Button cup relatives

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Button cup relatives
Orbilia xanthostigma

Orbilia xanthostigma

Systematics
Sub-kingdom : Dikarya
Department : Ascomycota mushrooms
Subdivision : Real ascent mushrooms (Pezizomycotina)
Class : Orbiliomycetes
Order : Button cup-like
Family : Button cup relatives
Scientific name of the  class
Orbiliomycetes
OE Erikss. & Baral
Scientific name of the  order
Orbiliales
Baral , OE Erikss. , G. Marson & E. Weber
Scientific name of the  family
Orbiliaceae
Nannf.

The button cup relatives (Orbiliaceae) are a family of ashlar mushrooms , which are in their own order of the button cup-like (Orbiliales) and their own class Orbiliomycetes . Some species are nematophagous .

features

The fruiting bodies (main fruit form, teleomorphs ) are small (usually between 0.2 and 2 mm in diameter), waxy apothecia that are translucent or slightly to noticeably pigmented (usually with yellow to red tones, rarely purple-violet or olive-black). They usually grow on wood and bark, also on herbaceous substrates, old mushroom fruit bodies, animal remains or on the ground. The excipulum (housing) consists of round to square or prismatic, translucent cells. At the edge, the apothecium is smooth or, more rarely, covered with hair of very different lengths, which can be glued together to form teeth. The hymenium consists of paraphyses , which often have a club-headed apex (tip). The asci are often 8-spore but also 16, 32, 64 or 128-spore, and their cell wall is nowhere amyloid . They are apical in the dead state (without turgor ) rounded or flattened to saddle-shaped indented, thin-walled or slightly to very thick-walled. Basally they vary from short and thick to long and thin stalked. The ascogenic hyphae are either simply septate or form hooks on the septa . The interascal tissue is sometimes connected by a gel matrix and is often covered on the outside by an amorphous exudate .

The ascospores are always hyaline and almost always not septate . Depending on the species, they are extremely diverse and of very different sizes (from 2 to 30 micrometers long). Shapes are: spherical, ellipsoid, club-shaped, tear-shaped, spindle-shaped, thread-shaped, rarely triangular or heart-shaped. It is not uncommon for the spores to have a more or less thin, tail-like extension at the lower end, which is sometimes swollen at the end. Depending on the species, elongated spores are often slightly to very strongly curved.

Living ascospores have an apical, highly refractive, KOH- soluble cytoplasmic enclosure of very different shape and unknown function. These “spore bodies” (spore bodies, SB) also occur in several species at both ends of the spore. They are a characteristic feature of the Orbiliomycetes and are not known from any other class of Ascomycetes . The spore bodies are formed like vacuoles by an indentation of the cell membrane (plasmalemma) at the end of the spore maturation and remain connected to the cell membrane via a membrane strand. They are filled with a light-refracting substance that turns dark blue when vital staining with brilliant cresyl blue. Shapes are: spherical, teardrop-shaped, thread-shaped, plug-shaped, lens-shaped. In some groups the membrane cord is greatly thinned and elongated. The shape and size of the ascospores and their spore bodies serve as the main feature of the button cup relatives for classification and species separation. In species with only one (apical) spore body, some of the spores in the ascus are almost always oriented the wrong way round.

Another characteristic feature of some species are the KOH-soluble cytoplasmic bodies (“soluble cytoplasmatic bodies”, SCB), which can only be observed vitally and which occur individually or in groups per cell in sterile tissue. These have a jagged shape, but are also often ring, horseshoe or keyhole-shaped, and often pale orange.

Most species form a secondary crop form ( anamorphic ), which is counted among the Hyphomycetes . This forms a hyaline or pale pink to orange mycelium and relatively large (about 10-100 µm long), holoblastic , hyaline, mostly septate conidia of very different shape, curvature and branching (ellipsoid, club-like, thread-like, C-, Y-, H-shaped etc.).

Distribution and way of life

The species occur worldwide. They live in the soil, on and in dead plant material and break down cellulose and lignin as destructive agents . Some representatives also train traps and use them to catch nematodes . A species not yet described is catching rotifers .

About 80 percent of the species grow on dry or at least temporarily drying plant material. In these so-called xerotolerant species, the fruiting bodies endure complete drying out, in many species for up to two to three years. Such species prefer extremely dry areas (bushland, semi-deserts ), where they are protected from competition from less adapted fungi. In Tibet representatives have been found at 3500 m above sea level.

Nematophagous representatives of the family are mostly found in temperate regions in permanently humid areas close to the ground, but are also found in extremely dry areas such as Oman . They were also found in the ground on some Antarctic islands ( Signy Island in the South Orkney Islands , and Galíndez Island in the Argentine Islands in the Wilhelm Archipelago near the Antarctic Peninsula ).

Systematics

Besides the Pezizomycetes, the class is the most basic group of the real ashlar mushrooms . Around 360 species are now known, many of which have not yet been described. A monograph on the Orbiliomycetes by Baral, Weber and Marson is in preparation.

Teleomorphs

Eriksson 2006 names two teleomorphic genres for the class :

  • Hyalorbilia : The ascus tip is hemispherical and thin-walled. The asci are usually short and thickly stalked and always arise from hooks. The excipulum usually consists of prismatic cells. In most species the spores have one or more, mostly spherical, spore bodies at each end. The tips of the paraphyses are not to moderately thickened. The hymenium is often gelatinous. The apothecia are mostly hyaline to yellowish-chlorine in color, less often orange.
  • Orbilia : The ascus tip is hemispherical or saddle-shaped, flattened, and thin to thick-walled. The asci are often long and thinly stalked, usually forked at the base, and do not arise from hooks. The excipulum usually consists of round to square cells. The spores have only one apical spore body of very different shape. The tips of the paraphyses are not to much thickened. The hymenium is not gelatinous. The apothecia are hyaline, pink, orange, red or yellow, rarely black. The species delimitations are often still artificial, sofour anamorphs are already knownfor the teleomorph Orbilia auricolor .
  • Pseudorbilia : In 2007 a new genus Pseudorbilia with the only species Pseudorbilia bipolaris was describedfor the time being: The ascus tip is flattened and thin-walled. The asci arise from hooks. The excipulum consists of square cells. The spores are cylindrical to slightly dumbbell-shaped and have a spore body at each end. The tips of the paraphyses are slightly thickened. The hymenium is not gelatinous. The apothecia are hyaline.

Anamorphic

There are around 12 genera among the anamorphs. Eriksson et al. 2003 names:

  • Anguillospora
  • Arthrobotrys
  • Dactylella
  • Dactylellina
  • Dicranidion
  • Drechslerella
  • Dwayaangam
  • Gamsylella : after Li et al. (2005) the name is a synonym for Dactylellina.
  • Helicoon (membership questionable)
  • Lecophagus
  • Tridentaria
  • Trinacrium

According to more recent work, there are at least two other genera:

  • Brachyphoris
  • Vermispora

Nematophage representatives

Some anamorphic genera are able to catch nematodes (roundworms). They make up the majority of the nematophagous fungi . Your traps are of three basic types: sticky buttons, sticky nets, and catch loops. The trap type has proven to be a taxonomically useful feature, in contrast to the long-used conidia and conidia carrier forms.

System of the nematophagous representatives

According to the studies by Li et al. Only three genera of nematophagous fungi are distinguished in the family, the genus Gamsylella is no longer considered valid. Based on molecular genetic investigations, they group the previously known forms into three genera, which have the following trap types:

  • Drechslerella : The trap is a constricting loop consisting of three cells that suddenly swell when touched by a nematode and a short, strong stalk.
  • Arthrobotrys : The trap is a sessile adhesive button that develops into an adhesive net, or there are only adhesive nets. The following two types have been identified by Li et al. rewritten in its scope:
    • Arthrobotrys arcuata
    • Arthrobotrys gephyropaga
  • Dactylellina : The trap is a sticky sticky button. Some have non-constricting loops or sessile adhesive buttons that grow into adhesive hyphae and rings. The following species were identified by Li et al. rewritten in its scope:
    • Dactylellina candidum
    • Dactylellina lobata
    • Dactylellina parvicolle
    • Dactylellina phymatopaga
    • Dactylellina robusta

evolution

The nematophagous representatives arose from non-nematophagous ancestors of the genus Orbilia . They form a monophyletic group within the button cup relatives . However, some non-nematophagous species of the genus Dactylella lie within their kin group, close to representatives with sticky buttons . One possible explanation is that these species have lost the adhesive buttons again. The loss of these structures is also known for nematophagous species in culture. The sticky- button species are also more closely related to the non-nematophagous Dactylella species than to those species with other traps.

Adhesive buttons are considered a more original feature, loops and adhesive nets as forms derived from them. These are much more effective in catching nematodes, since with adhesive buttons the nematodes are only held at one point. In the case of adhesive nets, they get stuck at several points, in the loop they are actively held and strangled by the swelling of the cells. These suspected evolutionary lines are supported on the one hand by the molecular genetic studies on the relationship, on the other hand by the ontogenetic development in species with several trap types. In several species it has been observed how catch nets or loops are formed from adhesive buttons.

The types of traps developed in two lines of evolution: One line led to the catch loops, the other to the adhesive traps. The line to the adhesive nets branched off from the latter early on, while the main line developed into the adhesive buttons. These led through an extension of the stem to the shapes with non-contracting loops.

swell

literature

  • OE Eriksson, H.-O. Baral, RS Currah, K. Hansen, CP Kurtzman, G. Rambold, T. Laessøe: Notes on ascomycete systematics. Myconet Volume 9, 2003, pp. 91-103. (online html) (features)
  • Yan Li et al .: Phylogenetics and evolution of nematode-trapping fungi (Orbiliales) estimated from nuclear and protein coding genes . In: Mycologia . Volume 97 (5), 2005, pp. 1034-1046. (Section Nematophage Representatives)

Individual evidence

  1. a b c d Ying Zhang, Ze-Fen Yu, H.-O. Baral, Min Qiao, Ke-Qin Zhang: Pseudorbilia gen. Nov. (Orbiliaceae) from Yunnan, China . In: Fungal Diversity . Volume 26, 2007, pp. 305-312. (PDF; 712 kB)
  2. ^ GL Barron: Predatory fungi, wood decay, and the carbon cycle . In: Biodiversity . Volume 4, 2003, pp. 3-9.
  3. a b c O.E. Eriksson et al .: Notes on ascomycete systematics. 2003.
  4. Mei-Lee Wu, Yu-Chih Su, Hans-Otto Baral, Shih-Hsiung Liang: Two new species of Hyalorbilia from Taiwan . In: Fungal Diversity Volume 25, 2007, pp. 233–244.
  5. ^ B. Liu, XZ Liu, WY Zhuang, HO Baral: Orbiliaceous fungi from Tibet, China . In: Fungal Diversity . Volume 22, 2006, pp. 107-120.
  6. AE Elshafie et al .: Diversity and trapping efficiency of nematophagous fungi from Oman . In: Phytopathologia Mediterranea . Volume 45, 2006, pp. 266-270 online .
  7. ^ NF Gray, CHE Wyborn, RIL Smith: Nematophagous Fungi from the Maritime Antarctic . In: Oikos . Volume 38, 1982, pp. 194-201.
  8. Joseph W. Spatafora et al .: A five-gene phylogeny of Pezizomycotina . In: Mycologia . Volume 98, 2006, pp. 1018-1028.
  9. ^ OE Eriksson (ed.): Outline of Ascomycota - 2006 In: Myconet . Volume 12, 2006, pp. 1-82. (online html)
  10. MingHe Mo, XiaoWei Huang, Wei Zhou, Ying Huang, Yu E Hao, KeQin Zhang: Arthrobotrys yunnanensis sp. nov., the fourth anamorph of Orbilia auricolor . In: Fungal Diversity . Volume 18, 2005, pp. 107-115.
  11. a b c Yan Li et al .: Phylogenetics and evolution of nematode-trapping fungi (Orbiliales) estimated from nuclear and protein coding genes . In: Mycologia . Volume 97, No. 5, 2005, pp. 1034-1046.
  12. Juan Chen, Ling-Ling Xu, Bin Liu, Xing-Zhong Liu: Taxonomy of Dactylella complex and Vermispora. III. A new genus Brachyphoris and revision of Vermispora . In: Fungal Diversity . Volume 26, 2007, pp. 127-142.
  13. ^ Markus Scholler, Gregor Hagedorn, Annemarthe Rubner: A reevaluation of predatory orbiliaceous fungi. II. A new generic concept . In: Sydowia 51, 1999, pp. 89-113.
  14. ^ Gregor Hagedorn, Markus Scholler: A reevaluation of predatory orbiliaceous fungi. I. Phylogenetic analysis using rDNA sequence data . In: Sydowia . 51, 1999, pp. 27-48.
  15. Ying Yang, Ence Yang, Zhiqiang An, Xingzhong Liu: Evolution of nematode-trapping cells of predatory fungi of the Orbiliaceae based on evidence from rRNA-encoding DNA and multiprotein sequences . In: Proceedings of the National Academy of Sciences . Volume 104, No. 20, 2007, pp. 8379-8384, doi : 10.1073 / pnas.0702770104

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