Biology:Graphidaceae

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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.[1] Although the family has a cosmopolitan distribution, most Graphidaceae species occur in tropical regions, and typically grow on bark. These lichens are characterized by their crust-like growth form and typically partner with orange-pigmented Trentepohlia algae. Many species produce distinctive elongated, slit-like fruiting bodies, and some have unusually large ascospores that can begin germinating almost immediately upon release. The family originated approximately 176 million years ago and has diversified steadily since, with major bursts of new species arising alongside the spread of flowering plants and tropical rainforests.

Taxonomy

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.[2]

In 2002, the German lichenologist Bettina Staiger revised the Graphidaceae 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.[3]

Two subfamilies are recognized in the Graphidaceae:[4]

  • Fissurinoideae Rivas Plata, Lücking & Lumbsch (2011)[5]
  • Graphidoideae Rivas Plata, Lücking & Lumbsch (2011)[5]

Subfamily Redonographoideae, proposed by Lücking and colleagues in 2013,[6] has since been promoted to familial status (as the monogeneric family Redonographaceae).[7]

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.[8] In 2018,[9] 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.[10] 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".[11]

Description

Closeup of the lirellae of Acanthothecis abaphoides

The photosynthetic partner ([[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]]) 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 [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] it grows on. The form of the ascomata of Graphidaceae are usually [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] (cup-shaped), [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] (narrow, and slit-like) or [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] (round or oval-shaped structure and embedded in the substrate), but can be rarely [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] (with a powdery mass of ascospores and [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] formed by the disintegration of the asci). They are usually [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] (with both a [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] and a [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]]), but are sometimes [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] (without a thallus margin) or [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] (surrounded by a pale thalline margin). The [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]], which is the hyphae and tissue that between the asci, consists of usually unbranched, sometimes [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]], rarely branched and anastomosing paraphyses. In some lineages, the paraphyses may be apically [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] (covered with small spines or projections at the tips), and there may be lateral [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] in some lineages. The asci are [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]], which means they have a ring structure protruding into the [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] and with an apical [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]]. They are non-amyloid or amyloid in some lineages, such as in the genus Diorygma. The asci are [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] to oblong or [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] in shape.[4]

The ascospores of Graphidaceae are most often produced eight per ascus, but in some lineages the number is frequently reduced to four, two, or even one. They range from simply transversely septate to [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] (divided into many compartments by both transverse and longitudinal septa), and are typically ellipsoid to oblong. In many species the spore wall is built as an inner [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] separated by [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]], giving lens-shaped to rounded lumina; this combination of features is often referred to as "graphidoid". Spores may remain hyaline or become brown with age, and the endospore commonly reacts as amyloid. Graphidaceae also form pycnidia that produce conidia; these asexual spores are usually hyaline, non-septate, and oblong.[4]

Some graphidaceous species have exceptionally large ascospores. In Ocellularia, the spores of O. subpraestans have been reported to reach about 880 × 65 micrometres in size, and can protrude through the apothecial pore and begin germinating almost immediately, with germ tubes produced from multiple cells of the muriform spore. Observations like these have been used to argue that very large, many-celled spores may favour establishment (and survival) even when dispersal is imperfect, including scenarios where spores are damaged or partly consumed during transport, potentially by invertebrates.[12]

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).[6] 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]

Diversification

Analyses of time-calibrated phylogenies indicate that Graphidaceae has diversified steadily over deep time. The family is estimated to have originated about 176 million years ago (Ma), with comparatively low, fairly uniform speciation rates (about 0.06–0.10 lineages per million years) and brief accelerations early in its history and again around about 110 and 65 Ma. Two hyper-diverse clades—Graphideae and Ocellularia sensu stricto—show the highest rates, and the most frequently inferred models include rate shifts near the stems of one or both of these groups. Cohort analyses recover pronounced heterogeneity across the family, resolving five broad diversification regimes rather than a single family-wide pattern. No clear correlation is evident between Graphidaceae speciation rates and long-term global temperature trends over the past ~180 million years.[16]

The observed pattern is consistent with lineage-specific and ecological drivers. Graphidaceae is largely tropical, and major clades tend to occupy different microhabitats—for example, Ocellularia s.str. is characteristic of shaded, protected tropical forest, whereas many Graphideae occur in more exposed sites—suggesting diversification linked to habitat differences rather than uniform climatic forcing. Pulses of diversification coincide with major events in plant and Earth history: Graphideae began diversifying soon after the origin of angiosperms (c. 140 Ma), and Ocellularia s.str. diversified around the Cretaceous–Paleogene boundary (c. 65 Ma), contemporaneous with the expansion of angiosperm-dominated tropical rainforests; a perithecial [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] in Ocellularia is also associated with higher diversification in trait-dependent analyses, consistent with a possible key innovation (pp. 5–6). Because species discovery is incomplete (recent estimates suggest the true species count may exceed 4,000) future taxonomic sampling may refine rate estimates and their timing.[16]

Genera

According to the Catalogue of Life, there are 94 genera and more than 2100 species in Graphidaceae.[1] 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).[17] The following list gives the genus name, its taxonomic authority, year of publication, and the number of species:

Diorygma antillarum
Dyplolabia afzelii
Fissurina insidiosa
Phaeographina obfirmata
Sarcographa glyphiza
Platythecium hypoleptum
Thecaria quassicola

The genus Medusulina, proposed by Johannes Müller Argoviensis in 1894,[63] is not recognised: its type species, Medusulina nitida, belongs in Fissurina, which renders the name unavailable. Other species once placed there have been reassigned to Redonographa or Jocatoa.[15]

References

  1. ↑ 1.0 1.1 "Graphidaceae". Species 2000: Leiden, the Netherlands. https://www.catalogueoflife.org/data/taxon/625CF. 
  2. ↑ Dumortier, B.C.J. (1822) (in fr). Commentationes botanicae. 1822. Tournay: C. Casterman-Dieu. pp. 69, 78. https://www.biodiversitylibrary.org/page/7469806. 
  3. ↑ 3.0 3.1 3.2 Staiger; B. (2002) (in de). Die Flechtenfamilie Graphidaceae: Studien in Richtung einer natürlichen Gliederung. Bibliotheca Lichenologica. 85. Berlin/Stuttgart: J. Cramer. p. 98. ISBN 978-3-443-58064-3. 
  4. ↑ 4.0 4.1 4.2 Jaklitsch, Walter; Baral, Hans-Otto; Lücking, Robert; Lumbsch, H. Thorsten (2016). Syllabus of Plant Families: Adolf Engler's Syllabus der Pflanzenfamilien. 1/2 (13 ed.). Berlin Stuttgart: Gebr. Borntraeger Verlagsbuchhandlung, Borntraeger Science Publishers. p. 141. ISBN 978-3-443-01089-8. OCLC 429208213. 
  5. ↑ 5.0 5.1 5.2 5.3 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. 
  6. ↑ 6.0 6.1 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. PMID 23594913. 
  7. ↑ Wijayawardene, Nalin; Hyde, Kevin; Al-Ani, LKT; Dolatabadi, S; Stadler, Marc; Haelewaters, Danny et al. (2020). "Outline of Fungi and fungus-like taxa". Mycosphere 11: 1060–1456 [1102]. doi:10.5943/mycosphere/11/1/8. 
  8. ↑ Mangold, Armin; Martín, María P.; Lücking, Robert; Lumbsch, H. Thorsten (2008). "Molecular phylogeny suggests synonymy of Thelotremataceae within Graphidaceae (Ascomycota: Ostropales)". Taxon 2 (2): 476–486. doi:10.2307/25066016. 
  9. ↑ 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. 
  10. ↑ Hodkinson, Brendan P. (2012). "An evolving phylogenetically based taxonomy of lichens and allied fungi". Opuscula Philolichenum 4: 4–10. https://sweetgum.nybg.org/images3/479/367/OP11_p2.pdf. 
  11. ↑ 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. doi:10.1080/07352689.2019.1650517. 
  12. ↑ Sweetwood, Garrett; Lücking, Robert; Nelsen, Matthew P.; Aptroot, André (2012). "Ascospore ontogeny and discharge in megalosporous Trypetheliaceae and Graphidaceae (Ascomycota: Dothideomycetes and Lecanoromycetes) suggest phylogenetic relationships and ecological constraints". The Lichenologist 44 (2): 277–296. doi:10.1017/S0024282911000740. 
  13. ↑ 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. 
  14. ↑ Lücking, Robert; Johnston, Mark K.; Aptroot, André; Kraichak, Ekaphan; Lendemer, James C.; Boonpragob, Kansri et al. (2014). "One hundred and seventy-five new species of Graphidaceae: closing the gap or a drop in the bucket?". Phytotaxa 189 (1): 7–38. doi:10.11646/phytotaxa.189.1.4. https://data.fs.usda.gov/research/pubs/iitf/ja_iitf_2014_Lucking001.pdf. 
  15. ↑ 15.0 15.1 15.2 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. 
  16. ↑ 16.0 16.1 Kraichak, Ekaphan; Divakar, Pradeep K.; Crespo, Ana; Leavitt, Steven D.; Nelsen, Matthew P.; Lücking, Robert; Lumbsch, H. Thorsten (2015). "A tale of two hyper-diversities: diversification dynamics of the two largest families of lichenized fungi". Scientific Reports 5 (1). doi:10.1038/srep10028. PMID 25944223. PMC 4421861. https://www.nature.com/articles/srep10028.pdf. 
  17. ↑ 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. 
  18. ↑ Clements, Frederic E. (1909). The Genera of Fungi (1 ed.). Minneapolis: H.W. Wilson Co.. p. 59. https://www.biodiversitylibrary.org/page/2892753. 
  19. ↑ 19.0 19.1 19.2 19.3 Frisch, A. (2006). The lichen family Thelotremataceae in Africa. Bibliotheca Lichenologica. 92. p. 77. ISBN 978-3-443-58071-1. 
  20. ↑ 20.0 20.1 20.2 Cáceres, Marcela; Aptroot, André; Parnmen, Sittiporn; Lücking, Robert (2014). "Remarkable diversity of the lichen family Graphidaceae in the Amazon rain forest of Rondônia, Brazil". Phytotaxa 189 (1): 87–136. doi:10.11646/phytotaxa.189.1.8. https://www.researchgate.net/publication/287460701. 
  21. ↑ Chevallier, F.F. (1824) (in fr). Histoire des Graphidées, accompagné d'un tableau analytique des genres. Ouvrage renfermant des observations anatomiques et physiologiques sur ces végétaux. Paris: Firmin Didot Père et fils. p. 16. https://archive.org/details/bub_gb_79QUAAAAYAAJ/page/n21/mode/2up. 
  22. ↑ Kalb, Klaus (2009). "New taxa and new records of thelotremoid Graphidaceae". Herzogia 22: 17–24. https://blam-bl.de/images/Herzogia_22/Downloads/H22-Kalb.pdf. 
  23. ↑ Kalb, K. (2004). "New or otherwise interesting lichens. II". Bibliotheca Lichenologica 88: 301–329. 
  24. ↑ 24.0 24.1 Medeiros, Ian D.; Kraichak, Ekaphan; Lücking, Robert; Mangold, Armin; Lumbsch, H. Thorsten (2017). "Assembling a taxonomic monograph of tribe Wirthiotremateae (lichenized Ascomycota: Ostropales: Graphidaceae)". Fieldiana Life and Earth Sciences 9: 1–31. doi:10.3158/2158-5520-16.9.1. https://www.biodiversitylibrary.org/bibliography/169194. 
  25. ↑ 25.0 25.1 25.2 25.3 Parnmen, Sittiporn; Lücking, Robert; Lumbsch, H. Thorsten (2012). "Phylogenetic classification at generic level in the absence of distinct phylogenetic patterns of phenotypical variation: a case study in Graphidaceae (Ascomycota)". PLOS ONE 7 (12). doi:10.1371/journal.pone.0051392. PMID 23251515. 
  26. ↑ 26.0 26.1 Mercado-Díaz, Joel A.; Lücking, Robert; Parnmen, Sittiporn (2014). "Two new genera and twelve new species of Graphidaceae from Puerto Rico: a case for higher endemism of lichenized fungi in islands of the Caribbean?". Phytotaxa 189 (1): 186–203. doi:10.11646/phytotaxa.189.1.14. 
  27. ↑ Massalongo, A.B. (1860). "Esame comparativo di alcune genere di licheni" (in it). Atti dell'Istituto Veneto Scienze 5 (3): 247–276. 
  28. ↑ Rivas Plata, Eimy; Lücking, Robert; Lumbsch, H. Thorsten (2012). "Molecular phylogeny and systematics of the Ocellularia clade (Ascomycota: Ostropales: Graphidaceae)". Taxon 61 (6): 1161–1179. doi:10.1002/tax.616001. 
  29. ↑ 29.0 29.1 29.2 Massalongo, A.B. (1860). "Esame comparativo di alcune genere di licheni" (in it). Atti dell'Istituto Veneto Scienze 5: 313–337. 
  30. ↑ Hyde, Kevin D.; Tennakoon, Danushka S.; Jeewon, Rajesh; Bhat, D. Jayarama; Maharachchikumbura, Sajeewa S. N.; Rossi, Walter et al. (2019). "Fungal diversity notes 1036–1150: taxonomic and phylogenetic contributions on genera and species of fungal taxa". Fungal Diversity 96 (1): 1–242. doi:10.1007/s13225-019-00429-2. https://backend.orbit.dtu.dk/ws/files/218608407/Hyde2019_Article_FungalDiversityNotes10361150Ta.pdf. 
  31. ↑ Kalb, Klaus; Archer, Alan W.; Sutjaritturakan, Jutarat; Boonpragob, Kansri (2009). "New or otherwise interesting lichens V". Bibliotheca Lichenologica 99: 225–246. 
  32. ↑ Norman, J.M. (1852). "Conatus praemissus redactionis novae generum nonnullorum Lichenum in organis fructificationes vel sporis fundatae" (in la). Nytt Magazin for Naturvidenskapene [New Magazine for the Natural Sciences] 7: 213–252. 
  33. ↑ Massalongo, A.B. (1854) (in la). Neagenea lichenum. Tip. Ramanzini. pp. 1–10. 
  34. ↑ Fée, A.L.A. (1824) (in fr). Essai sur les cryptogames des écorces exotiques officinales. pp. 35, 110. 
  35. ↑ 35.0 35.1 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. 
  36. ↑ 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. 
  37. ↑ 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. 
  38. ↑ 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. 
  39. ↑ Lücking, R. (2007). "Kalbographa: Monografie einer unerkannten Flechtengattung" (in de). Bibliotheca Lichenologica 96: 185–192. 
  40. ↑ Montagne, J.F.C. (1856) (in la). Sylloge generum specierumque plantarum cryptogamarum. p. 363. https://www.biodiversitylibrary.org/page/4324526. 
  41. ↑ 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. 
  42. ↑ 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. 
  43. ↑ 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. 
  44. ↑ Frisch, A.; Kalb, K. (2006). "A monograph of Thelotremataceae with a complex structure of the columella". Bibliotheca Lichenologica 92: 371–516 [382]. 
  45. ↑ 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. 
  46. ↑ Tibell, L. (1984). "A reappraisal of the taxonomy of Caliciales". Beihefte zur Nova Hedwigia 79: 597–713 [672]. 
  47. ↑ Meyer, G. (1825) (in de). Nebenstudien meiner Beschäftigungen im Gebiete der Pflanzenkunde. p. 327. 
  48. ↑ Lücking, Robert; Chaves, José Luis; Sipman, Harrie J.M.; Umaña, Loengrin; Aptroot, André (2008). "A first assessment of the ticolichen biodiversity inventory in Costa Rica: The genus Graphis, with notes on the genus Hemithecium (Ascomycota: Ostropales: Graphidaceae)". Fieldiana Botany 46: 1–126. doi:10.3158/0015-0746(2008)46[1:afaott2.0.co;2]. https://www.biodiversitylibrary.org/bibliography/169217. 
  49. ↑ Müller, J. (1882). "Lichenologische Beiträge. XV" (in la). Flora (Regensburg) 65 (25): 397–402. https://www.biodiversitylibrary.org/page/67046. 
  50. ↑ Aptroot, André; Diederich, Paul; Sérusiaux, Emmanuel; Sipman, Harrie J.M. (1997). Lichens and lichenicolous fungi from New Guinea. Bibliotheca Lichenologica. 64. Berlin/Stuttgart: J. Cramer. p. 129. ISBN 978-3-443-58043-8. 
  51. ↑ Müller, J. (1887). "Graphideae Féeanae" (in la). Mémoires de la Société de Physique et d'Histoire Naturelle de Genève 29 (8): 1–80 [10]. 
  52. ↑ Fée, A. (1874). "Materiaux pour une flore lichenologique du Bresil, II: Les Graphidées". Bulletin de la Société Botanique de France 21: 21–32. doi:10.1080/00378941.1874.10827705. 
  53. ↑ Clemente y Rubio, S.R. (1807). Ensayo sobre las variedades de la vid comun que vegetan en Andalucia. 1. 
  54. ↑ 54.0 54.1 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. 
  55. ↑ Rivas Plata, E.; Lücking, R.; Lumbsch, T. (2012). "A new classification for the lichen family Graphidaceae (Ascomycota: Lecanoromycetes: Ostropales)". Fungal Diversity 52 (1): 107–121. doi:10.1007/s13225-011-0135-8. https://www.researchgate.net/publication/257799861. 
  56. ↑ Kalb, Klaus (2001). "The lichen genus Topeliopsis in Australia and remarks on Australian Thelotremataceae". Mycotaxon 79: 319–328. http://www.cybertruffle.org.uk/cyberliber/59575/0079/0319.htm. 
  57. ↑ Vainio, Edvard A. (1921). "Lichenes insularum Philippinarum III" (in la). Annales Academiae Scientiarum Fennicae 15 (6): 184. 
  58. ↑ Lücking, Robert; Mangold, Armin; Rivas Plata, Eimy; Parnmen, Sittiporn; Kraichak, Ekaphan; Lumbsch, H. Thorsten (2015). "Morphology-based phylogenetic binning to assess a taxonomic challenge: a case study in Graphidaceae (Ascomycota) requires a new generic name for the widespread Leptotrema wightii". Botanical Journal of the Linnean Society 179 (3): 436–443. doi:10.1111/boj.12327. 
  59. ↑ Mangold, A.; Elix, J.A.; Lumbsch, H.T. (2009). "Thelotremataceae". Flora of Australia. 57. Australian Biological Resources Study/CSIRO Publishing. pp. 653–659. ISBN 978-0-643-09664-6. 
  60. ↑ Trevisan de Saint-Léon, V. (1853). Spighe e Paglie. Scritti Botanici Varj. 1. p. 13. 
  61. ↑ Nylander, W. (1865). "Lecideae quaedam europaeae novae" (in la). Flora (Regensburg) 48: 3–7. https://www.biodiversitylibrary.org/page/56112. 
  62. ↑ Rivas Plata, Eimy; Kalb, Klaus; Frisch, Andreas (2010). "Wirthiotrema: a new genus for the Thelotrema glaucopallens group (Ascomycota: Ostropales: thelotremoid Graphidaceae)". The Lichenologist 42 (2): 197–202. doi:10.1017/s0024282909990454. https://www.researchgate.net/publication/231961543. 
  63. ↑ Müller, J. (1894). "Conspectus systematicus lichenum Novae Zelandiae" (in la). Bulletin de l'Herbier Boissier 2 (1): 93. https://www.biodiversitylibrary.org/page/33739897. 

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