Graphidaceae | |
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Graphis desquamescens | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Fungi |
Division: | Ascomycota |
Class: | Lecanoromycetes |
Order: | Graphidales |
Family: | Graphidaceae Dumort. (1822) |
Type genus | |
Graphis Adans. (1763) | |
Subfamilies | |
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Synonyms [1] [2] [3] | |
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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. [4] Although the family has a cosmopolitan distribution, most Graphidaceae species occur in tropical regions, and typically grow on bark. [5]
Graphidaceae was originally proposed by French botanist Barthélemy Charles Joseph Dumortier in 1822 (as "Graphineae"). Graphis , Opegrapha , and Arthonia were included in the new family. [6]
In 2002, German lichenologist Bettina Staiger revised the Graphidaceae family in a monograph, proposing a new classification of genera that was widely accepted until molecular phylogenetic studies led to a further reorganization of the family. [7]
Two subfamilies are recognized in the Graphidaceae: [5]
Subfamily Redonographoideae, proposed by Lücking and colleagues in 2013, [8] has since been promoted to familial status (as the monogeneric family Redonographaceae). [9]
The family Thelotremataceae was placed in synonymy with Graphidaceae in 2008, after molecular phylogenetic analysis showed that the families formed several lineages within one strongly supported monophyletic lineage. This study also showed that many genera in Graphidaceae and Thelotremataceae were paraphyletic or polyphyletic. [10] In 2018, [11] Kraichak and colleagues, using a "temporal phylogenetic" approach to identify temporal bands for specific taxonomic ranks, proposed removing Fissurina from the Graphidaceae and instead as placing it as the type genus of Fissurinaceae, a family originally proposed by Brendan P. Hodkinson in 2012. [12] They similarly suggested recognizing Diploschistaceae and Thelotremataceae as independent families. This taxonomic proposal was rejected by Robert Lücking in a critical 2019 review of the temporal method for the classification of lichen-forming fungi, using these specific examples to highlight several drawbacks of this approach. He noted "there is substantial loss of information content in this alternative classification at the second most important rank, the level of family, and such a revised classification is practically meaningless. It makes more sense to apply such essentially phylogenetic classifications, with less information content, to infraranks, such as subfamily or tribe". [3]
The photosynthetic partner ( photobiont ) of Graphidaceae fungi is typically green algae from the genus Trentepohlia , or very rarely Trebouxia . The lichen thallus is typically crustose, which means it forms a crust-like structure on the substrate it grows on. The form of the ascomata of Graphidaceae are usually apothecioid (cup-shaped), lirellate (narrow, and slit-like) or perithecioid (round or oval-shaped structure and embedded in the substrate), but can be rarely mazaediate (with a powdery mass of ascospores and paraphyses formed by the disintegration of the asci). They are usually zeorine (with both a thalline exciple and a proper exciple ), but are sometimes lecideine (without a thallus margin) or lecanorine (surrounded by a pale thalline margin). The hamathecium , which is the hyphae and tissue that between the asci, consists of usually unbranched, sometimes capitate , rarely branched and anastomosing paraphyses. In some lineages, the paraphyses may be apically spinulose (covered with small spines or projections at the tips), and there may be lateral priphysoids in some lineages. The asci are annelasceous , which means they have a ring structure protruding into the lumen and with an apical tholus . They are non-amyloid or amyloid in some lineages, such as in the genus Diorygma . The asci are clavate to oblong or fusiform in shape. [5]
The ascospores of Graphidaceae number primarily 8 per ascus but can often be reduced to 2–4 or 1 per ascus. They are transversely septate to muriform (elongated and divided into multiple compartments by transverse and longitudinal septa), usually ellipsoid to oblong, and often have endospore forming distosepta and lens-shaped to rounded lumina. This particular set of features is known as "graphidoid". The spores can be hyaline (colourless) to (dark) brown, and the endospore is often amyloid. Graphidaceae also have pycnidia for producing conidia, which are non-septate, usually oblong, and hyaline. [5]
The vast majority of Graphidaceae species are restricted to the tropics. Most Graphidaceae species are epiphytic (i.e. they grow only on plants). [8] Forty-two species are known from the Galápagos Islands, where they are among the most diverse of the crustose lichens there. [13] Mexico is thought to be a biodiversity hotspot of undiscovered Graphidaceae species, with about 430 species predicted to occur in tropical regions, [14] compared to less than 200 recorded in the entire country. [15]
According to the Catalogue of Life, there are 94 genera and more than 2100 species in Graphidaceae. [4] In terms of number of species, Graphidaceae is the second-largest family of lichen-forming fungi, after the Parmeliaceae (2765 species) and ahead of the Verrucariaceae (943 species). [2] The following list gives the genus name, its taxonomic authority, year of publication, and the number of species:
Fissurina is a genus of lichenized fungi in the family Graphidaceae. It has about 160 species, most of which are found in tropical regions.
Diorygma is a genus of lichen-forming fungi in the family Graphidaceae. The genus was circumscribed by Franz Gerhard Eschweiler in 1824. Species of the genus are widely distributed in tropical and subtropical regions of the world.
Acanthotrema is a genus of lichens in the family Graphidaceae. The genus was circumscribed by German lichenologist Andreas Frisch in 2006, with Acanthotrema brasilianum assigned as the type species. Acanthotrema species are commonly found in rainforests ranging from lowland to montane environments.
Helge Thorsten Lumbsch is a German-born lichenologist living in the United States. His research interests include the phylogeny, taxonomy, and phylogeography of lichen-forming fungi; lichen diversity; lichen chemistry and chemotaxonomy. He is the Associate Curator and Head of Cryptogams and Chair of the Department of Botany at the Field Museum of Natural History.
Astrochapsa is a genus of lichen-forming fungi in the subfamily Graphidoideae of the family Graphidaceae. It has 28 species. The genus was circumscribed by Sittiporn Parnmen, Robert Lücking, and H. Thorsten Lumbsch in 2012, with Astrochapsa astroidea assigned as the type species. It was segregated from the genus Chapsa, from which it differs in having a more frequently densely corticate thallus, an apothecial margin that is mostly recurved, and the almost exclusively subdistoseptate, non-amyloid ascospores.
Wirthiotrema is a genus of lichen-forming fungi in the family Graphidaceae. The genus was circumscribed in 2010 by Eimy Rivas Plata, Klaus Kalb, Andreas Frisch, and H. Thorsten Lumbsch, with Wirthiotrema glaucopallens assigned as the type species. Wirthiotrema contains species that were formerly considered part of the Thelotrema glaucopallens species group. The genus name honours lichenologist Volkmar Wirth, "for his numerous outstanding contributions to lichenology".
Rhabdodiscus is a genus of script lichens in the family Graphidaceae. It has 36 species.
Schizotrema is a genus of lichen-forming fungi in the family Graphidaceae. The genus was circumscribed in 2009 by Armin Mangold and H. Thorsten Lumbsch.
Pseudochapsa is a genus of lichen-forming fungi in the family Graphidaceae. It has 19 species. It was circumscribed in 2012 by Sittiporn Parnmen, Robert Lücking, and Helge Thorsten Lumbsch, with Pseudochapsa dilatata as the type species. Pseudochapsa differs from Chapsa it that its excipulum is typically brown. Additionally, its ascospores are mostly discoseptate and amyloid. The generic name combines the Greek pseudo ("false") with the genus name Chapsa.
Cruentotrema is a genus of corticolous (bark-dwelling) lichens in the family Graphidaceae. It has seven species.
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, which are widely distributed in tropical and subtropical regions of the world.
Clandestinotrema is a genus of lichen-forming fungi in the family Graphidaceae. It has 17 species. They typically inhabit montane and cloud forest at higher elevations in the tropics.
Topeliopsis acutispora is a species of crustose lichen in the family Graphidaceae. It is found in New South Wales and southern Queensland (Australia), where it grows on bryophytes.
Gintarasia is a genus of lichen-forming fungi in the family Graphidaceae. It has seven species, all of which are found in Australia. Gintarasia species are corticolous (bark-dwelling), crustose lichens with a thelotremoid form.
Compositrema is a genus of lichen-forming fungi in the family Graphidaceae. It has four species. The genus was circumscribed in 2012 by lichenologists Eimy Rivas Plata, Robert Lücking, and Helge Thorsten Lumbsch, with C. cerebriforme assigned as the type species. The genus is distinguished by its unique, composite pseudostromatic ascomata, which sets it apart from the otherwise similar genus Stegobolus.
Pseudochapsa isidiifera is a species of corticolous (bark-dwelling), crustose lichen in the family Graphidaceae. Found in Brazil, it was first formally described as a new species in 2009 by the lichenologists Andreas Frisch and Klaus Kalb, as a member of the genus Chapsa. The type specimen was collected by Kalb in 1980 in a rainforest along the Rio Negro, between 100 and 200 km upstream from Manaus. The species epithet isidiifera refers to the presence of isidia on the thallus. Sittiporn Parnmen, Robert Lücking, and Helge Thorsten Lumbsch transferred the taxon to the genus Pseudochapsa in 2012.
Pycnotrema is a small genus of lichen-forming fungi in the family Graphidaceae. Its two species are characterised by their small, rounded apothecial pores.
Glaucotrema is a genus of lichen-forming fungi in the family Graphidaceae. It has five species.
Corticorygma is a monotypic fungal genus in the subfamily Graphidoideae of the family Graphidaceae. It contains a single species, the corticolous (bark-dwelling) crustose lichen Corticorygma stellatum. This script lichen is found in the shaded understory of rainforests in the Brazilian states of Rondônia and Paraná.