Graphidales
A–B Pallidogramme chrysenteron, formerly Phaeographina fukiensis. C–D Sarcographa glyphiza, formerly Graphis glyphiza. Scale bars = 1 mm
Scientific classification
Domain:	Eukaryota
Kingdom:	Fungi
Division:	Ascomycota
Class:	Lecanoromycetes
Order:	Graphidales Hazlinszky, F.A. (1884)[1]
Families
Diploschistaceae Fissurinaceae Gomphillaceae Graphidaceae Redonographaceae Thelotremataceae

Graphidales is an order of lichen-forming fungi in the class Lecanoromycetes. It contains 6 families, about 81 genera and about 2,228 species. Family Graphidaceae are the largest crustose family within Graphidales order comprising more than 2000 species,^[2] which are widely distributed in tropical and subtropical regions of the world.^[3]

History

The Graphidales were introduced in a 1884 publication by Frigyes Ákos Hazslinszky in Magyar Birodalom Zuzmó-Flórája on page 216 as family Graphideae.^[1] In 1907, they were established as an order by American botanist Bessey (1845–1915),^[4][5][6]

When the order was introduced, it contained just two families, the Graphidaceae and Thelotremataceae who were both mainly tropical based and each family had about 800–1000 species.^[7]

Sherwood in 1977 proposed to maintain a distinction between the Graphidales with mostly lichemised members and the Ostropales which included mostly non-lichenised fungi, based on different spore septation types.^[8]

Molecular data by Winka et al. in 1998, supported a close relationship between the two groups of species.^[9][10]

The Graphidales were then included in the Ostropales order (Lecanoromycetes) for a long time (Staiger 2002;^[11] Kalb et al. 2004;^[12] Hibbett et al. 2007;^[13] Lumbsch et al. 2007;^[14] Kirk et al. 2008;^[15] Baloch et al. 2010;^[16] Rivas Plata and Lumbsch 2011;^[17] Rivas Plata et al. 2012;^[18] Lumbsch et al. 2014;^[19] Lücking et al. 2017;^[20] Wijayawardene et al. 2018).^[21]

In 2004, the phylogenetic relationships of class Lecanoromycetes were examined by using mitochondrial small subunit ribosomal DNA sequencing which found that orders Graphidales and Ostropales were monophyletic.^[22]

Using molecular data (partial DNA sequencing) in 2012, it was also shown that Graphidaceae and Thelotremataceae were non-monophyletic and consequently Thelotremataceae was included in Graphidaceae as a synonym.^[23] Graphidaceae also included subfamilies Fissurinoideae and Graphidoideae.^[24][25]

However, Kraichak et al. in 2018,^[26] ranked Graphidales as a separate order based on a temporal approach, and accepted five families; Diploschistaceae, Fissurinaceae, Gomphillaceae, Graphidaceae and Thelotremataceae.^[27][28] Wijayawardene et al. 2020 agreed but also added family Redonographaceae to the order,^[2] Other authors have agreed on the use of reinstated order of Graphidales.^{[3][29][30][31]}

Description

Most species in the order are lichens which have a thallus (vegetative tissue) which is crustose and ascocarps (fruiting body) which are apothecioid (cup-shaped).^[32][33]

The Graphidaceae are mostly epiphytic lichens with trentepohlioid photobiont (i.e., filamentous, multicellular green algae from genus Trentepohlia),^[34] and graphidoid, distoseptate (forming a layer) ascospores.^[35][36]

Family Gomphillaceae was originally based on a single species, Gomphillus calycioides (Watson, 1929), which is an unusual taxon growing over bryophytes.^[37]

It includes a common asexual fungus Lawreya glyphidiphila (Teratosphaeriaceae family) which is described as growing on lichenized fungi Glyphis scyphulifera (Graphidaceae family).^[29]

Distribution

They are mainly found in warmer regions living on bark.^[32] Genera in the order of Graphidales has been found worldwide,^[38] from North America (including Florida,^[29][39]), South America (including Venezuela,^[40] Costa Rica,^[41] and Guianas,^[42]), Africa (including Kenya,^[43] and South Africa,^[44]), Asia (including China,^[31][45] Vietnam,^[46] Sri Lanka,^[47] India,^[3][48] and Thailand,^[49]) Australia,^[50][51] and also New Zealand.^[5][52]

Species of family Gomphillaceae are found in north-eastern Brazil, Mexico, Guatemala, Costa Rica, Panama and Cuba.^[37]

Families and genera

This is a list of the families and genera contained within the Graphidales, based on a 2020 review and summary of ascomycete classification.^[2] Following the taxon name is the taxonomic authority, year of publication, and (for genera) the number or estimated number of species:

References