Arthoniales

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Arthoniales
Cryptothecia.jpg
Cryptothecia rubrocincta
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Fungi
Division: Ascomycota
Class: Arthoniomycetes
Order: Arthoniales
Henssen ex D. Hawksw. & O.E. Erikss. (1986)
Families

Andreiomycetaceae
Arthoniaceae
Chrysotrichaceae
Lecanographaceae
Opegraphaceae
Roccellaceae
Roccellographaceae

Contents

The Arthoniales is the second largest order of mainly crustose lichens, but fruticose lichens are present as well. [1] The order contains around 1500 species, [2] while the largest order with lichenized fungi, the Lecanorales, contains more than 14000 species. [3]

Classification

The Arthoniales is one of two orders of the class Arthoniomycetes within the phylum Ascomycota. [2] The order includes seven families (Andreiomycetaceae, [4] Arthoniaceae, Chrysotrichaceae, Lecanographaceae, Opegraphaceae, Roccellaceae and Roccellographaceae). [2] Lecanographaceae, Roccellographaceae, Opegraphaceae and Roccellaceae are well-supported families within Arthoniales, and they were circumscribed in 2011. [5] Andreiomycetaceae was described as a new family by Hodkinson and Lendemer in 2013. [4]

The Arthoniales is the sister group to Dothideomycetes. [6]

Arthoniales

Andreiomycetaceae

Chrysothricaceae

Rocellaceae

Opegraphaceae

Rocellographaceae

Lecanographaceae

Arthoniaceae

Curvularia brachyspora

Cudonia circinans

Seynesia erumpens

Figure 1. Cladogram of the Arthoniales, rooted with Curvularia brachyspora, Cudonia circinans and Seynesia erumpens as the outgroup. The cladogram shows the division of Arthoniales into seven families, based on Hodkinson et al. 2013. [4]

Distribution and habitats

The Arthoniales is distributed in most habitats worldwide, as it ranges at latitudes from arctic to tropical regions. [2] They grow on different types of substrates like bark, wood, rocks, bryophytes and living leaves. The order has adapted to live in both humid forests and dry habitats like savannas and steppes, as well as varying altitudes from sea level to alpine regions. [7] The highest species diversity are known from subtropical coastal areas with a Mediterranean or dry climate. [8] [9] The species diversity of the Arthoniales is expected to be higher than currently reported from several areas worldwide like the tropical rain forests. [2]

Ecology

The majority of species in Arthoniales have a lichenized lifestyle, but lichenicolous and saprophytic species are presented as well. [10] The original state in the Arthoniales is believed to be the lichenized state, and the non-lichenized and parasitic states have evolved independently several times. [11] The lichenized fungi live in symbiosis with a photobiont that in most cases is a species of Trentepohliaceae, but photobionts from Chlorococcaleae are known from Chrysotrichaceae and a few species in Arthonia . [12]

Characters

It is difficult to mention typical morphological characters like fruit body, exiple, hypothecium, hymenium, chemistry, ascospore color and ascospore septation that can be used as uniting traits for delimiting families and genera in the Arthoniales. The reason is that the Arthoniales is an old order and taxa have evolved in parallel for a long time, which gives a high level of homoplasy. [5]

Ascomatal anatomy

The ascomata are usually apothecial and it produces bitunicate asci. The bitunicate asci are thick-walled, with an outer and inner layer. The outer and inner layer of the ascus wall are called exotunica and endotunica, and they separate during ascospore release. [6] The asci usually contains eight ascospores, while the shape and separation of the ascospores are more variating. [13]

Thallus

The majority of the species in Arthoniales are crustose lichens, with a thallus growing tight to the surface. Fruticose lichens have a bushy thallus and are known in Roccellaceae, where it has evolved and been lost multiple times. [14] The thalli among crustose lichens can either grow within the substrate, called endophloedal or it can grow at the surface of the substrate, called epiphloedal. [13]

Genera incertae sedis

Bactrospora brevispora, shown here growing on a cypress stump, is one of many taxa of uncertain familial placement in the Arthoniales. Bactrospora brevispora - Flickr - pellaea (1).jpg
Bactrospora brevispora , shown here growing on a cypress stump, is one of many taxa of uncertain familial placement in the Arthoniales.

The following genera have been tentatively classified in the Arthoniales but have not been placed with any certainty into a family. [15]

Related Research Articles

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

The Arthoniaceae are a family of lichenized, lichenicolous and saprobic fungi in the order Arthoniales. The Arthoniaceae is the largest family of Arthoniales, with around 800 species. Most species in Arthoniaceae belong in Arthonia which is the largest genus with 500 species. The second and third largest genus is Arthothelium with 80 species, and Cryptothecia with 60 species.

<span class="mw-page-title-main">Roccellaceae</span> Family of lichens

The Roccellaceae are a family of fungi in the order Arthoniomycetes. Most taxa are lichenized with green algae, although some are lichenicolous, growing on other lichens.

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

The Graphidaceae are a family of lichen-forming fungi in the order Graphidales. The family contains nearly a hundred genera and more than 2000 species. Although the family has a cosmopolitan distribution, most Graphidaceae species occur in tropical regions, and typically grow on bark.

<i>Enterographa</i> Genus of lichen

Enterographa is a genus of lichens in the family Roccellaceae.

<i>Lecanographa</i> Genus of lichens in the family Lecanographaceae

Lecanographa is a genus of about 40 species of lichens in the family Lecanographaceae. It was circumscribed in 1994 by José M. Egea and Pilar Torrente, with Lecanographa lyncea as the type species.

<i>Lecanactis</i> Genus of lichen

Lecanactis is a genus of crustose lichens, commonly called old wood rimmed lichen. The genus was circumscribed in 1855 by German lichenologist Gustav Wilhelm Körber, who assigned Lecanactis abietina as the type species.

<i>Syncesia</i> Genus of lichens

Syncesia is a genus of lichen-forming fungi in the family Roccellaceae.

<span class="mw-page-title-main">Trypetheliaceae</span> Family of mainly lichen-forming fungi

The Trypetheliaceae are a family of mainly lichen-forming fungi in the order Trypetheliales. The family consists almost exclusively of corticolous (bark-dwelling), crustose lichens with an almost strictly tropical distribution.

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

Malmideaceae is a family of crustose and corticolous lichens in the order Lecanorales. It contains eight genera and about 70 species.

<span class="mw-page-title-main">Opegraphaceae</span> Family of lichen

Opegraphaceae is a family of lichen-forming and lichenicolous fungi in the order Arthoniales. It was originally proposed by German lichenologist Ernst Stizenberger in 1862. It fell into disuse, but was resurrected in a molecular phylogenetic study of the order Arthoniales published in 2010. It now includes taxa that were previously referred to the family Roccellaceae, its sister group.

<span class="mw-page-title-main">Roccellographaceae</span> Family of lichen-forming fungi

Roccellographaceae is a family of lichen-forming fungi in the order Arthoniales. It contains three genera: Dimidiographa, Fulvophyton, and Roccellographa.

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

Rhizocarpales are an order of lichen-forming fungi in the subclass Lecanoromycetidae of the class Lecanoromycetes. It has two families, Rhizocarpaceae and Sporastatiaceae, which contain mostly crustose lichens.

Angiactis is a genus of crustose lichens of uncertain familial placement in the order Arthoniales. It has four species.

Vigneronia is a genus of lichen-forming fungi in the family Roccellaceae. It has five species. The genus was circumscribed in 2014 by Damien Ernst, with Vigneronia spieri assigned as the type species. This species, originally described as Schismatomma spieri from collections made in the Galápagos Islands, has since been recorded from mainland Ecuador and the Antilles (Curaçao). The genus is named after Ernst's wife, Nathalie Vigneron, who accompanied him on collecting trips.

Dirina catalinariae is a species of saxicolous (rock-dwelling), crustose lichen in the family Roccellaceae. It occurs in the southwestern United States, Mexico, and the Galápagos Islands.

Vigneronia spieri is a species of corticolous (bark-dwelling), crustose lichen in the family Roccellaceae. It is found in the Galápagos Islands, mainland Ecuador and the Antilles (Curaçao).

Fouragea is a genus of lichen-forming fungi in the family Opegraphaceae. It has nine species.

<i>Fulvophyton</i> Genus of lichens

Fulvophyton is a genus of lichen-forming fungi in the family Roccellographaceae. It has 11 species. Fulvophyton is characterised by its crust-like thallus, which is often pale yellowish-brown in colour. This genus features a photobiont from the green algal genus Trentepohlia and exhibits a unique arrangement of reproductive structures.

Neosergipea is a genus of lichen-forming fungi in the family Roccellaceae. It has four species, all of which are corticolous (bark-dwelling), crustose lichens that are found in the tropical forests of Brazil. This genus is related to the genera Dichosporidium, Enterographa, and Erythrodecton. It distinguishes itself through unique morphological characteristics and certain chemical constituents, in particular, its non-carbonised ascomata and the presence of a vivid orange anthraquinone compound.

<i>Coniocarpon</i> Genus of lichens

Coniocarpon is a genus of lichen-forming fungi in the family Arthoniaceae. It has eight species of corticolous (bark-dwelling) lichens. This genus is distinct for its crystalline orange, red, and purple quinoid pigments in the ascomata that turn purple in potassium hydroxide solution, its colourless, transversely septate ascospores with large apical cells, and its rounded to lirellate ascomata.

References

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  2. 1 2 3 4 5 Frisch A, Thor G, Ertz D, Grube M (August 2014). "The Arthonialean challenge: Restructuring Arthoniaceae". Taxon. 63 (4): 727–744. doi:10.12705/634.20.
  3. Ekman, Stefan; Andersen, Heidi L.; Wedin, Mats (2008-02-01). "The Limitations of Ancestral State Reconstruction and the Evolution of the Ascus in the Lecanorales (Lichenized Ascomycota)". Systematic Biology. 57 (1): 141–156. doi: 10.1080/10635150801910451 . ISSN   1063-5157. PMID   18300027.
  4. 1 2 3 Hodkinson, BP (2013). "Next-generation sequencing reveals sterile crustose lichen phylogeny". Mycosphere. 4 (6): 1028–1039. doi: 10.5943/mycosphere/4/6/1 .
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  9. Aptroot, André; Sparrius, Laurens B. (2008-12-01). "Crustose Roccellaceae in The Galapagos Islands, With The New Species Schismatomma spierii". The Bryologist. 111 (4): 659–666. doi:10.1639/0007-2745-111.4.659. ISSN   0007-2745. S2CID   84038943.
  10. Grube, Martin (1998). "Classification and Phylogeny in the Arthoniales (Lichenized Ascomycetes)". The Bryologist. 101 (3): 377–391. doi:10.2307/3244176. JSTOR   3244176.
  11. Nelsen MP, Lücking R, Grube M, Mbatchou JS, Muggia L, Plata ER, Lumbsch HT (2009). "Unravelling the phylogenetic relationships of lichenised fungi in Dothideomyceta". Studies in Mycology. 64: 135–144S4. doi:10.3114/sim.2009.64.07. PMC   2816970 . PMID   20169027.
  12. Smith CW, Aptroot A, Coppins BJ, eds. (2009). The Lichens of Great Britain and Ireland (2nd ed.). London: The Natural History Museum. ISBN   978-0-9540418-8-5.
  13. 1 2 Smith, C. W (2009). The lichens of Great Britain and Ireland. British Lichen society London.
  14. Theler, Irestedt, Anders, Martin (2007). "Parallel evolution of lichen growth forms in the family Roccellaceae (Arthoniales, Ascomycota)". Cladistics. 23 (5): 432–454. doi: 10.1111/j.1096-0031.2007.00156.x . S2CID   84702559.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  15. Wijayawardene, Nalin; Hyde, Kevin; Al-Ani, Laith Khalil Tawfeeq; Somayeh, Dolatabadi; Stadler, Marc; Haelewaters, Danny; et al. (2020). "Outline of Fungi and fungus-like taxa". Mycosphere. 11: 1060–1456. doi: 10.5943/mycosphere/11/1/8 . hdl: 10481/61998 .