Lichen Tuesday, June 30, 2009

Lichen-covered tree, Tresco, Isles of Scilly, UK. Grey, leafy Parmotrema perlatum on upper half of trunk; yellowy-green Flavoparmelia caperata on middle and lower half and running up the extreme right side; and the fruiticose Ramalina farinacea

"Lichenes" from Ernst Haeckel's Artforms of Nature, 1904

Lichens (pronounced /ˈlaɪkən/,^[1] sometimes /ˈlɪtʃən/^[2]) are composite organisms consisting of a symbiotic association of a fungus (the mycobiont) with a photosynthetic partner (the photobiont or phycobiont), usually either a green alga (commonly Trebouxia) or cyanobacterium (commonly Nostoc).^[3] The morphology, physiology and biochemistry of lichens are very different to that of the isolated fungus and alga in culture. Lichens occur in some of the most extreme environments on Earth—arctic tundra, hot deserts, rocky coasts and toxic slag heaps. However, they are also abundant as epiphytes on leaves and branches in rain forests and temperate woodland, on bare rock, including walls and gravestones and on exposed soil surfaces (e.g. Collema) in otherwise mesic habitats. Lichens are widespread and may be long-lived;^[4] however, many species are also vulnerable to environmental disturbance, and may be useful to scientists in assessing the effects of air pollution,^[5][6][7] ozone depletion, and metal contamination. Lichens have also been used in making dyes and perfumes, as well as in traditional medicines.

Overview

The body (thallus) of most lichens is quite different from those of either the fungus or alga growing separately, and may strikingly resemble simple plants in form and growth. The fungus surrounds the algal cells, often enclosing them within complex fungal tissues unique to lichen associations. In many species the fungus penetrates the algal cell wall, forming penetration pegs or haustoria similar to those produced by pathogenic fungi.^[3][8] Lichens are poikilohydric, capable of surviving extremely low levels of water content.^[9] However, the re-configuration of membranes following a period of dehydration requires several minutes at least. During this period a “soup” of metabolites from both the mycobiont and phycobiont leaks into the extracellar spaces. This is readily available to both bionts to take up essential metabolic products ensuring a perfect level of mutualism.^{[citation needed]} Other epiphytic organisms may also benefit from this nutrient rich leachate.^{[citation needed]} This phenomenon also points to a possible explanation of lichen evolution from its original phycobiont and mycobiont components with its subsequent migration from an aquatic environment to dry land.^{[citation needed]} During repeated periods of dehydration in an alga and the resultant leakage of beneficial metabolites to the adjacent aquatic fungus, the mutalistic “marriage” slowly becomes constant.^{[citation needed]}

The algal or cyanobacterial cells are photosynthetic, and as in higher plants they reduce atmospheric carbon dioxide into organic carbon sugars to feed both symbionts. Both partners gain water and mineral nutrients mainly from the atmosphere, through rain and dust. The fungal partner protects the alga by retaining water, serving as a larger capture area for mineral nutrients and, in some cases, provides minerals obtained from the substrate. If a cyanobacterium is present, as a primary partner or another symbiont in addition to green alga as in certain tripartite lichens, they can fix atmospheric nitrogen, complementing the activities of the green alga.

Algal and fungal components of some lichens have been cultured separately under laboratory conditions^{[citation needed]}, but in the natural environment of a lichen, neither can grow and reproduce without a symbiotic partner.^{[citation needed]} Indeed, although strains of cyanobacteria found in various cyanolichens are often closely related to one another, they differ from the most closely related free-living strains. ^[10] The lichen association is a close symbiosis: It extends the ecological range of both partners and is obligatory for their growth and reproduction in natural environments.^{[citation needed]} Propagules (diaspores) typically contain cells from both partners, although the fungal components of so-called "fringe species" rely instead on algal cells dispersed by the “core species.”

Lichen associations may be considered as examples of mutualism, commensalism or even parasitism, depending on the species. Cyanobacteria in laboratory settings can grow faster when they are alone rather than when they are part of a lichen. The same, however, might be said of isolated skin cells growing in laboratory culture, which grow more quickly than similar cells that are integrated into a functional tissue. However, from the work of Coxson (see above) mutualism would appear to best summarise our current knowledge.

Morphology and structure

Crustose and foliose lichens on a wall

Lichen on a stone.

Some lichens have the aspect of leaves (foliose lichens); others cover the substrate like a crust (crustose lichens) (illustration, right), others such as the genus Ramalina adopt shrubby forms (fruticose lichens), and there are gelatinous lichens such as the genus Collema.^[11] Although the form of a lichen is determined by the genetic material of the fungal partner, association with a photobiont is required for the development of that form. When grown in the laboratory in the absence of its photobiont, a lichen fungus develops as an undifferentiated mass of hyphae. If combined with its photobiont under appropriate conditions, its characteristic form emerges, in the process called morphogenesis (Brodo, Sharnoff & Sharnoff, 2001). In a few remarkable cases, a single lichen fungus can develop into two very different lichen forms when associating with either a green algal or a cyanobacterial symbiont. Quite naturally, these alternative forms were at first considered to be different species, until they were first found growing in a conjoined manner.

There is evidence to suggest that the lichen symbiosis is parasitic or commensalistic, rather than mutualistic (Ahmadjian 1993). However, this now needs to be re-examined in light of Coxon's work. The photosynthetic partner can exist in nature independently of the fungal partner, but not vice versa. Furthermore, photobiont cells are routinely destroyed in the course of nutrient exchange. The association is able to continue because photobiont cells reproduce faster than they are destroyed. (ibid.)

Under magnification, a section through a typical foliose lichen thallus reveals four layers of interlaced fungal filaments. The uppermost layer is formed by densely agglutinated fungal hyphae building a protective outer layer called the cortex, which can reach several hundred μm in thickness.^[12] This cortex may be further topped by an epicortex 0.6-1μm thick in some Parmeliaceae, which may be with or without pores, and is secreted by cells - it is not itself cellular.^[12] In lichens that include both green algal and cyanobacterial symbionts, the cyanobacteria may be held on the upper or lower surface in small pustules called cephalodia. Beneath the upper cortex is an algal layer composed of algal cells embedded in rather densely interwoven fungal hyphae. Each cell or group of cells of the photobiont is usually individually wrapped by hyphae, and in some cases penetrated by an haustorium. Beneath this algal layer is a third layer of loosely interwoven fungal hyphae without algal cells. This layer is called the medulla. Beneath the medulla, the bottom surface resembles the upper surface and is called the lower cortex, again consisting of densely packed fungal hyphae. The lower cortex often bears rootlike fungal structures known as rhizines, which serve to attach the thallus to the substrate on which it grows. Lichens also sometimes contain structures made from fungal metabolites, for example crustose lichens sometimes have a polysaccharide layer in the cortex. Although each lichen thallus generally appears homogeneous, some evidence seems to suggest that the fungal component may consist of more than one genetic individual of that species. This seems to also be true of the photobiont species involved.

Growth form

Lichens are informally classified by growth form into:

• crustose (paint-like, flat), e.g., Caloplaca flavescens
• filamentous (hair-like), e.g., Ephebe lanata
• foliose (leafy), e.g., Hypogymnia physodes
• fruticose (branched), e.g., Cladonia evansii, C. subtenuis, and Usnea australis
• leprose (powdery), e.g., Lepraria incana
• squamulose (consisting of small scale-like structures, lacking a lower cortex), e.g., Normandina pulchella
• gelatinous lichens, in which the cyanobacteria produce a polysaccharide that absorbs and retains water.

Reproduction and dispersal

Thalli and apothecia on a foliose lichen

Xanthoparmelia sp.

Many lichens reproduce asexually, either by vegetative reproduction or through the dispersal of diaspores containing algal and fungal cells. Soredia (singular soredium) are small groups of algal cells surrounded by fungal filaments that form in structures called soralia, from which the soredia can be dispersed by wind. Another form of diaspore are isidia, elongated outgrowths from the thallus that break off for mechanical dispersal. Fruticose lichens in particular can easily fragment. Due to the relative lack of differentiation in the thallus, the line between diaspore formation and vegetative reproduction is often blurred. Many lichens break up into fragments when they dry, dispersing themselves by wind action, to resume growth when moisture returns.

Many lichen fungi appear to reproduce sexually in a manner typical of fungi, producing spores that are presumably the result of sexual fusion and meiosis. Following dispersal, such fungal spores must meet with a compatible algal partner before a functional lichen can form. This may be a common form of reproduction in basidiolichens, which form fruitbodies resembling their nonlichenized relatives. Among the ascolichens, spores are produced in spore-producing bodies, the three most common spore body types are the apothecia, perithecia and the pycnidia.^[13]

For reproduction, lichen possess isidia, soredia, and undergo simple fragmentation. These structures are also composed of a fungal hyphae wrapped around cyanobacteria. (Eichorn, Evert, and Raven, 2005) While the reproductive structures are all composed of the same components(Mycobiont and Photobiont) they are each unique in other ways. Isidia are small outgrowths on the exterior of the lichen. Soredia are powdery propagules that are released from the top of the thallus.^[14] In order to establish the lichen, the soredia propagules must contain both the photobiont and the mycobiont^[15]

Growth and longevity

Lichenometry

Main article: Lichenometry

Lichenometry is a technique used to determine the age of exposed rock surfaces based on the size of lichen thalli. Introduced by Beschel in the 1950s,^[16] the technique has found many applications.

Ecology

Lichens must compete with plants for access to sunlight, but because of their small size and slow growth, they thrive in places where higher plants have difficulty growing. Lichens are often the first to settle in places lacking soil, constituting the sole vegetation in some extreme environments such as those found at high mountain elevations and at high latitudes.^{[citation needed]} Some survive in the tough conditions of deserts, and others on frozen soil of the Arctic regions.^{[citation needed]}

A major ecophysiological advantage of lichens is that they are poikilohydric (poikilo- variable, hydric- relating to water), meaning that though they have little control over the status of their hydration, they can tolerate irregular and extended periods of severe desiccation. Like some mosses, liverworts, ferns, and a few "resurrection plants", upon desiccation, lichens enter a metabolic suspension or stasis (known as cryptobiosis) in which the cells of the lichen symbionts are dehydrated to a degree that halts most biochemical activity. In this cryptobiotic state, lichens can survive wider extremes of temperature, radiation and drought in the harsh environments they often inhabit.

Lichens do not have roots and do not need to tap continuous reservoirs of water like most higher plants, thus they can grow in locations impossible for most plants, such as bare rock, sterile soil or sand, and various artificial structures such as walls, roofs and monuments. Many lichens also grow as epiphytes (epi- on the surface, phyte- plant) on other plants, particularly on the trunks and branches of trees. When growing on other plants, lichens are not parasites; they do not consume any part of the plant nor poison it. Some ground-dwelling lichens, such as members of the subgenus Cladina (reindeer lichens), however, produce chemicals which leach into the soil and inhibit the germination of plant seeds and growth of young plants. Stability (that is, longevity) of their substrate is a major factor of lichen habitats. Most lichens grow on stable rock surfaces or the bark of old trees, but many others grow on soil and sand. In these latter cases, lichens are often an important part of soil stabilization; indeed, in some desert ecosystems, vascular (higher) plant seeds cannot become established except in places where lichen crusts stabilize the sand and help retain water.

Pine forest with lichen ground-cover

The European Space Agency has discovered that lichens can survive unprotected in space. In an experiment led by Leopoldo Sancho from the Complutense University of Madrid, two species of lichen – Rhizocarpon geographicum and Xanthoria elegans – were sealed in a capsule and launched on a Russian Soyuz rocket on 31 May 2005. Once in orbit the capsules were opened and the lichens were directly exposed to the vacuum of space with its widely fluctuating temperatures and cosmic radiation. After 15 days the lichens were brought back to earth and were found to be in full health with no discernible damage from their time in orbit. ^[17][18]

Although lichens typically grow in naturally harsh environments, most lichens, especially epiphytic fruticose species and those containing cyanobacteria, are sensitive to manufactured pollutants. Hence, they have been widely used as pollution indicator organisms. When growing on mineral surfaces, some lichens slowly decompose their substrate by chemically degrading and physically disrupting the minerals, contributing to the process of weathering by which rocks are gradually turned into soil. While this contribution to weathering is usually benign, it can cause problems for artificial stone structures. For example, there is an ongoing lichen growth problem on Mount Rushmore National Memorial that requires the employment of mountain-climbing conservators to clean the monument.

Lichens may be eaten by some animals, such as reindeer, living in arctic regions. The larvae of a number of Lepidoptera species feed exclusively on lichens. These include Common Footman and Marbled Beauty. However, lichens are very low in protein and high in carbohydrates, making them unsuitable for some animals. Lichens are also used by the Northern Flying Squirrel for nesting, food, and a water source during winter.

Evolution

The evolution of lichens and the class Ascomycota is complex and not well understood, but because there are thirteen different orders of Ascomycetes, scientists generally believe that different lichens have evolved independently from one another through analogous evolution. Lichenized fungi have continued to evolve, developing differently than those that do not form lichens.^{[citation needed]}

Paleontology

The extreme habitats that lichens inhabit are not ordinarily conducive to producing fossils.^[19] Though lichens may have been among the first photosynthesizers to colonize land,^{[citation needed]} the oldest fossil lichens in which both symbiotic partners have been recovered date to the Early Devonian Rhynie chert, about 400 million years old.^[20] The slightly older fossil Spongiophyton has also been interpreted as a lichen on morphological^[21] and isotopic^[22] grounds, although the isotopic basis is decidedly shaky.^[23] It has been suggested - although not yet proven - that the even older fossil Nematothallus was a lichen.^[24]

It has also been claimed that Ediacaran fossils were lichens;^[25] although this claim was met with scepticism and has since been retracted by its author.^[24] A lichen-like symbiosis, however, has been observed in marine fossils from the Ediacaran, 600 million years ago.^[26]

Taxonomy and classification

Lichens are named based on the fungal component, which plays the primary role in determining the lichen's form. The fungus typically comprises the majority of a lichen's bulk, though in filamentous and gelatinous lichens this is not always the case. The lichen fungus is typically a member of the Ascomycota—rarely a member of the Basidiomycota, and then termed basidiolichens to differentiate them from the more common ascolichens. Formerly, some lichen taxonomists placed lichens in their own division, the Mycophycophyta, but this practice is no longer accepted because the components belong to separate lineages. Neither the ascolichens nor the basidiolichens form monophyletic lineages in their respective fungal phyla, but they do form several major solely or primarily lichen-forming groups within each phylum^[27]. Even more unusual than basidiolichens is the fungus Geosiphon pyriforme, a member of the Glomeromycota that is unique in that it encloses a cyanobacterial symbiont inside its cells. Geosiphon is not usually considered to be a lichen, and its peculiar symbiosis was not recognized for many years. The genus is more closely allied to endomycorrhizal genera.

The following table lists the orders and families of fungi within the Ascomycota that include lichen-forming species. Taxonomic classification is based on the "Outline of Ascomycota" (Dec 31st 2007).^[28]

Fungal Taxa
Order	Families
Acarosporales	Acarosporaceae
Agyriales	Agyriaceae, Anamylosporaceae
Arthoniales	Arthoniaceae, Chrysothricaceae, Melaspileaceae, Roccellaceae
Lecanorales	Catillariaceae, Cladoniaceae, Lecanoraceae, Parmeliaceae, Ramalinaceae, Stereocaulaceae
Lichinales	Gloeoheppiaceae, Heppiaceae, Lichinaceae, Peltulaceae
Ostropales	Gomphillaceae, Graphidaceae, Gyalectaceae, Stictidaceae, Thelotremataceae
Peltigerales	Collemataceae, Lobariaceae, Nephromataceae, Pannariaceae, Peltigeraceae, Placynthiaceae
Pertusariales	Megasporaceae, Pertusariaceae
Pyrenulales	Monoblastiaceae, Pyrenulaceae
Teloschistales	Letroutiaceae, Physciaceae, Teloschistaceae
Verrucariales	Verrucariaceae
Incertae sedis	Arthrorhaphidaceae (Ostropomycetidae), Arthopyreniaceae (Dothideomycetes), Elixiaceae (Lecanoromycetes), Microtheliopsidaceae (Dothideomycetes), Pyrenotrichaceae (Dothideomycetes), Lecideaceae (Lecanoromycetidae), Trypetheliaceae (Dothideomycetes)

Economic uses

Food

Iwatake (Umbilicaria esculenta) gathering at Kumano in Kishū by Hiroshige II

Umbilicaria esculenta (Japanese: Iwatake; Korean: Seogi) are collected from cliffs for use in a variety of traditional Korean and Japanese foods. Lichen flour is used by the Tarahumara as an ingredient of Tesguino.^[29]

Other Uses

Many lichens produce secondary compounds, including pigments that reduce harmful amounts of sunlight and powerful toxins that reduce herbivory or kill bacteria. These compounds are very useful for lichen identification, and have had economic importance as dyes such as cudbear or primitive antibiotics.

There are reports dating almost 2000 years of lichens being used to extract purple and red colors.^[30] Of great historical and commercial significance are lichens belonging to the family Roccellaceae, commonly called orchella weed or orchil. Orcein and other lichen dyes have largely been replaced by synthetic versions. The pH indicator litmus is a dye extracted from the lichen genus Rocella tinctoria by boiling.

Extracts from many Usnea^[31] species were used to treat wounds in Russia in the mid-twentieth century.

Gallery

Xanthoparmelia cf. lavicola, a foliose lichen, on basalt.	Usnea australis, a fruticose form, growing on a tree branch	Map lichen (Rhizocarpon geographicum) on rock	The cyanobacterium Hyella caespitosa with fungal hyphae in the lichen Pyrenocollema halodytes
Physcia millegrana (a foliose lichen), with an unlichenized polypore fungus (bottom right), on a fallen log.	Reindeer moss (Cladonia rangiferina)	Hypogymnia cf. tubulosa with Bryoria sp. and Tuckermannopsis sp. in the Canadian Rockies	Crustose lichens on limestone in Alta Murgia-Southern Italy
Cladonia cf. cristatella, a lichen commonly referred to as 'British Soldiers'. Notice the red tips.	Foliose lichens on rock growing outward and dying in the center. These lichens are at least several decades old.	Letharia sp. with Bryoria sp. on pine branches near Blackpine Lake, Washington	Xanthoria sp. lichen on volcanic rock in Craters of the Moon National Monument (Idaho, USA)
Lecanora cf. muralis lichen on the banks of the Bega canal in Timisoara	Tree covered with Yellow Lichen before and after treatment	Caloplaca marina, a marine lichen	Gravestone covered with lichen, Hingham, Massachusetts

Notes

1. ^ "Lichen". Oxford English Dictionary. Oxford University Press. 2nd ed. 1989.
2. ^ Cambridge Advanced Learner's Dictionary Second Edition, page 731. Cambridge University Press, 2005
3. ^ ^{a b} F.S. Dobson (2000) Lichens, an illustrated guide to the British and Irish species. Richmond Publishing Co. Ltd., Slough, UK
4. ^ Morris J, Purvis W. (2007). Lichens (Life). London: The Natural History Museum. p. 19. ISBN 0-565-09153-0.
5. ^ Ferry, B.W., Baddeley, M.S. & Hawkworth, D. L. (Editors) (1973) Air Pollution and Lichens. Athlone Press, London.
6. ^ D.L. Hawksworth and F. Rose(1976) Lichens as pollution monitors. Edward Arnold, Institute of Biology Series, No. 66. 60pp. ISBN 0713125543: 0713125551(pbk.)
7. ^ C.I. Rose & D.L. Hawksworth (1981) Lichen recolonization in London's cleaner air. Nature 289, 289-292.
8. ^ R. Honegger (1988) Mycobionts. Chapter 3 in T.H. Nash (ed.) (1996) Lichen Biology. Cambridge University Press. ISBN 0521 45368 2
9. ^ T.H. Nash (editor) (1996) Lichen Biology. Cambridge University Press. ISBN 0521 45368 2
10. ^ http://www.sciencemag.org/cgi/content/full/297/5580/357
11. ^ Smith, A.L. (1929). Lichens. Cambridge Botanical Handbooks. Cambridge University Press.
12. ^ ^{a b} Büdel, B.; Scheidegger, C. (1996). "Thallus morphology and anatomy". Lichen Biology: 37–64.
13. ^ [1]
14. ^ Eichorn, Susan E., Evert, Ray F., and Raven, Peter H. 2005. Biology of Plants. New York (NY):W.H. Freeman and Company. 289 p.1.
15. ^ Cook, Rebecca and McFarland, Kenneth. 1995. General Botany 111 Laboratory Manual. Knoxville (TN): University of Tennessee. 104 p.
16. ^ Beschel RE. (1950). "Flecten als altersmasstab Rezenter morainen."Zeitschrift für Gletscherkunde und Glazialgeologie 1: 152–161.
17. ^ http://www.esa.int/esaHS/SEMUJM638FE_index_0.html
18. ^ Sancho, L.G.; De La Torre, R.; Horneck, G.; Ascaso, C.; De Los Rios, A.; Pintado, A.; Wierzchos, J.; Schuster, M. (2007). "Lichens survive in space: results from the 2005 LICHENS experiment.". Astrobiology 7 (3): 443–54. doi:10.1089/ast.2006.0046. http://www.ncbi.nlm.nih.gov/sites/entrez?db=pubmed&uid=17630840&cmd=showdetailview&indexed=google. Retrieved on 2007-12-22.
19. ^ (University of California at Berkeley) Fossil Records of Lichens.
20. ^ Taylor, T.N.; Hass, H.; Remy, W.; Kerp, H. (1995). "The oldest fossil lichen". Nature 378 (6554): 244–244. doi:10.1038/378244a0. http://www.uni-muenster.de/GeoPalaeontologie/Palaeo/Palbot/nature.html.
21. ^ Wilson A. Taylor; Chris Free; Carolyn Boyce; Rick Helgemo; Jaime Ochoada (2004). "SEM Analysis of Spongiophyton Interpreted as a Fossil Lichen". Int. J Plant Sci 165: 875–881. doi:10.1086/422129. http://www.journals.uchicago.edu/doi/full/10.1086/422129?cookieSet=1.
22. ^ Jahren, A.H.; Porter, S.; Kuglitsch, J.J. (2003). "Lichen metabolism identified in Early Devonian terrestrial organisms". Geology 31 (2): 99–102. doi:10.1130/0091-7613(2003)031<0099:lmiied>2.0.CO;2.
23. ^ Fletcher, B.J.; Beerling, D.J.; Chaloner, W.G. (2004). "Stable carbon isotopes and the metabolism of the terrestrial Devonian organism Spongiophyton". Geobiology 2 (2): 107–119. doi:10.1111/j.1472-4677.2004.00026.x.
24. ^ ^{a b} Retallack, G.J. (2007). "Growth, decay and burial compaction of Dickinsonia, an iconic Ediacaran fossil" (PDF). Alcheringa: an Australasian Journal of Palaeontology 31 (3): 215–240. doi:10.1080/03115510701484705. http://www.informaworld.com/index/781217204.pdf. Retrieved on 2008-02-04.
25. ^ Retallack, G.J. (1994). "Were the Ediacaran Fossils Lichens?". Paleobiology 20 (4): 523–544. http://links.jstor.org/sici?sici=0094-8373(199423)20%3A4%3C523%3AWTEFL%3E2.0.CO%3B2-V. Retrieved on 2008-02-04.
26. ^ Yuan, X.; Xiao, S.; Taylor, T.N. (2005). "Lichen-Like Symbiosis 600 Million Years Ago". Science 308 (5724): 1017. doi:10.1126/science.1111347. PMID 15890881.
27. ^ Lutzoni et al. (2004). "Assembling the fungal tree of life: progress, classification, and evolution of subcellular traits". Amer J Bot 91: 1446–1480. doi:10.3732/ajb.91.10.1446.
28. ^ "Myconet". http://www.fieldmuseum.org/myconet/outline.asp#classTaph. Retrieved on 2009-02-04.
29. ^ http://www.utexas.edu/courses/stross/ant322m_files/tarahumara04.htm
30. ^ Casselman, Karen Leigh; Dean, Jenny (1999). Wild color: [the complete guide to making and using natural dyes]. New York: Watson-Guptill Publications. ISBN 0-8230-5727-5.
31. ^ http://www.tcbmed.com/Newsletters/Volume4-Issue4-Usnea.html|//www.tcbmed.com/Newsletters/Volume4-Issue4-Usnea.html

References

• Ahmadjian, V. (1993.). The Lichen Symbiosis.. New York: John Wiley & Sons.. pp. 250 pages.. ISBN 0-471-57885-1.
• Brodo, I.M., S.D. Sharnoff, and S. Sharnoff, 2001. Lichens of North America. Yale University Press, New Haven.
• http://www.newscientistspace.com/article/dn8297 Hardy lichen shown to survive in space
• http://www.lichen.com
• Gilbert, O. 2004. The Lichen Hunters. The Book Guild Ltd. England.
• Hawksworth, D.L. and Seaward, M.R.D. 1977. Lichenology in the British Isles 1568 - 1975. The Richmond Publishing Co. Ltd., Richomd, 1977.
• Kershaw, K.A. "Physiological Ecology of Lichens", 1985. Cambridge University Press Cambridge.
• Knowles, M.C. 1929. "The lichens of Ireland." Proceedings of the Royal Irish Academy 38:1 - 32.
• Purvis, O.W., Coppins, B.J., Hawksworth, D.L., James, P.W. and Moore, D.M. (Editors) 1992. The Lichen Flora of Great Britain and Ireland. Natural History Museum, London.
• Sanders, W.B. 2001. "Lichens: interface between mycology and plant morphology." Bioscience 51: 1025-1035.
• Seaward, M.R.D. 1984. "Census Catalogue of Irish Lichens." Glasra 81 - 32.