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The following outline provides an overview of and topical guide to lichens.

Lichen – composite organism made up of multiple species – a fungal partner, one or more photosynthetic partners, which can be either green algae or cyanobacteria, and, in at least 52 genera of lichens, a basidiomycete yeast.^[1]

What is a lichen?

A lichen can be described as all of the following:

Life form – an entity that is alive

Composite organism – symbiotic life form composed of multiple partners from different biological kingdoms or domains, and different phyla within those kingdoms, including:

Fungi (kingdom) – the fungal partner (known as the mycobiont) and any basidiomycete yeast partner

Ascomycota (phylum) and/or Basidiomycota (phylum)

Plantae (kingdom) – if the photosynthetic partner, or photobiont, is a green alga

Trebouxiophyceae (phylum)

Bacteria (domain) – if the photobiont is a cyanobacterium

Cyanobacteria (phylum)

Nature of lichens

Morphology

Apoplast – the symbiotic interface zone, outside the cell membranes or walls of the mycobiont and photobiont
Ascocarp – the fruiting body of a lichen, containing fungal hyphae and asci

Ascus (plural asci) – a sexual spore-bearing cell
Ascospore – a spore produced in an ascus

Epispore – a transparent bag-like outer covering on some spores^[2]

Cephalodium (plural cephalodia) – a gall-like structure that contains cyanobacteria^[3]
Hypha (plural hyphae) – a long, branching, thread-like structure composed of one or more fungal cells; typically densely packed in lichens, making up a large part of their structure

Appressorium (plural appressoria) – a flat hyphal structure that attaches to photobiont cells

Haustorium (plural haustoria) – a root-like structure which allows the fungal partner to extract nutrients from the photosynthetic partner^[2]

Pycnidium (plural pycnidia) – an asexual fruiting body possessed by some lichens

Conidium (plural conidia) – an asexual fungal spore produced in pycnidia by some lichens

Rhizine – a root-like structure that anchors a lichen to the substrate on which it grows
Soralia – a localized structure containing soredium

Soredium (plural soredia) – an asexual reproductive structure containing small amounts of fungal hyphae and algae

Thallus (plural thalli) – the vegetative body of a lichen, made up of both mybiont and photobiont components^[4]

Cortex – the lichen's outer layer(s), made up of tightly woven fungal filaments^[5]
Isidium (plural isidia) – outgrowths of the thallus which contain photobiont cells and provide means of vegetative reproduction for some lichens^[6]
Medulla – a loose layer of interwoven fungal hyphae within the thallus^[7]
Podetium (plural podetia) – an upright secondary thallus, which serves as the fruiting structure of Cladonia species^[8]

Ecology

Symbiosis in lichens – the relationship between the lichen partners can be complicated; while generally mutualistic, sometimes it is not
Sexual reproduction in lichens – many lichens reproduce sexually using ascospores
Lichens and nitrogen cycling – some lichens (in particular those with cyanobacteria as a photobiont) can fix nitrogen^[9]
Lichen biogeography – the study of the current distribution of extant lichens and the reasons for those distributions^[10]
Lichen resynthesis – lichens can be artificially "recreated" by combining partners in a lab
Lichens and pedogenesis – lichens contribute to the formation of soil by breaking down rock^[11]

Types of lichens

Lichen lists

Lichen taxonomical classifications

Although they are composite organisms, lichens have traditionally been classified on the basis of their fungal partner. These span eight different biological classes, 38 orders, 115 families, and nearly 1,000 genera.^[12]^[14]

Lichen systematics

Ascolichen – a lichen whose fungal partner is an ascomycete – that is, a member of the Ascomycota, one of the two main fungal divisions.^[15]
Basidiolichen – a lichen whose fungal partner is a member of the Basidiomycota, the other of the two main fungal divisions; these are far fewer in occurence than ascolichens^[16]

Classes

Lichens fall into eight fungal classes:^[13]

Orders

They are split across 38 orders. Those which cannot be assigned to a particular order are assigned instead to "incertae sedis" within the appropriate class.^[13]

Families

They fall into 115 families. Those which cannot be assigned to a particular family are assigned instead to "incertae sedis" within the appropriate order.^[14]

Genera

They are found in nearly 1000 genera:^[14]

Species

Nearly 20,000 species of lichen have been described, and taxonomists estimate that the number of as yet undescribed species may be as high as 8,000 more.^[17]

Lichens, by growth form

Lichen growth forms – These vary depending on the species:

Byssoid – wispy, with the appearance of teased wool
Crustose – paint-like appearance that adheres tightly to the underlying substrate
- Areolate – crustose, but divided into rounded or polygonal pieces by means of cracks^[19]
- Calicioid – crustose growth with small fruiting bodies which resemble sewing pins
- Placodioid – crustose in the centre and lobed at the periphery^[19]
Filamentous – thin, threadlike growth, often with a matted appearance
Foliose – flattened, leafy appearance^[19]
Fruticose – shrubby, bush-like or coral-like appearance^[19]
Gelatinous – jelly-like appearance
Leprose – powdery or granular appearance^[19]
Squamulose – scaly, sometimes leafy appearance; can resemble a foliose lichen but has no outer cortex
- Cladoniform – squamulose, but with erect podetia

Lichens, by substrate

Lichens can be classified based on what they grow on:

Bryophilous lichen – on mosses or liverworts^[20]
- Hepaticolous lichen – on liverworts^[22]
- Muscicolous lichen – on mosses^[23]
Corticolous lichen – on bark^[20]
- Ramicolous lichen – on twigs^[23]
Foliicolous lichen – on plant leaves^[20]
- Epiphyllous lichen – on the upper surface of a leaf^[24]
- Hypophyllous lichen – on the lower surface of a leaf^[24]
Lichenicolous lichen – on other lichens^[20]
Lignicolous lichen – on wood^[20]
Omnicolous lichen – on various different substrates, including manmade structures^[23]
Plasticolous lichen – on plastic^[25]
Saxicolous lichen – on stone^[20]
Terricolous lichen – on soil^[23]
Vagrant lichen – loose, on no substrate^[26]

Lichens, by region

Africa

Photobiont

Photobiont – the photosynthetic partner in a lichen^[28]

Cyanolichen – a lichen with a cyanobacteria photobiont^[29]
List of lichen photobionts

Lichen metabolites

Lichen product – organic products, known as secondary metabolites, produced by lichens; these provide a variety of protections for the lichen – from microbes, viruses, herbivores, radiation, oxidants and more^[30]

List of lichen products

Study of lichens

Lichenology – the study of lichens^[32]

Acharius Medal – awarded for lifetime achievement in lichenology
History of lichenology
Lichenometry – a process where measuring the growth of a lichen colony over time can be used to estimate the minimum age of the substrate on which it is growing^[33]
Spot test (lichen) – chemical tests used to aid in species identification^[34]

Threats

Lichenicolous fungus – parasitic fungus that uses lichens as a host^[35]

List of lichenicolous fungi

Lichens as bioindicators – lichens are sensitive to various pollutants and can be thus be used as bioindicators^[36]

Lichens and air pollution – many lichens are sensitive to various forms of air pollution^[37]
Lichens and climate change – the inability of algae to quickly evolve means that climate change may adversely impact lichens^[38]

Lichens in culture

Edible lichen – some lichens have traditionally been used as food^[40]
Ethnolichenology – a branch of ethnobotany that studies human usage of lichens^[41]
Lichens in popular culture

Lichen organizations

American Bryological and Lichenological Society (ABLS)

The Bryologist – peer-reviewed journal published by ABLS

British Lichen Society (BLS)

The Lichenologist – peer-reviewed journal published by the BLS

Bryological and Lichenological Association for Central Europe (BLAM)

Herzogia – peer-reviewed journal published by BLAM

International Association for Lichenology (IAL)

Citations

^ Van Hoose 2021.
^ ^a ^b Brodo, Sharnoff & Sharnoff 2001, p. 758.
^ Brodo, Sharnoff & Sharnoff 2001, p. 756.
^ Brodo, Sharnoff & Sharnoff 2001, p. 763.
^ Brodo, Sharnoff & Sharnoff 2001, p. 757.
^ Brodo, Sharnoff & Sharnoff 2001, p. 759.
^ Brodo, Sharnoff & Sharnoff 2001, p. 760.
^ Brodo, Sharnoff & Sharnoff 2001, p. 761.
^ Henriksson & Simu 1971, p. 119.
^ Galloway 2012, p. 315.
^ Syers & Iskandar 1973.
^ ^a ^b Wijayawardene et al. 2022.
^ ^a ^b ^c Lücking, Hodkinson & Leavitt 2017, p. 371.
^ ^a ^b ^c Lücking, Hodkinson & Leavitt 2017, p. 377–400.
^ Bendre 2010, p. 131.
^ Lepp.
^ Lücking et al. 2009.
^ Dobson 2011, p. 470.
^ ^a ^b ^c ^d ^e The British Lichen Society 2022a.
^ ^a ^b ^c ^d ^e ^f ^g The British Lichen Society 2022b.
^ Schultz, Zedda & Rambold 2009, p. 315.
^ Lendemer, Buck & Harris 2016, p. 441.
^ ^a ^b ^c ^d Upreti & Rai 2013, p. 2.
^ ^a ^b Lücking 2008, p. 4.
^ Lücking 1998, p. 287.
^ Rosentreter 1993, p. 333.
^ Purvis 2000, p. 9.
^ Purvis 2000, p. 5.
^ Rikkinen 2015, p. 973.
^ Goga et al. 2018, p. 1.
^ Dobson 2011, p. 11.
^ Merriam-Webster.
^ Brodo, Sharnoff & Sharnoff 2001, p. 84.
^ Brodo, Sharnoff & Sharnoff 2001, p. 103.
^ Lawrey & Diederich 2003, p. 80.
^ Nimis, Scheidegger & Wolseley 2002.
^ Oliver 2011, p. 1.
^ Field Museum 2022.
^ Turner 1977, p. 461.
^ Ivanova & Ivanov 2009, p. 11.
^ Vinayaka & Krishnamurthy 2012, p. 265.

References

Bendre, Ashok M. (2010). A Text Book of Practical Botany. New Dehli: Rastogi Publications. ISBN 9788171339235.

Brodo, Irwin M.; Sharnoff, Sylvia Duran; Sharnoff, Stephen (2001). Lichens of North America. New Haven, CT: Yale University Press. ISBN 0-300-08249-5.

Dobson, Frank S. (2011). Lichens: An Illustrated Guide to the British and Irish Species. Slough, UK: Richmond Publishing Co. ISBN 978-0-85546-316-8.

Field Museum (15 February 2022). "Lichens are in danger of losing the evolutionary race with climate change". Phys.org. Retrieved 23 August 2022.

Galloway, D. J. (2012). "Lichen biogeography". In Nash III, Thomas H. (ed.). Lichen Biology. Cambridge UK: Cambridge University Press. pp. 315–335. ISBN 978-0-521-69216-8.

"Glossary of Terms". The British Lichen Society. 2022b. Retrieved 18 August 2022.

Goga, Michal; Elečko, Ján; Marcinčinová, Margaréta; Ručová, Dajana; Bačkorová, Miriam; Bačkor, Martin (2018). "Lichen Metabolites: An Overview of Some Secondary Metabolites and Their Biological Potential". In Merillon, J. M.; Ramawat, K. (eds.). Co-Evolution of Secondary Metabolites. Reference Series in Phytochemistry. Springer, Cham. pp. 1–36. doi:10.1007/978-3-319-76887-8_57-1. ISBN 978-3-319-76887-8.

Henriksson, Elisabet; Simu, Barbro (1971). "Nitrogen Fixation by Lichens". Oikos. 22 (1): 119–121. JSTOR 3543371.

Lawrey, James D.; Diederich, Paul (Spring 2003). "Lichenicolous Fungi: Interactions, Evolution, and Biodiversity". The Bryologist. 106 (1): 80–120. JSTOR 3244800.

Lendemer, James C.; Buck, William R.; Harris, Richard C. (September 2016). "Two new host-specific hepaticolous species of Catinaria (Ramalinaceae)". The Lichenologist. 48 (5): 441–449. doi:10.1017/S0024282916000438.

Lepp, Heino. "Basidiolichens". Australian Lichens. Australian National Herbarium and Australian National Botanic Gardens. Retrieved 20 August 2022.

Ivanova, Diana; Ivanov, Dobri (2009). "Ethnobotanical use of lichens: lichens for food review". Scripta Scientifica Medica. 41 (1): 11–16. doi:10.14748/ssm.v41i1.456.

"Lichen Morphology". The British Lichen Society. 2022a. Retrieved 23 August 2022.

"Lichenology". Merriam-Webster. Retrieved 20 August 2022.

Lücking, Robert (May 1998). "'Plasticolous' Lichens in a Tropical Rain Forest at La Selva Biological Station, Costa Rica". The Lichenologist. 30 (3): 287–291. doi:10.1006/lich.1998.0124.

Lücking, Robert (2008). "Foliicolous Lichenized Fungi". Flora Neotropica. 103: 1–866. ISSN 0071-5794.

Lücking, Robert; Rivas-Plata, E; Chavez, JL; Umaña, L; Sipman, HJM (2009). "How many tropical lichens there are…really?". Bibliotheca Lichenologica. 100: 399–418.

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. JSTOR 44250015. S2CID 90258634.

Nimis, Pier Luigi; Scheidegger, Christoph; Wolseley, Patricia A., eds. (2002). Monitoring with Lichens - Monitoring Lichens. Vol. 7. Dordrecht: Springer Science+Business. ISBN 978-94-010-0423-7.

Oliver, Mary (March 2011). "Canaries in a coal mine: using lichens to measure nitrogen pollution" (PDF). Science Findings. 131. USDA Forest Service, Pacific Northwest Research Station.

Purvis, William (2000). Lichens. Life Series. London: The Natural History Museum. ISBN 978-0-565-09153-8.

Rikkinen, Jouko (April 2015). "Cyanolichens". Biodiversity and Conservation. 24 (4): 973–993. doi:10.1007/s10531-015-0906-8.

Rosentreter, Roger (Autumn 1993). "Vagrant Lichens in North America". The Bryologist. 96 (3): 333–338. JSTOR 3243861.

Schultz, Matthias; Zedda, Luciana; Rambold, Gerhard (2009). "New records of lichen taxa from Namibia and South Africa Bibliotheca Lichenologica". In Aptroot, André; Seaward, Mark R.D.; Sparrius, Laurens B. (eds.). Biodiversity and ecology of lichens – Liber Amicorum Harrie Sipman (PDF). Vol. 99. Berlin and Stuttgart: J. Cramer. pp. 315–334.

Syers, J.K.; Iskandar, I.K. (1973). "Pedogenetic Significance of Lichens". In Ahmadjian, Vernon; Hale, Mason E. (eds.). The Lichens. New York: Academic Press. pp. 225–248. doi:10.1016/B978-0-12-044950-7.50012-X. ISBN 0-12-044950-1.

Turner, Nancy J. (October 1977). "Economic importance of black tree lichen (Bryoria fremontii) to the Indians of western North America". Economic Botany. 31 (4): 461–470. JSTOR 4253875.

Upreti, Dalip K.; Rai, Himanshu, eds. (2013). Terricolous Lichens in India. Vol. 1: Diversity Patterns and Distribution Ecology. New York: Springer. ISBN 978-1-4614-8736-4.

Van Hoose, Natalie (21 July 2021). "Yeast emerges as hidden third partner in lichen symbiosis". Purdue University News. Purdue University.

Vinayaka, K. S.; Krishnamurthy, Y. L. (2012). "Ethno-lichenological Studies of Shimoga and Mysore Districts, Karnataka, India" (PDF). Advances in Plant Sciences. 25 (1): 265–267.

Wijayawardene, N.N.; Hyde, K.D.; Dai, D.Q.; Sánchez-García, M.; Goto, B.T.; Saxena, R.K.; et al. (2022). "Outline of Fungi and fungus-like taxa – 2021". Mycosphere. 13 (1): 53–453. doi:10.5943/mycosphere/13/1/2. S2CID 249054641.

External links

[FOOTNOTEVan_Hoose2021-1] Van Hoose 2021.

[FOOTNOTEBrodoSharnoffSharnoff2001758-2] Brodo, Sharnoff & Sharnoff 2001, p. 758.

[FOOTNOTEBrodoSharnoffSharnoff2001756-3] Brodo, Sharnoff & Sharnoff 2001, p. 756.

[FOOTNOTEBrodoSharnoffSharnoff2001763-4] Brodo, Sharnoff & Sharnoff 2001, p. 763.

[FOOTNOTEBrodoSharnoffSharnoff2001757-5] Brodo, Sharnoff & Sharnoff 2001, p. 757.

[FOOTNOTEBrodoSharnoffSharnoff2001759-6] Brodo, Sharnoff & Sharnoff 2001, p. 759.

[FOOTNOTEBrodoSharnoffSharnoff2001760-7] Brodo, Sharnoff & Sharnoff 2001, p. 760.

[FOOTNOTEBrodoSharnoffSharnoff2001761-8] Brodo, Sharnoff & Sharnoff 2001, p. 761.

[FOOTNOTEHenrikssonSimu1971119-9] Henriksson & Simu 1971, p. 119.

[FOOTNOTEGalloway2012315-10] Galloway 2012, p. 315.

[FOOTNOTESyersIskandar1973-11] Syers & Iskandar 1973.

[FOOTNOTEWijayawardeneHydeDaiSánchez-García2022-12] Wijayawardene et al. 2022.

[FOOTNOTELückingHodkinsonLeavitt2017371-13] Lücking, Hodkinson & Leavitt 2017, p. 371.

[FOOTNOTELückingHodkinsonLeavitt2017377–400-14] Lücking, Hodkinson & Leavitt 2017, p. 377–400.

[FOOTNOTEBendre2010131-15] Bendre 2010, p. 131.

[FOOTNOTELepp-16] Lepp.

[FOOTNOTELückingRivas-PlataChavezUmaña2009-17] Lücking et al. 2009.

[FOOTNOTEDobson2011470-18] Dobson 2011, p. 470.

[FOOTNOTEThe_British_Lichen_Society2022a-19] The British Lichen Society 2022a.

[FOOTNOTEThe_British_Lichen_Society2022b-20] ^ ^a ^b ^c ^d ^e ^f ^g The British Lichen Society 2022b.

[FOOTNOTESchultzZeddaRambold2009315-21] Schultz, Zedda & Rambold 2009, p. 315.

[FOOTNOTELendemerBuckHarris2016441-22] Lendemer, Buck & Harris 2016, p. 441.

[FOOTNOTEUpretiRai20132-23] Upreti & Rai 2013, p. 2.

[FOOTNOTELücking20084-24] Lücking 2008, p. 4.

[FOOTNOTELücking1998287-25] Lücking 1998, p. 287.

[FOOTNOTERosentreter1993333-26] Rosentreter 1993, p. 333.

[FOOTNOTEPurvis20009-27] Purvis 2000, p. 9.

[FOOTNOTEPurvis20005-28] Purvis 2000, p. 5.

[FOOTNOTERikkinen2015973-29] Rikkinen 2015, p. 973.

[FOOTNOTEGogaElečkoMarcinčinováRučová20181-30] Goga et al. 2018, p. 1.

[FOOTNOTEDobson201111-31] Dobson 2011, p. 11.

[FOOTNOTEMerriam-Webster-32] Merriam-Webster.

[FOOTNOTEBrodoSharnoffSharnoff200184-33] Brodo, Sharnoff & Sharnoff 2001, p. 84.

[FOOTNOTEBrodoSharnoffSharnoff2001103-34] Brodo, Sharnoff & Sharnoff 2001, p. 103.

[FOOTNOTELawreyDiederich200380-35] Lawrey & Diederich 2003, p. 80.

[FOOTNOTENimisScheideggerWolseley2002-36] Nimis, Scheidegger & Wolseley 2002.

[FOOTNOTEOliver20111-37] Oliver 2011, p. 1.

[FOOTNOTEField_Museum2022-38] Field Museum 2022.

[FOOTNOTETurner1977461-39] Turner 1977, p. 461.

[FOOTNOTEIvanovaIvanov200911-40] Ivanova & Ivanov 2009, p. 11.

[FOOTNOTEVinayakaKrishnamurthy2012265-41] Vinayaka & Krishnamurthy 2012, p. 265.

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@@ Line 1,349: / Line 1,349: @@
 * [[Lichenicolous fungus]] – parasitic fungus that uses lichens as a host{{sfn|Lawrey|Diederich|2003|p=80}}
 :* [[List of lichenicolous fungi]]
-* [[Lichens as bioindicators]] – lichens are sensitive to various pollutants and can be thus be used as bioindicators
+* [[Lichens as bioindicators]] – lichens are sensitive to various pollutants and can be thus be used as bioindicators{{sfn|Nimis|Scheidegger|Wolseley|2002}}
 :* [[Lichens and air pollution]] – many lichens are sensitive to various forms of air pollution{{sfn|Oliver|2011|p=1}}
 :* [[Lichens and climate change]] – the inability of algae to quickly evolve means that climate change may adversely impact lichens{{sfn|Field Museum|2022}}
@@ Line 1,409: / Line 1,409: @@
 * {{cite journal |last1=Lücking |first1=Robert |last2=Hodkinson |first2=Brendan P. |last3=Leavitt |first3=Steven D. |year=2017 |title=The 2016 classification of lichenized fungi in the Ascomycota and Basidiomycota–Approaching one thousand genera |journal=The Bryologist |volume=119 |issue=4 |pages=361–416 |doi=10.1639/0007-2745-119.4.361 |s2cid=90258634 |jstor=44250015}}
+* {{cite book |editor1-last=Nimis |editor1-first=Pier Luigi |editor2-last=Scheidegger |editor2-first=Christoph |editor3-last=Wolseley |editor3-first=Patricia A. |year=2002 |title=Monitoring with Lichens - Monitoring Lichens |publisher=Springer Science+Business |location=Dordrecht |volume=7 |isbn=978-94-010-0423-7}}
 * {{cite journal |last=Oliver |first=Mary |title=Canaries in a coal mine: using lichens to measure nitrogen pollution |journal=Science Findings |volume=131 |publisher=[[USDA Forest Service]], Pacific Northwest Research Station |date=March 2011 |url=https://www.fs.usda.gov/pnw/sciencef/scifi131.pdf}}