Arthoniales | |
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Cryptothecia rubrocincta | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Fungi |
Division: | Ascomycota |
Class: | Arthoniomycetes |
Order: | Arthoniales Henssen ex D. Hawksw. & O.E. Erikss. (1986) |
Families | |
Andreiomycetaceae 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]
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]
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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]
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]
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]
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]
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]
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]
The following genera have been tentatively classified in the Arthoniales but have not been placed with any certainty into a family. [15]
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.
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.
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.
Enterographa is a genus of lichens in the family Roccellaceae.
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.
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.
Syncesia is a genus of lichen-forming fungi in the family Roccellaceae.
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.
Malmideaceae is a family of crustose and corticolous lichens in the order Lecanorales. It contains eight genera and about 70 species.
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.
Roccellographaceae is a family of lichen-forming fungi in the order Arthoniales. It contains three genera: Dimidiographa, Fulvophyton, and Roccellographa.
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.
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.
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.
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