Outline of lichens

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 domains, families and kingdoms, and different phyla, classes and divisions within those domains and kingdoms

Eukaryote (domain) – organisms with a cell nucleus within a nuclear envelope^[2]

Fungi (kingdom) – the fungal partner (known as the mycobiont) and any basidiomycete yeast partner^[1]

Ascomycota (phylum) and/or Basidiomycota (phylum)^[3]

Chlorophyta (division) – if the photosynthetic partner, or photobiont, is a green alga^[4]

Trebouxiophyceae (class)

Trebouxiaceae (family)^[5]

Ulvophyceae (class)

Trentepohliaceae (family)^[5]

Prokaryote (domain) – organisms without a cell nucleus^[2]

Bacteria (domain) – if the photobiont is a cyanobacterium^[6]

Cyanobacteria (phylum)^[6]

Nature of lichens

Morphology

Lichen anatomy and physiology
Lichen morphology – a lichen's morphology is very different from that of its individual partners^[8]

Apoplast – the symbiotic interface zone, outside the cell membranes or walls of the mycobiont and photobiont^[9]
Ascocarp – the fruiting body of a lichen, containing fungal hyphae and asci

Ascus (plural asci) – a sexual, fungal spore-bearing structure, typically sac-like^[10]
Ascospore – a fungal spore, the product of meiosis, produced in an ascus^[11]

Epispore – a transparent bag-like outer covering on some spores,^[12] which helps to determine spore shape^[13]

Cephalodium (plural cephalodia) – a gall-like structure that contains cyanobacteria^[14]
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

Haustorium (plural haustoria) – a root-like structure which allows the fungal partner to extract nutrients from the photosynthetic partner^[12]

Pycnidium (plural pycnidia) – a flask-shaped, asexual fruiting body possessed by some lichens^[15]

Conidium (plural conidia) – an asexual fungal spore produced in pycnidia by some lichens^[16]

Rhizine – a root-like structure that anchors a lichen to the substrate on which it grows^[17]
Soralium (plural soralia) – a localized region or structure, typically a crack or pore, containing soredium^[18]

Soredium (plural soredia) – asexual reproductive propagules composed of loose aggregates of fungal hyphae and photobiont cells, produced in soralia^[18]

Thallus (plural thalli) – the vegetative body of a lichen, made up of both mycobiont and photobiont components^[19]

Cortex – the lichen's outer layer(s), made up of tightly woven fungal filaments^[20]
Isidium (plural isidia) – outgrowths of the thallus which contain photobiont cells and provide means of vegetative reproduction for some lichens^[21]
Medulla – a loose layer of interwoven fungal hyphae within the thallus^[22]
Podetium (plural podetia) – an upright secondary thallus, which serves as the fruiting structure of Cladonia species^[23]

Ecology

Symbiosis in lichens – the relationship between the lichen partners can be complicated; while generally mutualistic, sometimes it is not. Recent research also shows other partners, including bacteria and "accessory" fungi, may be involved^[24]
Reproduction in lichens – lichens reproduce both sexually and asexually, and both strategies are complicated by the presence of multiple partners
- Sexual reproduction in lichens – some lichens reproduce sexually using ascospores, which means they have to acquire their photobiont partners somehow after germinating^[25]
Lichens and nitrogen cycling – some lichens (in particular those with cyanobacteria as a photobiont) can fix nitrogen^[7]
Lichen biogeography – the study of the current distribution of extant lichens and the reasons for those distributions^[26]
Lichen resynthesis – lichens can be artificially "recreated" by combining partners in a lab^[27]
Lichens and pedogenesis – lichens contribute to the formation of soil by breaking down rock^[28]
- Biological soil crust – lichens are among the common dominant biota in biocrusts, one of the world's largest environmental community types in terms of area covered^[29]
Photosynthesis in lichens

Types of lichens

Lichen lists

Lichen taxonomical classifications

Lichen systematics – Although they are composite organisms, lichens have traditionally been classified on the basis of their fungal partner. These span eight different biological classes, 37 orders, 121 families, and around 1,000 genera.^[30]^[32]

Ascolichen – a lichen whose fungal partner is a member of the Ascomycota, one of the two main fungal divisions.^[33]
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 occurrence than ascolichens^[34]

Classes

Lichens fall into eight fungal classes and several subclasses:^[31]

Orders

They are split across 37 orders. Those which cannot be assigned to a particular order are assigned instead to "incertae sedis" within the appropriate class.^[31]

Families

They fall into 121 families. Those which cannot be assigned to a particular family are assigned instead to "incertae sedis" within the appropriate order.^[30]^[32]

Genera

Extant lichens are found in around 1000 genera. These were listed in Lücking, Hodkinson and Leavitt's 2016 treatise on the genera of lichenized fungi, except where otherwise noted.^[32]

Species

About 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.^[41]

Lichens, by growth form

Lichen growth forms – These vary depending on the species:

Byssoid – wispy, with the appearance of teased wool^[43]
Crustose – paint-like appearance that adheres tightly to the underlying substrate
- Areolate – crustose, but divided into rounded or polygonal pieces by means of cracks^[44]
- Calicioid – crustose growth with small fruiting bodies which resemble sewing pins
- Placodioid – crustose in the centre and lobed at the periphery^[44]
Filamentous – thin, threadlike growth, often with a matted appearance^[45]
Foliose – flattened, leafy appearance^[44]
Fruticose – shrubby, bush-like or coral-like appearance^[44]
Gelatinous – jelly-like appearance
Leprose – powdery or granular appearance^[44]
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 by the substrate on which they grow:

Bryophilous lichen – on mosses or liverworts^[46]
- Hepaticolous lichen – on liverworts^[48]
- Muscicolous lichen – on mosses^[49]
Corticolous lichen – on bark^[46]
- Ramicolous lichen – on twigs^[49]
Foliicolous lichen – on plant leaves^[46]
- Epiphyllous lichen – on the upper surface of a leaf^[50]
- Hypophyllous lichen – on the lower surface of a leaf^[50]
Lichenicolous lichen – on other lichens^[46]
Lignicolous lichen – on wood^[46]
Omnicolous lichen – on various different substrates, including manmade structures^[49]
Plasticolous lichen – on plastic^[51]
Saxicolous lichen – on stone^[46]
Terricolous lichen – on soil^[49]
Vagrant lichen – loose, on no substrate^[52]

Lichens, by region

Africa

Antarctica

Oceania

Pacific

South America

Photobiont

Photobiont – the photosynthetic partner in a lichen^[53]

Cyanolichen – a lichen with a cyanobacteria photobiont^[54]
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^[55]

List of lichen products

Study of lichens

Lichenology – the study of lichens^[57]

Acharius Medal – awarded for lifetime achievement in lichenology^[58]
Evolution of lichens – lichenization of fungi has occurred multiple times, and several pathways towards acquiring photobionts have arisen
- Fossil lichens
Exsiccata (plural exsiccatae) – a published set of preserved specimens, numbered and distributed with printed labels^[59]
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^[60]
Spot test (lichen) – chemical tests used to aid in species identification^[61]

Threats

Lichenicolous fungus – parasitic fungus that uses lichens as a host^[62]

List of lichenicolous fungi

Lichens as bioindicators – lichens are sensitive to various pollutants and can be thus be used as bioindicators^[63]

Lichens and air pollution – many lichens are sensitive to various forms of air pollution^[64]
Lichens and climate change – the inability of algae to quickly evolve means that climate change may adversely impact lichens^[65]

Lichens in culture

Edible lichen – some lichens have traditionally been used as food^[67]
Ethnolichenology – a branch of ethnobotany that studies human usage of lichens^[68]
Lichens in popular culture
- Trouble with Lichen – novel by John Wyndham

Lichen organizations

American Bryological and Lichenological Society (ABLS)

The Bryologist – peer-reviewed journal published by ABLS

Australasian Lichen Society

Australasian Lichenology – official publication of the Australasian Lichen Society

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

Bryological and Lichenological Working Group (Bryologische en Lichenologische Werkgroep, BLWG)

Buxbaumiella – peer-reviewed journal published by BLWG

Dutch Bryological and Lichenological Society

Lindbergia – peer-reviewed journal co-published by the Dutch Bryological and Lichenological Society and the Nordic Bryological Society

Independent lichenological journals

Asian Journal of Mycology – an international peer-reviewed journal published by Mae Fah Luang University in Thailand
Bibliotheca Lichenologica – scientific monographs on lichens and mosses
Hattoria – an international, peer-reviewed journal issued by Hattori Botanical Laboratory
International Journal of Mycology and Lichenology

Citations

^ ^a ^b Van Hoose 2021.
^ ^a ^b Favor 2005, p. 5.
^ Laundon 1986, p. 3.
^ Li et al. 2021, p. evab101.
^ ^a ^b ^c Purvis 2000, p. 9.
^ ^a ^b Laundon 1986, p. 2.
^ ^a ^b Henriksson & Simu 1971, p. 119.
^ Baron 1999, p. 14.
^ Honegger 1998, p. 197.
^ Silverstein, Silverstein & Silverstein 1996, p. 32.
^ Smith et al. 2009, p. 22.
^ ^a ^b Brodo, Sharnoff & Sharnoff 2001, p. 758.
^ Smith et al. 2009, p. 30.
^ Brodo, Sharnoff & Sharnoff 2001, p. 756.
^ Smith et al. 2009, p. 36.
^ Smith et al. 2009, p. 24.
^ Hawksworth, Sutton & Ainsworth 1983, p. 330.
^ ^a ^b Smith et al. 2009, p. 38.
^ 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.
^ Smith et al. 2020.
^ Bowler & Rundel 1975, p. 326–327.
^ Galloway 2012, p. 315.
^ Zakeri et al. 2022.
^ Syers & Iskandar 1973.
^ Green et al. 2018, p. 397.
^ ^a ^b ^c Wijayawardene et al. 2022.
^ ^a ^b ^c Lücking, Hodkinson & Leavitt 2016, p. 371.
^ ^a ^b ^c Lücking, Hodkinson & Leavitt 2016, p. 377–400.
^ Bendre 2010, p. 131.
^ Lepp.
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l ^m Lücking, Hodkinson & Leavitt 2017, p. 58.
^ Lücking & Kalb 2018.
^ Aptroot, Sparrius & Alvarado 2018, p. 12.
^ Wilk et al. 2021.
^ Kondratyuk et al. 2016.
^ Díaz-Escandón et al. 2021.
^ Lücking et al. 2009.
^ Dobson 2011, p. 470.
^ Kantvilas 1996, p. 229.
^ ^a ^b ^c ^d ^e The British Lichen Society 2022a.
^ Dobson 2011, p. 26.
^ ^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. 5.
^ Rikkinen 2015, p. 973.
^ Goga et al. 2018, p. 1.
^ Dobson 2011, p. 11.
^ Merriam-Webster.
^ International Association for Lichenology.
^ Pfister 1985, p. 1.
^ 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.

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External links

[FOOTNOTEVan_Hoose2021-1] Van Hoose 2021.

[FOOTNOTEFavor20055-2] Favor 2005, p. 5.

[FOOTNOTELaundon19863-3] Laundon 1986, p. 3.

[FOOTNOTELiHouXuShi2021evab101-4] Li et al. 2021, p. evab101.

[FOOTNOTEPurvis20009-5] Purvis 2000, p. 9.

[FOOTNOTELaundon19862-6] Laundon 1986, p. 2.

[FOOTNOTEHenrikssonSimu1971119-7] Henriksson & Simu 1971, p. 119.

[FOOTNOTEBaron199914-8] Baron 1999, p. 14.

[FOOTNOTEHonegger1998197-9] Honegger 1998, p. 197.

[FOOTNOTESilversteinSilversteinSilverstein199632-10] Silverstein, Silverstein & Silverstein 1996, p. 32.

[FOOTNOTESmithAptrootCoppinsFletcher200922-11] Smith et al. 2009, p. 22.

[FOOTNOTEBrodoSharnoffSharnoff2001758-12] Brodo, Sharnoff & Sharnoff 2001, p. 758.

[FOOTNOTESmithAptrootCoppinsFletcher200930-13] Smith et al. 2009, p. 30.

[FOOTNOTEBrodoSharnoffSharnoff2001756-14] Brodo, Sharnoff & Sharnoff 2001, p. 756.

[FOOTNOTESmithAptrootCoppinsFletcher200936-15] Smith et al. 2009, p. 36.

[FOOTNOTESmithAptrootCoppinsFletcher200924-16] Smith et al. 2009, p. 24.

[FOOTNOTEHawksworthSuttonAinsworth1983330-17] Hawksworth, Sutton & Ainsworth 1983, p. 330.

[FOOTNOTESmithAptrootCoppinsFletcher200938-18] Smith et al. 2009, p. 38.

[FOOTNOTEBrodoSharnoffSharnoff2001763-19] Brodo, Sharnoff & Sharnoff 2001, p. 763.

[FOOTNOTEBrodoSharnoffSharnoff2001757-20] Brodo, Sharnoff & Sharnoff 2001, p. 757.

[FOOTNOTEBrodoSharnoffSharnoff2001759-21] Brodo, Sharnoff & Sharnoff 2001, p. 759.

[FOOTNOTEBrodoSharnoffSharnoff2001760-22] Brodo, Sharnoff & Sharnoff 2001, p. 760.

[FOOTNOTEBrodoSharnoffSharnoff2001761-23] Brodo, Sharnoff & Sharnoff 2001, p. 761.

[FOOTNOTESmithDal_GrandeMuggiaKeuler2020-24] Smith et al. 2020.

[FOOTNOTEBowlerRundel1975326–327-25] Bowler & Rundel 1975, p. 326–327.

[FOOTNOTEGalloway2012315-26] Galloway 2012, p. 315.

[FOOTNOTEZakeriJunneJägerDostert2022-27] Zakeri et al. 2022.

[FOOTNOTESyersIskandar1973-28] Syers & Iskandar 1973.

[FOOTNOTEGreenPintadoRaggioSancho2018397-29] Green et al. 2018, p. 397.

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