Trypetheliaceae

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Trypetheliaceae
Bathelium carolinianum - Flickr - pellaea (1).jpg
Bathelium carolinianum
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Fungi
Division: Ascomycota
Class: Dothideomycetes
Order: Trypetheliales
Family: Trypetheliaceae
Eschw. (1824)
Type genus
Trypethelium
Spreng. (1804)
Synonyms [1] [2]

The Trypetheliaceae are a family of mainly lichen-forming fungi in the order Trypetheliales. [3] The family consists almost exclusively of corticolous (bark-dwelling), crustose lichens with an almost strictly tropical distribution. [4]

Contents

Taxonomy

Trypetheliaceae was circumscribed by German botanist Franz Gerhard Eschweiler in 1824. His diagnosis of the family was as follows (translated from Latin):"Thallus is crust-like. Apothecia are of varied shape, immersed in warts formed from the thallus's medullary substance." Eschweiler further notes that the family is almost exclusively found within the tropics. In his initial circumscription, he included the following genera: Arthonia , Porothelium , Medusula , Ophthalmidium , Trypethelium , Astrothelium , Glyphis , Chiodecton , and Conioloma . [5]

The family Arthopyreniaceae was placed into synonymy with Trypetheliaceae, and its type genus, Arthopyrenia , was transferred to Trypetheliaceae in 2021. [2] Other synonyms of Trypetheliaceae include Astrotheliaceae, Cryptotheliaceae, Hyalophragmiaceae, and Laureraceae. [1] The order Trypetheliales was proposed in 2008 to contain the family. [6] [7]

In their 2016 revisionary synopsis of the Trypetheliaceae, André Aptroot and Robert Lücking accepted 418 species, distributed amongst 15 genera. [4]

Description

The general features of the Trypetheliaceae include a thallus that is mostly crustose, though the growth form of some species are squamulose to foliose or even fruticose . The family displays a diverse array of appearances, ranging from clearly visible and areolate to evanescent , hidden under bark, or even barely discernible from the substrate . The growth habit and development of the thallus surface are influenced by a combination of environmental conditions and evolutionary factors. [4]

The internal anatomy of the ascomata, including the pseudostromata , in this family is still an emerging area of research with potential taxonomic and systematic implications. One noteworthy aspect is that the ascomata in Trypetheliaceae can develop either superficially or emerge from beneath the periderm, with some species even exhibiting both forms. This divergence is seen in species such as Architrypethelium grande , Astrothelium megaspermum , and others. The wall of the ascoma is typically multi-layered and may contain a medullary layer. In some species, like Astrothelium megaspermum, an additional brownish 'medullary' layer is present. The intricate anatomy of the ascoma provides an array of characters that are used in the development of genus and species concepts within Trypetheliaceae. [4]

Another consistent feature in most Trypetheliaceae is the hamathecium , comprising thin, branching paraphyses that form a network within a gelatinous matrix. However, the density and presence of the gelatinous matrix can vary across lineages. One feature, the hymenial inspersion, consists of oil droplets or infusions lining the paraphyses. The inspersion can be localized or spread throughout the hamathecium, and its taxonomic importance is still being uncovered. [4]

The asci of Trypetheliaceae are typically fissitunicate , a structure best observed in species with larger ascospores . Additionally, ascospore type is a significant character in this family. While septation has historically been a key differentiator, recent understandings emphasize the nature of the septa and walls as more critical taxonomic indicators. The ascospores themselves can vary considerably, from the typical astrothelioid type to those that are multiseptate or muriform , and this variation offers important insights into the relationships within the family. [4]

Chemistry

The Trypetheliaceae family's chemical profile is less intricate than some tropical crustose families like Graphidaceae. The family's secondary substances primarily encompass xanthones and pigments, predominantly anthraquinones. [4]

Xanthones

Lichexanthone is the most abundant xanthone, fluorescing a yellow hue under UV light, primarily on parts like the ascomata and thallus. Its presence, once a point of contention in taxonomy, is now considered species-specific due to phylogenetic studies. Another notable xanthone, coronatone, is less prevalent, showing an UV+ (orange) reaction. [4]

Pigments

Anthraquinones predominantly dominate the pigments in the Trypetheliaceae, manifesting in various parts of the lichen. Among these, parietin (also known as physcione) stands out as a yellow-orange substance. It is especially prevalent in species such as Astrothelium aeneum and Marcelaria cumingii , and has a purple reaction when tested with a K chemical spot test. Similarly, teloschistin, another yellow pigment present in Marcelaria benguelensis , also reveals a purple reaction to K. Xanthorin, on the other hand, offers a vibrant red hue and can be identified in Marcelaria purpurina , reacting purple with K+. While these are the more commonly noted pigments, there are several others like skyrin and emodin that appear in lesser quantities. The taxonomic significance of these lesser-found pigments, however, remains a subject of ongoing research. [4]

Habitat, distribution, and ecology

Trypetheliaceae species, primarily found in tropical regions, are predominantly epiphytic, meaning these lichens mainly grow on other plants without deriving nutrients from them. Although largely confined to the tropics, there are a few exceptions, such as Viridothelium virens , which is found in temperate areas and occasionally on substrates other than bark. This distribution and ecological pattern closely resemble that of the family Graphidaceae, but with notable differences: Graphidaceae species are more prevalent in extra-tropical regions and on a variety of substrates. [4]

The Trypetheliaceae are often observed in semi-exposed or fully sun-exposed environments, such as forest canopies, open savannahs, and dry forests. In these settings, they frequently contribute to exatensive crustose lichen communities, distinguished by their production of yellow to orange pigments. Species with green thalli and partly immersed or covered ascomata are more commonly found in the understory of rainforests. It has suggested that many undiscovered species may reside in these less studied, exposed canopy microhabitats. [4]

Genera

Astrothelium aeneum Astrothelium aeneum 1368620.jpg
Astrothelium aeneum

In 2020, Hongsanan and colleagues synonymized the type species of Distothelia and Novomicrothelia with Bogoriella and confirmed the placement of this genus in Trypetheliaceae. [17]

Related Research Articles

Aptrootia is a genus of fungi in the family Trypetheliaceae. It has three species. The genus was circumscribed by Robert Lücking and Harrie Sipman in 2007, with Aptrootia terricola assigned as the type species. This species, originally described by Dutch mycologist André Aptroot as a species of Thelenella, is known from Papua New Guinea and Costa Rica. Later molecular work showed that the species did not belong in Thelenella, but rather, in the Trypetheliaceae, with a sister taxon relationship to a branch including Bathelium and a lineage containing Trypethelium floridanum. The new genus name honours Aptroot, "in recognition of his numerous contributions to tropical lichenology".

<i>Architrypethelium</i> Genus of lichens

Architrypethelium is a genus of lichen-forming fungi in the family Trypetheliaceae.

<i>Astrothelium</i> Genus of lichens

Astrothelium is a large genus of corticolous (bark-dwelling) lichens in the family Trypetheliaceae. The genus is characterized by a corticate thallus and diverse ascomata structures, which can be simple, aggregated, or forming pseudostromata. Astrothelium is also notable for the carbonized walls of its ascomata, the so-called textura intricata arrangement of cells in these walls, and various forms of distoseptate, transparent spores.

<span class="mw-page-title-main">Trypetheliales</span> Order of fungi

The Trypetheliales are an order of fungi in the class Dothideomycetes. Most of the species in the order form lichens, although some are lichenicolous fungi. Trypetheliales contains two families, Polycoccaceae and Trypetheliaceae. The order was circumscribed in 2008 by lichenologists Robert Lücking, André Aptroot, and Harrie Sipman.

Constrictolumina is a genus of lichen-forming fungi in the family Trypetheliaceae. The genus was circumscribed in 2016 by Robert Lücking, Matthew Nelsen, and André Aptroot, with Constrictolumina cinchonae assigned as the type species. The genus contains tropical species, formerly placed in genus Arthopyrenia, with a unique hamathecium.

Astrothelium ecuadorense is a species of corticolous (bark-dwelling), crustose lichen in the family Trypetheliaceae. Found in Ecuador, it was formally described as a new species in 2016 by Dutch lichenologist André Aptroot. The type specimen was collected by Harrie Sipman in the Estacion Cientifico San Francisco in the South Ecuadorian Andes ; there, it was found in a mountainous rainforest growing on smooth tree bark. The lichen has a smooth and shiny, pale yellowish-green thallus with a cortex but lacking a prothallus, covering areas of up to 10 cm (4 in) in diameter. No lichen products were detected in the collected specimens. The main characteristics of the lichen distinguishing it from others in Astrothelium are the lack of pseudocyphellae on the thallus, and the solitary, exposed, black ascomata. It is also one of the few Astrothelium species with only two spores in its asci.

Astrothelium flavostiolatum is a species of corticolous (bark-dwelling), crustose lichen in the family Trypetheliaceae. Found in Ecuador, it was formally described as a new species in 2016 by Dutch lichenologist André Aptroot. The type specimen was collected by Harrie Sipman from the Reserva Biológica San Francisco in the Andes of southern Ecuador at an altitude of 2,020 m (6,630 ft); there, it was found in a rainforest growing on smooth tree bark. The lichen has an olive-green thallus with a cortex but lacking a prothallus, covering areas of up to 10 cm (4 in) in diameter. The thallus is covered with small wart-like structures, and consists of swollen and distended (bullate) areas that coalesce, forming a continuous structure. The presence of the lichen does not induce the formation of galls in its host. A yellow anthraquinone is the only lichen product that was detected from the species using thin-layer chromatography. The group of characteristics of the lichen distinguishing it from others in Astrothelium are its irregular, bullate thallus; its solitary to confluent ascomata with yellowish ostiolar rims, and the coarse thallus verrucae.

Astrothelium sexloculatum is a species of corticolous (bark-dwelling), crustose lichen in the family Trypetheliaceae. Found in Guyana and Papua New Guinea, it was formally described as a new species in 2016 by Dutch lichenologist André Aptroot. The type specimen was collected by Harrie Sipman on the Dadadanawa ranch at an altitude of 120 m (390 ft); there, it was found growing on smooth tree bark in a savanna. The lichen has a smooth and somewhat shiny, pale yellowish-grey thallus with a cortex and a thin black prothallus line. It covers areas of up to 9 cm (3.5 in) in diameter. Both the thallus and the pseudostromata contain lichexanthone, a lichen product that causes these structures to fluoresce yellow when lit with a long-wavelength UV light. The combination of characteristics of the lichen that distinguish it from others in Astrothelium are the indistinctly pseudostromatic ascomata, with erumpent to prominent pseudostromata that are covered by thallus. The species epithet sexloculatum refers to the ascospores, which are divided into six chambers (locules) by five transverse septa.

Astrothelium septemseptatum is a species of corticolous (bark-dwelling), crustose lichen in the family Trypetheliaceae. Formally described as a new species in 2016 by Dutch lichenologist André Aptroot, it is found in Guyana and Venezuela, where it grows on trees in savanna and rainforest. The type specimen was collected by Harrie Sipman about 3 km (1.9 mi) south of Kuyuwini Landing at an altitude of 230 m (750 ft); there, it was found growing on smooth tree bark. The lichen has a smooth and somewhat shiny, pale yellowish-grey thallus with a cortex and a thin black prothallus line. It covers areas of up to 9 cm (3.5 in) in diameter. The presence of the lichen does not induce the formation of galls in its host plant. Both the thallus and the pseudostroma contain lichexanthone, a lichen product that causes these structures to fluoresce yellow when lit with a long-wavelength UV light. The combination of characteristics of the lichen that distinguish it from others in Astrothelium are the presence of lichexanthone in thallus and ascomata, and the ascospores, which are divided by from 7 to 9 septa and have dimensions of 50–55 by 12–17 μm.

Astrothelium pseudodissimulum is a species of corticolous (bark-dwelling), crustose lichen in the family Trypetheliaceae. Found in Papua New Guinea, it was formally described as a new species in 2016 by Dutch lichenologist André Aptroot. The type specimen was collected by Harrie Sipman in Brahman Mission at an altitude of 100 m (330 ft); there, it was found in a rainforest growing on smooth tree bark. The lichen has a smooth, pale yellowish-green thallus with a cortex but lacking a prothallus. It covers areas of up to 15 cm (6 in) in diameter. The only lichen product detected in collected samples of the lichen using thin-layer chromatography was an anthraquinone. The combination of characteristics of the lichen that distinguish it from others in Astrothelium are its inspersedhamathecium; pigment in the medulla that reacts K+ (red); and the pseudostromatic form of the ascomata, with whitish-coloured, erumpent to prominent pseudostromata.

Astrothelium rimosum is a species of corticolous (bark-dwelling), crustose lichen in the family Trypetheliaceae. Found in Guyana and Colombia, it was formally described as a new species in 2016 by Dutch lichenologist André Aptroot. The type specimen was collected by Harrie Sipman in Kaieteur National Park at an altitude of 400 m (1,300 ft); there, in a sclerophyllous forest, it was found growing on smooth tree bark. The lichen has an olive-green to yellowish-green thallus with a thick, hyaline cortex, and a yellow medulla. The thallus texture is chinked and fissured (rimose); its species epithet rimosum alludes to this characteristic. the lichen covers areas of up to 9 cm (3.5 in) in diameter. Its thallus and medulla contain an anthraquinone that results in a K+ (red) chemical spot test. The combination of characteristics of the lichen that distinguish it from others in Astrothelium are the external and internal yellow pigment of its ascomata, and the form and dimensions of its ascospores.

Astrothelium megochroleucum is a species of corticolous (bark-dwelling), crustose lichen in the family Trypetheliaceae. Found in El Salvador, it was formally described as a new species in 2016 by Dutch lichenologist André Aptroot. The type specimen was collected by Harrie Sipman in the El Imposible National Park (Ahuachapán) at an altitude of 1,300 m (4,300 ft); there, it was found in a coffee plantation growing on the smooth tree bark of Leucaena trichandra.

Astrothelium neoinspersum is a species of corticolous (bark-dwelling), crustose lichen in the family Trypetheliaceae. Found in El Salvador, it was formally described as a new species in 2016 by Dutch lichenologist André Aptroot. The type specimen was collected by Harrie Sipman from Montecristo National Park ; there, it was found in a rainforest growing on smooth tree bark. The lichen has a smooth and somewhat shiny, olive-green thallus with a cortex but without a prothallus. It covers areas of up to 5 cm (2 in) in diameter. The presence of the lichen does not induce the formation of galls in the host plant. The only lichen product detected from collected specimens using thin-layer chromatography was an anthraquinone compound. The combination of characteristics of the lichen that distinguish it from others in Astrothelium are the yellow pigment in the ascomata; the form of the ascomata ; and the arrangement of the ascomata. The bright yellow pseudostromata help distinguish Astrothelium neoinspersum from the otherwise similar A. aenascens, found in Papua New Guinea.

Architrypethelium hyalinum is a species of corticolous (bark-dwelling), crustose lichen in the family Trypetheliaceae. Found in Costa Rica and Brazil, it was formally described as a new species in 2008 by lichenologist André Aptroot. The type specimen was collected by Harrie Sipman from the Las Cruces Biological Station in Puntarenas. The lichen has a smooth to uneven, olive-green thallus. Its ascomata occur solitarily, have an apical ostiole, and measure 0.7–1.5 mm in diameter. Ascospores number 4 to 8 per ascus, have an oblong to ellipsoid shape with 3 septa, and measure 100–150 by 30–50 μm. These spores are among the largest of the 3-septate lichens in the Trypetheliaceae. Both the thallus and ascomata contain lichexanthone, a lichen product that causes these structures to glow yellow when lit with a long-wavelength UV light; A. hyalinum is the only species in genus Architrypethelium that contains lichexanthone.

<i>Nigrovothelium</i> Genus of lichens

Nigrovothelium is a genus of lichen-forming fungi in the family Trypetheliaceae. It has three species. The genus was circumscribed in 2016 by lichenologists Robert Lücking, Matthew Nelsen, and André Aptroot, to contain species formerly in the Trypethelium tropicum species group. The type species, Nigrovothelium tropicum, was originally described by Erik Acharius in 1810, as a species of Verrucaria.

Astrothelium aurantiacocinereum is a species of corticolous (bark-dwelling), crustose lichen in the family Trypetheliaceae. It occurs in New Caledonia.

Astrothelium carassense is a species of corticolous (bark-dwelling), crustose lichen in the family Trypetheliaceae. Found in Brazil, it was formally described as a new species in 2016 by Robert Lücking, Matthew Nelsen, and Marcelo Marcelli. The type specimen was collected by the first author from the Santuário do Caraça, at an altitude between 1,300 and 1,400 m. The lichen has an uneven to bumpy, pale olive-yellow thallus that covers areas of up to 10 cm (4 in) in diameter. The pseudostromata are covered with an orange anthraquinone compound. The species epithet carassense refers to the type locality. The characteristics of the lichen that distinguish it from others in genus Astrothelium are its pseudostromatic ascomata, which are erumpent, with an orange cover; and the dimensions and form of its ascospores, which are muriform and measure 100–170 by 30–40 μm. Astrothelium purpurascens is somewhat similar in appearance to A. carassense, but it lacks pseudostromatic pigment and has slightly smaller ascospores.

Astrothelium cryptolucens is a species of corticolous (bark-dwelling), crustose lichen in the family Trypetheliaceae. Found in Panama, it was formally described as a new species in 2016 by Robert Lücking, Matthew Nelsen, and Noris Salazar Allen. The type specimen was collected by the first author from the Altos de Campana National Park at an altitude of 500–600 m (1,600–2,000 ft); there, in a submontane rainforest, it was found growing on the bark of roadside trees. The lichen has an uneven to coarsely bumpy, olive-yellow thallus that covers areas of up to 10 cm (4 in) in diameter. The species epithet cryptolucens makes reference both to the habit of the immersed perithecia and the presence of lichexanthone. The characteristics of the lichen that distinguish it from others in genus Astrothelium are the finely and densely cracked pseudostromata that dominate the thallus, with dispersed thallus bumps; and the hardly visible ascomata that are completely immersed in pseudostromata. A. cryptolucens is somewhat similar to Astrothelium carrascoense, but that species does not have an inspersed hamathecium.

Astrothelium macrostomoides is a species of corticolous (bark-dwelling), crustose lichen in the family Trypetheliaceae. Found in Brazil, it was formally described as a new species in 2016 by Robert Lücking, Matthew Nelsen, Michel Navarro Benatti. The type specimen was collected by the first author from the Santuário do Caraça at an altitude between 1,300 to 1,400 m ; there, in the Atlantic Forest biome, it was found growing on bark in a gallery forest along a river. The lichen has an uneven to coarsely bullate, olive-green thallus that covers areas of up to 7 cm (2.8 in). Ascomata are in the form of perithecia, which are arranged in aggregated groups of 5–10 in pseudostromata. The pseudostromata contain lichexanthone, which is a lichen product that causes these structures to fluoresce yellow when lit with a long-wavelength UV light. The species epithet macrostomoides alludes to the similarity of the lichen with Astrothelium macrostomum, from which it differs by having larger ascospores. The characteristics of the lichen that distinguish it from others in genus Astrothelium are the diffusely pseudostromatic ascomata with erumpent pseudostromata, covered by thallus, each with a single group of fused ascomata; and the dimensions and form of the ascospores, which measure 80–110 by 17–25 μm, and have from five to seven septa.

Dictyomeridium is a genus of corticolous (bark-dwelling), crustose lichens in the family Trypetheliaceae. It has eight species.

References

  1. 1 2 Jaklitsch, 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. pp. 76–78. ISBN   978-3-443-01089-8. OCLC   429208213.
  2. 1 2 Thiyagaraja, V.; Lücking, R.; Ertz, D.; Coppins, B.J.; Wanasinghe, D.N.; Karunarathna, S.C.; Suwannarach, N.; To-Anun, C.; Cheewangkoon, R.; Hyde, K.D. (2021). "Sequencing of the type species of Arthopyrenia places Arthopyreniaceae as a synonym of Trypetheliaceae". Mycosphere. 12 (1): 993–1011. doi: 10.5943/mycosphere/12/1/10 .
  3. ↑ 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. doi: 10.5943/mycosphere/11/1/8 . hdl: 10481/61998 .
  4. 1 2 3 4 5 6 7 8 9 10 11 Aptroot, André; Lücking, Robert (2016). "A revisionary synopsis of the Trypetheliaceae (Ascomycota: Trypetheliales)". The Lichenologist. 48 (6): 763–982. doi: 10.1017/s0024282916000487 .
  5. 1 2 Eschweiler, F.G. (1824). Systema Lichenum, Genera Exhibens rite distincta, Pluribus Novis Adaucta (in Latin). Nuremberg: J.L. Schrag. p. 17.
  6. ↑ Aptroot, André; Lücking, Robert; Sipman, Harrie J.M.; Umana, Loengrin; Chaves, José Luis (2008). Pyrenocarpous lichens with bitunicate asci. A first assessment of the lichen biodiversity inventory in Costa Rica. Bibliotheca Lichenologica. Vol. 97. Berlin; Stuttgart: J. Cramer. p. 13. ISBN   978-3-443-58076-6.
  7. ↑ Hyde, Kevin D.; Jones, E. B. Gareth; Liu, Jian-Kui; Ariyawansa, Hiran; Boehm, Eric; Boonmee, Saranyaphat; et al. (2013). "Families of Dothideomycetes". Fungal Diversity. 63 (1): 1–313. doi:10.1007/s13225-013-0263-4.
  8. ↑ Hyde, Kevin D.; Hongsanan, Sinang; Jeewon, Rajesh; Bhat, D. Jayarama; McKenzie, Eric H.C.; Jones, E.B. Gareth; et al. (2016). "Fungal diversity notes 367–490: taxonomic and phylogenetic contributions to fungal taxa" (PDF). Fungal Diversity. 80 (1): 1–270 [80]. doi:10.1007/s13225-016-0373-x.
  9. ↑ Lücking, Robert; Sipman, Harrie J.M.; Umaña, Loengrin; Chaves, Jose-Luis; Lumbsch, H.Thorsten (2007). "Aptrootia (Dothideomycetes: Trypetheliaceae), a new genus of pyrenocarpous lichens for Thelenella terricola". The Lichenologist. 39 (2): 187–193. doi:10.1017/s0024282907006445.
  10. ↑ Massalongo, A.B. (1852). Ricerche sull'autonomia dei licheni crostosi (in Italian). p. 165.
  11. ↑ Aptroot, A. (1991). A Monograph of the Pyrenulaceae (Excluding Anthracothecium and Pyrenula) and the Requienellaceae, with Notes on the Pleomassariaceae, the Trypetheliaceae, and Mycomicrothelia (Lichenized and Non-lichenized Ascomycetes). Bibliotheca Lichenologica. Vol. 44. Berlin/Stuttgart: J. Cramer. p. 120. ISBN   978-3-443-58023-0.
  12. ↑ Acharius, E. (1803). Methodus qua Omnes Detectos Lichenes Secundum Organa Carpomorpha ad Genera, Species et Varietates Redigere atque Observationibus Illustrare Tentavit Erik Acharius (in Latin). Stockholm: impensis F.D.D. Ulrich. p. 111.
  13. ↑ Zahlbruckner, A. (1928). "Neue und ungenügend beschriebene javanische Flechten". Annales de Cryptogamie Exotique (in German). 1: 109–212 [111].
  14. ↑ Sambo, M. (1940). "Licheni del Brasile". Annali di Botanica. 22: 19–41.
  15. 1 2 3 4 Lücking, Robert; Nelson, Matthew P.; Aptroot, André; Barillas de Klee, Roselvira; Bawingan, Paulina A.; Benatti, Michel N.; et al. (2016). "A phylogenetic framework for reassessing generic concepts and species delimitation in the lichenized family Trypetheliaceae (Ascomycota: Dothideomycetes)". The Lichenologist. 48 (6): 739–762. doi: 10.1017/S0024282916000505 .
  16. ↑ Reichenbach, Heinrich Gottlieb Ludwig (1841). Der deutsche Botaniker, Erster Band. Das Herbarienbuch (in German). Dresden; Leipzig: In der Arnoldischen Buchh. p. 15.
  17. 1 2 Hongsanan, Sinang; Hyde, Kevin D.; Phookamsak, Rungtiwa; Wanasinghe, Dhanushka N.; McKenzie, Eric H.C.; Sarma, V. Venkateswara; et al. (2020). "Refined families of Dothideomycetes: orders and families incertae sedis in Dothideomycetes". Fungal Diversity. 105 (1): 17–318 [184]. doi: 10.1007/s13225-020-00462-6 .
  18. ↑ Aptroot, A.; Nelsen, M.P.; Parnmen, S. (2013). "Marcelaria, a new genus for the Laurera purpurina group in the Trypetheliaceae (Ascomycota: Dothideomycetes)". Glalia. 5 (2): 1–14.
  19. ↑ Aptroot, A.; Sipman, H. (1993). "Musaespora, a genus of pyrenocarpous lichens with campylidia, and other additions to the foliicolous lichen flora of New Guinea". The Lichenologist. 25 (2): 121–135. doi:10.1006/lich.1993.1021.
  20. ↑ Hawksworth, D.L. (1985). "A redisposition of the species referred to the ascomycete genus Microthelia". Bulletin of the British Museum of Natural History. 14 (2): 43–181 [165].
  21. ↑ Müller, J. (1883). "Lichenologische Beiträge XVIII". Flora (Regensburg) (in Latin). 66 (16): 243–249.
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