordem Entorrhizales, que por sua vez contém uma única família Entorhizaceae, um pequeno grupo de parasitas de raízes teliospóricos que formam galhas em plantas das famílias Juncaceae e Cyperaceae.
Entorrhizomycetes é uma classe de fungos do filo Basidiomycota. Contém uma única ordem Entorrhizales, que por sua vez contém uma única família Entorhizaceae, um pequeno grupo de parasitas de raízes teliospóricos que formam galhas em plantas das famílias Juncaceae e Cyperaceae.
根肿黑粉菌属 Entorrhiza
Talbotiomyces
根肿黑粉菌纲(学名:Entorrhizomycetes)是担子菌门黑粉菌亚门下的一个纲。该纲仅含一个目(根肿黑粉菌目,Entorrhizales),该目下也仅含一个科(根肿黑粉菌科,Entorrhizales)。此科中的真菌可寄生于灯心草科及莎草科植物上并形成菌瘿,内生冬孢子。
根肿黑粉菌纲(学名:Entorrhizomycetes)是担子菌门黑粉菌亚门下的一个纲。该纲仅含一个目(根肿黑粉菌目,Entorrhizales),该目下也仅含一个科(根肿黑粉菌科,Entorrhizales)。此科中的真菌可寄生于灯心草科及莎草科植物上并形成菌瘿,内生冬孢子。
エントリザ菌綱 (Entorrhizomycetes)は、担子菌門の綱の一つ。それぞれ1つの目、科からなる。冬胞子の菌で、イグサ科イグサ属とカヤツリグサ科スゲ属植物の根に寄生しコブを作る。
엔토르리자과(Entorrhizaceae)는 쌍핵균류 과의 하나이다. 엔토르리자강(Entorrhizomycetes), 엔토르리자목(Entorrhizomycetes)의 유일한 과이다. 겨울포자인 깜부기포자를 만드는 분류군으로 골풀과와 사초과 식물 위에 충영(蟲癭, 벌레혹, gall)을 형성한다.
다음은 2018년 테더수 등(Tedersoo et al. 2018)의 진균류 계통 분류이다.[3]
진균류 호상균아계블라스토클라디아균아계 / 블라스토클라디아균문
바시디오볼루스균아계 / 바시디오볼루스균문
올피디움균아계 / 올피디움균문
포충균아계 쌍핵균아계