Biology:Heterodermia

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Heterodermia is a genus of foliose lichens in the family Physciaceae.[1] The genus has a widespread distribution, especially in tropical regions, and contains about 70 species. Heterodermia was historically confused with the related genus Anaptychia, but was distinguished in 1965 based on differences in spore structure and chemical composition. These lichens can be identified in the field using simple chemical spot tests that produce distinctive colour changes when applied to the thallus. The lichens in this genus are small- to medium-sized, usually pale grey in colour, comprising narrow lobes with widened tips fringed with cilia. The lichens can be identified by their thick-walled ascospores and the presence of specific chemical compounds (lichen products) that produce colour reactions when spot tested. Most species are found in tropical and subtropical regions around the world, commonly growing on tree bark in mountain forests. Heterodermia species support a rich community of specialized parasitic fungi, with over 20 different species known to grow specifically on these lichens. Some species have traditional uses in medicine and cooking, particularly in India, Nepal, and Brazil.

Taxonomy

According to Fernanda de Souza and colleagues,[2] the first scientific study of the genus began in 1847 when Thomas Taylor described Parmelia diademata,[3] a lichen that would eventually end up classified as a species of Heterodermia. The genus was formally circumscribed in 1868 by Italian botanist Vittore Benedetto Antonio Trevisan de Saint-Léon.[4] The generic name, which combines the Greek heteros (other, different) and derma (a skin or hide), refers to the presence or absence of a lower cortex.[5]

Prior to this classification, Heterodermia species were placed in genus Anaptychia, an idea proposed by Syo Kurokawa in his 1962 monograph on the genus,[6] until some studies showed that the presence of thick-walled spores and the presence of atranorin could be used as characters to separate the genera.[7][8] In 1965 Josef Poelt split Anaptychia into two genera. Lichens that remained in Anaptychia had thin-walled spores with sculptured surfaces, while those transferred to Heterodermia had thick-walled spores with smooth surfaces. Only nine species remained in the original Anaptychia classification.[7] Although most contemporary lichenologists accepted Poelt's classification,[9][10][8][11] Kurokawa initially rejected these changes in 1973.[12] He later reversed his position and in 1998 accepted Heterodermia as distinct from Anaptychia, transferring several species accordingly.[13]

In North America, members of Heterodermia are colloquially known as fringe lichens or centipede lichens.[14]

Description

Lower surface of Heterodermia leucomela at 30x magnification, showing the lack of cortex characteristic of this genus.

Heterodermia typically forms a foliose thallus—occasionally shading towards a somewhat fruticose habit—that is continuous, lobate and irregular, or arranged in loose rosettes 2–10 cm (0.8–3.9 in) across. Neighbouring thalli can merge, producing broad radiating mats or tangled clumps. The lobes may lie separate or touch; they range from closely adnate and appressed to partly ascending and loosely attached, and are linear to linear-cuneate or spoon-shaped (spathulate). Branching is mostly dichotomous but can be irregular, and the margins are often fringed with cilia that are simple or densely branched.[5]

The upper surface is whitish-, grey- or yellow-grey, flat to convex—sometimes concave near the edge—and may be dull or glossy. It can carry isidia, soredia, [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] or [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]], but never pseudocyphellae. The cortex consists of longitudinally aligned hyphae (a [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]], superficially like plant collenchyma). A [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] is absent. The photobiont forms a continuous band above a well-defined white medulla that may be tinged yellow, orange or brown. The lower surface, which may lack a cortex or bear a prosoplectenchymatous one, is white to whitish grey and often darkens to purple-grey, grey-black, or partly yellow, orange or brown. Rhizines are white to black, simple to densely branched—sometimes long enough to project beyond the lobe margins—and only rarely absent.[5]

The ascomata (fruiting bodies) of Heterodermia are apothecial and [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] (ringed by thallus-like tissue). They sit on the thallus surface ([[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]]) and are rounded, either [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] or borne on a short stalk. The exposed hymenial surface, or [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]], ranges from pale to dark brown or black; it can be concave or nearly flat and may appear frosted ([[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]]) or smooth. A [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] exciple rims the disc—prominent or [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]]—and remains distinct throughout the apothecium's life.[5]

In section, the epihymenium is pale brown to brown-black, while the underlying hymenium is colourless. The [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] is usually colourless, only rarely tinged pale yellow. Paraphyses branch toward the top, where their terminal cells broaden and turn brown. The asci are cylindrical to somewhat club-shaped—Lecanora-type—with eight ascospores. Their apex is amyloid and thick-walled, enclosing a clear axial body. The spores develop one or more [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] (small vacuoles in the wall). They turn grey-brown to dark brown, are ellipsoidal to oblong or fusiform, and have a single septum that often causes a slight constriction. Walls are very thick; internal apical thickenings only appear after the septum forms. A [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] is thin or absent, and the surface remains smooth.[5]

Conidiomata lie immersed in the thallus at first, later becoming emergent. Their [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] cells form short, branched chains and produce conidia enteroblastically (inner-wall budding). The resulting conidia are bacilliform to short-cylindrical.[5]

The genus Physcia most closely resembles Heterodermia. Unlike Heterodermia, however, it has a differently structured upper cortex consisting of [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] (which gives it a uniform, unoriented appearance), and its ascospores are different.[14]

Species

Heterodermia granulifera
Heterodermia obscurata
Heterodermia pseudospeciosa
Heterodermia verrucifera

Recent estimates have placed the number of species in Heterodermia at about 115.[15] As of July 2025, Species Fungorum accepts 66 species of Heterodermia, but does not yet account for several recent studies that have added many new species.[16]

  • Heterodermia adunca P.Jung & L.Werner (2023)[17]
  • Heterodermia africana (Kurok.) M.F.Souza & Aptroot (2022)[2]
  • Heterodermia amphilacinulata M.F.N.Martins & M.P.Marcelli (2022)[2] – Brazil
  • Heterodermia andina Moberg (2011)[18]
  • Heterodermia angustiloba (Müll.Arg.) D.D.Awasthi (1973) – Asia; Australia
  • Heterodermia antillarum (Vain.) Swinscow & Krog (1976) – Australia; Central America; Caribbean; Africa; Galapagos Islands
  • Heterodermia apicalis Aptroot & M.F.Souza (2022)[2] – Brazil
  • Heterodermia archeri Elix (2011)[19]
  • Heterodermia arvidssonii Moberg (2011)[18]
  • Heterodermia badia Moberg (2011)[18]
  • Heterodermia barbifera (Nyl.) Kr.P.Singh (1981)
  • Heterodermia borphyllidiata (Kalb & Meesim) M.F.Souza & Aptroot (2022)[2]
  • Heterodermia caesiosora J.L.Watts, E.A.Tripp & Raynor (2025)[20]
  • Heterodermia caneziae M.F.Souza & Aptroot (2022)[2] – Brazil
  • Heterodermia comosa (Eschw.) Follmann & Redón (1972)
  • Heterodermia coralloides Elix (2011)[19] – Asia; Australia
  • Heterodermia corcovadensis (Kurok.) Elix (2011)
  • Heterodermia delicatula M.F.N.Martins & M.P.Marcelli (2022)[2] – Brazil
  • Heterodermia diademata (Taylor) D.D.Awasthi (1973) – Australia; North America; Central America; South America; Africa; Asia
  • Heterodermia dissecta (Kurok.) D.D.Awasthi (1973) – Asia; Australia; Réunion
  • Heterodermia dissecticodiademata M.F.Souza & Aptroot (2022)[2] – Brazil
  • Heterodermia dissecticoflabellata M.F.Souza & Aptroot (2022)[2] – Brazil
  • Heterodermia domingensis (Ach.) Trevis. (1869)
  • Heterodermia erecta Lendemer (2009)[21]
  • Heterodermia erinacea (Ach.) W.A.Weber (1987)
  • Heterodermia exuberans J.L.Watts, E.A.Tripp & Raynor (2025)[20]
  • Heterodermia flavodactyliza M.F.Souza & Aptroot (2022)[2] – Brazil
  • Heterodermia flavulifera M.F.Souza & Aptroot (2022)[2] – Brazil
  • Heterodermia follmannii Sipman (1995)
  • Heterodermia fragmentata Weerakoon & Aptroot (2016)[22]
  • Heterodermia galactophylla (Tuck.) W.L.Culb. (1967)
  • Heterodermia granulifera (Ach.) W.L.Culb. (1967)
  • Heterodermia guzmaniana (Guzmán-Guillermo, Díaz-Escandón & Medel) J.L.Watts, E.A.Tripp & Raynor (2025)[20]
  • Heterodermia himalayana Y.Joshi, K.Chandra & M.Tripathi (2014)[23]
  • Heterodermia hybocarponica Elix (2010)[24] – Australia
  • Heterodermia isidiophora (Nyl.) D.D.Awasthi (1973)
  • Heterodermia isidiophorella Elix (2011)[19] – Australia
  • Heterodermia kalbii M.F.N.Martins & Marcelli (2007)[25] – Brazil
  • Heterodermia koyana (Kurok.) Elix (2010)
  • Heterodermia koyanoides Elix (2011)[19] – Australia
  • Heterodermia labiata M.F.N.Martins & M.P.Marcelli (2022)[2] – Brazil
  • Heterodermia langdoniana Lendemer & E.Tripp (2018)[26]
  • Heterodermia linearis Moberg & T.H.Nash (1999)[27] – Mexico
  • Heterodermia macrosoraliata M.F.N.Martins & M.P.Marcelli (2022)[2] – Brazil
  • Heterodermia minor M.F.Souza & Aptroot (2022)[2] – Brazil
  • Heterodermia mobergiana Michlig, M.P.Rodríguez & Aptroot (2017)[28]
  • Heterodermia namaquana Brusse (1992)
  • Heterodermia neocomosa M.P.Rodríguez, L.I.Ferraro & Aptroot (2017)[28]
  • Heterodermia neocrocea M.F.Souza & Aptroot (2022)[2] – Brazil
  • Heterodermia neoleucomelaena (Kurok.) D.D.Awasthi (1973)
  • Heterodermia nigromarginata M.F.Souza & Aptroot (2022)[2] – Brazil
  • Heterodermia obscurata (Nyl.) Trevis. (1869)
  • Heterodermia orientalis J.B.Chen & D.P.Wang (2001)[29] – China
  • Heterodermia papuana Aptroot & Sipman (1991)
  • Heterodermia parva Moberg (2011)[18]
  • Heterodermia phyllalbicans M.F.Souza & Aptroot (2022)[2] – Brazil
  • Heterodermia pindurae Eb.Fisch., Killmann, Ertz & Sérus. (2017)[30] – Rwanda
  • Heterodermia pinnata Sipman (1995)
  • Heterodermia podocarpa (Bél.) D.D.Awasthi (1973)
  • Heterodermia pseudospeciosa (Kurok.) W.L.Culb. (1967)
  • Heterodermia queensberryi Weerakoon & Aptroot (2014)[31]
  • Heterodermia ramosociliata M.P.Rodríguez, Michlig & Aptroot (2017)[28]
  • Heterodermia rubrotricha Weerakoon & Aptroot (2013)[32]
  • Heterodermia sinocomosa J.B.Chen (2001)[29] – China
  • Heterodermia sorediosa Michlig, L.I.Ferraro & Aptroot (2017)[28]
  • Heterodermia spathulifera Moberg & Purvis (1997)
  • Heterodermia speciosa (Wulfen) Trevis. (1868)
  • Heterodermia spielmannii M.F.Souza & Aptroot (2022)[2] – Brazil
  • Heterodermia subcitrina Moberg (2004)[33]
  • Heterodermia subcomosa (Nyl.) Elix (1985)
  • Heterodermia sublinearis M.F.N.Martins & M.P.Marcelli (2022)[2] – Brazil
  • Heterodermia tabularis Elix (2011)[19] – Australia
  • Heterodermia tasmanica Elix (2011)[19] – Australia
  • Heterodermia tremulans (Müll.Arg.) W.L.Culb. (1967)
  • Heterodermia upretii Y.Joshi, S.Upadhyay & K.Chandra (2014)[34]
  • Heterodermia urtasuni Chaves, L.Umaña & Sipman (2006)[35]
  • Heterodermia velata Marcelli & Benatti (2018)[36] – Brazil
  • Heterodermia verdonii Elix (2011)
  • Heterodermia verrucifera (Kurok.) W.A.Weber (1981)

Several former Heteroderma species have been moved to Polyblastidium. This includes P. appendiculatum, P. casarettianum, P. chilense, P. corallophorum, P. dendriticum, P. fragilissimum, P. hypocaesium, P. propaguliferum, P. japonicum, P. magellanicum, P. microphyllum, P. neglectum, P. queenslandicum, P. squamulosum, P. subneglectum, P. togashii, and P. violostriatum.[15]

Habitat and distribution

Regional monographs of Heterodermia have been published for several tropical regions, including Africa (26 species),[37][33] Australia (42 species),[5] South America (33 species),[18] Brazil (68 species),[2] Thailand (39 species),[15] Most species have a pantropical or subtropical distribution, although a few have ranges that extend into temperate or oceanic regions.[15] Tropical montane forest are a common habitat for many species, and many grow on tree bark or twigs. A few species however, also grow on rock or compacted soil, while fewer yet occur only on these substrates.[2]

Ecology

Heterodermia species serve as hosts for a diverse community of lichenicolous fungi—specialized fungi that grow parasitically or commensally on lichens. These fungi represent one of the most species-rich groups of organisms associated with Heterodermia, with more than 20 documented species: Lichenostigma heterodermiae, Phoma heterodermiae, Polycoccum heterodermiae, Lichenotubeufia heterodermiae, Lichenopeltella heterodermiae, Sphaerellothecium episoralium, Sphaerellothecium gallowayi, Sphaerellothecium heterodermiae, Capronia muellerelloides, Capronia solitaria, Endococcus sipmanii, Stigmidium heterodermiae, Sclerococcum heterodermiae, Nanostictis heterodermiae, Hyalopeziza heterodermiae, Pleoscutula arsenii, Epicladonia heterodermiae, Nectriopsis heterodermiae, Nectriopsis lichenophila, Pronectria pycnidioidea, and Neobarya ciliaris.[38]

Chemistry

Careful application of the K spot test can be used to detect most of the diagnostic lichen products in Heterodermia, although thin-layer chromatography is usually essential for a definite identification. Because they occur in all species of Heterodermia, atranorin and zeorin have no diagnostic value. Norstictic acid (K+ yellow → orange-red), with or without connorstictic acid (or rarely only connorstictic), salazinic acid (K+ yellow → blood red), or dissectic acid are occasionally present, and their presence is a useful character at species level. Pigments are also diagnostic, even though the certain identification of the emodin derivatives is not always possible. The pigments are clustered in chemosyndromes (a biogenetically related set of major and minor natural metabolic products produced by a species) and there are only three different main chemosyndromes that can be distinguished by colour and K reaction. Additional terpenoids have also been recorded, including (spathulene, japonene, leucotylin, and several hopane derivatives.[2]

Uses

There are some Heterodermia species that are used as components of traditional medicine, and as spices or flavouring agent. Heterodermia diademata is used by the Nepalese of Sikkim, who apply the thalli of this lichen to cuts to protect them from wetting and infection. This species is also used in Uttar Pradesh, India as a perfume ingredient.[39] Several ethnic groups in Madhya Pradesh (a state in middle India) use the thallus of Heterodermia tremulans as a spice and flavouring agent in various dishes of vegetables and meat.[40] The Pankararu people of Pernambuco State, Brazil, use H. galactophylla for treating digestive system-related problems such as diarrhea and vomiting and for treating epilepsy.[41]

References

  1. ↑ Wijayawardene, Nalin; Hyde, Kevin; Al-Ani, Laith Khalil Tawfeeq; Somayeh, Dolatabadi; 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. 
  2. ↑ 2.00 2.01 2.02 2.03 2.04 2.05 2.06 2.07 2.08 2.09 2.10 2.11 2.12 2.13 2.14 2.15 2.16 2.17 2.18 2.19 2.20 2.21 Fernanda de Souza, Maria; Aptroot, André; Spielmann, Adriano Afonso (2022). "Key to Heterodermia (Physciaceae, Teloschistales) in Brazil, with 15 new species". The Lichenologist 54 (1): 25–44. doi:10.1017/s0024282921000499. Bibcode: 2022ThLic..54...25F. 
  3. ↑ Taylor, T. (1847). "New lichens, principally from the herbarium of Sir William J. Hooker". London Journal of Botany 6: 148–197 [165]. https://www.biodiversitylibrary.org/page/2926139. 
  4. ↑ Trevisan, V. (1868). "Sul genere Dimelaena di Norman" (in it). Atti della Società Italiana di Scienze Naturali 11: 604–630 [613]. https://www.biodiversitylibrary.org/page/39827534. 
  5. ↑ 5.0 5.1 5.2 5.3 5.4 5.5 5.6 Elix, J.A. (2011). "Heterodermia". Australian Physciaceae (Lichenised Ascomycota). https://www.anbg.gov.au/abrs/lichenlist/HETERODERMIA.pdf. 
  6. ↑ Kurokawa, Syo (1962). A Monograph of the Genus Anaptychia. Beihefte zur Nova Hedwigia. 6. Weinheim: J. Cramer. pp. 1–115. 
  7. ↑ 7.0 7.1 Poelt, J. (1965). "Zur Systematik der Flechtenfamilie Physciaceae" (in de). Nova Hedwigia 9: 21–32. 
  8. ↑ 8.0 8.1 Swinscow, T.D.V.; Krog, Hildur (2007). "The genera Anaptychia and Heterodermia in East Africa". The Lichenologist 8 (2): 103–138. doi:10.1017/S0024282976000212. 
  9. ↑ Culberson, W.L. (1966). "Chemistry and taxonomy of the lichen genera Heterodermia and Anaptychia in the Carolinas". The Bryologist 69 (4): 472–487. doi:10.1639/0007-2745(1966)69[472:CATOTL2.0.CO;2]. 
  10. ↑ Awasthi, D.D. (1973). "On the species of Anaptychia and Heterodermia from India and Nepal". Geophytology 3: 113–116. 
  11. ↑ Trass, H. (1992). "Synopsis of the lichen genus Heterodermia (Ascomycotina, Physciaceae sive Pyxinaceae)". Folia Cryptogamica Estonica 29: 1–24. 
  12. ↑ Kurokawa, Syo (1973). "Supplementary notes on the genus Anaptychia". Journal of the Hattori Botanical Laboratory 37: 563–607. 
  13. ↑ Kurokawa, Syo (1998). "A catalogue of Heterodermia". Folia Cryptogica Estonica 32: 21–25. 
  14. ↑ 14.0 14.1 Brodo, Irwin M.; Sharnoff, Sylvia Duran; Sharnoff, Stephen (2001). Lichens of North America. Yale University Press. p. 334. ISBN 978-0300082494. 
  15. ↑ 15.0 15.1 15.2 15.3 Mongkolsuk, Pachara; Meesim, Sanya; Poengsungnoen, Vasun; Buarung, Kawinnat; Schumm, Felix; Kalb, Klaus (2015). The lichen family Physciaceae in Thailand—II. Contributions to the genus Heterodermia sensu lato. Phytotaxa. Monograph. 235. New Zealand: Magnolia Press. pp. 1–66. doi:10.11646/phytotaxa.235.1.1. ISBN 978-1-77557-840-6. http://publication.plazi.org/id/FFD60E49FFFF6A62FFCF4960FF8EFF81. 
  16. ↑ "Heterodermia". Species 2000: Leiden, the Netherlands. https://www.catalogueoflife.org/data/taxon/KNV62. 
  17. ↑ Jung, Patrick; Werner, Lina; Briegel-Williams, Laura; Emrich, Dina; Lakatos, Michael (2023). "Roccellinastrum, Cenozosia and Heterodermia: Ecology and phylogeny of fog lichens and their photobionts from the coastal Atacama Desert". MycoKeys (98): 317–348. doi:10.3897/mycokeys.98.107764. PMID 37564324. 
  18. ↑ 18.0 18.1 18.2 18.3 18.4 Moberg, Roland (2011). "The lichen genus Heterodermia (Physciaceae) in South America – a contribution including five new species". Nordic Journal of Botany 29 (2): 129–147. doi:10.1111/j.1756-1051.2009.00519.x. Bibcode: 2011NorJB..29..129M. 
  19. ↑ 19.0 19.1 19.2 19.3 19.4 19.5 Elix, J.A. (2011). "Further new species and new records of Heterodermia (Physciaceae, Ascomycota) from Australia". Australasian Lichenology 69: 12–24. https://www.anbg.gov.au/abrs/lichenlist/AL_69.pdf. 
  20. ↑ 20.0 20.1 20.2 Watts, Jacob L.; Raynor, Seth J.; Manzitto-Tripp, Erin A. (2025). "Character evolution in Heterodermia s.l. (Physciaceae; Caliciales) and two new species from the southern Rocky Mountains, USA". Phytotaxa 698 (2): 61–81. doi:10.11646/phytotaxa.698.2.1. 
  21. ↑ Lendemer, James C. (2009). "A synopsis of the lichen genus of Heterodermia (Physciaceae, lichenized Ascomycota) in eastern North America". Opuscula Philolichenum 6: 1–36. doi:10.5962/p.381963. https://sweetgum.nybg.org/images3/250/135/op6p1.pdf. 
  22. ↑ Weerakoon, Gothamie; Aptroot, André (2016). "Nine new lichen species and 64 new records from Sri Lanka". Phytotaxa 280 (2): 152–162. doi:10.11646/phytotaxa.280.2.5. Bibcode: 2016Phytx.280..152W. http://publication.plazi.org/id/FFAFFFFAFFF0FF8BFFF531541728FFFC. 
  23. ↑ Joshi, Y.; Chandra, K.; Tripathi, M. (2014). "A new species of Heterodermia Trevis. (Ascomycota, Physciaceae) from India, along with a new record and range extension of lichenized fungi in India". Phytotaxa 170 (1): 49–52. doi:10.11646/phytotaxa.170.1.7. 
  24. ↑ Elix, John A. (2010). "Two new species, a new combination and new chemical data for Heterodermia (Physciaceae: Ascomycota)". Australasian Lichenologist 67: 3–7. https://www.anbg.gov.au/abrs/lichenlist/AL_67.pdf. 
  25. ↑ Marcelli, Marcelo P.; Jungbluth, Patrícia; Benatti, Michel N.; Spielmann, Adriano A.; Canêz, Luciana S.; Cunha, Iane P.R.; Martins, Milton F.N. (2007). "Some new species and combinations of Brazilian lichenized fungi". Bibliotheca Lichenologica 96: 209–228. https://archive.org/details/SomeNewSpeciesAndCombinationsOfBrazilianLichenizedFungi. 
  26. ↑ Tripp, Erin A.; Lendemer, James C. (2019). "Highlights from 10+ Years of Lichenological Research in Great Smoky Mountains National Park: Celebrating the United States National Park Service Centennial". Systematic Botany 44 (4): 943–980. doi:10.1600/036364419X15710776741332. Bibcode: 2019SysBo..44..943T. 
  27. ↑ Moberg, Roland; Nash III, Thomas H. (1999). "The genus Heterodermia in the Sonoran Desert area". The Bryologist 102 (1): 1–14. doi:10.2307/3244452. 
  28. ↑ 28.0 28.1 28.2 28.3 Michlig, Andrea; Rodríguez, Maria Pia; Aptroot, André; Niveiro, Nicolás; Ferraro, Lidia Itatí (2017). "New species of the Heterodermia comosa-group (Physciaceae, lichenized Ascomycota) from southern South America". Cryptogamie, Mycologie 38 (2): 155–167. doi:10.7872/crym/v38.iss1.2017.155. 
  29. ↑ 29.0 29.1 Chen, J.B. (2001). "The lichen family Physciaceae (Ascomycota) in China II. Two new species of Heterodermia". Mycotaxon 77: 101–105. http://www.cybertruffle.org.uk/cyberliber/59575/0077/0101.htm. 
  30. ↑ Fischer, E.; Killmann, D.; Ertz, D.; Sérusiaux, E. (2017). "Heterodermia pindurae (Physciaceae)—a new foliose lichen from Rwanda". Phytotaxa 311 (3): 277–282. doi:10.11646/phytotaxa.311.3.8. Bibcode: 2017Phytx.311..277F. 
  31. ↑ Weerakoon, Gothamie; Aptroot, André (2014). "Over 200 new lichen records from Sri Lanka, with three new species to science". Cryptogamie, Mycologie 35 (1): 51–62. doi:10.7872/crym.v35.iss1.2014.51. 
  32. ↑ Weerakoon, Gothamie; Aptroot, André (2013). "Some new lichen species from Sri Lanka, with a key to the genus Heterodermia in Sri Lanka". Cryptogamie, Mycologie 34 (4): 321–328. doi:10.7872/crym.v34.iss4.2013.321. 
  33. ↑ 33.0 33.1 Moberg, Roland (2004). "Notes on foliose species of the lichen family Physciaceae in southern Africa". Contributions to lichen taxonomy and biogeography: Dedicated to Leif Tibell. Symbolae Botanicae Upsalienses. 34. Uppsala: Acta Universitatis Upsaliensis. pp. 257–288. ISBN 91-554-6025-9. 
  34. ↑ Yogesh, Joshi; Upadhyay, Shashi; Chandra, Krishna (2014). "Heterodermia upretii, a new species from India (Physciaceae, Ascomycota)". Phytotaxa 175 (2): 117–120. doi:10.11646/phytotaxa.175.2.7. Bibcode: 2014Phytx.175..117J. 
  35. ↑ Chaves, José Luis; Umaña, Loengrin; Lücking, Robert; Sipman, Harrie J.M.; Aptroot; André (2006). "Una nueva especie de Heterodermia de Costa Rica y Venezuela, en honor al embajador de España en Costa Rica, Juan José Urtasun Erro" (in es). Cryptogamie, Mycologie 27 (4): 335–339. https://sciencepress.mnhn.fr/fr/periodiques/mycologie/27/4/una-nueva-especie-de-heterodermia-de-costa-rica-y-venezuela-en-honor-al-embajador-de-espana-en-costa-rica-juan-jose-urtasun-erro. 
  36. ↑ Benatti, Michel Navarro; Marcelli, Marcelo Pinto (2017). "Physciaceae foliosas do Parque Estadual da Cantareira, Estado de São Paulo, Brasil. II. Gêneros Heterodermia e Polyblastidium" (in es). Iheringia 72 (2): 255–266. doi:10.1590/2175-7860201970061. 
  37. ↑ Swinscow, Thomas Douglas Victor; Krog, Hildur (1988). Macrolichens of East Africa. London: British Museum (Natural History). ISBN 978-0-565-01039-3. 
  38. ↑ Diederich, Paul; Lawrey, James D.; Ertz, Damien (2018). "The 2018 classification and checklist of lichenicolous fungi, with 2000 non-lichenized, obligately lichenicolous taxa". The Bryologist 121 (3): 340. doi:10.1639/0007-2745-121.3.340. 
  39. ↑ Singh, S.; Upreti, D.K.; Lehri, A.; Paliwal, A.K. (2015). "Quantification of lichens commercially used in traditional perfumery industries of Uttar Pradesh, India". Indian Journal of Plant Sciences 4 (1): 29–33. 
  40. ↑ Upreti, Dalip K.; Divakar, Pradeep K.; Nayaka, Sanjeeva (2005). "Commercial and ethnic use of lichens in India". Economic Botany 59 (3): 269–273. doi:10.1663/0013-0001(2005)059[0269:CAEUOL2.0.CO;2]. https://www.researchgate.net/publication/227205410. 
  41. ↑ Londoño-Castañeda, P.A.; Buril, M.L.L.; Rego-Cunha, I.P.; Silva, N.H.; Honda, N.K.; Pereira, E.C.; Andrade, L.H.C. (2017). "Lichens used in the traditional medicine by the Pankararu Indigenous community, Pernambuco-Brazil". Global Journal of Science Frontier Research: C Biological Science 17 (4): 15–22. 
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