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]
Synonymy
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]
Closeup of the lirellae of Acanthothecis abaphoides
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:
Rivas Plata, Eimy; Lücking, Robert; Lumbsch, H. Thorsten (2011). "A new classification for the family Graphidaceae (Ascomycota: Lecanoromycetes: Ostropales)". Fungal Diversity. 52 (1): 107–121. doi:10.1007/s13225-011-0135-8.
Lücking, Robert; Hodkinson, Brendan P.; Leavitt, Steven D. (2017). "The 2016 classification of lichenized fungi in the Ascomycota and Basidiomycota–Approaching one thousand genera". The Bryologist. 119 (4): 361–416. doi:10.1639/0007-2745-119.4.361. JSTOR44250015.
Lücking, Robert (2019). "Stop the abuse of time! Strict temporal banding is not the future of rank-based classifications in fungi (including lichens) and other organisms". Critical Reviews in Plant Sciences. 38 (3): 199–253. Bibcode:2019CRvPS..38..199L. doi:10.1080/07352689.2019.1650517.
Staiger; B. (2002). Die Flechtenfamilie Graphidaceae: Studien in Richtung einer natürlichen Gliederung[The lichen family Graphidaceae: studies towards a natural organization]. Bibliotheca Lichenologica (in German). Vol.85. Berlin/Stuttgart: J. Cramer. p.98. ISBN978-3-443-58064-3.
Lücking, Robert; Tehler, Anders; Bungartz, Frank; Rivas Plata, Eimy; Lumbsch, H. Thorsten (2013). "Journey from the west: did tropical Graphidaceae (lichenized Ascomycota: Ostropales) evolve from a saxicolous ancestor along the American Pacific Coast?". American Journal of Botany. 100 (5): 844–856. doi:10.3732/ajb.1200548. PMID23594913.
Kraichak, Ekaphan; Huang, Jen-Pan; Nelsen, Matthew; Leavitt, Steven D.; Lumbsch, H. Thorsten (2018). "A revised classification of orders and families in the two major subclasses of Lecanoromycetes (Ascomycota) based on a temporal approach". Botanical Journal of the Linnean Society. 188 (3): 233–249. doi:10.1093/botlinnean/boy060.
Bungartz, Frank; Lücking, Robert; Aptroot, André (2010). "The family Graphidaceae (Ostropales, Lecanoromycetes) in the Galapagos Islands". Nova Hedwigia. 90 (1–2): 1–44. doi:10.1127/0029-5035/2010/0090-0001.
Miranda-González, Ricardo; Lücking, Robert; Barcenas-Peña, Alejandrina; Ángeles Herrera-Campos, María de los (2020). "The new genus Jocatoa (Lecanoromycetes: Graphidaceae) and new insights into subfamily Redonographoideae". The Bryologist. 123 (2): 127–143. doi:10.1639/0007-2745-123.2.127.
Kalb, Klaus; Archer, Alan W.; Sutjaritturakan, Jutarat; Boonpragob, Kansri (2009). "New or otherwise interesting lichens V". Bibliotheca Lichenologica. 99: 225–246.
Norman, J.M. (1852). "Conatus praemissus redactionis novae generum nonnullorum Lichenum in organis fructificationes vel sporis fundatae" [Preliminary attempt at a new arrangement of certain genera of lichens based on fruiting bodies or spores]. Nytt Magazin for Naturvidenskapene [New Magazine for the Natural Sciences] (in Latin). 7: 213–252.
Fée, A.L.A. (1824). Essai sur les cryptogames des écorces exotiques officinales[Essay on cryptogams of exotic medicinal barks] (in French). pp.35, 110.
Kraichak, Ekaphan; Parnmen, Sittiporn; Lücking, Robert; Lumbsch, H. Thorsten (2013). "Gintarasia and Xalocoa, two new genera to accommodate temperate to subtropical species in the predominantly tropical Graphidaceae (Ostropales, Ascomycota)". Australian Systematic Botany. 26 (6): 466. doi:10.1071/sb13038.
Lücking, Robert; Rivas Plata, Eimy; Kalb, Klaus; Common, Ralph S.; Barcenas Peña, Alejandrina; Duya, Melizar V. (2011). "Halegrapha (Ascomycota: Graphidaceae), an enigmatic new genus of tropical lichenized fungi dedicated to Mason E. Hale Jr". The Lichenologist. 43 (4): 331–343. doi:10.1017/s0024282911000302.
Nelsen, Matthew P.; Lücking, Robert; Plata, Eimy Rivas; Mbatchou, Joelle S. (2010). "Heiomasia, a new genus in the lichen-forming family Graphidaceae (Ascomycota: Lecanoromycetes: Ostropales) with disjunct distribution in Southeastern North America and Southeast Asia". The Bryologist. 113 (4): 742–751. doi:10.1639/0007-2745-113.4.742.
Lücking, R. (2007). "Kalbographa: Monografie einer unerkannten Flechtengattung" [Kalbographa: Monograph of an unrecognised lichen lenus]. Bibliotheca Lichenologica (in German). 96: 185–192.
Cáceres, Marcela Eugenia da Silva; Rivas Plata, Eimy; Lücking, Robert (2012). "Malmographina, a new genus for Graphina malmei (Ascomycota: Ostropales: Graphidaceae)". The Lichenologist. 44 (1): 115–120. doi:10.1017/S0024282911000697.
Lücking, R.; Sittiporn Parnmen, H.; Lumbsch, H. Thorsten (2012). "Mangoldia, a new lichen genus in the family Graphidaceae (Ascomycota: Ostropales)". Phytotaxa. 69. doi:10.11646/PHYTOTAXA.69.1.1.
Frisch, A.; Kalb, K. (2006). "A monograph of Thelotremataceae with a complex structure of the columella". Bibliotheca Lichenologica. 92: 371–516 [382].
Parnmen, Sittiporn; Cáceres, Marcela E.S.; Lücking, Robert; Lumbsch, H. Thorsten (2013). "Myriochapsa and Nitidochapsa, two new genera in Graphidaceae (Ascomycota: Ostropales) for chroodiscoid species in the Ocellularia clade". The Bryologist. 116 (2): 127–133. doi:10.1639/0007-2745-116.2.127.
Aptroot, André; Diederich, Paul; Sérusiaux, Emmanuel; Sipman, Harrie J.M. (1997). Lichens and lichenicolous fungi from New Guinea. Bibliotheca Lichenologica. Vol.64. Berlin/Stuttgart: J. Cramer. p.129. ISBN978-3-443-58043-8.
Kantvilas, Gintaras; Vězda, Antonín (2000). "Studies on the lichen family Thelotremataceae in Tasmania. The genus Chroodiscus and its relatives". The Lichenologist. 32 (4): 325–357. doi:10.1006/lich.2000.0274.
Vainio, Edvard A. (1921). "Lichenes insularum Philippinarum III" [Lichens of the Philippine Islands III]. Annales Academiae Scientiarum Fennicae (in Latin). 15 (6): 184.
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