Erik Acharius, known as the "father of lichenology," coined many lichen terms still in use today around the turn of the 18th century. Before that, only a couple of lichen-specific terms had been proposed. Johann Dillenius introduced scyphus in 1742 to describe the cup-shaped structures associated with genus Cladonia, while in 1794 Michel Adanson used lirella for the furrowed fruitbodies of the genus Graphis. Acharius introduced numerous terms to describe lichen structures, including apothecium, cephalodium, cyphellae, podetium, proper margin, soredium, and thallus. In 1825, Friedrich Wallroth published the first of his multi-volume work Naturgeschichte der Flechten ("Natural History of Lichens"), in which he proposed an alternative terminology based largely on roots from the Greek language. His work, presented as an alternative to that of Acharius (of whom he was critical) was not well received, and the only terms he proposed to gain widespread acceptance were epi- and hypophloeodal, hetero- and homoiomerous, and gonidium, the last of which remained in use until the 1960s.[3] Until about 1850, there were 21 terms for features of the lichen thallus that remain in use today.[3]
The increasing availability of the optical microscope as an aid to identifying and characterizing lichens led to the creation of new terms to describe structures that were previously too small to be visualized. Contributions were made by Julius von Flotow (e.g. epithecium), Edmond Tulasne (e.g pycnidium), and William Nylander (e.g. pseudocyphella, thecium). Gustav Wilhelm Körber, an early proponent of using spore structure as a character in lichen taxonomy, introduced amphithecium, muriform, and "polari-dyblastae", later anglicized to "polari-bilocular" and then shortened to polarilocular.[4] In the next five decades that followed, many other additions were made to the repertoire of lichen terms, subsequent to the increased understanding of lichen anatomy and physiology made possible by microscopy. For whatever reasons, there were not any new terms (still currently used) introduced from the period 1906 to 1945, when Gustaf Einar Du Rietz proposed replacing epi- and hypothecium with epi- and subhymenium; all four terms remain in use.[4] In some cases, older terminology became obsolete as better understanding of the nature of the fungal–algal relationship led to changes in their terminology. For example, after Gunnar Degelius objected to the use of gonidia for the algal partner, George Scott proposed the use of mycobiont and phycobiont for lichen components, recommendations that were generally accepted by lichenologists.[5]
This glossary includes terms defining features of lichens unique to their composite nature, such as the major components the two major components of lichens (mycobiont and photobiont); specialized structures in lichen physiology; descriptors of types of lichens; two- and three-dimensional shapes used to describe spores and other lichen structures; terms of position and shape; prefixes and suffixes commonly used to form lichen terms; terminology used in methods for the chemical identification of lichens; the names of 22 standard insoluble lichen pigments and their associated reference species; and "everyday" words that have a specialized meaning in lichenology. The list also includes a few historical terms that have been supplanted or are now considered obsolete. Familiarity with these terms is helpful for understanding older literature in the field.
Referring to a developmental process in lichens where certain structures, such as spores or reproductive organs, fail to reach full development or maturity, often resulting in non-viable or malformed structures.[7]
A lichen product that is sometimes present, sometimes not present in a species. In literature, these are usually indicated with a ± symbol, e.g. ±usnic acid.[9]
Referring to fruticose lichens, a branching pattern that is unusual or abnormal, like that which sometimes occurs after the original branches are damaged in Cladonia.[8]
-al
A suffix used to indicate a relation to, or having the form and character of something.[20]
Plural alcobioses. A form of symbiosis involving algae and corticioid fungi, primarily occurring on bark and wood surfaces. In this relationship, algae form a layer beneath the fungal basidiomata—structures akin to the photobiont layer in lichens. This association, unlike in lichens, does not render the fungal partner nutritionally dependent on the algae, thus all involved fungal species are capable of surviving without the algal partner. Alcobiosis represents a diverse interaction, seen in various stages of coevolution, involving multiple species across the Agaricomycetes fungal group and three algal species from the class Trebouxiophyceae.[24]
alveolate
Used to describe a surface that has a pattern similar to a honeycomb (i.e. with more or less 6-sided hollows), where the surface appears to be composed of small pits or cavities like alveoli.[25]Compare: faveolate, foveolate, scrobiculate.
amphi-
A prefix used to indicate on both sides, or on all sides.[26]
A morphotype of corticolousthelotremoid lichens used to describe characteristics of apothecial and thallus morphology. Ampliotremoid lichens have
prominent apothecia with wide pores, black walls (viewed in microscopic section), and a smooth, more or less shiny thallus; this morphotype occurs in the genera Ampliotrema and Ocellularia.[30]
ampulliform
Bottle-shaped, i.e., with a narrow neck and swollen base.[31]
Also anisotomous.[33] Having branches of unequal length;[34] if the branching is anisotomic, one branch is typically stouter than the other, forming a main stem while the other appears like a lateral branch, as in the species Alectoria ochroleuca.[33]Contrast: dichotomous.
annulotremoid
A morphotype of corticolousthelotremoid lichens used to describe characteristics of apothecial and thallus morphology. Annulotremoid lichens have prominent apothecia with wide pores, pores with an inner ring, and a smooth, more or less shiny thallus; this morphotype occurs in the genera Myriotrema and Thelotrema.[30]
anticlinal
Perpendicular to a surface;[35] used to refer to hyphal alignment.
apical
Located at the highest point (the apex), the tip, or the end of something.[36]
apiculate
Having a short projection (an apicule) at one end; typically used to describe spore morphology.[37]
apothecium
Plural apothecia. A type of ascocarp that is open, saucer-shaped or cup-shaped, and in which the hymenium is exposed at maturity.[38] The term was first used by Erik Acharius in 1803.[3]
Also araneose, araneous.[40] Having a cobweb-like form, like that of the irregularly oriented and loosely interwoven hyphae of the medullary layer of some lichens.[41]
A shape or structure that is curved or arched like a bow.[43]
ardella
Plural ardelae. A type of apothecium, typical of lichens in the family Arthoniaceae, which is small and round. Elongated ardellae are called lirellae.[44] The term was first used by William Allport Leighton in 1854, who described an ardella as resembling a "sprinkled spot".[4]
Plural areolae. A small area, typically rounded to polygonal or irregular in shape, and often with a distinct texture. In a lichen thallus, the areolae are often separated from the rest of the thallus by fissures or cracks.[45]
Also ascogenic. Producing or supporting the growth of an ascus.[47]
ascolichen
A lichen in which the fungal partner (the mycobiont) is a member of the Ascomycota. About 98% of lichens are ascolichens.[50]See related: basidiolichen.
A morphotype of corticolousthelotremoid lichens used to describe characteristics of apothecial and thallus structure. This term refers to a morphotype of lichen where the apothecia are partially embedded and partially protruding, having a dark, hardened thalline margin that forms irregular cracks. This morphotype is uniquely seen in "Thelotrema" dislaceratum, a species with uncertain taxonomic placement.[58]
A lichen in which the fungal partner (the mycobiont) is a member of the Basidiomycota. About 0.4% of lichens are basidiolichens.[50]See related: ascolichen.
Referring to a type of ascus in which the ectotunica splits at the top and exposes the endotunica by forming an opening with a lip on each side; bilabiate asci occur in the genus Pertusaria.[63]
Also defined: endotunica, ectotunica. A type of ascus that has two functional layers, the internal layer, the endotunica, and the external layer, the ectotunica. Bitunicate asci are characteristic of the historical class Loculoascomycetes.[67]
A symbiotic interaction where either green algae or cyanobacteria are enveloped by fungal tissue, but without forming the discrete layers that occur in most lichens.[70]
botryose
Resembling rounded, bead-like structures or clusters resembling grapes.[43]
A lateral growth of the main stem of a thallus in usneoid lichens; various features of a branch are diagnostically valuable in distinguishing species.[71]
Also defined: hepaticolous lichen; muscicolous lichen. A lichen that grows on a moss or liverwort – i.e. on a bryophyte.[73] A hepaticolous lichen is found only on liverworts, while a muscicolous lichen is found only on mosses.[74][75]
bullate
Having blister-like or bubble-like swellings on a surface.[76]
Also caespitous, cespitose.[79] Growing in dense clusters or tufts, often used to describe fungi that arise from a common base or grow closely together without fusing.[18]
A lichen that prefers acidic soils and tends to avoid, or is intolerant of, alkaline conditions often associated with high calcium carbonate content; opposite of calcicole.[81]
Having one or more longitudinal grooves or channels.[83]
capitate
Having a well-formed head, usually spherical or hemispherical in shape.[68]See related: fuscocapitate.
capitulum
Plural capitula;also sphaeridium/sphaeridia. A more-or-less spherical or cup-shaped apothecium on the top of a stalk, found in the genera Calicium and Chaenotheca.[84][68]See related: mazaedium.
carbonized
Also carbonised, carbonaceous. Blackened and brittle tissue resulting from the accumulation of pigments.[85]
cariose
A lichen thallus or structure that is cracked, split, spongy, or otherwise decayed or in a state of disintegration.[43]
cartilaginous
Also cartilagineus. A term used to describe the texture of certain parts of a lichen. Cartilaginous structures have a texture similar to animal cartilage – firm but somewhat pliable, not brittle or soft.[86]
Plural cephalodia. A small gall-like structure that contains cyanobacteria, found in some lichens. These structures can be located on the lichen's upper or lower surface, or within the thallus itself.[68] These structures are found in most lichens that contain both algal and cyanobacterial photobionts.[91] The term was first used by Erik Acharius in 1803.[3]
cerebriform
Having a surface texture that is deeply wrinkled or convoluted in a manner resembling the structure of a brain. The term is used to characterize the appearance of certain lichens with a complex, highly folded thallus.[92]
cetrarioid lichen
An informal growth form category used for lichens with erect, foliosethalli, and apothecia and pycnidia on the margins of the lobes; characteristic of lichens previously classified in the genus Cetraria (in the broad sense).[93]
chalaroplectenchyma
Plural chalaroplectenchymata. A type of plectenchyma comprising loosely interwoven hyphae with holes; found in the medulla of some lichens.[92]
character
A distinguishing feature that is characteristic for an organism; equivalent to phenotypic trait.[92]
chasmoendolith
Also chasmoendolithic. A type of organism, typically a lichen or fungus, that lives within cracks and fissures of rocks.[94]See related: endolith.
checklist
A list of all of the species (sometimes including subspecies, varieties and forms) that occur within a particular region.[95]
chemosyndrome
A set of lichen products produced by a species; this typically includes one or more major compounds and a set of biosynthetically related minor compounds.[96]
A morphotype of corticolousthelotremoid lichens used to describe characteristics of apothecial and thallus structure. Chroodiscoid lichens have open apothecia with recurved lobules and a smooth and more or less shiny thallus; this morphotype occurs in the genera Acanthotrema and Chapsa.[30]
A shape or form that resembles a lemon or citrus fruit. The term is used to describe structures, particularly ascospores or other components of a lichen, that have an elongated, oval shape with tapered ends.[100]
CK test
A seldom-used spot test performed with an application of C followed immediately by K.[101]
cladoniiform lichen
Also cladoniform lichen, dimorphic lichen.Also defined: primary thallus and secondary thallus. A lichen with a two-fold growth form that includes both a crustose, squamulose, or foliose form and a fruticose form; the thallus differentiates into both horizontal (primary thallus) and vertical (secondary thallus, or podetium) structures. Cladoniiform lichens occur in the families Cladoniaceae and Baeomycetaceae.[102]
clavate
Also claviform. A shape resembling a club, broader at one end and tapering towards the other; typically used to refer to ascospores and asci.[103]
Having a finely scalloped edge; similar to crenate but with smaller notches.[119]
cruentodiscoid
A morphotype of corticolousthelotremoid lichens used to describe characteristics of apothecial and thallus morphology. Cruentodiscoid lichens have open apothecia with erect lobules and a pigmented disc, and a smooth, more or less shiny thallus; this morphotype occurs in the genus Chapsa.[30]
A form of growth where the lichen is pressed so tightly against the substrate upon which it grows that it is impossible to remove without destroying either it or part of the substrate. Crustose lichens have a cortex only on their upper surface.[120]
cryptoendolith
Also cryptoendolithic. A type of organism, particularly certain lichens and fungi, that live inside rocks or in the microscopic spaces within the mineral grains of rocks.[94]See related: endolith.
Closeup of the thallus underside of a Sticta lichen showing its crater-like cyphellae; scale bar = 1 mm
cyphella
Plural cyphellae. A sharply defined, rounded, ovate, or shapeless pore in the lower thallus surface (typically the lower cortex), which is lined with a "pseudocortex" made of loosely connected, non-gelatinized hyphae (often with globular cells, formed from the medulla) and bounded by a pale ring; known to occur in the genera Sticta and Oropogon.[129] The term was first used by Erik Acharius in 1799.[3]
Having small tooth-like projections or serrations along the edge.[134]
dextrinoid
A chemical property referring to a substance's ability to turn reddish-brown in the presence of Melzer's reagent or iodine due to the presence of dextrins. This color reaction is also called hemiamyloid or pseudoamyloid.[135]
determinate
Having well-defined or clearly marked edges.[136]Contrast: effuse.
diagnosis
A brief account of a taxon describing the essential characteristics that distinguish it from its relatives.[137]
diahypha
Plural diahyphae. A type of conidium formed from hyphae that split apically in several branches, with prominent constrictions at the septa, resulting in the appearance of chain links; found in the family Gomphillaceae.[138][139]
Also: disciform. A shape that is flat and circular, resembling a disk. In lichenology, this term often refers to the apothecia of lichens that have a flat, disk-like shape.[143]
Plural discothecia. The fruiting body of certain types of lichens, with cylindrical, bitunicate asci. It is distinguished from a hysterothecium, which is another type of fruiting body, by not opening through a slit but by expanding the asci to weather or push apart the typically thin upper stromatal layer.[145] The term was introduced by Richard P. Korf in 1962.[146]
disk
Also: disc. The curved or flat upper surface of the hymenium in an apothecium, often pigmented and surrounded by a margin or rim.[143]
distal
Positioned away from a point of origin or from the center of a body.[147]
distoseptum
Plural distosepta. A type of septum found in some conidia and ascospores, which is located within but distinct from the outer wall and surrounds the internal lumina. Structures with distosepta are said to be distoseptate.[148]
Referring to crustose, areolate lichens with marginal areoles that are extended and arranged radially;[154] also defined more generally as "obscurely lobed".[155][156]
A fungus that lives within the thallus of a lichen without producing any visible symptoms of disease; these fungi are transmitted horizontally.[165]
Lecidea laboriosa is an endolithic lichen; the thallus, hardly visible, grows under and around the rock crystals, while the apothecia are visible on the surface.
endolith
Also endolithic. A crustose lichen that grows in the interior of rocks (under and around the rock crystals), typically with little or no visible thallus on the outer rock surface.[166]Contrast: epilithic.See related: chasmoendolith, cryptoendolith, euendolith.
endophloeodal
Also endophloeodic, endophloeic, endophloic. Refers to crustose lichens whose thalli are more or less immersed in tree bark.[167]Contrast: epiphloedal.
Refers to margins or edges of lichen structures (such as apothecia, lobes, or thalli) that are smooth and unbroken, without any notches, teeth, or irregularities.[168]
epi-
Also ep-. A prefix meaning "upon" or "above".[169][170]
epibryophyllous
Also epibryophytic. Referring to organisms, particularly lichens or fungi, that grow on the surface of mosses (bryophytes).[169]
A thin tissue layer of interwoven hyphae situated directly above the hymenium, which can contain pigments and sometimes plays a role in the coloration of the lichen.[171]Compare: epithecium.
epilithic
Also petricolous, rupicolous, saxicolous. A crustose lichen that grows on the surface of rocks.[166]Contrast: endolithic.
Also perrumpent. Breaking through a surface.[178] A more precise definition has been suggested by Aptroot and Lücking, who propose that the term applies to ascomata and pseudostromata that are more than 1/2 to 3/4 above the level of the thallus.[179]
Plural eucortices or eucortexes. A cortex made of well-differentiated tissue.[181] Another sense of the term, used by Josef Poelt, refers to cortical tissue made entirely of fungal cells originating from a cambium-like tissue layer in or above the algal layer.[182] The term eucortex was first used by Gunnar Degelius in 1954.[5]
euendolith
Also euendolithic. A type of organism, often a lichen or microbe, that actively bores into and resides within the mineral matrix of rocks or other hard substrates.[94]See related: endolithic.
Plural exsiccata, exsiccatae, exsiccati. A dried and labeled herbarium specimen, often part of a numbered set.[183]
excipulum
Alsoexciple. Plural excipula. The cup-shaped or ring-shaped layer of tissue supporting the hymenium in an apothecium; this tissue sometimes develops into a distinct margin, as in the lecanorine apothecia.[184]See related: proper margin, thallin margin.
A region of the cortex where the hyphae are aligned perpendicularly to the main axis of the thallus.[190] The term was first used by Auguste-Marie Hue in 1906.[5]See related: palisade cell.
A form of bitunicate ascus in which the flexible layers of the inner wall (the endotunica) and the more rigid layers of the outer wall (the ectotunica) are physically separated; as a consequence, the inner walls extend past the outer walls before the spores are released.[198]
fissurine
Also fissurate. A term used characterize a structure or surface displaying a pattern of narrow, elongated cracks or fissures.[199]
fissurinoid
A morphotype of corticolousthelotremoid lichens used to describe characteristics of apothecial and thallus structure. This morphotype is somewhat similar to the chroodiscoid, but it differentiates by the way the apothecia open through irregular thallus cracks, finally resembling chroodiscoid apothecia. It often has a unique elongated form at maturity and can be seen in species such as Acanthotrema brasilianum and various Chapsa species.[200]
Leaf-like; a type of lichen thallus comprising numerous small leafy lobes, often extending in a roughly circular pattern from a center of growth, on a lower cortex that is attached to the substrate by rhizines or at a base.[205][206]
forage lichen
Lichens that serve as important food sources for fauna. For example, species from the genera Alectoria, Bryoria, and Cladonia are winter forage lichens for caribou in northern North America.[207][208]
The outer gelatinous layer, also known as the g-layer, found on the exterior of an ascus, often exhibiting a gelatinous consistency and staining blue in iodine. Typically present in all asci, the fuzzy coat usually forms a thin layer along the ascus sides but may also appear as an apically thickened cap.[216][217]
A morphotype of corticolousthelotremoid lichens used to describe characteristics of apothecial and thallus morphology. Glaucescentoid lichens have open apothecia with erect lobules, and a rough thallus containing crystals; this morphotype occurs in the species Leucodecton glaucescens.[30]
glaucophaenoid
A morphotype of corticolousthelotremoid lichens used to describe characteristics of apothecial and thallus morphology. Glaucophaenoid lichens have prominent apothecia with wide pores, pale walls (viewed in microscopic section), and a smooth and more or less shiny thallus; this morphotype occurs in the genus Myriotrema.[30]
A now-obsolete, historical term for the algal layer in a lichen.[225]
gonidium
Plural gonidia. A now-obsolete, historical term for a lichen photobiont.[225] The term was first used by Friedrich Wallroth in 1825, and supplanted in the 1960s.[3]
gonimium
Plural gonimia. A now-obsolete, historical term for a lichen cyanobiont.[225]
A term for the general appearance (the habit) of a lichen.[227]
grumose
Having a granular or crumbly texture or appearance.[43]
guttulate
Referring to structures containing small oil droplets (guttules); often used to describe spores. More precisely, spores can be described as uni-, bi-, tri-, or multiguttulate.[228]
Also gyrate. Curved backward and forward; with folds and undulations.[230]
gyrotremoid
A morphotype of corticolousthelotremoid lichens used to describe characteristics of apothecial and thallus morphology. Gyrotremoid lichens have open apothecia with recurved lobules, a disc that is pigmented with concentric rings, and a smooth, more or less shiny thallus; this morphotype occurs in the genus Gyrotrema.[30]
A flattened attachment point formed through direct contact of a lichen thallus to its substrate; associated with foliose and fruticose lichens that lack other attachment organs, such as Hypogymnia.[231]
halonate
Also defined: halo. Referring to a spore that is surrounded by a transparent outer layer or a gelatinous, translucent sheath. This sheath is readily observed when the spore is stained with India ink, as the ink does not penetrate the mucilage of the sheath, creating a light-transparent halo that is visible against a blackened background.[232][233]
Also heter-. A prefix meaning "other" or "different".[235]
heterocyst
A specialized type of cell found in some cyanobacteria; heterocysts are thought to be involved in the fixation of nitrogen by the lichen thallus, as well as in the multiplication of cyanobacteria.[236]
heteromerous
A lichen thallus that is organized into discrete layers or strata; the term applies to the majority of foliose, squamulose, and crustose lichens.[237]
Also hormocyte. Also defined: hormocystangium, plural hormocystangia. An asexual propagule, produced in a cup-shaped structure called the hormocystangium, comprising heavy, gelatinous fungal hyphae enclosing a few cyanobiont cells; these structures occur in some gelatinous lichens in the family Collemataceae.[242] Both terms, hormocyst and hormocystangium, were introduced by Gunnar Degelius in 1945.[5] According to one source, hormocyte and hormocytangium (spelled without the "s") are more accurate terms, because the cells (indicated by the ending -cyte) are not sexual propagules (which is implied by the ending -cyst).[243]
Embedded or sunken into the surface; in lichens, often used to describe perithecia.[253] A more precise definition has been suggested by Aptroot and Lücking, who propose that the term applies to ascomata and pseudostromata that are more than 3/4 beneath the level of the thallus.[179]
Also defined: inspersion.Also inspers, interspersed. Terms used to describe the presence of minute, scattered oil droplets or granules within a tissue, typically observed in section with a compound microscope.[256] An inspersion refers to the degree of obstruction caused by the sprinkling of small particles within different sections of a given structure.[257]
A structure that resembles an isidium, but is formed as an outgrowth of the medulla rather than the cortex; associated with soralia of species in the genus Usnea.[260]
Plural isidia. A propagule that is an outgrowth of the thallus; it has a cortex and contains photobiont cells.[259] The term was first used in the sense it is used now by Georg Meyer in 1825, and adopted by Elias Fries in 1831.[3]
isodiametric
Having roughly equal dimensions in all directions, resulting in a roughly spherical or cube-like shape; often used to describe cells that are not elongated or flattened but maintain a uniform size across their width, height, and depth.[261]
Plural jugae. A tiny carbonized structure made of hyphal tissue, visible as a black dot, line, or ridge, on or in a thallus; associated with the genus Verrucaria.[264]
Also LCB. A histologicalstain commonly used to prepare semi-permanent slides. With this reagent, fungal hyphae stain blue, and algal cells stain deep blue to blue-green.[101]
lacunose
Also lacunar, lacunous. A texture that appears pitted or containing gaps or holes, giving it a somewhat rough or uneven appearance.[269]
lageniform
Flask-shaped; with a swollen base tapering to a narrow top.[270]
lamella
Plural lamellae. In the genus Umbilicaria, lamellae are flattened plate- or strap-like structures that project downward from the thallus undersurface.[271]
Also lecanoroid. An apothecium in which the disk is surrounded by a pale thalline margin, which has both algal and fungal cells, as in the genus Lecanora. The term is also used more generally to refer to crustose lichens of the order Lecanorales that have rounded apothecia with thick, protruding margins.[274]
lecideine
Also lecideoid. An apothecium in which the disk lacks a thalline margin, as in the genus Lecidea. The term is also used more generally to refer to apothecia with a blackened (carbonaeous) ring and a blackish disk.[275]
leiodisk
Also leiodisc. A disk of an apothecia that is smooth and without folds or protrusions.[276] The term was introduced by George Llano in 1950.[5]
A morphotype of corticolousthelotremoid lichens used to describe characteristics of apothecial and thallus morphology. Leprocarpoid lichens have open apothecia with erect lobules and a mealy and more or less matte thallus; this morphotype occurs in the genus Chapsa.[30]
A morphotype of corticolousthelotremoid lichens used to describe characteristics of apothecial and thallus morphology. Leptotremoid lichens have immersed apothecia with small pores, and a rough thallus containing crystals; this morphotype occurs in the genus Leptotrema.[30]
leucodectonoid
A morphotype of corticolousthelotremoid lichens used to describe characteristics of apothecial and thallus morphology. Leucodectonoid lichens have closed apothecia with tiny pores, and a rough thallus containing crystals; this morphotype occurs in the genus Leucodecton.[30]
The name of a historical class that contained all of the lichen-forming fungi. This name was used when it was still erroneously believed that these fungi were quite separate from the non lichen-forming fungi; now obsolete.[279]
Plural lirellae.Also hysterothecia, lirelline apothecia. A linear ascocarp, which may be straight, curved, branched, or flexuous, with a longitudinal slit; characteristic of lichens in the genus Graphis.[282] The term was first proposed by Michel Adanson in 1794.[3]
A lichen with a thallus large enough that its main characteristics can be identified without the use of viewing magnification; generally refers to foliose, squamulose, and fruticose species.[289]
Cuff-shaped; the term is used to describe soralia that break open to form a central perforation revealing a duct to the medullary cavity. Maniciform soralia occur in the genera Hypogymnia and Menegazzia.[291]
A morphotype of corticolousthelotremoid lichens describing a type of apothecium structure resembling topeliopsidoid apothecia but characterized by a minimized thallus margin. The distinctive feature of melanotopelioid apothecia is the presence of "teeth" that are black and carbonized, at least on the external surface.[296]
melanotremoid
A morphotype of corticolousthelotremoid lichens used to describe characteristics of apothecial and thallus morphology. Melanotremoid lichens have prominent apothecia with wide pores and a black margin; pore filled with broad "stump" (columella); black walls (viewed in microscopic section); and a smooth, more or less shiny thallus. This morphotype occurs in the genera Melanotrema, Ocellularia, Clandestinotrema, and Trinathotrema.[30]
Also microcrystal test. A method used to identify some lichen products that involves re-crystallization on a microscope slide from a range of solvents and the formation of crystals with characteristic shapes; the crystals are examined microscopically for identification. Although this technique has largely been supplanted by the more reliable and sensitive technique of thin-layer chromatography, there are certain situations where it is still useful.[298]
microlichen
A small lichen whose physical features cannot be distinguished without the aid of a 10X or greater viewing magnification;[299] it generally refers to crustose and foliicolous species.[300] The prefix "micro-" is also used to indicate small versions of particular growth forms, e.g. microfruticose or microfoliose.
Also defined: mycophycobiont. A symbiosis where an ascomycete fungus is housed inside multicellular algae; the algae and fungus involved in this association are called mycophycobionts.[309] Contrary to a lichen symbiosis, the fungal partner is the inhabitant, and the algal partner dominates.[310]
myriotremoid
A morphotype of corticolousthelotremoid lichens used to describe characteristics of apothecial and thallus morphology. Myriotremoid lichens have immersed apothecia with small pores, and a smooth and more or less shiny thallus; this morphotype occurs in the glaucopallensgroup of genus Myriotrema.[30]
Egg-shaped, with the narrower end at the base.[311]
obpyriform
Shaped like an inverted pear.[312]See related: pyriform.
ocellularioid
A morphotype of corticolousthelotremoid lichens used to describe characteristics of apothecial and thallus morphology. Ocellularioid lichens have prominent apothecia with wide pores; pores with a "finger" (columella); black walls (viewed in microscopic section); and a smooth, more or less shiny thallus. This morphotype occurs in the genus Ocellularia.[30]
-oid
A suffix meaning "like" or "having the form of".[313]
A small pore or opening; in lichens, it is used to refer to the paraphysis-lined cavity in a parathecium that ends in a pore, or more generally to any pore from which spores are released from an ascus-bearing fruit body.[315]
ostropalean
Referring to asci that are unitunicate with a thickened apex and a narrow canal ending in a pore; associated with species in the order Ostropales.[316]
Plural palisade plectenchymata. Also palisadoplectenchyma, plural palisadoplectenchymata. A type of plectenchyma in a cortex where the hyphae are arranged perpendicularly to the plane of the thallus.[319]
pallidostegoboloid
A morphotype of corticolousthelotremoid lichens used to describe characteristics of apothecial and thallus morphology. Pallidostegoboloid lichens have prominent apothecia with wide pores; pore filled with irregular structures; pale walls (viewed in microscopic section); and a smooth, more or less shiny thallus. This morphotype occurs in wrightiigroup of the genus Stegobolus.[30]
An organism that lives in close association with a host species, deriving benefits at the host's expense but not causing immediate harm; an intermediate state between symbiosis and parasitism.[267]
An informal growth form category used for lichens that are mostly foliose, often closely attached to the substrate, and have laminalapothecia and pycnidia; this group of features is characteristic of lichens previously classified in the genus Parmelia (in the broad sense).[93]
Also clypeate, scutiform. Referring to a rounded structure attached on the lower side at a single central point (often on a short stalk), with free edges.[330]
peltidiangium
A basket-like structure in which peltidia are produced.[331]
Plural periphyses. A short, sterile hypha that develops from above the ascus and grows down a short distance, typically lining the internal walls of the ostiole in a perithecium.[335]
Also defined: exospore, epispore, mesospore, endospore. The colorless and usually gelatinous outermost layer of a spore.[336] This transparent layer determines the spore's shape.[337] The other four layers of a spore, going inward, are the exospore, epispore, mesospore, and endospore.[336]
Plural perithecia. A spherical or flask-shaped ascocarp that is sessile or partly immersed in the thallus, with a single opening (ostiole) and enclosed by a distinct wall; a characteristic of pyrenolichens.[338] Although it was in 1831 that Elias Fries first applied the term perithecium to lichen fruit bodies, the word was originally coined by Christiaan Hendrik Persoon in 1794.[339]
Plural phenocortices, phenocortexes. A structure, similar to a cortex, containing hyphal fragments and dead, collapsed algal cells sloughed off from the algal layer.[341]
photobiont
Also defined: phycobiont, cyanobiont. The photosynthetic component of a lichen. This can be either a green alga (known as a phycobiont) or a cyanobacteria (known as a cyanobiont).[342] The term "phycobiont" was proposed by George Scott in 1957.[5]
Plural phyllidia. A small leaf-like or scale-like propagule that is corticate and has distinct upper and lower sides (i.e., it is dorsiventral); it originates from the margins or on the upper surface of thallus.[346] Phyllidia occur in some species of the Lecanorales and the Peltigerales.[347]
Characterised by longitudinal folds forming pleats, often used to describe closely adjacent, markedly convex thallus lobes or elongated areoles that display a "folded" appearance.[355]
plurilocular
Also multilocular. Having many cavities or locules; used to describe spore structure.[356]
A lichen test performed by shining a polarized light at a lichen structure in microscopic view; in the genus Hypogymnia, the presence (POL+) or absence (POL−) of POL-sensitive crystals in the hypothecium is a useful character to help distinguish species.[357]
polarilocular
Also placodiomorphic,[252]polar-diblastic, polaribilocular, polocellate. A spore divided into two components (locules) separated by a central septum with a perforation or isthmus.[358][359] The term was first used by Wilhelm Körber in 1855 (as "polari-dyblastae") to describe the spores of Rhizocarpon and Umbilicaria. It was anglicized to "polari-bilocular" by William Mudd in 1861, and finally shortened to polarilocular by the Henri Olivier in 1882.[4]
A morphotype of corticolousthelotremoid lichens used to describe characteristics of apothecial and thallus morphology. Praestantoid lichens have large and prominent apothecia with small pores; pores with "finger" (columella); black walls (viewed in microscopic section); and a smooth, more or less shiny thallus. This morphotype occurs in the praestansgroup of the genus Ocellularia.[30]
Sticking out from the surface of the thallus.[363] A more precise definition has been suggested by Aptroot and Lücking, who propose that the term applies to ascomata and pseudostromata that are more than 1/2 above the level of the thallus, and have a base that is expanded outwards.[179]
proper margin
Also proper exciple, ectal excipulum, medullary excipulum, proprium. A ring of tissue around the disk of a lecanorineapothecium; this tissue, which originates from the medulla, is not lichenized, and is internal to the thalline margin (if present).[364] The term "proper margin" was first used by Erik Acharius in 1803; in 1825 Elias Fries changed the noun and called it "proper exciple".[3]
Plural prosenchymata. A type of plectenchyma in which the constituent fungal hyphae are arranged parallel to each other, such that individual hyphae can be clearly distinguished using microscopy.[365]
prosoplectenchyma
Plural prosoplectenchymata. A type of plectenchyma, common in the thalluscortex of lichens, in which the constituent fungal hyphae are aligned in a particular direction.[365]
A fungal layer upon which an algae-containing thallus may develop, lacking photobiont; usually white, brown, or black, and found between the areoles and at the growing margins of crustose lichens.[366] The term was first used by Georg Meyer in 1825.[3]
prototunicate
A form of unitunicateascus in which the wall breaks down before maturity (thus releasing its ascospores), and which lacks differentiated apical structures.[367]
proximal
Positioned close to a point of origin or near the center of a body.[368]
pruina
A powdery, frost-like or flour-like deposit on a surface. In lichens, pruina is often the result of the accumulation of crystalline hydrates of calcium oxalate, of lichen products, or sometimes of the dead or dying cells of the epinecral layer.[369]
Also pseud-. A prefix meaning "false";[370] used in terminology to denote something is false, or that one structure resembles something else, such as the pseudocyphella resembling the cyphella.[368]
Plural pseudocortices, pseudocortexes. A boundary layer of the thallus containing distinct hyphae that are not organized into a regular tissue structure;[371] sometimes used to refer to the false cortex present on the outer layer of pseudopodetia, such as those found in the lichen Pycnothelia papillaria.[372]
Plural pseudocyphellae. Small openings in the cortex of a lichen, where the medulla is exposed to air, and there are no specialized cells surrounding the cavity.[373] The term was first used by William Nylander in 1858.[4]
Also cataphysis. A paraphysis-like hypha that forms in the locule or perithecial cavity before the formation of the ascus; it grows downward from the top of the cavity to the base of the ascomata.[375]
A lichen with flask-shaped fruiting bodies (perithecia) that develop from the fungal partner. Originally thought to form a natural group, molecular studies have shown pyrenocarpous lichens to be highly polyphyletic, evolving independently in multiple fungal lineages. Most belong to Chaetothyriomycetidae (e.g., Verrucariales, Pyrenulales), with some in Dothideomycetes (e.g., Arthopyreniaceae, Trypetheliaceae) and others unusually placed in Lecanoromycetes (e.g., Porinaceae, Protothelenellaceae, Thelenellaceae). Pyrenocarpous lichens are notably absent from the classical pyrenomycete class Sordariomycetes. This diverse group demonstrates that ascoma morphology alone is insufficient for determining evolutionary relationships, with convergent evolution of perithecial fruiting bodies occurring multiple times across fungal lineages.[386]
Also recurvate, reflexed.[391] Curved or bent back; in lichens, these terms are used to describe the tips of branches or lobes that are curved up or down, or back onto themselves.[392]
redingerioid
A morphotype of corticolousthelotremoid lichens used to describe characteristics of apothecial and thallus morphology. Redingerioid lichens have immersed apothecia with linear slit; slit filled with irregular structures; black walls (viewed in microscopic section); and a smooth, more or less shiny thallus. This morphotype occurs in the genera Redingeria and Stegobolus.[30]
reimnitzioid
A morphotype of corticolousthelotremoid lichens used to describe characteristics of apothecial and thallus morphology. Reimnitzioid lichens have open apothecia with erect lobules, and a rough thallus containing crystals; this morphotype occurs in the genus Reimnitzia.[30]
A single hyphal strand on the thallus underside that serves as an attachment organ.[396]
rhodostromoid
A morphotype of corticolousthelotremoid lichens used to describe characteristics of apothecial and thallus morphology. Rhodostromoid lichens have large and prominent apothecia with small pores; pore with “finger” (columella); black walls and pigment (viewed in microscopic section); and a smooth, more or less shiny thallus. This morphotype occurs in the rhodostromagroup of the genus Ocellularia.[30]
A morphotype of corticolousthelotremoid lichens used to describe characteristics of apothecial and thallus morphology. Schizotremoid lichens lack apothecia and have schizidia, and a smooth, more or less shiny thallus; this morphotype occurs in the genus Stegobolus.[30]
Plural scleroplectenchymata. A type of plectenchyma comprising thick-walled hyphae that are stuck closely together; present as a component of the stereoma tissue supporting the thallus in the genera Cladonia and Alectoria.[408]
scrobiculate
Having large, shallow depressions that are narrowly separated by rounded ridges.[191]Compare: faveolate, foveolate.
Lacking a stem.[414] A more precise definition has been suggested by Aptroot and Lücking, who propose that the term applies to ascomata and pseudostromata that are more than 3/4 above the level of the thallus, with a constricted base.[179]
sibling species
Closely related species that are morphologically indistinguishable but can be distinguished by non-morphological traits, such as chemistry or genetic differences. Initially defined as species recognized mainly through cryptic or non-morphological discontinuities, the concept has evolved to specifically refer to cryptic species that form a monophyletic group, meaning they share a common ancestor not shared with any other species. This concept is a subset of the broader terms species pair and "cryptic species".[415]
sigmoid
Curved upon itself twice, like the letter "S".[416]
simple
Lacking branches or divisions; in lichenology, it is used to describe structures such as thalli or rhizines, or spores without septa.[417]
Plural soralia. A part of the thallus where the cortex has cracked or broken down and soredia are produced. Soralia can be further characterized as diffuse if they are spread out on the upper thallus surface as a continuous layer, or delimited if they are confined to a more restricted area. If soralia originate in tubercules they are tuberculate, while they are fissural if they are created in fissures.[420] The term was proposed by Johannes Reinke in 1895.[5]
Plural soredia. A powdery to granular reproductive propagule that is not covered with a well-defined cortex (in contrast to isidia, and contains both algal (photobiont) and fungal (mycobiont) components.[420] The term was first used by Erik Acharius in 1803.[3]
Two lichen species that are identical morphologically, anatomically, and chemically, but can be distinguished by their sexual versus asexual reproductive strategies; the fertile taxon is known as the primary species, while the vegetatively reproducing taxon is known as the secondary species.[362] The use of molecular methods to analyze putative species pairs has shown that the underlying phylogenetic situation is more complex than had been assumed, and not necessarily correlated with reproductive strategy.[422]See related: sibling species.
spermogonium
Also spermagone, spermagonium. In lichenology, an obsolete term for pycnidium.[423]
A small spine; in some fruticose lichens of the Lecanoromycetes, it refers to a small cylindrical outgrowth, with a narrow base, in which the central axis is not connected with the central axis of the main branch.[424]
spinulose
Also spinulous. Covered with or having small spines (spinules) or spiny projections.[424]
A lichen species that is extremely similar (or identical) in external morphology, anatomy, chemistry, and spore size to another, but that is placed in a different genus solely based on differences in spore septation and/or spore colour. The term was introduced by Michael Wirth and Mason Hale in their 1978 monograph about the Graphidaceae, a family in which sporomorphs are common.[428]
A spot analysis used to help identify lichens; it is performed by placing a drop of a reagent on different parts of the lichen and noting any color change associated with application of the reagent. The four most common tests are C, K, KC, and PD.[430]
Brush-like, with many short, more or less perpendicular lateral branches. In lichenology, used to refer to rhizine structure.[432]
stegoboloid
A morphotype of corticolousthelotremoid lichens used to describe characteristics of apothecial and thallus morphology. Stegoboloid lichens have prominent apothecia with wide pores; pore filled with irregular structures; black walls (viewed in microscopic section); and a smooth, more or less shiny thallus. This morphotype occurs in the genus Stegobolus.[30]
Plural stromata. A dense mass of vegetativehyphae that supports spore-bearing structures.[434] In lichens, the stroma is often hard and carbonaceous.[396]
sub-
A prefix meaning "below", "under", "somewhat, or "almost".[435] Also used in front of names of taxonomic ranks to indicate intermediate categories, e.g. subspecies or subgenus.[436]
Also subicule.Plural subicula. A layer of loosely-compacted mycelia that covers the substrate and cushions fruiting bodies such as apothecia and perithecia.[437] The texture of the subiculum can be described as net-like, wool-like, or crust-like.[435]
Also substratum; plural substrata. The surface or base upon which a lichen grows or is attached. Although the terms substratum and substrate are often used equivalently in lichenology, the latter term has different meanings in microbiology and in enzymology.[438]
subulate
Slender and narrowing to a fine point; awl-shaped.[439]
A morphotype of corticolousthelotremoid lichens used to describe characteristics of apothecial and thallus morphology. Tenuitremoid lichens have immersed apothecia with small pores and a black margin; pore with “finger” (columella); black walls (viewed in microscopic section); and a smooth, more or less shiny thallus. This morphotype occurs in the genus Clandestinotrema.[30]
tegulicolous lichen
A lichen that lives on tiles.[441] In general, these are calcicolous lichens or lichens that are indifferent as to their substrate.[442]
Describing a cylindrical or rod-like structure that is round in cross-section.[419]
teretiform
Describing a shape that is nearly cylindrical or rod-like, similar to but not exactly terete; circular in cross-section, gradually tapering towards one end.[444]
Also thalline exciple, excipulum thallinum. A rim of tissue around the disk of a lecanorineapothecium; this tissue, external to the inner proper margin, is made of tissue with a structure similar to that of the thallus.[446] The term "thalline margin" was first used by Erik Acharius in 1803; in 1825 Elias Fries changed the noun and called it "thalline exciple".[3]
Plural thalloconidia. A dark brown, smooth to wrinkled propagule arising directly from a thallus, particularly the lower cortex and/or the rhizines. They are found in some species of Umbilicaria, and similar structures arise from the prothallus of some species in the genera Protoparmelia, Rhizoplaca, and Sporastatia. Thalloconidia have distinct cell layers in their walls, and comprise between 1 and about 2500 cells.[448][449]
Plural thalli. The body of a lichen,[451] made up of both fungal and algal or cyanobacterial cells.[452] The term was first used by Erik Acharius in 1803.[3]
A morphological group of lichens within the Graphidaceae, the largest family of crustose lichens.[454] Thelotremoid lichens are characterized by immersed-erumpent, rounded ascomata, non-branched to slightly branched paraphyses, mostly distoseptate ascospores, and mostly a prosoplectenchymatousexcipulum. Thelotremataceae, a traditional family of lichens, has been included in Graphidaceae, and its species are now informally accepted as thelotremoid lichens.[454]
tholus
Plural tholi.Also dome. The apical, often thickened part of the inner wall in a bitunicate ascus.[455]
tomentum
Plural tomenta. Also defined: tomentose. A layer of short interwoven or coiled fungal hyphae with a texture similar to velvet. In lichens, the tomentum projects from the lower cortex and serves to help it attach to its substrate. Structures with this type of hyphae are called tomentose.[456] Tomentose surfaces are found in foliose genera such as Lobaria, Pseudocyphellaria, and Sticta.[457]
topeliopsidoid
A morphotype of corticolousthelotremoid lichens used to describe characteristics of apothecial and thallus structure. It pertains to lichens where the apothecia are either prominent or sessile, often hidden between the substratum, and they open with multiple, typically regular "teeth" that stay relatively curved over the hardly visible disk. The margins tend to peel off, meaning the overlaying thallus cortex separates from the underlying marginal thallus tissue, but no distinct, clean split between thallus margin and excipulum is formed. This morphotype is seen in species like Chapsa meridensis and Topeliopsis.[458]
trabecula
Plural trabeculae. In the genus Umbilicaria, they are rib- or strap-shaped structures radiating outward from the umbilicus that merge towards the mid-zone of the thallus undersurface.[271]
trentepohlioid
Resembling or belonging to the green algal genus Trentepohlia; trentepohlioid cells are filamentous (elongated and cylindrical), multicellular, and have a yellow to orange colour.[459]
Also defined: funiculus, umbilicus, umbilical cord. A lichen with a concave, circular, leafy thallus that is joined to its substrate only by its central part, called an umbilicus, umbilical cord or funiculus.[462][463]
A type of ascus with a single functional layer; the rigid internal and external wall layers do not separate during release of the ascospores. Most ascomycetes have unitunicate asci.[467]
Deeply cup-shaped or urn-shaped; in lichens, the term is used to describe some apothecia with a sunken hymenium and elevated parathecium that forms a narrow mouth.[469]
A lichen test performed by shining a long-wavelength ultraviolet light (350 nm) at a lichen structure to check if it fluoresces; a positive test (abbreviated as UV+) indicates the presence of certain lichen products. Xanthone compounds in the cortex tend to fluoresce yellow, orange, or red, while depsides and depsidones in the medulla fluoresce blue to white.[471]
A lichen not attached to a substrate, typically able to be blown around by wind.[472]
vegetative
Also assimilative. Having to do with the growth phase of an organism before reproduction, including spore germination, growth, development and asexual multiplication.[473]
"Vitricolous lichens – Lichen website". Australian National Botanic Gardens - Botanical Web Portal. Australian National Herbarium. 20 October 2009. Retrieved 18 October 2022.
Ahti, T. (1982). "The morphological interpretation of cladoniiform thalli in lichens". The Lichenologist. 14 (2): 105–113. doi:10.1017/s0024282982000255.
Brodo, Irwin (2016). "Glossary". Keys to Lichens of North America: Revised and Expanded. New Haven, Connecticut: Yale University Press. pp. 369–382. ISBN978-0-300-19573-6.
Fjelde, Markus Osaland; Timdal, Einar; Haugan, Reidar; Bendiksby, Mika (2024). "Paraphyly and cryptic diversity unveils unexpected challenges in the "naked lichens" (Calvitimela, Lecanoromycetes, Ascomycota)". Molecular Phylogenetics and Evolution. 190: e107944. doi:10.1016/j.ympev.2023.107944. hdl:10852/106762.
Fraser, Robert H.; Pouliot, Darren; van der Sluijs, Jurjen (2021). "UAV and high resolution satellite mapping of forage lichen (Cladonia spp.) in a rocky Canadian Shield landscape". Canadian Journal of Remote Sensing. 48 (1): 5–18. doi:10.1080/07038992.2021.1908118.
Goward, Trevor (1986). "Brodoa, a new lichen genus in the Parmeliaceae". The Bryologist. 89 (3): 219–223. doi:10.2307/3243288. JSTOR3243288.
Grube, Martin; Hawksworth, David L. (2007). "Trouble with lichen: the re-evaluation and re-interpretation of thallus form and fruit body types in the molecular era". Mycological Research. 111 (9): 1116–1132. doi:10.1016/j.mycres.2007.04.008. PMID17698333.
Hafellner, Josef; Kalb, Klaus (1995). "Studies in Trichotheliales ordo novus". Studies in Lichenology with Emphasis on Chemotaxonomy, Geography and Phytochemistry. Festschrift Ch. Leuckert. Bibliothecia Lichenologica. Vol. 57. Berlin-Stuttgart: J. Cramer in der Gebrüder Borntraeger Verlagsbuchhandlung. pp. 161–186. ISBN978-3-44-358036-0.
Hawksworth, D. L. (1988). "The variety of fungal-algal symbioses, their evolutionary significance, and the nature of lichens". Botanical Journal of the Linnean Society. 96 (1): 3–20. doi:10.1111/j.1095-8339.1988.tb00623.x.
Hertel, Hannes (2009). "A new key to cryptothalline species of the genus Lecidea (Lecanorales)". In Aptroot, A.; Seaward, M.R.D.; Sparrius, L.B. (eds.). Biodiversity and Ecology of Lichens – Liber Amicorum Harrie Sipman. Bibliotheca Lichenologica. Vol. 99. Berlin/Stuttgart: J.Cramer in der Gebrüder Borntraeger Verlagsbuchhandlung. pp. 185–204. ISBN978-3-443-58078-0.
Kantvilas, G. & Jarman, S.J. (1999). Lichens of rainforest in Tasmania. Canberra, ACT, Australia: Australian Biological Resources Study. ISBN0-642-56802-2.
Kantvilas, Gintaras (2023). "Three new species of siphuloid lichens, with a first key to the genus Parasiphula". The Lichenologist. 55 (1): 17–25. doi:10.1017/s0024282922000421.
Kirk, Paul M.; Cannon, Paul F.; Minter, David W.; Stalpers, Joost A., eds. (2008). Dictionary of the Fungi. Vol. 10. CAB International. ISBN978-1-84593-933-5.
Kistenich, Sonja; Timdal, Einar; Bendiksby, Mika; Ekman, Stefan (2018). "Molecular systematics and character evolution in the lichen family Ramalinaceae (Ascomycota: Lecanorales)". Taxon. 67 (5): 871–904. doi:10.12705/675.1. hdl:10852/67955.
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.
MacKenzie Lamb, I. (1968). "Appendix II. Glossary of Technical Terms". Antarctic Lichens. II. The Genera Buellia and Rinodina (Report). British Antarctic Survey Scientific Reports. British Antarctic Survey.
Meyer, Barbara; Printzen, Christian (2000). "Proposal for a standardized nomenclature and characterization of insoluble lichen pigments". The Lichenologist. 32 (6): 571–583. doi:10.1006/lich.2000.0294.
Orange, A.; James, P.W.; White, F.J. (2001). Microchemical Methods for the Identification of Lichens. British Lichen Society. ISBN978-0-9540418-0-9.
Øvstedal, D.O.; Lewis Smith, R.I. (2001). "Glossary". Lichens of Antarctica and South Georgia. A Guide to Their Identification and Ecology. Cambridge, UK: Cambridge University Press. pp. 367–374. ISBN978-0-521-66241-3.
Nash, T.H. III (2008). "1. Introduction". In Nash III, Thomas H. (ed.). Lichen Biology (2nd ed.). Cambridge, UK: Cambridge University Press. pp. 1–8. ISBN978-0-521-69216-8.
Honegger, R. (2008). "3.Mycobionts". In Nash III, Thomas H. (ed.). Lichen Biology (2nd ed.). Cambridge, UK: Cambridge University Press. pp. 27–39. ISBN978-0-521-69216-8.
Büdel, B.; Scheidegger, C. (2008). "4.Thallus morphology and anatomy". In Nash III, Thomas H. (ed.). Lichen Biology (2nd ed.). Cambridge, UK: Cambridge University Press. pp. 40–68. ISBN978-0-521-69216-8.
Nelsen, Matthew P.; Lücking, Robert; Aptroot, André; Andrew, Carrie J.; Cáceres, Marcela; Plata, Eimy Rivas; Gueidan, Cécile; da Silva Canêz, Luciana; Knight, Allison; Ludwig, Lars R.; Mercado-Díaz, Joel A.; Parnmen, Sittiporn; Lumbsch, H. Thorsten (2014). "Elucidating phylogenetic relationships and genus-level classification within the fungal family Trypetheliaceae (Ascomycota: Dothideomycetes)". Taxon. 63 (5): 974–992. doi:10.12705/635.9.
Papong, Khwanruan; Boonpragob, Kansri; Mangold, Armin; Divakar, Pradeep K.; Lumbsch, H. Thorsten (2010). "Thelotremoid lichen species recently described from Thailand: a re-evaluation". The Lichenologist. 42 (2): 131–137. doi:10.1017/s0024282909990405.
Randlane, Tiina; Tõrra, Tiiu; Saag, Andres; Saag, Lauri (2009). "Key to European Usnea species". In Thell, Arne; Seaward, Mark R. D.; Feuerer, Tassilo (eds.). Diversity of Lichenology – Anniversary Volume. Bibliotheca Lichenologica. Vol. 100. Stuttgart: J. Kramer. pp. 419–462. ISBN978-3-443-58079-7.
Rapai, Sean B.; McMullin, Richard Troy; Newmaster, Steven G. (2012). "The importance of macrolichen traits and phylogeny in forest community assemblage along a high elevation gradient in southwestern British Columbia". Forest Ecology and Management. 274: 231–240. Bibcode:2012ForEM.274..231R. doi:10.1016/j.foreco.2012.02.018.
Rivas Plata, Eimy; Lücking, Robert; Sipman, Harrie J. M.; Mangold, Armin; Kalb, Klaus; Lumbsch, H. Thorsten (2010). "A world-wide key to the thelotremoid Graphidaceae, excluding the Ocellularia-Myriotrema-Stegobolus clade". The Lichenologist. 42 (2): 139–185. doi:10.1017/s0024282909990491.
Sanders, William B. (2004). "Bacteria, algae, and phycobionts: maintaining useful concepts and terminology". The Lichenologist. 36 (5): 269–275. doi:10.1017/s0024282904014343.
Sérusiaux, E.; Lücking, R. (2007). "Gallaicolichen, a new genus of foliicolous lichen with unique diaspores". In Kärnefelt, Ingvar; Thell, Arne (eds.). Lichenological Contributions in Honour of David Galloway. Bibliotheca Lichenologica. Vol. 95. Berlin/Stuttgart: J. Cramer. pp. 509–516. ISBN978-3-443-58074-2.
Smith, C.W.; Aptroot, A.; Coppins, B.J.; Fletcher, A.; Gilbert, O.L.; James, P.W.; Wolseley, P.A., eds. (2009). "Glossary". The Lichens of Great Britain and Ireland. London: British Lichen Society. pp. 21–39.
Upreti, Dalip K.; Rai, Himanshu, eds. (2013). Terricolous Lichens in India. Vol. 1: Diversity Patterns and Distribution Ecology. New York: Springer. ISBN978-1-4614-8736-4.
Silverstein, Alvin; Silverstein, Virginia B.; Silverstein, Robert A. (1996). Fungi. New York: Twenty-First Century Books. ISBN0-8050-3520-6.
Thüs, Holger; Schultz, Matthias (2009). "Glossary". In Büdel, Burkhard; Gärtner, Georg; Krienitz, Lothar; Preisig, Hans R.; Schagerl, Michael (eds.). Fungi. Part 1: Lichens. Süßwasserflora von Mitteleuropa [Freshwater Flora of Central Europe]. Vol. 21/1. Heidelberg: Spektrum Akademischer Verlag. pp. 14–23. doi:10.1007/978-3-8274-2299-6_2. ISBN978-3-8274-1594-3.
Tripp, Erin (2017). Field Guide to the Lichens of White Rocks (Boulder, Colorado). University Press of Colorado. ISBN978-1-60732-554-3.
Ulloa, Miguel; Hanlin, Richard T. (2012). Illustrated Dictionary of Mycology (2nd ed.). St. Paul, Minnesota: The American Phytopathological Society. ISBN978-0-89054-400-6.
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