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]
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]
Distribution and ecology
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]
Genera
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:
^
abcdeRivas 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.
^
abLücking, 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.
^
abLücking, 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.
^
abcdJaklitsch, Walter; Baral, Hans-Otto; Lücking, Robert;
Lumbsch, H. Thorsten (2016). Frey, Wolfgang (ed.). Syllabus of Plant Families: Adolf Engler's Syllabus der Pflanzenfamilien. Vol. 1/2 (13 ed.). Berlin Stuttgart: Gebr. Borntraeger Verlagsbuchhandlung, Borntraeger Science Publishers. p. 141.
ISBN978-3-443-01089-8.
OCLC429208213.
^Dumortier, B.C.J. (1822).
Commentationes botanicae [Botanical Commentaries] (in French). Vol. 1822. Tournay: C. Casterman-Dieu. pp. 69, 78.
^
abcStaiger; 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.
^
abLücking, 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.
^
abMiranda-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.
^Clements, Frederic E. (1909).
The Genera of Fungi (1 ed.). Minneapolis: H.W. Wilson Co. p. 59.
^
abcdFrisch, A. (2006). The lichen family Thelotremataceae in Africa. Bibliotheca Lichenologica. Vol. 92. p. 77.
ISBN978-3-443-58071-1.
^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.
^Massalongo, A.B. (1854). Neagenea lichenum (in Latin). Tip. Ramanzini. pp. 1–10.
^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.
^
abKraichak, 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.
^Rivas Plata, E.; Lücking, R.; Lumbsch, H.T. (2012). "Molecular phylogeny and systematics of the Ocellularia clade (Ascomycota: Ostropales: Graphidaceae)". Taxon. 61 (6): 1161–1179.
doi:
10.1002/TAX.616001.
^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.
^Mangold, A.; Martín, M.P.; Kalb, K.; Lücking, R.; Lumbsch, H.T. (2008). "Molecular data show that Topeliopsis (Ascomycota, Thelotremataceae) is polyphyletic". The Lichenologist. 40 (1): 39–46.
doi:
10.1017/S0024282908007366.
^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.
^Tibell, L. (1984). "A reappraisal of the taxonomy of Caliciales". Beihefte zur Nova Hedwigia. 79: 597–713 [672].
^Meyer, G. (1825). Nebenstudien meiner Beschäftigungen im Gebiete der Pflanzenkunde (in German). p. 327.
^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.
^Müller, J. (1887). "Graphideae Féeanae". Mémoires de la Société de Physique et d'Histoire Naturelle de Genève (in Latin). 29 (8): 1–80 [10].
^Clemente y Rubio, S.R. (1807). Ensayo sobre las variedades de la vid comun que vegetan en Andalucia. Vol. 1.
^
abKantvilas, 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.
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]
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]
Distribution and ecology
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]
Genera
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:
^
abcdeRivas 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.
^
abLücking, 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.
^
abLücking, 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.
^
abcdJaklitsch, Walter; Baral, Hans-Otto; Lücking, Robert;
Lumbsch, H. Thorsten (2016). Frey, Wolfgang (ed.). Syllabus of Plant Families: Adolf Engler's Syllabus der Pflanzenfamilien. Vol. 1/2 (13 ed.). Berlin Stuttgart: Gebr. Borntraeger Verlagsbuchhandlung, Borntraeger Science Publishers. p. 141.
ISBN978-3-443-01089-8.
OCLC429208213.
^Dumortier, B.C.J. (1822).
Commentationes botanicae [Botanical Commentaries] (in French). Vol. 1822. Tournay: C. Casterman-Dieu. pp. 69, 78.
^
abcStaiger; 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.
^
abLücking, 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.
^
abMiranda-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.
^Clements, Frederic E. (1909).
The Genera of Fungi (1 ed.). Minneapolis: H.W. Wilson Co. p. 59.
^
abcdFrisch, A. (2006). The lichen family Thelotremataceae in Africa. Bibliotheca Lichenologica. Vol. 92. p. 77.
ISBN978-3-443-58071-1.
^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.
^Massalongo, A.B. (1854). Neagenea lichenum (in Latin). Tip. Ramanzini. pp. 1–10.
^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.
^
abKraichak, 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.
^Rivas Plata, E.; Lücking, R.; Lumbsch, H.T. (2012). "Molecular phylogeny and systematics of the Ocellularia clade (Ascomycota: Ostropales: Graphidaceae)". Taxon. 61 (6): 1161–1179.
doi:
10.1002/TAX.616001.
^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.
^Mangold, A.; Martín, M.P.; Kalb, K.; Lücking, R.; Lumbsch, H.T. (2008). "Molecular data show that Topeliopsis (Ascomycota, Thelotremataceae) is polyphyletic". The Lichenologist. 40 (1): 39–46.
doi:
10.1017/S0024282908007366.
^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.
^Tibell, L. (1984). "A reappraisal of the taxonomy of Caliciales". Beihefte zur Nova Hedwigia. 79: 597–713 [672].
^Meyer, G. (1825). Nebenstudien meiner Beschäftigungen im Gebiete der Pflanzenkunde (in German). p. 327.
^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.
^Müller, J. (1887). "Graphideae Féeanae". Mémoires de la Société de Physique et d'Histoire Naturelle de Genève (in Latin). 29 (8): 1–80 [10].
^Clemente y Rubio, S.R. (1807). Ensayo sobre las variedades de la vid comun que vegetan en Andalucia. Vol. 1.
^
abKantvilas, 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.